Dissertations / Theses on the topic 'Prey species'

Many biological problems can be reduced to the description of a food chain model or a food web. In these systems, the biodiversity and coexistence of all species are vital issues to discuss. Three ecological models have been proposed in case of the existence of a reserved area, in order to understand multi-species interactions so as to prevent the slow extinction of some endangered species and to test the stability when the length of the food chain and size of the web models are increased. It is taken that the environment has been divided into two disjoint regions, namely, unreserved and reserved zones, where a predator is not allowed to enter the latter. The first model describes a four species food chain predator-prey model with prey refuge (prey in the reserved zone, prey in the unreserved zone, predator and top predator), with the predator being entirely dependent on the prey in the unprotected area. The second model addresses the same problem, but in addition, a third component in the chain partially depends on the prey in the unreserved zone. Finally, the last model investigates a four species food web system with a prey refuge and in this case, the fourth component can also feed directly on the prey in the unreserved zone. The boundedness, existence and uniqueness of the solutions of the proposed models are established. The local and global dynamical behaviours are investigated, with the persistence conditions of the models being elicited. The local bifurcation near each of the equilibrium points is obtained. The numerical simulations in MATLABR are used to study the influence of the existence of the reserved zone on the dynamical behaviour of the proposed models. It has been concluded that the role of the reserved area could be beneficial for the survival and stabilising of multi-species interactions.

The use of resources by otters (Lutra lutra L.) living along coasts and streams in Southwest Portugal was described, and related to their spatial and temporal patterns of availability. The use of freshwater along coasts was inferred from the spatial distribution of spraints, and from radio-tracking. Spraints were mostly concentrated at the mouth of streams that provided freshwater throughout the dry season, and where rest-sites were found within dense thickets of brambles. Rest-sites were considered a scarce resource, because they seemed to be associated with the few permanent sources of freshwater. Along the coast prey availability was highest in winter and lowest in later summer and autumn. The corkwing wrasse was the dominant prey, and the seasonal variation in diet was primarily related to fluctuations in its abundance: wrasse were consumed when they were most abundant, other prey being taken to compensate for lows in their availability. In streams there were large year-to-year fluctuations in fish abundance, and these were related to the severity of summer droughts. Spring was probably the period of maximum prey abundance. Eels and crayfish were the dominant prey, but cyprinids, toads and frogs were also important seasonally. Prey were consumed relative to availability, except eels, which were consumed less than expected during the peak abundance of crayfish. Crayfish and eels were more associated in spraints that expected by chance. The introduction of the American crayfish to Iberia contributed to increase food abundance from spring to early autumn. Breeding of otters occurred seasonally, but births were earlier on the coast (October-December) than inland (January-March), most probably as a consequence of the seasonal patterns of food availability. Litter sizes inland were larger than on the coast. Differences between otter habitats in Southwest Portugal and elsewhere were discussed. A conservation strategy for otters in this region was developed.

This thesis explores the use of multi-element isotope analysis and intra-tooth sampling to reconstruct prey species ecology and biogeography. Modern caribou (Rangifer tarandus ssp.) and bison (Bison bison) from North America are used to assess the relationship between known lifetime movements and feeding habits, and those reconstructed through the stable isotope analysis of teeth and bone, including strontium (87Sr/86Sr), oxygen (d18O), carbon(d13C), nitrogen (d15N) and sulphur (d34S). Teeth (enamel and dentine) were sequentially-sampled in order to reconstruct time-series isotopic profiles at an intra- and inter-individual scale, allowing an assessment of the applicability of these methods to archaeological materials. The 87Sr/86Sr and d18O data indicate the clear potential for these methods to identify faunal movements, and to discern ranging behaviours from true migrations. d34S values of bone collagen compliment enamel 87Sr/86Sr data in the same individuals, and suggest the use of this approach for the identification of geographical origin.d13C and d15N from sequentially-sampled dentinal collagen allows the identification of some seasonal foraging behaviours, most notably winter lichen consumption in the caribou. The same stable isotope and sampling techniques are then applied to fauna from the late Pleistocene site of Jonzac (Chez-Pinaud), France, in order to investigate the biogeography and feeding ecology of Middle Palaeolithic prey-species. The elevated d13C values in reindeer bone collagen compared to the other species indicates lichen feeding and the prevalence of this niche feeding behaviour in this ancestral species. The sequential-sampling and strontium isotope analysis of herbivore enamel from the site clearly demonstrates seasonal migratory behaviour in reindeer (Rangifer tarandus sp.), and allows the identification of a non-migratory taxon, bison (Bison sp.) This is the first such evidence for migration in Pleistocene reindeer, allowing greater insight into the palaeoecology of this prey animal, and the palaeoenvironment in which Neanderthals lived and hunted.

Much of the world's species diversity is located in tropical and sub-tropical ecosystems, and a better understanding of the ecology of these systems is necessary to stem biodiversity loss and assess community- and ecosystem-level responses to anthropogenic impacts. In this dissertation, I endeavored to broaden our understanding of complex ecosystems through research conducted on the Cinaruco River, a floodplain river in Venezuela, with specific emphasis on how a human-induced perturbation, commercial netting activity, may affect food web structure and function. I employed two approaches in this work: (1) comparative analyses based on descriptive food web characteristics, and (2) experimental manipulations within important food web modules. Methodologies included monthly sampling of fish assemblages using a variety of techniques, large-scale field experiments, extensive stomach content and stable isotope analyses. Two themes unite the information presented: (1) substantial spatial and temporal variability in food web structure, and (2) how body-size can be used to generalize species-interactions across this complexity. Spatial variability occurred at various scales, from among small fish assemblages on seemingly homogeneous sand banks, to differences among landscape scale units (e.g. between lagoons and main river channel). Seasonal variability was apparent in predation patterns, with relative prey availability and body size primarily resulting in decreasing prey sizes with falling water levels. Body size was also related to functional outcomes of species interactions, for example, a size-based response of prey fishes to large-bodied piscivore exclusion. This pattern was further substantiated at the landscape-scale, as differences in assemblage structure among netted and un-netted lagoons were largely size-based. Trophic position of fish and body size was not found to be related, likely due to the diversity of prey available to consumers, and may signify that commercial netting activity will not decrease food chain lengths. In sum, by describing human impacts within a food web context, I endeavor to provide predictive power regarding a specific human-induced environmental problem, yet still allowing for generality that will broaden the theoretical foundations and applications of food web ecology.

Fish are one of the most imperiled groups of vertebrates worldwide. Threats to fish fall into one of four general categories: physical habitat loss or degradation, chemical pollution, overfishing, and nonnative species introductions. Nonnative predatory fish often have a devastating impact on native prey, especially with endemic fish, whose restricted distribution and often limited evolutionary history with predators make them particularly susceptible to nonnative predators. One reason nonnative fish are often so efficient predators is that the native fish do not recognize the predator as a threat. Although many studies have examined the role of predator odor recognition, no fish have been shown to possess an innate recognition of the odor of predators unless they share a close co-evolutionary history. Many fish learn to identify novel predators through exposure to a predator's odor in conjunction with a conspecific alarm cue. Alarm cues are substances that are stored in the epidermis of many fish and are released when skin cells are broken. Exposure to novel predator odor in conjunction with alarm cue does not necessarily require a fish to survive a close encounter with a predator to be able to learn to recognize the predator. Here I show that it is possible to train hatchery-raised fish (June sucker, Chasmistes liorus) to recognize a nonnative predator odor (largemouth bass, Micropterus salmoides) through exposure to the odor of a predator that has eaten June sucker. I also show that this training can translate into higher survival in subsequent encounters with predators. I propose that training hatchery-raised fish prior to stocking may increase survival of hatchery-raised fish through anti-predator behavior.

Evolution can occur rapidly, along timescales that are traditionally regarded as 'ecological'. Despite growing acceptance among biologists of rapid evolution, a strong paradigm of contemporary evolution is still absent in many sub-disciplines. Here I apply a contemporary evolution viewpoint to conservation biology. Specifically, I examine the impact of cane toads (Bufo marinus) on Australian snakes. Toads were introduced into Australia in 1935, have spread rapidly and represent a novel, extremely toxic prey item to na�ve Australian predators (including snakes). Based on dietary preferences and geographic distributions I find that 49 species of Australian snake are potentially at risk from the invasion of the toad. Furthermore, examination of physiological resistance to toad toxin in 10 of these �at risk� species strongly suggests that most species of Australian snake are poorly equipped to deal with a likely dose of toad toxin. Even species that are highly resistant to toad toxin (such as the keelback, Tropidonophis mairii) face indirect fitness costs associated with consuming toads. Within a population of snakes however, the impact of toads is unlikely to be random. For example, the examination of several component allometries describing the interaction between snakes and toads revealed that, within a species, smaller snakes are more likely to ingest a fatal dose of toad toxin than are larger snakes. Further consideration of the interaction between snakes and toads suggests that toads will not only be exerting differential impact on snakes based upon morphology, but also exert non-random selection on prey preference and resistance to toad toxin in snake populations. To examine the possibility of a morphological response by snakes to toads, I examined changes in the body size and relative head size of four species of snake as a consequence of time since exposure to toads. Two of the species (green treesnakes and red-bellied blacksnakes) are predicted to face strong impacts from toads. These two species showed an increase in mean body size and a decrease in relative head size as a consequence of time since exposure to toads; both changes in an adaptive direction. In contrast, the other two species (keelbacks and swampsnakes) are predicted to face much lower impact from toads, and these two species showed little or no evidence of morphological change associated with time since exposure to toads. These results indicate an adaptive change in morphology at a rate that is proportional to the predicted level of impact for each species, strongly suggesting an evolved response. Red-bellied blacksnakes (a toad-vulnerable species) were further assessed for evolved responses in prey preference and toxin resistance. Comparisons between toad-exposed and toad-na�ve populations of blacksnakes revealed that snakes from toad-exposed populations exhibited slightly higher resistance to toad toxin and a much-reduced tendency to eat toads, when compared with toad-na�ve snakes. Na�ve snakes exhibited no tendency to learn avoidance of toxic prey, nor were they able to acquire resistance to toxin as a result of several sub-lethal doses, suggesting that the observed differences between populations is evolved rather than acquired. Together, these results strongly suggest that blacksnakes are exhibiting an evolved shift in prey preference and toxin resistance as a consequence of exposure to toads. Thus, it appears that snakes are exhibiting adaptation at multiple traits in response to exposure to toads. Given the high likelihood that these adaptive shifts have an evolved basis, it appears that the impact of toads will decrease with time in many snake populations. But what about toads? Because the outcome of the interaction between a toad and a snake is also mediated by the body size and relative toxicity of toads, it is important to understand how these traits vary in space and time. Exploratory analysis revealed that toads exhibit a decrease in body size and a decrease in relative toxicity as a consequence of time since colonisation, indicating that their impact on native predators decreases with time. Additionally, there appears to be meaningful spatial variation in toad relative toxicity, indicating that some populations of native predators are facing higher impact from toads than others. Overall, these results clearly indicate the importance of assessing the potential for rapid evolutionary response in impacted systems. Doing so may provide evidence that some species are in less trouble than originally thought. Additionally, and as more data accumulate, it may be possible to characterise certain categories of environmental impact by their potential for eliciting adaptive response from �impacted� species. This approach has strong implications for the way conservation priorities are set and the way in which conservation dependent populations are managed.

This study examined the physiological and behavioural effects of a stress-inducing stimulus (predator odour) on potential prey species (Australian native and exotic). The aim was to determine if differences in the response of prey were related to the scent of evolutionary known predators compared to unfamiliar or short-term introduced ones. In laboratory experiments, responses were always restricted to changes in respiratory variables, with brushtail possums (Trichosurus vulpecula), rabbits (Oryctolagus cuniculus) and tammar wallabies (Macropus eugenii) showing no variation in metabolic rate after exposure to predator odours.Brushtail possums showed strong changes in ventilatory rate only when faced with the scent of an historical predator, increasing respiratory frequency (ratio after/before exposure = 4.55 ± 1.007) and decreasing tidal volume (ratio after/before exposure = 0.38 ± 0.113 ) in response to stale dingo urine. The changes were short-lived, lasting for only one minute of exposure. For this reason, it is unclear if the response observed could be considered as a fear reaction. However, there was no habituation after three exposures and this may indicate that possums were initially displaying an investigative approach to a predator scent and then relaxing once assessment was completed.The effect of predator and novel odours on the ventilation of rabbits appeared to be in general mediated by anxiety, as shown by the responses elicited by both predator and control scents. However, stronger reactions were observed in response to feral cat and quoll odours for both respiratory frequency (ratio after/before exposure to feral cat = 4.39 ± 0.721 and to quoll = 3.75 ± 0.486) and tidal volume (ratio after/before exposure to feral cat = 0.47 ± 0.065 and to quoll = 0.64 ± 0.129). This could be due to different intensity of the olfactory stimuli. Nevertheless, the effects of noxious odours on rabbits clearly demonstrate that they become highly vigilant at any sudden change in their environment.Tammar wallabies appeared to possess a mechanism for the recognition of predator odours as ventilatory responses were restricted to particular predator scents. After investigation, fox and cat odour provoked a stronger and more prolonged change in respiratory frequency (ratio after/before exposure to fox = 3.58 ± 0.918 and to feral cat = 2.44 ± 0.272) and tidal volume (ratio after/before exposure to fox = 0.84 ± 0.110 and to feral cat = 0.98 ± 0.155) compared to the other scents, suggesting that wallabies may have perceived these species as more immediate threats.For wild, free-living brushtail possums and southern brown bandicoots (Isodoon obesulus), there was no pattern of avoidance of historical or introduced predator odours, with no difference in number of animals captured in predator and control scented traps. This may indicate that predator odour avoidance has not evolved in these species and that they have poor possibilities of escaping potential predators. However it may also be explained by the long-term predator-free environment in which the study was conducted, and suggests loss of anti-predator behaviour in populations without predation risk.Wild, free-ranging western grey kangaroos (Macropus fuliginosus) reacted to the odour of both historical (dingo) and novel (fox) predators by reducing number of feeding events (5.2±2.08 for fox and 5.9±1.33 for dingo) and time spent foraging (17.7±7.2 sec for fox and 22.2±4.6 sec for dingo) when predator scents were present and by escaping areas tainted with predator odours (41.4±17.5 sec for fox and 33.8±13 sec for dingo). Clearly these results suggest that kangaroos are scared of predator odours. However, a close investigation of predator scents was necessary before a response could be elicited and feeding areas were not completely abandoned.It is still unclear if small Australian prey, such as southern brown bandicoots and brushtail possums, respond to olfactory cue of predation in the wild, as different results were obtained in the studies conducted in the laboratory and in free populations. However, this research showed that medium and large-sized macropods respond to both native, long-term and introduced predator odours. This indicates that at least some Australian prey species can recognise the odour of potential predators, although responses usually occur after a period of investigation, and do not always result in avoidance behaviour. Differential responses appear to be based on the perceived risk. Use of predator odour is unlikely to be an effective mechanism of deterring herbivores from preferred feeding locations.

Invasive predators are major drivers of global biodiversity loss and their impacts may be worsened by other disturbances such as fire. I examined how the fire history of shrublands influences the ecology of feral cats Felis catus, dingoes Canis dingo and their prey species in Western Australia’s northern Wheatbelt region. A review of the literature revealed that feral cats inhabit a diverse range of ecosystems worldwide, but are generally recorded most often in habitat types characterised by a mixture of plant growth forms close to ground level. Cat habitat use is influenced by predation/competition, prey availability, shelter availability and anthropogenic resource subsidies. Relatively few studies were available for review and the strength of evidence contained within them was generally low, which highlighted the need for more rigorous field studies. I examined overlap in resource use between cats and dingoes using remote camera surveys and dietary analysis of scats. Both carnivores were recorded in all four major habitat types: recently burnt shrublands (10 to 14 years since last fire), long unburnt shrublands (34 to ~49 years), very long unburnt shrublands (> 50 years), and woodlands. Dingoes and cats preferred woodlands and very long unburnt shrublands respectively, but spatial overlap between the two species was still common. Mean diurnal activity time for feral cats was two and a half hours later than that of dingoes. The diet of feral cats was more diverse than that of dingoes and dietary overlap between the two carnivores was relatively low. Rabbit remains did occur relatively frequently in both cat and dingo scats, but small mammals, reptiles and birds were also common in cat scats, and macropods in dingo scats. Nine of the 15 prey species studied showed a preference for either recently burnt or long unburnt shrublands. Two small mammals and three reptiles were most abundant in recently burnt areas, while the abundance of one small mammal and three reptiles was highest in long unburnt areas. Using giving up density experiments, I showed that rodents exhibited differential foraging behaviour in the two vegetation fire ages. The rodents foraged for longer in sheltered compared to open microhabitats, but this pattern only occurred in recently burnt, not long unburnt shrublands, probably because the higher density of understorey vegetation in recently burnt areas provided the rodents with extra cover to hide and escape from predators. I also developed a new framework for conceptualising interactions between invasive predators and other ecological disturbances, such as fire, habitat fragmentation, and top -predator decline. The impacts of invasive predators can be classified as either functional (density -independent) or numerical (density -dependent), and they interact with other threats through both habitat -mediated (fire, grazing, land clearing) and community -mediated (top -predator decline, altered prey populations, anthropogenic resource subsidies) interaction pathways. The key findings of this thesis show that both old and young shrublands can be suitable habitat for feral cats; predator -prey dynamics are influenced by successional habitat stages; small mammals show behavioural, as well as population -level responses to fire; and that invasive predator management is likely to benefit from addressing multiple threats in unison.

Thesis (Ph. D.)--Illinois State University, 2007.
Title from title page screen, viewed on February 11, 2008. Dissertation Committee: Steven A. Juliano (chair), Diane L. Byers, L. Philip Lounibos, Charles F. Thompson, William L. Perry. Includes bibliographical references (leaves 152-163) and abstract. Also available in print.

This thesis examines the foraging behaviour of Carcinus maenas and Cancerpagurus when presented with bivalves of contrasting morphology: Mytilus edulis, Ostrea edulis, Crassostrea gigas and Cerastoderma edule. Because foraging may be influenced by the relative abundance and morphological characteristics of both predator and prey, these aspects are also considered. Chelal size and strength of these crabs and prey shell shape largely determined handling techniques. When offered a size range of these bivalves individually, crabs attacked all encountered prey but rejected those that remained unbroken after several opening attempts, thus, emphasising the passive nature of their size-selective feeding. When offered paired combinations of mussels, oysters and cockles, larger crabs selected species in the ranked order of their profitability. Species-related preferences exhibited by crabs feeding on prey at or near the optimal size suggest that foraging decisions are partly based on evaluations of overall prey shape and volume, and that shell width constitutes an important feature which crabs recognise and associate with prey value. Variations in crab strength relative to size accounted for most intra-specific differences in foraging behaviour. Juvenile C maenas are limited in their choice of prey size, and are thus less species-selective. Adult C maenas are not so constrained, and exhibit a higher degree of species-selectivity. C. pagurus possesses powerful monomorphic chelae that operate at higher mechanical advantage than the cbelae of C maenas, and readily crushed larger mussels relative to their size. Differences in prey size selection between crab species varied with the species of prey offered, suggesting that certain shell features of these bivalves constitute effective barriers to even the powerful chelae of C pagurus. These results are relevant in the context of aquaculture, since predatory impact on commercially reared bivalves might be reduced by combining different prey species that offer predators alternative or preferred sources of food.

This study considers the dynamical interaction of two predatory carnivores (Lions (Panthera leo) and Spotted Hyaenas (Crocuta crocuta)) and three of their common prey (Buffalo (Syncerus caffer), Warthog (Phacochoerus africanus) and Kudu (Tragelaphus strepsiceros)). The dependence on spatial structure of species’ interaction stimulated the author to formulate reaction-diffusion models to explain the dynamics of predator-prey relationships in ecology. These models were used to predict and explain the effect of threshold populations, predator additional food and prey refuge on the general species’ dynamics. Vital parameters that model additional food to predators, prey refuge and population thresholds were given due attention in the analyses. The stability of a predator-prey model for an ecosystem faced with a prey out-flux which is analogous to and modelled as an Allee effect was investigated. The results highlight the bounds for the conversion efficiency of prey biomass to predator biomass (fertility gain) for which stability of the three species ecosystem model can be attained. Global stability analysis results showed that the prey (warthog) population density should exceed the sum of its carrying capacity and threshold value minus its equilibrium value i.e., W >(Kw + $) −W . This result shows that the warthog’s equilibrium population density is bounded above by population thresholds, i.e., W < (Kw+$). Besides showing the occurrence under parameter space of the so-called paradox of enrichment, early indicators of chaos can also be deduced. In addition, numerical results revealed stable oscillatory behaviour and stable spirals of the species as predator fertility rate, mortality rate and prey threshold were varied. The stabilising effect of prey refuge due to variations in predator fertility and proportion of prey in the refuge was studied. Formulation and analysis of a robust mathematical model for two predators having an overlapping dietary niche were also done. The Beddington-DeAngelis functional and numerical responses which are relevant in addressing the Principle of Competitive Exclusion as species interact were incorporated in the model. The stabilizing effect of additional food in relation to the relative diffusivity D, and wave number k, was investigated. Stability, dissipativity, permanence, persistence and periodicity of the model were studied using the routine and limit cycle perturbation methods. The periodic solutions (b 1 and b 3), which influence the dispersal rate (') of the interacting species, have been shown to be controlled by the wave number. For stability, and in order to overcome predator natural mortality, the nutritional value of predator additional food has been shown to be of high quality that can enhance predator fertility gain. The threshold relationships between various ecosystem parameters and the carrying capacity of the game park for the prey species were also deduced to ensure ecosystem persistence. Besides revealing irregular periodic travelling wave behaviour due to predator interference, numerical results also show oscillatory temporal dynamics resulting from additional food supplements combined with high predation rates.

Nesting behaviour and prey selection was compared in eight species of Ammophila. The frequent sympatry of these species was a central point of interest in the study. Nest siting differed markedly in several species. A. ferrugineipes, A. dolichodera, A. dolichocephala and, probably, A. conifera nested mostly in open habitats, this being regarded as the most common situation for the majority of Ammophila. A. beniniensis was unusual in nesting in well vegetated habitats, A. vulcania doing so amongst small clumps of vegetation in otherwise open habitats. A. insignis nested in vertical banks or old animal burrows and A. braunsi utilised abandoned burrows of other wasps situated in non-friable clay soils. This is the first time such nest siting has been reported for Ammophila and each of these two species is consequently unique. Many aspects of nest construction behaviour, in particular methods of soil waste disposal, sealing of nests and their final coverage, differed interspecifically, in some cases intraspecifically, but often apparently in response to the habitat. With regard to nest provisioning strategies, all species hunted after digging their nests, except A. dolichodera which is the first Afrotropical Ammophila reported to hunt before excavating its nest. This brings the total number of the world's species of Ammophila showing this behaviour to three. This species, A. beniniensis, A. braunsi and A. vulcania provided one caterpillar per nest; the others supplied several, A. insignis being the only mass provisioner. A. ferrugineipes showed a form of progressive provisioning and maintained several nests at a time; it is the only Afrotropical species known so far to do so. Only two extra-limital species have been reported showing similar behaviour. The observed variations in provisioning strategies can most easily be explained in relation to the habitat. Investigation of prey selection showed that there was very little interspecific overlap in the species of caterpillars used for provisioning nests. This was shown to be due mainly to differing hunting habitat preferences. The various overall patterns of nesting behaviour, and variations in the different components within these patterns, were considered most likely to have resulted from allopatric speciation in different types of vegetation, the more advanced patterns arising in open habitats. The frequent occurrence of sympatry appears to be a reflection of a shifting distribution of mosaic patterns of vegetation types and the post-speciation spread of species. This influence of habitat on behaviour and prey selection has emphasised the need for caution in drawing phylogenetic conclusions from ethology. Further, these considerations provide an alternative to competition in attempting to explain the observed behaviour patterns and distribution of these wasps.

Microtine rodents are a key component in the boreal ecosystem. Due to unique yearly fluctuation in population numbers, in addition to their central placement within boreal foodwebs, microtine rodent population dynamics control a wide variety of predator populations. Despite their grouping as microtine rodents, many individual species exhibit very different cycles, such as voles and lemmings. As such, the identification of species composition and ratio in each given habitat is of utmost importance. The purpose of this study was to compare the composition of species of small rodents between four different habitats (grassland, wetland, spruce dominated forest and pine dominated forest) and to statistically analyze if the different biotopes hosted significant differences in species composition. The project also aimed to examine differences in weight and numbers between populations of the same species in the different biotopes. To accomplish this, rodents were captured using non-lethal traps, and the resulting data was analyzed using RxC tables and ANOVA. The species captured were A. sylvaticus, C. glareolus and M. agrestis. The result of the analysis showed that the only difference in species composition existed between the grassland habitat and the rest. In addition, there existed no difference between different populations of A. sylvaticus in regards to weight. However, a difference was found between the populations of C.glareolus, with the individuals inhabiting spruce dominated forest being larger than those found in other habitats.

The harbor porpoise is considered a threatened species in Swedish waters. A full understanding of its ecology and relationship to other species is important when designing effective conservation plans. This thesis aims to investigate the relationships between harbor porpoises and their prey by comparing the acoustic activity patterns of harbor porpoises and the movement patterns of some of their prey species available at Kullaberg, Sweden. For this purpose, the scientific literature of porpoise and fish activity patterns has been reviewed. Results show that harbor porpoises may follow vertically migrating herring, but further studies are needed to confirm this. It is also possible that the activity pattern of porpoises does not follow exactly the movements of a certain species of prey, but that they rather feed upon several species, and therefore change their behavior accordingly throughout the diurnal cycle.

Weltweit sind viele Populationen großer Raubtiere bedroht. Die Suche nach Wegen zur Erhaltung und Wiederherstellung von Populationen großer Raubtiere ist daher ein zentrales Naturschutzziel. In einigen Regionen erholen sich die Populationen jedoch auch, was der Wissenschaft eine hervorragende Möglichkeit bietet potentielle Wege zum Schutz von Raubtieren zu identifizieren. Das Ziel dieser Arbeit war es, die Voraussetzungen, die eine Erholung von Raubtierpopulationen begünstigen, besser zu verstehen. Am Beispiel der sich erholenden Population des Persischen Leoparden im Kaukasus beantwortet diese Arbeit die folgenden Forschungsfragen: (1) Bietet die Ökoregion des Kaukasus einen geeigneten, sicheren und zusammenhängenden Lebensraum für Persische Leoparden und ihre Beutetiere? (2) Was sind wirksame Schutzstrategien, die eine Erholung der Population des Persischen Leoparden im Kaukasus begünstigen? Die Ergebnisse dieser Arbeit lassen den Schluss zu, dass für Leoparden und ihre Beutetiere im Kaukasus großflächig geeigneter Lebensraum vorhanden ist. Trotz dieser günstigen Ausgangsposition für den Artenschutz zeigt diese Arbeit jedoch auch klar auf, dass Schutzmaßnahmen erforderlich sind, um direkte Bedrohungen abzumildern. Als wichtigsten Faktor für eine Erholung der Leopardenpopulation identifiziert diese Arbeit das Verhindern von Tötungen. Diese Arbeit verdeutlicht somit die wichtige Rolle von Konfliktminderung zwischen Menschen und Raubtieren, um die daraus resultierende Tötung von Raubtieren zu verhindern. Darüber hinaus unterstreicht diese Dissertation die Notwendigkeit der Vernetzung sicherer Lebensräume und den Wert von internationaler Zusammenarbeit zur Förderung der Genesung von Populationen großer Raubtiere. Zusammengefasst liefert diese Arbeit Einblicke in Bedingungen, die die Genesung von Großraubtieren fördern, und umreißt mögliche zukünftige Wege eines vom Aussterben bedrohten Raubtiers in einem globalen Biodiversitäts-Hotspot.
Large carnivore populations are threatened globally. This is worrisome because of their intrinsic value and their importance for ecosystems and humans. Finding ways to maintain and restore large carnivore populations is therefore a central goal for conservation. Interestingly, large carnivores are also returning to areas where they were extirpated before, which provides a great opportunity for conservation science to identify pathways to safeguard large carnivore populations. The overall goal of this thesis was to better understand the conditions that foster large carnivore recoveries. Gaining a better understanding of large carnivore recoveries is challenging though, because it requires working across broad areas, where data on environmental characteristics and prey species is often limited and inconsistent. Using the Persian leopard recovery in the Caucasus ecoregion as an example, the specific research questions of this thesis were: (1) Does the Caucasus ecoregion have suitable, safe, and connected habitat for Persian leopards and their prey? (2) What are effective management strategies to foster the recovery of Persian leopards in the Caucasus? Results revealed that suitable habitat for Persian leopards and their prey is widespread in the Caucasus. However, habitat patches are often not safe and conservation efforts are needed to mitigate threats and restore, protect, and connect populations of leopards and their prey. Reducing the persecution of leopards seems the most important strategy to foster leopard recovery in the Caucasus. For large carnivore recoveries in general, this thesis underpins the importance of human-carnivore conflict mitigation to prevent killings of carnivores, particularly in areas where populations are small. Further, this thesis underlines the need of improving connectivity among safe habitat patches and the value of international cooperation. Overall, this thesis provides insights into conditions that foster large carnivore recovery and outlines potential future pathways for a critically endangered apex predator in a global biodiversity hotspot.

The European Commission manages fish stocks by applying a fishing mortality based on the maximum sustainable yield concept. So far most Baltic Sea fishing maximum sustainable yieldmodels have focused on one species at a time. The few existing multi-species models have assumed that a species’ maturity and growth is dependent on the availability of food. Our two-species models make it possible to investigate if there is a conflict between fishing maximum sustainable yield for cod and herring in the Baltic Sea. This two-species model of cod, as a predator and herring as prey, takes into account environmental drivers on cod and herring recruitment. Reproductive volume together with year-growth, (a year specific effect on growth of external variables like food availability) and predation by grey seals was included in the cod model. The herring model was dependent on cod spawning stock biomass and year-growth. The result shows that the reproductive volume is the main factor that affects the maximum sustainable yield for cod. The spawning stock biomass at maximum sustainable yield is more sensitive to reproductive volume than year-growth. When predation from seals is added in mortality and high environmental factors occurs the spawning stock biomass would be 50% compared to the spawning stock biomass at high environmental effects without seal predation. Four simulations of high cod spawning stock biomass were devastating for the herring population that was eradicated with high predation pressure. The herring maximum sustainable yield depends on the amount of cod spawning stack biomass i.e. the effect of high or low reproductive volume. Two analyses were made on a current environmental state for both species. The first analysis had a natural mortality of 0.2 for cod, which gave an fishing mortality of 0.20 and maximum sustainable yield of 410 000 tons. The herring had a fishing mortality of 0.03 and maximum sustainable yield of 11 000 tons. The second simulation included seal predation in cod mortality which decreased the cod maximum sustainable yield by 98% at a fishing mortality of 0.02, which gave a fishing mortality of 0.19 and maximum sustainable yield of 275 000 tons for herring. This gives a 25 times increase of herring maximum sustainable yield compared to the result without predation on cod. The cod population dynamics is vulnerable to environmental changes and to secure a healthy and productive cod population the target fishing mortality should be kept in phase with current reproductive volume.
Europeiska kommissionen förvaltar fiskbestånden genom att tillämpa fiskekvoter baserat på konceptet maximalt hållbart uttag. Hittills har de flesta maximalt hållbara fiske-fångst modeller för Östersjön fokuserat på en art i taget. De få befintliga fler-arts-modeller har antagit att en arts mognad och tillväxt är beroende av tillgången på föda. Vår två-arts-modell gör det möjligt att undersöka om det finns en konflikt mellan maximal hållbar fiske-fångst på torsk och sill i Östersjön. Denna två-arts-modell med torsk som ett rovdjur och sill som byte, tar hänsyn till miljön som drivkraft på deras rekrytering. I torskmodellen ingick reproduktiv volym tillsammans med årlig tillväxt (ett års specifika effekt på tillväxten beroende av externa variabler som tillgången till föda) och predation av gråsäl. Sill-modellen var beroende av årlig tillväxt och lekbeståndets biomassa hos torsk. Resultaten visar att den viktigaste faktorn som påverkar maximalt hållbart uttag för torsk är reproduktiv volym. Lekbeståndets biomassa vid maximalt hållbart uttag är mer känsligt för förändringar i reproduktiv volym än årlig tillväxt. När predation från säl tillsätts och höga gynnsamma miljöfaktorer råder är lekbeståndets biomassa 50 % jämfört med lekbeståndets biomassa vid höga gynnsamma miljöeffekter utan säl predation. Fyra simuleringar gav hög lekbestånds biomassa för torsk vilket var förödande för sillpopulationen som utrotades pga. högt predationstryck. Sillens maximala hållbara uttag beror på mängden lekbestånds biomassa hos torsk, d.v.s. effekten av hög eller låg reproduktiv volym. Två analyser gjordes på nuvarande miljömässiga nivåer för båda arterna. Den första analysen hade en naturlig dödlighet på 0,2 för torsk, vilket gav en fiske-mortalitet på 0,20 och maximalt hållbart uttag på 410 000 ton. Sillen hade en fiske-mortalitet på 0,03 och maximalt hållbart uttag på 11 000 ton. I den andra simuleringen ingår sälpredation på torsk vilket minskade torskens maximala hållbara uttag med 98 % vid en fiske-mortalitet på 0,02, vilket gav en fiske-mortalitet på 0,19 och maximalt hållbart uttag på 275 000 ton för sill. Detta ger en ökning av maximalt hållbart uttag för sill 25 gånger jämfört med resultatet utan predation på torsk. Torskens populationsdynamik är sårbar för miljöförändringar och för att säkra ett sunt och produktivt torskbestånd bör fiskemortaliteten hållas i fas med nuvarande reproduktiva volym.

The spread of non-native species is a global threat and the rate at which biological invasions occur is likely to increase in the future. This thesis examines the implications of the invasive signal crayfish, Pacifastacus leniusculus (Dana), for instream communities and abiotic processes within lowland rivers in England. The potential effects on lotic macroinvertebrate communities and fine sediment dynamics are investigated at a range of spatial and temporal scales, from the examination of national long-term datasets through to short-term detailed mesocosm experiments. Interrogation of macroinvertebrate community data from three English regions was undertaken to understand the temporal and spatial extent of signal crayfish effects. Invasive crayfish had significant long-term and persistent effects on resident macroinvertebrate communities regardless of the lithology or other environmental characteristics of the region. The resultant modifications to community composition had repercussions for several widely employed freshwater biomonitoring tools which employ faunal abundance in their derivation. A reach-scale field study was undertaken at two sites, one invaded by crayfish and one which did not support populations of crayfish (control), to assess the potential contribution of signal crayfish for fine sediment dynamics (ingress, fluxes and ultimately the overall implications for sediment budgets). Reach-scale field experiments examining the effect of crayfish invasion on resident macroinvertebrate communities over a 126-day period indicated that once crayfish populations are well-established their effects are persistent. However, separating the effects of crayfish from wider macroinvertebrate community dynamics and life-history characteristics of the invader and resident taxa means that attributing the effects is far from clear. The thesis presents the results of a series of novel mesocosm experiments which examined the dynamic and two-way interactions of predator-prey relationships and potential zoogeomorphological effects of signal crayfish and the freshwater shrimp, Gammarus pulex. Signal crayfish had a significant effect on the mass of fine sediment infiltrating into the gravel matrix associated with foraging and predatory activity; however this was strongly controlled by prey availability. Finally, through the development of conceptual models, the thesis illustrates the need for greater integration of ecological and geomorphological theories, at relevant environmental scales (temporal and spatial) to achieve truly interdisciplinary research.

Collective intelligence (CI) emerges from local coordination, collaboration and competition among the individuals within a social group. CI mainly results in a global intelligent behavior. One of the fundamental interactional channels within a CI system is energy flow. Each agent within an artificial or physical ecosystem must absorb energy in order to survive, evolve, breed, and reshape its local environment. In addition because the energy resources are limited in the environment, each agent has to compete with other agents to reach the required level of energy. Understanding the internal energy flow can potentially provide a deep insight into internal activities and external emergent behaviors of a given complex system. This study proposes a stochastic scheme for modeling a multi-species prey-predator artificial ecosystem with two levels of food chain. This will enable us to investigate the influence of energy flow on the ecosystem’s lifetime. The proposed model consists of a stationary hosting environment with dynamic weather condition and fruit trees. The inhabitants of this ecosystem are herbivore and carnivore birds each consisting of species. In our model, the collective behavior emerges in terms of flocking with more added rules consist of breeding, competing, resting, hunting, escaping, seeking and foraging behaviors. Using multi-species scheme, we define the ecosystem as a combination of prey and predator species with inter-competition among species within same level of food chain and intra-competition among those belonging to different levels of food chain. Furthermore, in order to model the energy within the ecosystem, some energy variables as functions of behaviors are incorporated in to the model. Finally, a simulation and visualization structure for implementing the proposed model is developed in this study. The experimental results of 11,000 simulations analyzed by Cox univariate analysis and hazard function suggest that only five out of eight behaviors can statistically significant influence the ecosystem’s lifetime. Furthermore, the results of survival analysis show that out of all possible interactions among energy factors, only two of them, interaction between flocking and seeking energies, and interaction between flocking and hunting energies, have statistically significant impact on the system’s lifetime. In addition, software implementation of the proposed framework validates the stability of simulation and visualization architecture. At last regression results using Nelson-Aalen cumulative hazard function and Cox-Snell variable and scaled Schoenfeld residuals test strongly validate our experimental results. To the best of our knowledge, there are three contributions in this research: First, the high level of complexity in the structure of the proposed model in comparison with the other systems which mostly contains only one species of prey, one species of predator and a kind of resource. While this study introduces two species of prey, capability of competition among species, dynamic weather condition with two element of wind and rain and dynamic resources, various behavioral rules such as escaping, breeding, hunting, resting, etc. Energy flow analysis within an artificial ecosystem is the second contribution. To the best of author’s knowledge there is no similar comprehensive model in the previous literature that investigates the life span of a stochastic multi-species predator-prey artificial ecosystem based on energy flow using Survival Analysis method. Lastly, the simulation results show that the flocking and seeking energy and flocking and hunting energy interactions are the most significant interactions which match with the Thompson iii et al. [ 65] observations in the real life. Their findings indicate that in the real life, birds use flocking behavior for better movement, more efficient food searching and social learning. Flocking motion also decrease predation risk as much as the flock size increases.

The Chandeleur Islands (Louisiana) contain nursery habitats for lemon sharks that provide abundant prey and protection from predation. Other local shark species (Atlantic sharpnose, bull, and blacktip sharks) co-occur with lemon sharks in the same region, including the nearby Biloxi Marshes. To better assess how lemon sharks use these nursery habitats, I measured diet and prey availability of young of the year and juvenile lemon sharks from 2009 to 2010. Young lemon sharks at the Chandeleur Islands have a relatively reduced diet breadth in comparison to those from nurseries in Bimini (Bahamas) and the Florida Keys. At the Chandeleur Islands, young lemon sharks appear to be opportunistically feeding on the most abundant prey items, resulting in high prey abundance and low diversity in their diet. Opportunistic feeding by young lemon sharks suggests minimal dietary overlap with other local shark species, resulting in minimal competition for resources.

Piscivorous birds are an integrated part of lake and coastline ecosystems. Despite decades of research it is yet unclear what influence fish eating birds have on the fish community. The aim of this literature study was to focus on how two fish eating birds, Great Cormorant and Great Crested Grebe, influence the fish community in shallow and eutrophic lakes. Different types of analysis methods have been used when doing research on the subject, including pellet analysis, stomach content analysis, tagging of fishes by “PIT”-techniques and analysis by observation. Results show that conclusions by studies tend to vary, but generally higher bird density, lower water temperature and a relatively high turbidity contribute to a higher influence on the fish community. Both bird species are opportunistic in their choice of food and catch prey of the species that occur locally. Both species also limit themselves in the prey size, but the choice vary over seasons because of water temperature and the birds requirements in association with breeding and migration. The significant level of influence seems to be when predation is conducted on younger and smaller fishes. The influence on the fish size can also be indirect, where predation on smaller individuals prevent fishes from becoming older and bigger. As studies tend to deviate in conclusion, no general answer to this issue can be given at present. Comprehensive research with several years of full control over both fish-and bird population is needed to find the proper conclusion.

Eland Tragelaphus oryx are the largest and slowest African antelope, falling within the preferred weight range of the prey of lions Panthera leo. Little is known, however, about the antipredatory strategies employed by eland. Rapid population declines in several small reserves following the reintroduction of lions suggest that the antipredatory strategies of eland may be compromised in spatially limited areas. This emphasises the need for research aimed at investigating how eland respond to lions, particularly in the spatial context, given that lions are increasingly being reintroduced into small reserves in South Africa. This study aimed to investigate lion-eland interactions at multiple scales and determine where on the landscape eland were vulnerable to predation. Adult female eland (n = 3 per site) in breeding herds were collared with GPS satellite collars in two study sites; the Nyathi section of Addo Elephant National Park (Nyathi) and Mountain Zebra National Park (MZNP) in the Eastern Cape, South Africa. A reintroduced population of lions, comprising two males and one female, was present in each study site, with individuals fitted with GPS satellite collars. At a broad spatiotemporal scale, eland did not avoid lions and there was considerable overlap between eland and lion home ranges in both study sites. At a finer spatiotemporal scale however, eland and lion space use was asynchronous in time, except in the case of the female lion in Nyathi where there was considerable overlap at the finest temporal scale investigated (i.e. monthly). Herd-associated eland in Nyathi were killed most frequently in areas associated with high long-term use by the female lion, showing that failure to avoid the female lion at a fine scale increased predation risk for eland. Secondly, eland in Nyathi were killed most frequently in habitat types of intermediate structural density (i.e. mixed habitats), which limited the viewshed of eland but presumably offered increased forage. This habitat type was selected for by eland, despite it being associated with a higher level of predation risk. This suggested that eland in Nyathi traded off safety for forage. In MZNP, predation of herd-associated eland was low which reflected the limited spatial overlap between eland and lion at a fine scale in this reserve. In Nyathi, but not in MZNP, eland responded to lions within 1.5 km by increasing their movement rate, with this response particularly strong at night. The findings of this study show that eland responses to lions are scale-dependent and site-specific. Furthermore, eland responses to lions were shown to be lion gender-specific. This study has increased our understanding of the antipredatory behaviours of eland and demonstrated the importance of investigating prey responses to predation risk at multiple scales.

Large carnivores play many vital biological, economic, and conservation roles, however, their biological traits (low population densities, cryptic behavior) make them difficult to monitor. Pumas have been particularly difficult to study because the lack of distinctive markings on their coats prevents individual identification, precluding mark-recapture and other similar analyses. Further, compared to temperate areas, research on the interspecific interactions of Central American felids is particularly lacking. I used single- and multi-season, single-species occupancy models and two-species co-occurrence models to analyze camera trapping and habitat data collected at eight study sites across Belize. Puma occupancy was positively influenced by jaguar trap success, understory density, canopy cover, and human trap success, and negatively influenced by stream density. Jaguar trap success was the best predictor of where pumas occurred, while prey species were not found to influence puma occupancy. Mean occupancy was 0.740 (0.013) and ranged from 0.587 (0.042) to 0.924 (0.030). Over time, puma occupancy rates were generally high (> 0.90) and stable. Puma occupancy was higher in logged areas, suggesting that current levels of natural resource extraction at those sites were not detrimental to the species. Co-occurrence modeling showed little evidence for interactions between the carnivores, suggesting that jaguars may be acting as an umbrella species and that conservation efforts directed at jaguars are likely to benefit the other carnivores, including pumas. Overall, these findings are positive for puma conservation, but human-induced land use change is expanding and further monitoring will give us insight into how pumas respond to human encroachment.
Master of Science

Foraging forms the cornerstone of an animal’s life-history. An individual's foraging success shapes the demography and health of a population. Understanding key facets of maternal foraging behaviour are crucial to get a holistic picture of both regional and local environmental factors that drive foraging behaviour. This study aimed to measure the maternal foraging behaviour of a marine top predator, the Subantarctic fur seal (Arctocephalus tropicalis), from Marion Island (MI) over a range of spatial and temporal scales. Arctocephalus tropicalis females from MI have one of the longest duration foraging trips for the species. They are most similar to conspecifics at temperate Amsterdam Island, but differ considerably from those at subantarctic Îles Crozet and Macquarie Island. Hitherto, no diving data existed for MI females. I illustrate how their diving behaviour is more similar to individuals from Îles Crozet despite their differences in foraging trip parameters. Together with Îles Crozet, MI females have one of the deepest mean diving depths (34.5 ± 2.2 m , 45.2 ± 4.8 m summer and winter respectively) and longest dive durations (70.2 ± 3 s , 104.3 ± 7.8 s summer and winter respectively) for the species. In summer, females follow the diel vertical migration of their myctophid prey. Counter intuitively, during the winter, females performed short and shallow crepuscular dives, possibly foraging on different prey. Considering that these individuals dive in deep waters, this is most likely related to myctophids occupying lower depths in the water column during winter. At dusk and dawn they are inaccessible to diving fur seals. At-sea data from multiple foraging trips per female illustrated that females have both a colony- and individual preferred foraging direction which varied seasonally. Individuals travelled consistently in the same direction regionally, but locally appear to track prey in a heterogeneous environment. The few trips in the winter to the west of MI suggest that this is a short-term response to varying prey availability rather than a long-term foraging tactic. Six years of observer-based attendance cycle data were used to augment telemetry data. Multi-state mark-recapture models were used to determine the probability of a female being missed when she was present (detection probability). Attendance data were corrected accordingly. Neither El Niño (EN) nor anomalous seasurface temperature (SSTa) influenced any of the attendance cycle parameters, as foraging trip duration is a poor predictor of weak environmental change. Only season and pup sex had a significant impact on female provisioning rates. Foraging trip duration was longer during winter than during summer. Females spent a higher percentage of time on land when they had female pups rather than male pups. Although observational attendance data remain useful it ideally requires concomitant data on pup growth, production and female body condition to elucidate changes in female provisioning rates. Temporally, season had the most influence on female foraging behaviour. Spatially, it appears that a lack of prominent local bathymetrical features overshadows MI's favourable position in the productive Polar Frontal Zone. Arctocephalus tropicalis females from MI work harder at foraging than at any other island population of conspecifics.
Dissertation (MSc)--University of Pretoria, 2013.
gm2014
Zoology and Entomology
Unrestricted

Interacting species systems have complex dynamics that are often subject to change due to internal and external factors. Quantitative modelling approaches to capture demographic fluctuations can be insufficient in the presence of stochastic variation and uncertainty. This thesis establishes new modelling techniques to account for such demographic variations and develops novel numerical simulation tools for solving these systems. These explorations are extended for solving invasive species management problems where robust management actions and efficient use of allocated budgets are necessary.

A variety of prey species are able to detect predators by odours emanating from their urine, feces, fur and anal glands. However, it is unknown whether the odour of a predator’s blood also contains information signalling “predator” to a prey. The aim of the present study was therefore to assess if blood odour from a cat elicits avoidance or anxiety responses in CD-1 mice (Mus musculus). A two-compartment test arena was used to assess place preference, motor activity and fecal excretions when the mice were simultaneously presented with cat blood and a blank control. Additionally, the mice were tested with horse blood and N-pentyl acetate, a fruity odour. The mice did not show avoidance of any of the three odours. Nevertheless, the mice were significantly less active when exposed to cat blood in comparison to horse blood, but did not increase defecation when exposed to cat blood. This suggests that the information mice get by the odour of cat blood did not contain the signal “predator”.

It is essential for prey species to be able to detect predators to avoid them. The sense of smell is used by a number of prey species for this purpose. The aim of the present study was to assess if one of the odourants that make up a predator odour is sufficient to induce a behavioural response in mice (Mus musculus). Two predator odourants were used, 2,2-dimethylthietane and methyl-2-phenylethyl sulfide, which are both found in the secretions of natural predators of mice. An odourant found in fruits, n-pentyl acetate, was also used. All three odourants were presented at a concentration that was a factor of 100 above the olfactory detection threshold of mice. Ten adult predator-naïve CD-1 mice were individually put in a two-compartment chamber one of which contained an odourant while the other contained a near-odourless solvent (diethyl phthalate). The results indicated that methyl-2-phenylethyl sulfide was actively avoided by the mice. Towards 2,2-dimethylthietane and n-pentyl acetate, in contrast, the mice behaved indifferent. Further, the results suggest a significant correlation between the number of switches between the two compartments of the test chamber and the test sessions when the animals were presented with n-pentyl acetate, but not when they were presented with the predator odourants. The results support the notion of an innate fear response towards the predator odourant methyl-2-phenylethyl sulfide in mice, but further studies with more animals and different concentrations of the odourants are necessary.

Seabirds are often used as indicator species for changes in marine ecosystems due to the species visibility and sensitivity to changing conditions, such as changes in prey abundance. They often reside in habitats affected by anthropogenic impacts such as large-scale fisheries and pollution. Understanding the connection between seabirds and their surrounding environment can give us important insight about the ecology of the ocean and how anthropogenic pressures affects it. Studying feeding habits, and foraging behavior especially, is useful for understanding seabird´s responses to changing environments. Feeding studies are commonly used in seabird monitoring and requires a lot of time and resources. Monitoring of seabirds are also logistically challenging, and the risk of disturbing bird colonies must be considered. It can be especially complicated when studying cliff nesting seabirds such as the common murre, Uria aalge, the study species for this thesis. Photography as a method for monitoring seabirds may limit the need of people on site, hence minimize disturbance and save time and resources. This study provides insight in whether it is a viable option to use photography instead of on-site field studies when monitoring sea birds. This was accomplished by installing cameras and monitor a feeding study at Stora Karlsö, Sweden, parallel with performing the usual monitoring in the field. This thesis makes it clear that a camera study very well could replace the field study without any larger concerns, although, improvements need to be considered if the study is to maintain a high quality and for results to be reliable.

It has been frequently mentioned that Colombia is a \"megadiverse\" country, containing a larger number of species than other countries, some of which with much larger territories. However, very few people know what this means and about the benefits that can be obtained from that high diversity, with its proper exploration. One of the benefits refers to the sustainable use of biological resources. Colombia can take advantage of the discovery, conservation or production, and commercialization of species that serve as biological control agents of species that harm certain economic activities. Applied biological control involves the possibility to exploit native biodiversity in the form of conservation or biocommerce for use in activities such as agriculture and livestock. Thus, the studies composing this thesis aimed at initiating the recognition of the richness of predatory mites in the soil of the major rose-producing region of the country, and the evaluation of potential representative predators for use in the management of thrips, the most important pest group on rose cultivation in Colombia. Chapter 1 addressed the importance of the crop, the pest groups and the possible natural enemies to be found in Colombia. In Chapter 2, the results of a general survey of the soil mites in four rose fields in greenhouses during one year and a half was presented. Additionally, the population dynamics of soil the mites and thrips was studied, relating the variation in population levels with environmental factors, including climatic and soil factors. Knowing the variation of populations over time under the usual conditions of crop management can help to understand how a predator can respond in the field when used in pest management programs. To complement those data, especially in what refers to the evaluation of the native fauna, surveys were carried out on patches of secondary vegetation located near two of the rose fields, in a rose field in another area and a patch of secondary vegetation nearby, and in two more extensive and less disturbed patches of the Bogota plateau. With this work, it was possible to verify the great richness of species of soil non-Uropodina Mesostigmata found in the soil of the Bogotá plateau. Only representative mites of the cohort Cercomegistina of the suborder Trigynaspida and of the cohort Gamasina of the suborder Monogynaspida were found. As expected, species and dominant families resemble those reported in temperate areas, given the similar climatic conditions. Both densities and number of species are lower in rose fields than in secondary vegetation patches nearby, which also was expected from the much more uniform ecosystem in rose fields. Additionally, the exclusive presence of some species in rose fields suggests either the introduction of those species with material related to the cultivation system or a drastic change in faunistic composition, turning rare field species abundant in rose fields because of the cultivation system. The variation in densities and presence of species was shown to be more related to soil, especially organic matter and pH, than to climatic characteristics. Therefore, patches of secondary vegetation, whose soils had higher content of organic matter, are more similar to each other, in terms of composition of species, than to the nearby rose fields. The results of the Chapter 2 indicated the need for morphological and taxonomic studies of several of the species found, either because of the poorly detailed descriptions currently available or because of the recognition of new species for science. Thus, in Chapters 3 and 4, the morphological characterization and the description of some of the new species found in the Ascidae, Blattisociidae and Melicharidae (Chapter 3) and Laelapidae (Chapter 4) were presented. It was also observed that many of the previously described species found in this study also need similar studies to be conducted in the future. However, through these two chapters, initial steps were taken to contribute to the recognition of scarcely known mites groups in Colombia to facilitate future ecological studies and the use of those species in pest management programs. The results of Chapter 2 also served as base to select species to be considered in an initial evaluation of the potential of members of the Colombian mesostigmatic fauna as control agents of Frankliniella occidentalis (Pergander), the most numerous thrips species found in this study. Two of the most frequently found predator species, Gaeolaelaps aculeifer (Canestrini) and Parasitus bituberosus Karg, were selected to carry out the second part of this work, whose results were presented in Chapters 5 and 6, respectively. For each predator, the study consisted of the evaluation the predation capacity on F. occidentalis, the oviposition rates on this and other prey species, and of the determination of their life table parameters. Gaeolaelaps aculeifer is presently used in other countries for the control of different pests including thrips. The results showed that both predators are able to feed and reproduce when pre-pupae and pupae of F. occidentalis were part of their diets. The biological characteristics of the Colombian population of G. aculeifer are comparable to those reported for other populations of the same predator, suggesting its potential for use to control F. occidentalis in Colombia. Although the presence of a complementary prey (Aleuroglyphus ovatus (Troupeau)) in the system led to a small reduction of its predation rate of F. occidentalis pre-pupae and pupae, the results show that A. ovatus can be used as factitious food for mass rearing or as complementary food in periodic releases, when the pest in not abundant. It was shown that P. bituberosus deutonymph needs a stimulus for the emergence of adults, which, in this case, was the pairing with a mite of the opposite sex. Nematodes are not only necessary in the diet in the immature stage of this predator, but also improve its performance, as indicated by the higher fecundity and the net and intrinsic reproductive rates on this prey. The results indicated as warranted the conduction of larger scale investigations on the possible use of G. aculeifer and P. bituberosus for thrips control, by periodic releases, as well as the evaluation of other predator species found less frequently. Larger-scale evaluations would initially consist of experiments in pots and later on at the semi-field and field conditions. Results of the relation between predator density and environmental factors also suggest to be important the conduction of studies to evaluate the possibility to increase predator performance by increasing the level of soil organic matter content or pH level.
Tem sido freqüentemente mencionado que a Colômbia é um país \"megadiverso\", contendo um número maior de espécies que outros países, alguns dos quais com territórios muito mais extensos. No entanto, poucos sabem o que isso significa e os benefícios que podem ser obtidos dessa alta diversidade, com sua exploração adequada. Um dos benefícios refere-se ao uso sustentável de recursos biológicos. A Colômbia pode aproveitar a descoberta, a conservação ou a produção e a comercialização de espécies que servem como agentes de controle biológico de espécies que prejudicam certas atividades econômicas. O controle biológico aplicado envolve a possibilidade de explorar a biodiversidade nativa na forma de conservação ou biocomércio para uso em atividades como agricultura e pecuária. Assim, os estudos que compõem esta tese visam a iniciar o reconhecimento da riqueza de ácaros predadores no solo da região produtora de roseiras da Colômbia, e a avaliação de potenciais predadores representativos para uso no manejo de tripes, o mais importante grupo de pragas no cultivo de rosas na Colômbia. O Capítulo 1 abordou a importância da cultura, dos grupos de pragas e dos possíveis inimigos naturais encontrados na Colômbia. No Capítulo 2, os resultados de um levantamento geral dos ácaros de solo em quatro campos de cultivo de rosas em estufas durante um ano e meio foram apresentados. Além disso, a dinâmica populacional de ácaros e tripes no solo foi estudada, relacionando a variação nos níveis populacionais com fatores ambientais, incluindo fatores climáticos e do solo. Conhecendo a variação das populações ao longo do tempo sob as condições usuais de manejo de culturas pode ajudar a entender como um predador pode responder no campo quando usado em programas de controle de pragas. Para complementar esses dados, especialmente no que se refere à avaliação da fauna nativa, foram realizados levantamentos em fragmentos de vegetação secundária localizados próximos a dois dos campos de cultivo de rosas, num cultivo de rosa em outra área e um fragmento de vegetação secundária nas proximidades, e em dois fragmentos mais extensos e menos perturbados do planalto de Bogotá. Com este trabalho, foi possível verificar a grande riqueza de espécies de ácaros Mesostigmata (não incluindo os Uropodina) encontradas no solo do planalto de Bogotá. Apenas ácaros representativos da coorte Cercomegistina da subordem Trigynaspida e da coorte Gamasina da subordem Monogynaspida foram encontrados. Como esperado, espécies e famílias dominantes se assemelham aos grupos relatados em áreas temperadas, dadas as condições climáticas semelhantes. Tanto as densidades quanto o número de espécies são menores nos campos de cultivo de rosas do que nos fragmentos de vegetação secundária próximos, o que também era esperado tendo em vista o ecossistema muito mais uniforme nos campos cultivo de rosas. Além disso, a presença exclusiva de algumas espécies nestes campos sugere que estas tenham sido introduzidas com material relacionado ao sistema de cultivo ou que tenha havido uma mudança drástica na composição faunística, tornando abundantes em campos de cultivo de rosas as espécies que são raras na vegetação secundária, devido às características do sistema de cultivo. A variação nas densidades e presença de espécies mostaram-se mais relacionadas às características do solo, especialmente níveis de matéria orgânica e pH, do que às características climáticas. Portanto, fragmentos da vegetação secundária, cujos solos tinham maior conteúdo de matéria orgânica, são mais semelhantes entre si, em termos de composição das espécies, do que aos campos de rosas próximos. Os resultados do Capítulo 2 indicaram a necessidade de estudos morfológicos e taxonômicos de várias das espécies encontradas, pelas descrições pouco detalhadas atualmente disponíveis ou pelo reconhecimento da ocorrência de espécies novas para a ciência. Assim, nos Capítulos 3 e 4, uma caracterização morfológica e a descrição de algumas das espécies novas encontradas em Ascidae, Blattisociidae e Melicharidae (Capítulo 3) e Laelapidae (Capítulo 4) foram apresentadas. Também foi observado que muitas das espécies descritas anteriormente e encontradas neste estudo necessitam de estudos semelhantes, a serem realizados no futuro. No entanto, através destes dois capítulos, os passos iniciais foram dados, para contribuir para o reconhecimento de grupos de ácaros pouco conhecidos na Colômbia, para facilitar futuros estudos ecológicos e o uso destas espécies em programas de controle de pragas. Os resultados do Capítulo 2 também serviram como base para a seleção de espécies a serem consideradas em uma avaliação inicial do potencial de membros da fauna mesostigmática colombiana como agentes de controle de Frankliniella occidentalis (Pergander), a espécie mais numerosa de tripes encontrada neste estudo. Duas das espécies de predadores mais encontradas, Gaeolaelaps aculeifer (Canestrini) e Parasitus bituberosus Karg, foram selecionadas para realizar a segunda parte deste trabalho, cujos resultados foram apresentados nos Capítulos 5 e 6, respectivamente. Para cada predador, o estudo consistiu na avaliação da capacidade de predação em F. occidentalis e das taxas de oviposição quando alimentando-se desta e de outras presas, assim como na determinação dos parâmetros de sua tabela de vida. Gaeolaelaps aculeifer é atualmente usado em outros países para o controle de diferentes pragas, incluindo tripes. Os resultados mostraram que ambos os predadores são capazes de se alimentar e se reproduzir quando pré-pupas e pupas de F. occidentalis fazem parte de suas dietas. As características biológicas da população colombiana de G. aculeifer são comparáveis àquelas relatadas para outras populações do mesmo predador, sugerindo seu potencial para uso no controle de F. occidentalis na Colômbia. Embora a presença de uma presa complementar (Aleuroglyphus ovatus (Troupeau)) no sistema tenha levado a uma pequena redução de sua taxa de predação de pupas e pupas de F. occidentalis, observou-se que A. ovatus pode ser usado como alimento factício para a criação massal ou como alimento complementar em liberações periódicas, quando a praga não seja suficientemente abundante para manter a população deste predador. Foi demonstrado que a deutoninfa de P. bituberosus precisa de um estímulo para a emergência dos adultos, que, neste caso correspondeu ao seu pareamento com um ácaro do sexo oposto. Os nematoides não apenas são necessários na dieta na fase imatura desse predador, mas também melhoram seu desempenho, como indicado pela maior fecundidade e pelas taxa liquida de reproducao e capacidade innata de aumentar em numero dessa presa. Os resultados indicaram como justificável a realização de estudos em maior escala sobre o possível uso de G. aculeifer e P. bituberosus no controle de tripes, por meio de liberações periódicas, bem como a avaliação de outras espécies predadoras encontradas com menor frequência. As avaliações em larga escala consistiriam inicialmente em experimentos em vasos e, posteriormente, em semi-campo e campo. Os resultados da relação entre densidade de predadores e fatores ambientais também sugerem ser importante a realização de estudos que avaliem a possibilidade de aumentar o desempenho dos predadores, aumentando o nível de matéria orgânica do solo ou o pH.

The Cape Horn Biosphere Reserve is located at the southern tip of South America. This large archipelago is considered one of the last pristine areas left on the world. Despite it being an unpopulated area with most of the native forest cover intact, it has not been exempt from biological invasions, one of the major drivers of biodiversity loss. Three species that naturally interact in their native range in North America – American beavers (Castor canadensis), muskrats (Ondatra zibethicus), and American mink (Neovison vison) – were independently introduced in this remote region. In my dissertation, I investigated (i) the hypothesis of niche expansion in the invasive mink population on Navarino Island towards terrestrial habitats; (ii) potential mink impact on breeding success of forest-bird populations; (iii) habitat selection of small-rodent species and their perception on the mink's novel predation risk; and (iv) the dynamics of multiple-species invasions under the hypothesis of an invasional meltdown. Additionally, I worked within the framework of environmental philosophy. I provide an example of combining ecological and cultural dimensions within the International Long-Term Ecological Research network to disentangle the ethical dilemmas that surround the management of invasive species. I finally proposed a management plan based on the idea of multiple invasive species management, discussing potential solutions to overcome the challenges that the control of invasive species represent and to more effectively protect the biological integrity of the native ecosystems.

This dissertation demonstrates the construction of the Panthera Population Persistence (PPP), an individual-based model for the Sumatran tiger (Panthera tigris sumatrae) which provides proper theoretical and application frameworks for the conservation of this tiger sub-species in central Sumatra. The PPP model was developed to gain insight into tiger-preyhabitat relationships as well as the effect of human impacts on the persistence of tiger populations. The model addresses three main problems for the survival of the Sumatran tiger: tiger poaching, prey depletion, and habitat loss. The description of the PPP model serves as an in-depth study of existing literature and covers the most important factors of existing models for tiger conservation. Existing modelling approaches have been improved by the inclusion of finer description of individual-level traits and behaviours in the PPP model. The modelling approach allows a direct inter-relationships between individuals and their environment. The relationship between individual behaviours, intrinsic states, and external factors are simulated spatially explicitly in a bottom-up approach where the emergence of the population dynamics of tiger and prey can be observed under different scenarios. The integration between the PPP model and geographical information system (GIS) has provided a much more meaningful spatial data by revealing the mechanism of the response of individuals to the present land-use types. The relative importance of the parameters within the PPP model was tested using two modes of sensitivity analysis: The Morris Method and the traditional One-factor-at-a-time method. The results provided guidance for the application of reasonable sensitivity analysis during the development of individual-based models. The Morris Method suggested that the overall output of the PPP model showed a high sensitivity on the change of time required by a tigress to take care of cubs. The analysis also revealed that the number of dispersers was sensitive toward perceptual distance of individuals to detect the presence of prey. Comparison with a similar predator-prey models provided insight into the predator-prey relationship. The comparison also suggested that perceptual distance of the individual is important for any spatially explicit individual-based model involving predator-prey relationships. The parameterization of the individual perceptual distance of tigers was tested by using existing literature on prey consumption by tigers as a benchmark. The simulation results were within the range of scientific acceptance for the number of prey killed by a tiger. Thus, further use of the set of parameters for a tiger’s perceptual distance is less uncertain for the output of the PPP model. The effect of habitat quality and landscape configuration on the mortality and migration of prey were evaluated through the use of virtual habitats and landscapes. The findings suggested that a good habitat quality enables prey survival, increases the population available for predation by tigers. When a low-quality habitat is combined with a high-quality habitat, the number of migrating prey was high, reducing resources for tigers. This suggested that landscape composition should be considered when predicting population persistence of the Sumatran tiger. Optimal movement of two different prey resulted in a high density of prey in high-quality habitat, providing a concentration of prey in a tiger’s habitat, but resulted in a lower tiger predation rate than random movement and species specific movement. The PPP model has been applied to evaluate the effect of poaching, prey depletion, and their combination for the probability of extinction of a tiger population. The results from the evaluation showed that prey depletion, tiger poaching, and a combination of both, created a 100% probability of extinction within 20 years if the density and frequency of those threats at high rates. However, the duration of those threats in the system caused a 100% probability of extinction from tiger poaching. The results are able to contribute to optimize anti-poaching programs in future, to reduce significantly the probability of total extinction of Sumatran tiger. Furthermore, various landscape configurations have been tested against the probability and time of extinction for the Sumatran tiger population. The integration of spatial GIS-data in the model provides an insight into the relationship between tiger-prey-habitat. The results suggested that habitat quality surrounding a protected area plays an important role for the persistence of the Sumatran tiger population. This study also recommends agroforestry systems as reasonable land-use type in the vicinity of protected areas. They provide not only positive effects for tiger conservation purpose but they also appear as adaptable to the current land-use situation in Sumatra island
Die vorliegende Dissertation beschreibt die Entwicklung des Panthera Populations Persistence (PPP) Modells, eines individuenbasierten Simulationsmodells für den Sumatra-Tiger (Panthera tigris sumatrae). Dieses stellt einen geeigneten theoretischen und anwendungsbezogenen Rahmen für den Schutz dieser Tiger-Unterart in Zentralsumatra bereit. Das PPP-Modell wurde entwickelt, um Einblicke in die Tiger-Beute-Habitat-Beziehungen zu gewinnen, sowie um den Effekt anthropogener Einflüsse auf den Fortbestand von Tigerpopulationen abzuschätzen. Dabei werden die drei Hauptprobleme für das Überleben des Sumatra-Tigers analysiert: die Wilderei, der Rückgang von Beutetieren und der Verlust von geeigneten Habitaten. Die Beschreibung des PPP-Modells gibt zunächst einen umfassenden Überblick zum aktuellen Wissensstand auf dem Gebiet des Tigerschutzes und integriert die wichtigsten Faktoren bereits existierender Modellansätze. Diese konnten durch die Einbeziehung einer detaillierten Beschreibung von individuellen Merkmalen und Verhalten verbessert werden. Das PPPModell stellt somit das Individuum in einen direkten Zusammenhang mit dessen Umwelt. Die Beziehung zwischen individuellem Verhalten, intrinsischen Merkmalen und externen Faktoren werden räumlich-explizit in einem bottom-up Ansatz simuliert. Damit kann sowohl die Populationsdynamik des Tigers als auch die seiner Beutetiere unter verschiedenen Annahmen beobachtet werden. Die Verknüpfung des PPP-Modells mit Geographischen Informationssystemen (GIS) bietet die Möglichkeit, die Reaktionsmechanismen der Individuen basierend auf der gegenwärtigen Landnutzungssituation zu simulieren und somit realitätsnahe räumliche Daten zu generieren. Die relative Bedeutung der Modell-Parameter auf die Simulationsergebnisse kann durch Sensitivitätsanalysen ermittelt werden. Hier wurden zwei verschiedene Ansätze verwendet: die Morris-Methode und die herkömmliche One-factor-at-a-time Methode. Der Vergleich beider methodischen Ansätze zeigte somit beispielhaft die Eignung unterschiedlicher Sensitivitätsanalysen für individuenbasierte Modelle auf. Die Morris-Methode zeigte, dass das Gesamtergebnis des PPP-Modells eine hohe Sensitivität gegenüber der Veränderung der Zeit aufweist, die ein Tigerweibchen braucht, um ihre Jungen aufzuziehen. Die Analyse zeigt auch, dass die Anzahl an abwandernden Tigern sensitiv gegenüber der IndividuellenWahrnehmungsdistanz von Beute ist. Der Vergleich mit einem ähnlichen Räuber-Beute-Modell lässt vermuten, dass diese Wahrnehmungsdistanz eines Individuums generell als ein entscheidender Faktor für Räuber-Beute-Beziehungen in räumlich-expliziten Individuenmodellen an- gesehen werden kann. Die Parametrisierung der IndividuellenWahrnehmungsdistanz des Tigers wurde so gewahlt, dass die damit ermittelten Simulationsergebnisse den Beutekonsum des Tigers, wie in der Literatur beschrieben, weitgehen widerspiegeln. Sie ist somit für die weitere Anwendung im PPP-Modell ausreichend gut beschrieben. Simulationsszenarien, welche verschiedene Habitatqualitäten sowie Landnutzungsmuster berücksichtigen, zeigen auch deren Bedeutung für die Mortalität und Migration der Beutetiere. Eine gute Habitatqualität hat eine geringe Mortalität der Beutetiere zur Folge, welche dann wiederum für den Tiger in ausreichender Zahl zur Verfügung stehen. Treten geringe Habitatqualitäten angrenzend an ein Habitat mit hoher Qualität auf, führte dies zu einer hohen Anzahl an abwandernden Beutetieren, womit sich die Ressourcen für den Tiger verringern. Die Landschaftsmerkmale sollten also bei der Vorhersage des Populationsfortbestandes des Sumatra-Tigers berücksichtigt werden. Die optimale Bewegung von zwei verschiedenen Beutetieren ergab eine hohe Beutedichte in einem Habitat mit hoher Qualität und stellte konzentriert Beute in einem Tigerhabitat bereit. Allerdings resultierte dies auch in einer geringeren Prädationsrate des Tigers, verglichen mit zufälligen oder artenspezifischen Bewegungen. Das PPP-Modell wurde angewandt, um die Auswirkungen von Wilderei, Beutetierrückgang sowie die Kombination beider Faktoren auf die Aussterbewahrscheinlichkeit einer Tigerpopulation zu bewerten. Die Ergebnisse zeigen, dass die genannten Faktoren eine 100-prozentige Aussterbewahrscheinlichkeit innerhalb von 20 Jahren zur Folge haben, wenn die Dichte und Häufigkeit dieser Bedrohungen hoch sind. Die Dauer dieser Bedrohungen im System verursachte allerdings eine 100-prozentige Aussterbewahrscheinlichkeit nur für die Wilderei von Tigern. Betrachtet man unabhängig von Dichte und Häufigkeit einzig die Dauer der Bedrohung, führt lediglich die Wilderei zum 100%-igen Aussterben. Diese Ergebnisse können maßgeblich dazu beitragen, zukünftig Schutzprogramme gegen die Wilderei zu optimieren, um das Aussterben des Sumatra-Tigers zu verhindern. DesWeiteren wurde der Einfluss von unterschiedlichen Landnutzungsmustern auf die Aussterbewahrscheinlichkeit und -zeit einer Sumatra-Tigerpopulation aufgezeigt. Die Integration von räumlichen GIS-Daten in das Modell ermöglichte einen Einblick in die Beziehungen zwischen Tiger, Beutetieren und Habitat. Die Ergebnisse zeigen, dass die Habitatqualität um Schutzgebiete herum eine wichtige Rolle für den Fortbestand der Population spielt. Die vorliegende Arbeit empfiehlt Agroforstsysteme als eine geeignete Landnutzungsform in der Nähe von Schutzgebieten, welche sowohl positive Effekte für den Tigerschutz bietet als auch mit den gegenwärtigen Landnutzungsmustern in Sumatra vereinbar erscheint

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Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES
The destruction of habitats is one of the most important factors leading to species extinction and loss of diversity, which is one of the most studied issues in biological and conservation in recent years. Since the human intervention on ecosystems is growing, resulting in fragmentation and loss of habitats, it is important to analyze the effects of these changes in spatial configuration have on ecological processes. In this work we aimed to study the dynamics of a model in which a predator and prey interact in a homogeneous environment with spatial structure when a proportion of the system is destroyed. We conducted this analysis using techniques from computer simulations as well as analytical tools. We obtained three different regimes depending on the values of model parameters in which there is coexistence of prey and predator, extinction of the predator with survival of prey and the extinction of prey leading to the extinction of the predator. We also studied the effects of destruction of a proportion of habitat on the system, and compared the results with those obtained for the case without fragmentation. We observed that increasing the number of destroyed sites leads to a reduction of the population of predators. We also observed that for a destruction of more than 15% of the sites, the regime of extinction of two species is no longer observed.
A destruição de habitats é um dos fatores mais relevantes que levam à extinção das espécies e perda da diversidade biológica e que é um dos assuntos de conservação mais estudados e debatidos nos últimos anos. Uma vez que a taxa de modificações humanas sobre os ecossistemas vem crescendo, resultando na fragmentação e perda de habitat, é importante analisar os efeitos que essas mudanças na configuração espacial têm nos processos ecológicos. Nesse trabalho tivemos como objetivo estudar a dinâmica de um modelo em que um predador e sua presa interagem em um ambiente homogêneo com estrutura espacial quando uma proporção do sistema é destruído. Realizamos essa análise através de técnicas de simulações computacionais e também ferramentas analíticas. Obtivemos três regimes diferentes dependentes dos valores dos parâmetros do modelo: coexistência de presa e predador, extinção do predador com sobrevivência da presa e extinção da presa levando a extinção do predador. Estudamos também os efeitos da destruição de uma proporção do habitat sobre o sistema, e comparamos os resultados com aqueles obtidos para o caso sem fragmentação. Observamos que o aumento na proporção de sítios destruídos leva a uma diminuição da população de predadores. Também observamos que para uma destruição de mais que 15% dos sítios, o regime de extinção das duas espécies não é mais observado.