Horizontal viewsheds of large herbivores as a function of woodland structure.

There is great potential for the use of terrestrial laser scanning (TLS) to quantify aspects of habitat structure in the study of animal ecology and behaviour. Viewsheds-the area visible from a given position-influence an animal's perception of risk and ability to respond to potential danger. The management and conservation of large herbivores and their habitats can benefit greatly from understanding how vegetation structure shapes viewsheds and influences animal activity patterns and foraging behaviour. This study aimed to identify how woodland understory structure influenced horizontal viewsheds at deer eye height. Mobile TLS was used in August 2020 to quantify horizontal visibility-in the form of Viewshed Coefficients (VC)-and understory leaf area index (LAI) of 71 circular sample plots (15-m radius) across 10 woodland sites in North Wales (UK) where fallow deer (Dama dama) are present. The plots were also surveyed in summer for woody plant size structure, stem density and bramble (Rubus fruticosus agg.). Eight plots were re-scanned twice in winter to compare seasonal VC values and assess scan consistency. Sample plots with higher densities of small stems had significantly reduced VC 1 m from the ground. Other stem size classes, mean percentage bramble cover and understory LAI did not significantly affect VC. There was no difference in VC between summer and winter scans, or between repeated winter scans. The density of small stems influenced viewsheds at deer eye height and may alter behavioural responses to perceived risk. This study demonstrates how TLS technology can be applied to address questions in large herbivore ecology and conservation.

A reinterpretation of the gap fraction of tree crowns from the perspectives of computer graphics and porous media theory.

The gap fraction (GF) of vegetative canopies is an important property related to the contained bulk of reproductive elements and woody facets within the tree crown volume. This work was developed from the perspectives of porous media theory and computer graphics techniques, considering the vegetative elements in the canopy as a solid matrix and treating the gaps between them as pores to guide volume-based GFvol calculations. Woody components and individual leaves were extracted from terrestrial laser scanning data. The concept of equivalent leaf thickness describing the degrees of leaf curling and drooping was proposed to construct hexagonal prisms properly enclosing the scanned points of each leaf, and cylinder models were adopted to fit each branch segment, enabling the calculation of the equivalent leaf and branch volumes within the crown. Finally, the volume-based GFvol of the tree crown following the definition of the void fraction in porous media theory was calculated as one minus the ratio of the total plant leaf and branch volume to the canopy volume. This approach was tested on five tree species and a forest plot with variable canopy architecture, yielding an estimated maximum volume-based GFvol of 0.985 for a small crepe myrtle and a minimal volume-based GFvol of 0.953 for a sakura tree. The 3D morphology of each compositional element in the tree canopy was geometrically defined and the canopy was considered a porous structure to conduct GFvol calculations based on multidisciplinary theory.

Shortwave Radiation Calculation for Forest Plots Using Airborne LiDAR Data and Computer Graphics.

Forested environments feature a highly complex radiation regime, and solar radiation is hindered from penetrating into the forest by the 3D canopy structure; hence, canopy shortwave radiation varies spatiotemporally, seasonally, and meteorologically, making the radiant flux challenging to both measure and model. Here, we developed a synergetic method using airborne LiDAR data and computer graphics to model the forest canopy and calculate the radiant fluxes of three forest plots (conifer, broadleaf, and mixed). Directional incident solar beams were emitted according to the solar altitude and azimuth angles, and the forest canopy surface was decomposed into triangular elements. A ray tracing algorithm was utilized to simulate the propagation of reflected and transmitted beams within the forest canopy. Our method accurately modeled the solar radiant fluxes and demonstrated good agreement (R 2 ≥ 0.82) with the plot-scale results of hemispherical photo-based HPEval software and pyranometer measurements. The maximum incident radiant flux appeared in the conifer plot at noon on June 15 due to the largest solar altitude angle (81.21°) and dense clustering of tree crowns; the conifer plot also received the maximum reflected radiant flux (10.91-324.65 kW) due to the higher reflectance of coniferous trees and the better absorption of reflected solar beams. However, the broadleaf plot received more transmitted radiant flux (37.7-226.71 kW) for the trees in the shaded area due to the larger transmittance of broadleaf species. Our method can directly simulate the detailed plot-scale distribution of canopy radiation and is valuable for researching light-dependent biophysiological processes.

Increased efficiency in the second-hand tire trade provides opportunity for dengue control.

Dengue fever is increasing in geographical range, spread by invasion of its vector mosquitoes. The trade in second-hand tires has been implicated as a factor in this process because they act as mobile reservoirs of mosquito eggs and larvae. Regional transportation of tires can create linkages between rural areas with dengue and disease-free urban areas, potentially giving rise to outbreaks even in areas with strong local control measures. In this work we sought to model the dynamics of mosquito transportation via the tire trade, in particular to predict its role in causing unexpected dengue outbreaks through vertical transmission of the virus across generations of mosquitoes. We also aimed to identify strategies for regulating the trade in second-hand tires, improving disease control. We created a mathematical model which captures the dynamics of dengue between rural and urban areas, taking into account the movement and storage time of tires, and mosquito diapause. We simulate a series of scenarios in which a mosquito population is introduced to a dengue-free area via movement of tires, either as single or multiple events, increasing the likelihood of a dengue outbreak. A persistent disease state can be induced regardless of whether urban conditions for an outbreak are met, and an existing endemic state can be enhanced by vector input. Finally we assess the potential for regulation of tire processing as a means of reducing the transmission of dengue fever using a specific case study from Puerto Rico. Our work demonstrates the importance of the second-hand tire trade in modulating the spread of dengue fever across regions, in particular its role in introducing dengue to disease-free areas. We propose that reduction of tire storage time and control of their movement can play a crucial role in containing dengue outbreaks.

Asymmetric competition causes multimodal size distributions in spatially structured populations.

Plant sizes within populations often exhibit multimodal distributions, even when all individuals are the same age and have experienced identical conditions. To establish the causes of this, we created an individual-based model simulating the growth of trees in a spatially explicit framework, which was parametrized using data from a long-term study of forest stands in New Zealand. First, we demonstrate that asymmetric resource competition is a necessary condition for the formation of multimodal size distributions within cohorts. By contrast, the legacy of small-scale clustering during recruitment is transient and quickly overwhelmed by density-dependent mortality. Complex multi-layered size distributions are generated when established individuals are restricted in the spatial domain within which they can capture resources. The number of modes reveals the effective number of direct competitors, while the separation and spread of modes are influenced by distances among established individuals. Asymmetric competition within local neighbourhoods can therefore generate a range of complex size distributions within even-aged cohorts.

Spatial complementarity and the coexistence of species.

Coexistence of apparently similar species remains an enduring paradox in ecology. Spatial structure has been predicted to enable coexistence even when population-level models predict competitive exclusion if it causes each species to limit its own population more than that of its competitor. Nevertheless, existing hypotheses conflict with regard to whether clustering favours or precludes coexistence. The spatial segregation hypothesis predicts that in clustered populations the frequency of intra-specific interactions will be increased, causing each species to be self-limiting. Alternatively, individuals of the same species might compete over greater distances, known as heteromyopia, breaking down clusters and opening space for a second species to invade. In this study we create an individual-based model in homogeneous two-dimensional space for two putative sessile species differing only in their demographic rates and the range and strength of their competitive interactions. We fully characterise the parameter space within which coexistence occurs beyond population-level predictions, thereby revealing a region of coexistence generated by a previously-unrecognised process which we term the triadic mechanism. Here coexistence occurs due to the ability of a second generation of offspring of the rarer species to escape competition from their ancestors. We diagnose the conditions under which each of three spatial coexistence mechanisms operates and their characteristic spatial signatures. Deriving insights from a novel metric - ecological pressure - we demonstrate that coexistence is not solely determined by features of the numerically-dominant species. This results in a common framework for predicting, given any pair of species and knowledge of the relevant parameters, whether they will coexist, the mechanism by which they will do so, and the resultant spatial pattern of the community. Spatial coexistence arises from complementary combinations of traits in each species rather than solely through self-limitation.

Herbivory of tropical rain forest tree seedlings correlates with future mortality.

Tree seedlings in tropical rain forests are subject to both damage from natural enemies and intense interspecific competition. This leads to a trade-off in investment between defense and growth, and it is likely that tree species specialized to particular habitats tailor this balance to correspond with local resource availability. It has also been suggested that differential herbivore impacts among tree species may drive habitat segregation, favoring species adapted to particular resource conditions. In order to test these predictions, a reciprocal transplant experiment in Sabah, Malaysia, was established with seedlings of five species of Dipterocarpaceae. These were specialized to either alluvial (Hopea nervosa, Parashorea tomentella) or sandstone soils (Shorea multiflora, H. beccariana), or were locally absent (S. fallax). A total of 3000 seedlings were planted in paired gap and understory plots in five sites on alluvial and sandstone soils. Half of all seedlings were fertilized. Seedling growth and mortality were recorded in regular samples over 3.5 years, and rates of insect herbivore damage were estimated from censuses of foliar tissue loss on marked mature leaves and available young leaves. Greater herbivory rates on mature leaves had no measurable effects on seedling growth but were associated with a significantly increased likelihood of mortality during the following year. In contrast, new-leaf herbivory rates correlated with neither growth nor mortality. There were no indications of differential impacts of herbivory among the five species, nor between experimental treatments. Herbivory was not shown to influence segregation of species between soil types, although it may contribute toward differential survival among light habitats. Natural rates of damage were substantially lower than have been shown to influence tree seedling growth and mortality in previous manipulative studies.

Mating behaviour in Lymnaea stagnalis pond snails is a maternally inherited, lateralized trait.

Lateralization of the brain has traditionally been considered a specialization that is confined to the vertebrates, but recent studies have revealed that a range of invertebrates also have a brain that is structurally asymmetric and/or each side performs a different set of functions. Here, we show that the precopulatory mating behaviour of the pond snail Lymnaea stagnalis is lateralized. We present evidence that the asymmetry of the behaviour corresponds to the sinistral or dextral shell coil, or chirality, of the snail, and is apparently also controlled by a maternal effect locus. As sinistral snails also tend to have mirror image brains, these findings suggest that the lateralized sexual behaviour of L. stagnalis is set up early in development, and is a direct consequence of the asymmetry of the entire body.