Microclimate influence in a physiological model of cattle-fever tick (Boophilus spp.) population dynamics

Since their official eradication from the US in 1943, the cattle-tick species Boophilus microplus and Boophilus annulatus, vectors of bovine babesiosis, frequently have penetrated a quarantine zone established along the Texas–Mexico border designed to exclude them. Inspection and quarantine procedures have eradicated reinfestations successfully within the US, but increasing acaricide resistance in Mexican B. microplus populations poses a threat to future eradication efforts. Better understanding of interrelationships among Boophilus populations, their hosts, and vegetation communities in south Texas could improve prediction of the behavior of reintroduced Boophilus populations and increase management options. To this end, we constructed a simulation model to evaluate how microclimate, habitat (i.e. vegetation) heterogeneity, and within-pasture cattle movement may influence dynamics of Boophilus ticks in south Texas. Unlike previous Boophilus tick models, this model simulates dynamics at an hourly time-step, calculates all off-host dynamics as functions of temperature and relative humidity, and runs with ground-level microclimate data collected bi-hourly in three different habitat types. Sensitivity analysis of the model showed that temperatures and relative humidities created by habitat type, as well as engorged female mass, influenced tick population dynamics most strongly. Host habitat selection, initial number of larvae per cow, and the number of cells into which the simulated pasture was divided also had a strong influence. Population dynamics appeared moderately sensitive to the proportion of Bos indicus in cattle genotypes and the larval attachment rate, while appearing relatively insensitive to factors such as mortality rate of engorged females. When used to simulate laboratory experiments from the literature, the model predicted most observed life-history characteristics fairly well; however, it tended to underestimate oviposition duration, incubation duration, and egg mortality and overestimate larval longevity, especially at low temperatures and high humidities. Use of the model to predict Boophilus population dynamics in hypothetical south Texas pastures showed that it reasonably generated qualitative patterns of stage-wise abundances but tended to overestimate on-host tick burdens. Collection and incorporation of data that appear not to exist for Boophilus ticks, such as larval lipid content and lipid-use rates, may improve model accuracy. Though this model needs refinements such as a smaller spatial resolution, it provides insight into responses of B. microplus or B. annulatus populations to specific weather patterns, habitat heterogeneity, and host movement.     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

Quantitative composition of the defensive secretion ofBledius species (Coleoptera: Staphylinidae: Oxytelinae) is adapted to naturally occurring predators

Summary The adaptation of defensive secretions to their target organisms was examined for the abdominal gland secretions ofBledius furcatus, B. spectabilis andB. arenarius. Therefore the target organisms of the secretion of theseBledius species (i.e. their predators) had to be identified. At the collection sites examined these were the earwigLabidura riparia, the antCataglyphis bicolor, the flyLispe candicans, different carabids of the generaPogonus, Dichirotrichus, Dyschirius, Bembidion andCalathus and the wading birdsHaematopus ostralegus andCalidris alba. The secretion of the abdominal glands contains the toxin ptoluquinone dissolved in either-dodecalactone and 1-undecene (B. furcatus andB. spectabilis) or in octanoic acid and octyloctanoate (B. arenarius). The ratio of these solvents is species-specific. Application experiments using some of the natural insect predators (L. riparia, C. bicolor, Pogonus, Di. gustavii, Dyschirius) revealed that these solvent ratios provided a more effective deterrent than other possible ratios. Thus by combining the solvents in certain ratios, the capability of cuticular penetration and therefore the effectiveness of the defensive secretions are adapted to their natural targets.     

Orientation ofRhizophagus grandis (Coleoptera: Rhizophagidae) to oxygenated monoterpenes in a species-specific predator-prey relationship

Summary The predator,Rhizophagus grandis, is linked to its specific prey,Dendroctonus micans, by semiochemical signals regulating oviposition and long range orientation. A mixture of simple oxygenated monoterpenes [(–)—fenchone, (–)—pinocamphone, rac. camphor, terpinene-4-ol, borneol, fenchol and verbenone], identified from the frass ofD. micans has been found to be extremely active in a flight windtunnel as a long—range attractant for the predator. The mixture elicited 84 % of the response to larval frass of the prey. Excluding pinocamphone from the mixture did not influence its activity. Also, changing the absolute configurations of some of the components (fenchol, terpinene-4-ol and borneol) did not influence the predators' response. However, the addition of (–)—-terpineol increased the attractivity of the synthetic blend to almost that of larval frass of the prey. Ecological implications of the identified semiochemicals and their use in pest management are discussed.     

A concept model to estimate the potential distribution of the Asiatic citrus psyllid (Diaphorina citri Kuwayama) in Australia under climate change—A means for assessing biosecurity risk

Increasing global temperatures as a result of climate change are widely considered inevitable for Australia. Despite this, the specific effects of climate change on Australian agriculture are little studied and the effects on agricultural pests and diseases are virtually unknown. In this paper we consider the impact of climate change on the Asiatic citrus psyllid (Diaphorina citri Kuwayama [Hemiptera: Psyllidae]); one of two known vectors of huanglongbing (citrus greening); a debilitating disease which is caused in Asia by a phloem-limited bacterium ‘Candidatus Liberibacter asiaticus’ (α-Proteobacteria). D. citri does not occur in Australia, but if introduced would pose a major threat to the viability of the Australian citrus industry and to native Citrus species. This paper presents an approach developed to understand how climate change may influence the behaviour, distribution and breeding potential of D. citri. Here we developed and describe an initial dynamic point model of D. citri biology in relation to its citrus host and applied it to a scenario of increasing temperatures, as indicators of climate change, on a continental scale. A comparison between model outputs for the three time frames considered (1990, 2030 and 2070) confirms that increasing temperatures projected under climate change will affect the timing and duration of new citrus growth (flush) necessary for psyllid development throughout Australia. Flushing will start progressively earlier as the temperature increases and be of shorter duration. There will also be a gradual southward expansion of shorter durations of the occurrence of flush. Increasing temperatures will impact on D. citri both directly through alteration of its temperature dependant development cycle and indirectly through the impact on the host flushing cycle. For the whole of Australia, a comparison between model outputs for the three scenarios considered indicates the seasonality of D. citri development will change to match changes in citrus flush initiation. Results indicate that the risk of establishment by D. citri is projected to decrease under increasing temperatures, mainly due to shortened intervals when it can feed on new leaf flushes of the host. However, the spatially heterogeneous results also suggest that regions located on the southern coastline of Australia could become more suitable for D. citri than projected under current temperatures. These results confirm the value of a linked host-pest approach as based on D. citri climatic requirements alone the model would have accounted only for shorter development periods and predicted an increased risk of potential distribution.     

Behavior and ecology of Old World Luperini beetles of the genusAulacophora (Coleoptera: Chrysomelidae)

Summary The attraction of Old World leaf beetles in the genusAulacophora to kairomones and parakairomones which are effective lures for New WorldDiabrotica andAcalymma were investigated. Beetles captured on sticky traps baited with single and multicomponent lures were no different from the control traps for two species ofAulacophora. Yellow colored traps and squash blossoms are attractive toAulacophora beetles which detect sub-microgram quantities of cucurbitacins on silica gel. Leaf feeding behavior and flight activity data are correlated with varietal preference of threeAulacophora species. The common response byDiabrotica andAulacophora to cucurbitacins reinforces the two groups' coevolutionary association with the Cucurbitaceae. The apparent lack of a common response toCucurbita blossom volatiles suggests recent evolutionary pathways are substantially different for these two groups of beetles.     

Sequestration of prenylated benzoic acid and chromenes by Naupactus bipes (Coleoptera: Curculionidae) feeding on Piper gaudichaudianum (Piperaceae)

The curculionid beetle Naupactus bipes (Germar, 1824) (Coleoptera: Curculionidae: Brachycerinae) has shown feeding preference for leaves of Piper gaudichaudianum, demonstrating an unexpected specificity for an insect considered to be a generalist. The leaves of P. gaudichaudianum contain the prenylated chromenes gaudichaudianic acid (4, major compound) and its methyl ester (5) in addition to a chromene (3) lacking one prenyl residue. In addition to 4, roots contain the chromone methyl ester (1) and methyl taboganate (2, major compound). Feeding on roots, larvae of N. bipes sequester exclusively the root-specific compounds 1 and 2. Adult beetles sequester the leaf-specific chromenes 3 and 4, but were found to also contain compounds 1 and 2 that are absent in leaves. Therefore, it is suggested that 1 and 2 are sequestered by larvae and can be found in the body of adult insects after long-term storage. In addition, 3 and 4, the major compounds in leaves were found to be associated with the eggs.     

A method for building spatial model of annual weed seed dispersal from experimental data and its application to simulating Bromus sterilis population dispersal

Spatial model of annual weed seed dispersal, in this article, was theoretically derived. According to the requirements of building the spatial model, we designed and done an indoor experiment of weed seed dispersal by wind. In the experiment, the seeds of Bromus sterilis were released at 100 cm height under different wind velocity conditions. Based on the experimental data, the spatial models of seed dispersal of the weed species were built, which were divided into three types according to the coefficient β < 0, β = 0, β > 0. The results showed that dispersal of annual weed seed in any direction obeyed an approximate Gaussian distribution; under the experimental conditions, spatial distribution type of weed seed dispersal changed with variation of wind velocity. Well-known Howard et al.'s model (Howard et al., 1991) of Bromus sterilis seed dispersal is an especial example of the model built in this article. The result of model analysis indicated that the distribution type described by Howard's model was similar to that of seed dispersal of the weed species at the height of 100 cm under the condition of lower wind velocity (about 2.18 m/s). Using CA simulation analysis we found that mean control agent applying to a cell with weed should have a decrease with an increase of wind velocity to prevent weed with the initial configuration from spreading, which implicated less herbicide needs spraying in every cell with weed on average when wind velocity increases.     

A simulation model of rhizome networks for Fallopia japonica (Japanese knotweed) in the United Kingdom

Fallopia japonica (Japanese knotweed) is an aggressively invasive herbaceous perennial that causes substantial economic and environmental damage in the United Kingdom (UK). As such, it is of considerable concern to councils, environmental groups, private landowners and property developers. We construct a 3D correlated random walk model of the development of the subterranean rhizome network for a single stand of F. japonica. The formulation of this model uses detailed knowledge of the morphology and physiology of the plant, both of which differ in the UK to that of its native habitat due to factors including a lack of predation and competition, longer growth seasons and favourable environmental conditions in the UK. Field data obtained as a part of this study are discussed and used in the model for parameterisation and validation. The simulation captures the field data well and predicts, for example, quadratic growth in time for the stand area. Furthermore, the role of a selection of parameters on long-term stand development are discussed, highlighting some key factors affecting vegetative spread rates.     

Aggregation pheromone ofCarpophilus freemani (Coleoptera: Nitidulidae): A blend of conjugated triene and tetraene hydrocarbons

Summary Males ofCarpophilus freemani Dobson (Coleoptera: Nitidulidae) produce an aggregation pheromone to which both sexes fly in a wind-tunnel bioassay. The major pheromone component (ca. 30 ng per male per day in volatile collections) was identified as (2E,4E,6E)-5-ethyl-3-methyl-2,4,6-nonatriene. A minor component, (2E,4E,6E,8E)-7-ethyl-3,5-dimethyl-2,4,6,8-undecatetraene, was 3–10% as abundant as the major triene and was 5–20% as active when compared at relative doses ranging from natural proportions to 1:1. These compounds act synergistically: a mixture of major and minor components in natural proportions attracted more than twice as many beetles as the major component alone, and the mixture fully accounted for the activity of male-derived volatile collections. Six other male-derived conjugated hydrocarbons, ranging from 2% down to 0.04% as abundant as the major component, were also identified. These are (in order of decreasing bioassay activity when compared on an equal-weight basis): (3E,5E,7E)-6-ethyl-4-methyl-3,5,7-decatriene, (2E,4E,6E)-5-ethyl-3-me-thyl-2,4,6-octatriene, (3E,5E,7E,9E)-8-ethyl-4,6-dimethyl-3, 5,7,9-dodecatetraene, (2E,4E,6E,8E)-3,5,7-trimethyl-2,4,6, 8-undecatetraene, (3E,5E,7E)-5-ethyl-7-methyl-3,5,7-undecatriene, and (2E,4E,6E)-3,5-dimethyl-2,4,6-nonatriene. All structure identifications were confirmed by synthesis. In the wind tunnel, the pheromone acted synergistically with host-type volatiles such as propyl acetate, valeric acid, and ethanol. This concept was verified by fields tests in California, in which there was dramatic synergism between the pheromone and fermenting host materials. Pheromone biosynthesis is discussed.     

The aggregation pheromone ofIps duplicatus and its role in competitive interactions withI. typographus (Coleoptera: Scolytidae)

Summary Ips duplicatus withI. typographus co-inhabiting Norway spruce (Picea abies (L.) Karst.) would benefit from a pheromone blend distinct from that of the larger competitorI. typographus. GC-MS analysis showed thatI. duplicatus males feeding in the host produced ipsdienol (Id),cis-verbenol (cV),trans-verbenol (tV), myrtenol (Mt), andE-myrcenol (EM) and traces of 2-methyl-3-buten-2-ol (MB).I. duplicatus produced Id in approximately racemic form (48.9-54.5% (+)-(S)-isomer). The amounts of Id and EM released over a 9 day period had a maximum of 250 and 5 ng/h/male, respectively, on day 2. Exposure ofI. duplicatus males to myrcene and -pinene resulted in the production of small amounts of Id, cV, tV, Mt, andtrans-pinocarveol, but not of EM. In laboratory bioassays with walking beetles, the pheromone component Id alone was weakly attractive while EM was inactive, but in binary combination with Id strongly synergized attraction. A combination of EM and Id at a release rate equivalent to 100–200 males was more attractive in the field than 70 unmated males in a spruce log. The addition of myrcene ( a suggested pheromone precursor of Id) to Id did not enhance trap caches, while addition of EM increased catches > 10-fold. Subtracting EM from a blend of Id, EM, cV and MB drastically reduced trap catches while subtraction of cV or MB or both had no significant effect. Addition of EM over a wide concentration range to the synthetic pheromone ofI. typographus did not reduce the attraction of females of this species in the laboratory. A two-species pheromone interaction field test releasingI. typographus pheromone components (MB + cV) at 10–1000 male equivalents (ME) andI. duplicatus pheromone (Id + EM) at 0, 10–1000 ME in all possible combinations showed both positive intraspecific dose-response effects and an interspecific inhibition. Higher release rates of EM appeared to inhibitI. typographus, especially males. In a tree colonization model, the response of the two competing species to their respective pheromones show a good separation during the mass-attack with a small initial cross-attraction. It remains to be shown whether either of the two pheromone systems have in fact evolved in the present sympatry, or if they are an incidental effect of ancestry of these phylogenetically distantIps.     

D-Pulegone: A feeding deterrent to deer mice (Peromyscus maniculatus) and prairie voles (Microtus ochrogaster)

Summary D-Pulegone, a monoterpene present in pennyroyålMentha sp. is repellent to a number of vertebrates, decreasing consumption of feed and granular particles. In the present study, several tests were used to determine the reliability of this substance as a feeding deterrent. D-Pulegone (10mM—1M) applied as a coating to apple pieces significantly (P < 0.001) decreased ingestion by deer mice,Peromyscus maniculatus, and prairie voles,Microtus ochrogaster. The deterrent effect of d-pulegone was not diminished following multiple exposures suggesting that this compound may be a promising candidate for use a a general vertebrate repellent. The inhibitory effect of 1M d-pulegone on apple ingestion in voles was abolished when contact was prevented by encasing the stimulus solution in plastic mesh capsules. We conclude that while d-pulegone has repellent properties following direct contact, volatile exposure alone may not be sufficient to elicit avoidance behavior.     

Sequestration of pyrrolizidine alkaloids by larvae ofTellervo zoilus (Lepidoptera: Ithomiinae) and their role in the chemical protection of adults against the spiderNephila maculata (Araneidae)

Summary Life stages of the primitive Australian ithomiine butterflyTellervo zoilus and its larval hostplant, the apocynaceous vineParsonsia straminea, were quantitatively assayed for pyrrolizidine alkaloids (PAs). PAs were found in all stages, mainly as N-oxides, being most concentrated in larvae and freshly-emerged adults. Although adults feed at various confirmed PA sources this probably does not compensate for losses, as wild-caught adults had considerably lower concentrations of PAs. The main alkaloid present in both freshly-emerged adults and in leaves of the host-plant was lycopsamine (1b), stored by butterflies in the N-oxide form. Its presence in higher proportion, in relation to intermedine (1a), in larvae, pupae and adults ofTellervo in relation to the host-plants suggests the inversion of intermedine to lycopsamine by the insects. No 14-member ring macrocyclic PAs were detected in either food-plant or butterflies. Several other PAs were found in wild-caught adults reflecting visits to other PA sources. PAs were also found in high concentrations in freshly-emerged individuals of the danaineEuploea core bred onParsonsia straminea. Wild-caughtDanaus affinis had high PA levels acquired from adult feeding. Freshly emergedEuploea raised onIschnocarpus frutescens andDanaus raised onIschnostemma carnosum (both PA-free) were preyed on by the orb weaving spiderNephila maculata, and showed no PAs. In all cases where PAs were present, most butterflies were liberated, usually cut out of the web unharmed, byNephila. The spider's response was not closely linked to PA concentration, however, and may also depend on hunger levels and previous experience with PA-containing butterflies. All control and other non-PA containing butterflies were consumed although rejection of some body parts of freshly-emergedDanaus affinis suggests that compounds other than PAs may be involved.     

Volatile substances in the pygidial secretion of gyrinid beetles (Coleoptera: Gyrinidae)

Summary Gyrinid beetles are common in freshwater habitats. They have paired pygidial glands with a secretion that contains high molecular weight substances rendering them toxic for predatory fish. In this paper we report on a laboratory study on volatile components released by three different gyrinid species when irritated. The volatile pattern had a clear difference between the different species.Gyrinus substriatus andG. aeratus, both produced 3-methyl-1-butanal and 3-methyl-1-butanol when irritated, but in quite different amounts. The third tested species,G. minutus, did not produce any substances above the detection level. It is suggested that the volatile compounds may be part of the beetles' communication and/or defence system.     

Offensive and defensive sperm competition roles in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae)

Sperm competition is predicted to generate opposing selection pressures on males. On one hand, selection should favour ‘defensive’ adaptations that protect a male’s ejaculate from displacement, while, on the other hand, selection should favour ‘offensive’ adaptations that overcome paternity assurance mechanisms of rivals. Here, we use the sterile male technique to assess sperm precedence when a male dung beetle Onthophagus taurus mates in both a defensive (first male) and an offensive (second male) role. Significant variation in a male’s sperm precedence (both P ₁ and P ₂) was detected, and an individual’s defensive (P ₁) and offensive (P ₂) abilities were positively correlated. Thus, it appears that sexual selection simultaneously selects for ‘defensive’ and ‘offensive’ adaptations in O. taurus. We discuss a variety of male traits in O. taurus that potentially contribute to a male’s ability to be successful when mating in an ‘offensive’ and a ‘defensive’ role.     

Persistence of individually distinctive display patterns in fatigued side-blotched lizards ( Uta stansburiana)

Iguanian lizards perform conspicuous species-typical push-up displays, which are used in territory advertisement, threat, and courtship contexts. Subtle individually distinctive differences in push-up characteristics have been suggested to play a role in the recognition of social partners. However, if the structure of push-ups is responsive to changing physiological states then their capacity to promote recognition may be limited. The current study evaluated whether the push-ups performed while in an experimentally imposed state of fatigue by male side-blotched lizards (Uta stansburiana), retain the individually distinctive characteristics apparent in rested lizards. We found repeatable among-individual differences in the duration and the relative height of push-up components. Repeatability values did not change consistently between the rested and fatigued conditions, nor when both conditions were pooled, indicating that these push-up characteristics do not change with fatigue. Similarly, discriminant functions that were generated using push-ups from one state assigned push-ups performed in an alternate state to the correct individuals. Furthermore, when analyzed independently of individual identity, the values of display parameters examined in the current study did not change significantly between states, and discriminant function analysis could not reliably classify push-ups to the correct state. Taken together the results show that individually distinctive push-up characteristics are robust to effects of fatigue, consistent with their suggested role in social recognition. In the future, video playback experiments can test whether lizards utilize the signature-like characteristics of push-ups to discriminate among individuals.Communicated by T. Czeschlik     

A comparison of four process-based models and a statistical regression model to predict growth of Eucalyptus globulus plantations

In forest management and ecological research, consideration of the impacts and risks of climate change or management optimisation is complex. Computer models have long been applied as tools for these tasks. Process-based forest growth models claim to overcome the limitations of empirical statistical models, but the capacity of different process-based models and modelling approaches have rarely been compared directly. This study evaluates stepwise multiple regression models in comparison to four process-based modelling approaches (3-PG, 3-PG+, CABALA and Forest-DNDC) for greenfield predictions of Eucalyptus globulus plantation growth from 2 to 8 years after planting throughout southern Australia.     

Response of the predatorRhizophagus grandis to host plant derived chemicals inDendroctonus micans larval frass in wind tunnel experiments (Coleoptera: Rhizophagidae,Scolytidae)

Summary Adults of the predatory beetleRhizophagus grandis are strongly attracted to both adult and larval frass of its specific prey,Dendroctonus micans, in walking bioassays. Spruce bark and resin are relatively unattractive. Solvent extracts of larval frass that were attractive toR. grandis adults in a flight wind tunnel contained a mixture of monoterpenes of host plant origin. A synthetic mixture of these monoterpenes, (+)—-pinene, (–)—-pinene, -phellandrene,dl limonene and 3-carene, was responsible for 70–80% of the activity of the most attractive extract. We suggest that a blend of monoterpenes in frass acts as a kairomone forR. grandis but that attraction to monoterpenes is only the first of a sequence of behavioural responses that ensures successful host location and identification.     

Modelling the spatial distribution of red grouper (Epinephelus morio) at Campeche Bank, México, with respect substrate

The red grouper (Epinephelus morio) has historically been one of the most important fisheries of the Campeche Bank area in the Gulf of México, where two fleets from México and one from Cuba participate. Current stocks have been estimated to be a third or less as abundant as in the 1970s; hence, the fishery has been declared to be over-exploited. The federal government of México has expressed the need to investigate the potential use of protected areas as a management tool for recovery of this species. Based on artificial neural network (ANN) modelling, we analysed the spatial distribution of red grouper with respect substrate, considering the demographic structure of the stock. We found a significant relationship between the type of substrate and the three stages of development (juveniles, pre-adults and adults). Juveniles are distributed in shallow waters close to the coast on coral substrates; pre-adults are also associated with coral substrates, but at an intermediate depth; adults are found in deeper waters on sandy substrates. We also identified seasonal reproductive aggregation patterns, always in relation to substrate types. Reproductive aggregation during the winter and early spring in the north-eastern continental shelf and the almost permanent (exception in autumn) concentration of juveniles in coastal waters around the central and eastern coast of Yucatan are of particular importance for management. These patterns represent seasons and areas of high fish vulnerability, which is the basic criterion for further analysis of protected areas.     

A patch use model to separate effects of foraging costs on giving-up densities: an experiment with white-tailed deer (<Emphasis Type="Italic">Odocoileus virginianus</Emphasis>)

The giving-up density of food (GUD), the amount of food remaining in a patch when a forager ceases foraging there, can be used to compare the costs of foraging in different food patches. But, to draw inferences from GUDs, specific effects of foraging costs (predation risk, metabolic and missed opportunities costs) on GUDs have to be identified. As high predation risk, high metabolic costs and abundant food all should produce high GUDs, this does not allow us to infer directly the quality of a habitat. In order to separate the effect of each foraging cost, we developed an optimal foraging model based on food supplementation. We illustrate the use of our model in a study where we assessed the impact of a power line right-of-way in a white-tailed deer (Odocoileus virginianus) winter yard by determining whether the negative effects of cover loss outweigh the positive effects of browse regeneration.