Network analyses show horizontal and vertical distribution of vascular epiphytes on their hosts in a fragment of cloud forest in Central Mexico.

Cloud forests figure as one of the most diverse ecosystems, accounting not only for a high number of plant species but also with a great variety of interactions among them. A common interaction in these forests is the one between vascular epiphytes and their hosts. However, few studies have used the network approach to analyze them. Here, we analyze the horizontal and vertical structure of the vascular epiphyte - host network in a cloud forest in central Mexico. We quantified the number of epiphyte stands on each host both total and per-stratum. Complete network, group, and species metrics were estimated at both levels of analysis. The host - epiphyte networks had relatively low network size but were highly connected; moderately nested, with low specialization, and modularity; but higher vulnerability than generality, and high niche overlap. The community was composed by a high number of generalist species. To our knowledge this is the first study in which network analyses are conducted with standardized data and including all host and epiphyte species in the community. The analyses suggest that the networks are robust, and that functional redundancy might be probable, two advantageous characteristics in a very fragmented and threatened cloud forest.

Ecological patterns and processes in the vertical dimension of terrestrial ecosystems.

Climatic gradients such as latitude and elevation are considered primary drivers of global biogeography. Yet, alongside these macro-gradients, the vertical space and structure generated by terrestrial plants form comparable climatic gradients but at a fraction of the distance. These vertical gradients provide a spectrum of ecological space for species to occur and coexist, increasing biodiversity. Furthermore, vertical gradients can serve as pathways for evolutionary adaptation of species traits, leading to a range of ecological specialisations. In this review, we explore the ecological evidence supporting the proposition that the vertical gradient serves as an engine driving the ecology and evolution of species and shaping larger biogeographical patterns in space and time akin to elevation and latitude. Focusing on vertebrate and invertebrate taxa, we synthesised how ecological patterns within the vertical dimension shape species composition, distribution and biotic interactions. We identify three key ecological mechanisms associated with species traits that facilitate persistence within the vertical environment and draw on empirical examples from the literature to explore these processes. Looking forward, we propose that the vertical dimension provides an excellent study template to explore timely ecological and evolutionary questions. We encourage future research to also consider how the vertical dimension will influence the resilience and response of animal taxa to global change.

The role of morphological traits in predicting the functional ecology of arboreal and ground ants in the Cerrado-Amazon transition.

There is often a vertical stratification of the vegetation in tropical forests, where each forest stratum has a unique set of environmental conditions, including marked differences in habitat heterogeneity, physical complexity, and microclimate. Additionally, many tropical forests are highly seasonal, and we need to consider the temporal variation in environmental conditions when assessing the functional aspects of their organisms. Here, we tested the hypothesis that vertical stratification and seasonality shape tropical ants' functional ecology and that there are differences in the functional trait diversity and composition between arboreal and ground-dwelling ant communities. We collected ants in the arboreal and ground strata in the rainy and dry seasons in six different areas, measuring seven morphological traits to characterize their functional ecology and diversity. Irrespective of the season, we found a distinct functional composition between arboreal and ground-dwelling ants and a higher functional richness on the ground. However, ground ants were more functionally redundant than arboreal ants. The differences in functional richness and redundancy between ant inhabiting strata and season could also be observed in the community-weighted mean traits: arboreal and ground ant traits can be distinguished in Weber's length, mandible length, eye length, and eye position on the head capsule. The differences in these functional traits are mainly related to the ants' feeding habits and the complexity of their foraging substrates. Overall, by providing the first systematic comparison of continuous traits between arboreal and ground-dwelling ants, our study opens new investigation paths, indicating important axes of functional diversification of tropical ants.

A mosaic of induced and non-induced branches promotes variation in leaf traits, predation and insect herbivore assemblages in canopy trees.

Forest canopies are complex and highly diverse environments. Their diversity is affected by pronounced gradients in abiotic and biotic conditions, including variation in leaf chemistry. We hypothesised that branch-localised defence induction and vertical stratification in mature oaks constitute sources of chemical variation that extend across trophic levels. To test this, we combined manipulation of plant defences, predation monitoring, food-choice trials with herbivores and sampling of herbivore assemblages. Both induction and vertical stratification affected branch chemistry, but the effect of induction was stronger. Induction increased predation in the canopy and reduced herbivory in bioassays. The effects of increased predation affected herbivore assemblages by decreasing their abundance, and indirectly, their richness. In turn, we show that there are multiple factors contributing to variation across canopies. Branch-localised induction, variation between tree individuals and predation may be the ones with particularly strong effects on diverse assemblages of insects in temperate forests.

Darker ants dominate the canopy: Testing macroecological hypotheses for patterns in colour along a microclimatic gradient.

Gradients in cuticle lightness of ectotherms have been demonstrated across latitudes and elevations. Three key hypotheses have been used to explain these macroecological patterns: the thermal melanism hypothesis (TMH), the melanism-desiccation hypothesis (MDH) and the photo-protection hypothesis (PPH). Yet the broad abiotic measures, such as temperature, humidity and UV-B radiation, typically used to detect these ecogeographical patterns, are a poor indication of the microenvironment experienced by small, cursorial ectotherms like ants. We tested whether these macroecological hypotheses explaining cuticle lightness held at habitat and microclimatic level by using a vertical gradient within a tropical rainforest. We sampled 222 ant species in lowland, tropical rainforest across four vertical strata: subterranean, ground, understory and canopy. We recorded cuticle lightness, abundance and estimated body size for each species and calculated an assemblage-weighted mean for cuticle lightness and body size for each vertical stratum. Abiotic variables (air temperature, vapour pressure deficit and UV-B radiation) were recorded for each vertical stratum. We found that cuticle lightness of ant assemblages was vertically stratified: ant assemblages in the canopy and understory were twice as dark as assemblages in ground and subterranean strata. Cuticle lightness was not correlated with body size, and there was no support for the TMH. Rather, we attribute this cline in cuticle lightness to a combination of the MDH and the PPH. Our findings indicate that broad macroecological patterns can be detected at much smaller spatial scales and that microclimatic gradients can shape trait variation, specifically the cuticle lightness of ants. These results suggest that any changes to microclimate that occur due to land-use change or climate warming could drive selection of ants based on cuticle colour, altering assemblage structure and potentially ecosystem functioning.

Broken promise of college? New educational sorting mechanisms for intergenerational association in the 21st century.

Previous studies have shown that intergenerational socioeconomic association becomes weaker as children's education level increases and is negligible among college graduates. A college degree is known as the great equalizer for intergenerational socioeconomic mobility. Recent studies, however, reported that the strong intergenerational association reemerges among advanced degree holders although it stays weak among BA-only holders. Despite the substantial theoretical importance and policy implications, the mechanisms behind the reemergence of the intergenerational association at the post-baccalaureate level have been less studied. In this paper, we examine the association between parents' education and children's earnings using the 2010, 2013, 2015, and 2017 National Survey of College Graduates data. Our results show that the strong intergenerational socioeconomic immobility among advanced degree holders is fully attributable to three educational sorting mechanisms: children from high-SES families (1) obtain expensive and financially rewarding advanced degrees, (2) attend selective institutions and major in hyper-lucrative fields of study such as law and medicine in graduate school, and (3) complete their education at a younger age and enjoy income growth over more years in the labor market. Implications of these findings are discussed.

Proximity to canopy mediates changes in the defensive chemistry and herbivore loads of an understory tropical shrub, Piper kelleyi.

Phytochemical traits are a key component of plant defense theory. Chemical ecology has been biased towards studying effects of individual metabolites even though effective plant defenses are comprised of diverse mixtures of metabolites. We tested the phytochemical landscape hypothesis, positing that trophic interactions are contingent upon their spatial location across a phytochemically diverse landscape. Specifically, intraspecific phytochemical changes associated with vertical strata in forests were hypothesised to affect herbivore communities of the neotropical shrub Piper kelleyi Tepe (Piperaceae). Using a field experiment, we found that phytochemical diversity increased with canopy height, and higher levels of phytochemical diversity located near the canopy were characterised by tradeoffs between photoactive and non-photoactive biosynthetic pathways. For understory plants closer to the ground, phytochemical diversity increased as direct light transmittance decreased, and these plants were characterised by up to 37% reductions in herbivory. Our results suggest that intraspecific phytochemical diversity structures herbivore communities across the landscape, affecting total herbivory.

Forest disturbance and vector transmitted diseases in the lowland tropical rainforest of central Panama.

OBJECTIVE: To explore possible changes in the community attributes of haematophagous insects as a function of forest disturbance. We compare the patterns of diversity and abundance, plus the behavioural responses of three epidemiologically distinct vector assemblages across sites depicting various levels of forest cover. METHODS: Over a 3-year period, we sampled mosquitoes, sandflies and biting-midges in forested habitats of central Panama. We placed CDC light traps in the forest canopy and in the understorey to gather blood-seeking females. RESULTS: We collected 168 405 adult haematophagous dipterans in total, including 26 genera and 86 species. Pristine forest settings were always more taxonomically diverse than the disturbed forest sites, confirming that disturbance has a negative impact on species richness. Species of Phlebotominae and Culicoides were mainly classified as climax (i.e. forest specialist) or disturbance-generalist, which tend to decrease in abundance along with rising levels of disturbance. In contrast, a significant portion of mosquito species, including primary and secondary disease vectors, was classified as colonists (i.e. disturbed-areas specialists), which tend to increase in numbers towards more disturbed forest habitats. At pristine forest, the most prevalent species of Phlebotominae and Culicoides partitioned the vertical niche by being active at the forest canopy or in the understorey; yet this pattern was less clear in disturbed habitats. Most mosquito species were not vertically stratified in their habitat preference. CONCLUSION: We posit that entomological risk and related pathogen exposure to humans is higher in pristine forest scenarios for Culicoides and Phlebotominae transmitted diseases, whereas forest disturbance poses a higher entomological risk for mosquito-borne infections. This suggests that the Dilution Effect Hypothesis (DEH) does not apply in tropical rainforests where highly abundant, yet unrecognised insect vectors and neglected zoonotic diseases occur. Comprehensive, community level entomological surveillance is, therefore, the key for predicting potential disease spill over in scenarios of pristine forest intermixed with anthropogenic habitats. We suggest that changes in forest quality should also be considered when assessing arthropod-borne disease transmission risk.

OBJECTIF: Explorer les changements possibles dans les attributs communautaires des insectes hématophages en fonction de la perturbation des forêts. Nous comparons les modèles de diversité et d'abondance, ainsi que les réponses comportementales de trois assemblages de vecteurs épidémiologiquement distincts sur des sites illustrant divers niveaux de couverture forestière. MÉTHODES: Au cours d'une période de trois ans, nous avons échantillonné des moustiques, des phlébotomes et des moucherons piqueurs dans les habitats forestiers du centre de Panama. Nous avons placé des pièges à lumière CDC dans la canopée de la forêt et dans le sous-étage pour recueillir les femelles en quête de sang. RÉSULTATS: Nous avons collecté un total de 168.405 diptères hématophages adultes, dont 26 genres et 86 espèces. Les environnements de forêt intacts étaient toujours plus diversifiés du point de vue taxonomique que les sites forestiers perturbés, confirmant que les perturbations avaient un impact négatif sur la richesse en espèces. Les espèces de phlébotome et Culicoïdes étaient principalement classés comme climax (spécialiste de la forêt) ou généralistes de perturbation, qui ont tendance à diminuer en abondance parallèlement aux niveaux croissants de perturbation. En revanche, une partie importante des espèces de moustiques, y compris les vecteurs primaires et secondaires de maladies, a été classée dans la catégorie des colons (c'est-à-dire spécialistes des zones perturbées), qui ont tendance à se multiplier vers des habitats forestiers plus perturbés. Dans la forêt vierge, les espèces de phlébotomes et Culicoïdes les plus répandues cloisonnaient la niche verticale en étant actives dans la canopée de la forêt ou dans le sous-étage; pourtant, cette tendance était moins nette dans les habitats perturbés. La plupart des espèces de moustiques n'étaient pas stratifiées verticalement dans leur préférence d'habitat. CONCLUSION: Nous estimons que le risque entomologique et l'exposition associée des agents pathogènes à l'homme est plus élevé dans les scénarios de forêt vierge pour les maladies transmises par les phlébotomes et Culicoïdes, alors que la perturbation des forêts pose un risque entomologique plus élevé pour les infections transmises par les moustiques. Cela suggère que l'hypothèse de l'effet de dilution ne s'applique pas dans les forêts tropicales humides où se reproduisent très abondamment les insectes vecteurs, mais non reconnus, et où des maladies zoonotiques négligées surviennent. Une surveillance entomologique approfondie au niveau de la communauté est donc la clé pour prédire le potentiel de propagation des maladies dans des scénarios de forêt vierge mélangée à des habitats anthropiques. Nous suggérons que les changements dans la qualité des forêts soient également pris en compte lors de l'évaluation du risque de propagation de maladies transmises par les arthropodes.

Distance-decay differs among vertical strata in a tropical rainforest.

Assemblage similarity decays with geographic distance-a pattern known as the distance-decay relationship. While this pattern has been investigated for a wide range of organisms, ecosystems and geographical gradients, whether these changes vary more cryptically across different forest strata (from ground to canopy) remains elusive. Here, we investigated the influence of ground vs. arboreal assemblages to the general distance-decay relationship observed in forests. We seek to explain differences in distance-decay relationships between strata in the context of the vertical stratification of assemblage composition, richness and abundance. We surveyed for a climate-sensitive model organism, amphibians, across vertical rainforest strata in Madagascar. For each tree, we defined assemblages of ground-dwelling, understory, or canopy species. We calculated horizontal distance-decay in similarity across all trees, and across assemblages of species found in different forest strata (ground, understory and canopy). We demonstrate that within stratum comparisons exhibit a classic distance-decay relationship for canopy and understory communities but no distance-decay relationships for ground communities. We suggest that differences in horizontal turnover between strata may be due to local scale habitat and resource heterogeneity in the canopy, or the influence of arboreal traits on species dispersal and distribution. Synthesis. Biodiversity patterns in horizontal space were not consistent across vertical space, suggesting that canopy fauna may not play by the same set of "rules" as their conspecifics living below them on the ground. Our study provides compelling evidence that the above-ground amphibian assemblage of tropical rainforests is the primary driver of the classical distance-decay relationship.

Vertical differentiation in tropical forest butterflies: a novel mechanism generating insect diversity?

Many tropical fruit-feeding nymphalid butterflies are associated with either the forest canopy or the understorey; however, the exceptions offer insights into the origins of tropical diversity. As it occurs in both habitats of tropical forests in Ecuador and Peru, Archaeoprepona demophon is one such exception. We compared patterns of occurrence of A. demophon in the canopy and understorey and population genomic variation for evidence of ecological and genetic differentiation between habitats. We found that butterfly occurrences in the canopy were largely uncorrelated with occurrences in the understorey at both localities, indicating independent demographic patterns in the two habitats. We also documented modest, significant genome-level differentiation at both localities. Genetic differentiation between habitat types (separated by approx. 20 m in elevation) was comparable to levels of differentiation between sampling locations (approx. 1500 km). We conclude that canopy and understorey populations of A. demophon represent incipient independent evolutionary units. These findings support the hypothesis that divergence between canopy and understorey-associated populations might be a mechanism generating insect diversity in the tropics.

Phlebotominae in peri-domestic and forest environments inhabited by Alouatta caraya in northeastern Argentina.

Multiple species of Phlebotominae are vectors of Leishmania (Protozoa: Trypanosomatidae), which causes visceral leishmaniasis (VL) and cutaneous leishmaniasis (CL). To describe the Phlebotominae (Diptera: Psychodidae) related to the environments of black and gold howler monkeys Alouatta caraya (Humbodlt, 1812) (Primates: Atelidae), potential vectors were sampled in different landscapes and vertical strata of sleeping trees. Phlebotomine captured between December 2011 and March 2012 (2365 individuals) belonged to eight species, of which Nyssomyia neivai (Pinto, 1926) (61.4%) and Migonemyia migonei (França, 1920) (18.73%) were the most abundant, and Ny. withmani was recorded for the first time in the Chaco province. In the 'peri-domestic' landscape, the phlebotomine were mainly captured in henhouses (78.7%), whereas the tree canopy in 'rural' and 'wild' landscapes yielded 31.2% and 29.1% of the phlebotomine, respectively. A significant association between the type of landscape and the species of phlebotomine was observed by multivariate analysis. Lutzomyia longipalpis (Lutz & Neiva, 1912) and Mg. migonei were associated with 'peri-domestic' landscape, and Ny. neivai was associated with the 'wild' landscape. The results of this prospective study suggest that the interaction between phlebotomine and A. caraya could be a key factor with respect to understanding the epidemiology of leishmaniasis.

Niche partitioning of intertidal seagrasses: evidence of the influence of substrate temperature.

The influence of soil temperature on rhizome depths of four intertidal seagrass species was investigated in central Queensland, Australia. We postulated that certain intertidal seagrass species are soil temperature-sensitive and vertically stratify rhizome depths. Below-ground vertical stratification of intertidal seagrass rhizome depths was analysed based upon microclimate (soil temperature) and microhabitat (soil type). Soil temperature profiles exhibited heat transfer from surface layers to depth that varied by microhabitat, with vertical stratification of rhizome depths between species. Halodule uninervis rhizomes maintain a narrow median soil temperature envelope; compensating for high surface temperatures by occupying deeper, cooler soil substrates. Halophila decipiens, Halophila ovalis and Zostera muelleri rhizomes are shallow-rooted and exposed to fluctuating temperatures, with broader median temperature envelopes. Halodule uninervis appears to be a niche specialist, with the two Halophila species considered as generalist niche usage species. The implications of niche use based upon soil temperature profiles and rhizome rooting depths are discussed in the context of species' thermal tolerances and below-ground biomass O2 demand associated with respiration and maintenance of oxic microshields. This preliminary evidence suggests that soil temperature interaction with rhizome rooting depths may be a factor that influences the distribution of intertidal seagrasses.

Exposure to the leaf litter microbiome of healthy adults protects seedlings from pathogen damage.

It is increasingly recognized that microbiota affect host health and physiology. However, it is unclear what factors shape microbiome community assembly in nature, and how microbiome assembly can be manipulated to improve host health. All plant leaves host foliar endophytic fungi, which make up a diverse, environmentally acquired fungal microbiota. Here, we experimentally manipulated assembly of the cacao tree (Theobroma cacao) fungal microbiome in nature and tested the effect of assembly outcome on host health. Using next-generation sequencing, as well as culture-based methods coupled with Sanger sequencing, we found that manipulating leaf litter exposure and location within the forest canopy significantly altered microbiome composition in cacao. Exposing cacao seedlings to leaf litter from healthy conspecific adults enriched the seedling microbiome with Colletotrichum tropicale, a fungal endophyte known to enhance pathogen resistance of cacao seedlings by upregulating host defensive pathways. As a result, seedlings exposed to healthy conspecific litter experienced reduced pathogen damage. Our results link processes that affect the assembly and composition of microbiome communities to their functional consequences for host success, and have broad implications for understanding plant-microbe interactions. Deliberate manipulation of the plant-fungal microbiome also has potentially important applications for cacao production and other agricultural systems in general.

Diversity and Flight Patterns of Caddisflies (Trichoptera) in an Atlantic Forest Fragment: Implications for Species Conservation in Threatened Ecosystems.

Anthropogenic activities have decimated the Atlantic Forest domain (AF) and increased the pressure on freshwater biota, such as Trichoptera, which is the most affected order by the current insect decline. Adult mobility is crucial for the colonisation of new environments unconnected by water sources. In this article, we describe the assemblage of caddisflies in a preserved AF fragment related to their functional feeding group and provide empirical data on the patterns of horizontal and vertical flight. Adults were collected using white sheet and light attraction traps, placed at different distances and heights from a stream in Pernambuco, Brazil. A total of 2934 specimens of 15 species from five families were collected, mostly collector-filterers. Horizontal flight was limited, with 80% of the abundance concentrated up to 20 m. Vertical stratification was also concentrated at lower heights. A female-biased proportion was observed at higher strata. The richness and abundance of species decreased with increasing distances and heights from the stream. Overall, Chimarra sp. and Macrostemum scharfi were the dominant species. Trichoptera is a key taxon used as a biological indicator of water quality, and here, knowledge on the diversity and flight patterns of adults is expanded. At the risk of intensive pollution of rivers in Atlantic forests, data on the adult dispersal can be incorporated in the assessment of endangerment status and in conservation strategies.

Patterns of avian tree usage in the primeval temperate forests of Bialowieza National Park.

Species distribution and resource utilization are a fundamental aspect of ecology. By analyzing the tree space usage by birds and determining the species composition of birds across different parts of trees, our study could shed light on the mechanisms shaping co-occurrence patterns in bird communities. Therefore, our study aimed to determine the species composition of birds across different parts of trees. We investigated whether species richness differs between positions on a tree and how these positions influence the probability of occurrence of the 10 most frequently observed bird species. To achieve this, we observed birds within permanent plots in Bialowieza National Park (BNP) and analyzed the distribution patterns of birds within six vertical and three horizontal sectors of trees. The compositional dissimilarity between tree sectors was assessed using detrended correspondence analysis. We employed generalized linear mixed-effects models to examine differences in species richness. The majority of the BNP bird community was associated with the branches, while other birds occupied the tree crown trunks and the understory trunks. Species richness was the highest on branches in the crown part of trees, followed by lower species richness on trunks associated with crowns, and the lowest richness was observed on branches and trunks in the understory. These results indicate that branches in the middle and lower parts of the crown serve as avian diversity hotspots on trees, likely due to the abundance of various food sources. The differing patterns of tree usage by specific bird species may suggest the avoidance of interspecific competition for resources. The study results of tree usage by bird species obtained in the primeval forest provides a reference point for studies conducted in human-altered woods.

Increasing arboreality with altitude: a novel biogeographic dimension.

Biodiversity is spatially organized by climatic gradients across elevation and latitude. But do other gradients exist that might drive biogeographic patterns? Here, we show that rainforest's vertical strata provide climatic gradients much steeper than those offered by elevation and latitude, and biodiversity of arboreal species is organized along this gradient. In Philippine and Singaporean rainforests, we demonstrate that rainforest frogs tend to shift up in the rainforest strata as altitude increases. Moreover, a Philippine-wide dataset of frog distributions shows that frog assemblages become increasingly arboreal at higher elevations. Thus, increased arboreality with elevation at broad biogeographic scales mirrors patterns we observed at local scales. Our proposed 'arboreality hypothesis' suggests that the ability to exploit arboreal habitats confers the potential for larger geographical distributions because species can shift their location in the rainforest strata to compensate for shifts in temperature associated with elevation and latitude. This novel finding may help explain patterns of species richness and abundance wherever vegetation produces a vertical microclimatic gradient. Our results further suggest that global warming will 'flatten' the biodiversity in rainforests by pushing arboreal species towards the cooler and wetter ground. This 'flattening' could potentially have serious impacts on forest functioning and species survival.

Dynamic characteristics of total and microcystin-producing Microcystis in a large deep reservoir.

Microcystis, one of the common cyanobacteria, often causes blooms in reservoirs, which has seriously threatened the safety of drinking water worldwide. To identify the growth characteristic of total and microcystin-producing Microcystis in large deep reservoirs, we used Quantitative PCR (qPCR) to measure the cell density of total and microcystin-producing Microcystis and monitored water quality in the water samples collected in Dongzhang Reservoir once a month. Microcystis blooms occurred in Dongzhang Reservoir in April 2017, which was composed of microcystin-producing and non-microcystin-producing Microcystis. Water temperature, dissolved oxygen, pH, and chlorophyll-a showed significant vertical stratification during Microcystis blooms. Total and microcystin-producing Microcystis grew rapidly under the high concentration of total phosphorus and rising water temperatures. Nitrate-nitrogen had a significant linear correlation with the abundance of microcystin-producing Microcystis. Our results indicated that nutrients and water temperature could be key triggers of Microcystis blooms and nitrate-nitrogen potentially regulates the competition between microcystin-producing and non-microcystin-producing Microcystis. This study improves our understanding of the characteristics of Microcystis blooms and the competition between microcystin-producing and non-microcystin-producing Microcystis in large deep reservoirs.

Analyses of three-dimensional species associations reveal departures from neutrality in a tropical forest.

The study of community spatial structure is central to understanding diversity patterns over space and species co-occurrence at local scales. Although most analytical approaches consider horizontal and vertical dimensions separately, in this study we introduce a three-dimensional spatial analysis that simultaneously includes horizontal and vertical species associations. Using tree census data (2000-2016) and allometries from the Luquillo forest plot in Puerto Rico, we show that spatial organization becomes less random over time as the forest recovered from land-use legacy effects and hurricane disturbance. Tree species vertical segregation is predominant in the forest with almost all species that co-occur in the horizontal plane avoiding each other in the vertical dimension. Horizontal segregation is less common than vertical, whereas three-dimensional aggregation (a proxy for direct tree competition) is the least frequent type of spatial association. Furthermore, dominant species are involved in more non-random spatial associations, implying that species co-occurrence is facilitated by species segregation in space. This novel three-dimensional analysis allowed us to identify and quantify tree species spatial distributions, how interspecific competition was reduced through forest structure, and how it changed over time after disturbance, in ways not detectable from two-dimensional analyses alone.

Species turnover in ant assemblages is greater horizontally than vertically in the world's tallest tropical forest.

Abiotic and biotic factors structure species assembly in ecosystems both horizontally and vertically. However, the way community composition changes along comparable horizontal and vertical distances in complex three-dimensional habitats, and the factors driving these patterns, remains poorly understood. By sampling ant assemblages at comparable vertical and horizontal spatial scales in a tropical rainforest, we tested hypotheses that predicted differences in vertical and horizontal turnover explained by different drivers in vertical and horizontal space. These drivers included environmental filtering, such as microclimate (temperature, humidity, and photosynthetic photon flux density) and microhabitat connectivity (leaf area), which are structured differently across vertical and horizontal space. We found that both ant abundance and richness decreased significantly with increasing vertical height. Although the dissimilarity between ant assemblages increased with vertical distance, indicating a clear distance-decay pattern, the dissimilarity was higher horizontally where it appeared independent of distance. The pronounced horizontal and vertical structuring of ant assemblages across short distances is likely explained by a combination of microclimate and microhabitat connectivity. Our results demonstrate the importance of considering three-dimensional spatial variation in local assemblages and reveal how highly diverse communities can be supported by complex habitats.

The drivers of avian-haemosporidian prevalence in tropical lowland forests of New Guinea in three dimensions.

Haemosporidians are among the most common parasites of birds and often negatively impact host fitness. A multitude of biotic and abiotic factors influence these associations, but the magnitude of these factors can differ by spatial scales (i.e., local, regional and global). Consequently, to better understand global and regional drivers of avian-haemosporidian associations, it is key to investigate these associations at smaller (local) spatial scales. Thus, here, we explore the effect of abiotic variables (e.g., temperature, forest structure, and anthropogenic disturbances) on haemosporidian prevalence and host-parasite networks on a horizontal spatial scale, comparing four fragmented forests and five localities within a continuous forest in Papua New Guinea. Additionally, we investigate if prevalence and host-parasite networks differ between the canopy and the understory (vertical stratification) in one forest patch. We found that the majority of Haemosporidian infections were caused by the genus Haemoproteus and that avian-haemosporidian networks were more specialized in continuous forests. At the community level, only forest greenness was negatively associated with Haemoproteus infections, while the effects of abiotic variables on parasite prevalence differed between bird species. Haemoproteus prevalence levels were significantly higher in the canopy, and an opposite trend was observed for Plasmodium. This implies that birds experience distinct parasite pressures depending on the stratum they inhabit, likely driven by vector community differences. These three-dimensional spatial analyses of avian-haemosporidians at horizontal and vertical scales suggest that the effect of abiotic variables on haemosporidian infections are species specific, so that factors influencing community-level infections are primarily driven by host community composition.