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Epiphytes on Lithocarpus xylocarpus Tree in Mixed Forest at Nat Ma Taung National Park, Myanmar (2700 m Above Sea Level). Photograph: L.S. 

Epiphytes on Lithocarpus xylocarpus Tree in Mixed Forest at Nat Ma Taung National Park, Myanmar (2700 m Above Sea Level). Photograph: L.S. 

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Forest canopies are dynamic interfaces between organisms and atmosphere, providing buffered microclimates and complex microhabitats. Canopies form vertically stratified ecosystems interconnected with other strata. Some forest biodiversity patterns and food webs have been documented and measurements of ecophysiology and biogeochemical cycling have a...

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... Vegetation structure may be an especially important vertical axis as it contributes to strong gradients in 53 micro-climate (e.g., temperature, (Nakamura et al., 2017)), resources (e.g., high energy fruits, (Fleming 54 et al., 1987;Schaefer et al., 2002), and risk (e.g., predation, (Shattuck and Williams, 2010). 55 Consequently, some of the wide array of functional traits possessed by animals likely evolved to exploit 56 ecosystem resources across vegetation vertical gradients, thereby determining the types of ecosystem 57 resources that are available to a given animal as well as the influence that animals may have on 58 ecosystem structure and function (Lundgren et al., 2021;Oliveira and Scheffers, 2019). ...
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... In tropical forests, it is likely that the midstory, with its abundant young foliage, favourable climatic conditions and increased overlap of host plant species and microclimates (Basset 2001;Hirao et al. 2009) leads to an increase in caterpillar species richness, diversity, and density, although this has never been explicitly studied. In contrast, the harsh weather conditions (Ulyshen 2011;Nakamura et al. 2017), and reduced foliage quality (Coley and Barone 1996;Murakami et al. 2005) in the upper canopy, lead to a decline in caterpillars populations. Additionally, in tropical adult lepidopteran assemblages, neighbouring strata have the highest similarity likely due to the overlap in biotic (e.g. ...
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Insect herbivores and their parasitoids play a crucial role in terrestrial trophic interactions in tropical forests. These interactions occur across the entire vertical gradient of the forest. This study compares how caterpillar communities, and their parasitism rates, vary across vertical strata and between caterpillar defensive strategies in a semi deciduous tropical forest in Nditam, Cameroon. Within a 0.1 ha plot, all trees with a diameter at breast height (DBH) ≥ 5 cm were felled and systematically searched for caterpillars. We divided the entire vertical gradient of the forest into eight, five-metre strata. All caterpillars were assigned to a stratum based on their collection height, reared, identified, and classified into one of three defensive traits: aposematic, cryptic and shelter-building. Caterpillar species richness and diversity showed a midstory peak, whereas density followed the opposite pattern, decreasing in the midstory and then increasing towards the highest strata. This trend was driven by some highly dense shelter-building caterpillars in the upper canopy. Specialisation indices indicated decreasing levels of caterpillar generality with increasing height, a midstory peak in vulnerability, and increasing connectance towards the upper canopy, although the latter was likely driven by decreasing network size. Both aposematic and shelter-building caterpillars had significantly higher parasitism rates than cryptic caterpillars. Our results highlight nuanced changes in caterpillar communities across forest strata and provide evidence that defences strategies are important indicators of parasitism rates in caterpillars and that both aposematic and shelter-building caterpillars could be considered a “safe haven” for parasitoids.
... The altitude, habitat type, and complexity of the understory vegetation structure, tree canopy cover and physical environmental factors such as temperature, humidity, light intensity at each location are very different. The presence of different environmental disturbances in each habitat is also strongly suspected to have a significant influence on the number of butterfly species in each habitat (Pardonnet et al. 2013;Widhiono 2015;Nakamura et al. 2017). ...
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... Forest ecosystems have a gigantic, complex, threedimensional structure (Ehbrecht et al., 2021;Walter et al., 2021), and available biodiversity data are strongly biased towards the portion near the ground. Consequently, knowledge related to conservation of canopy ecosystems is extremely limited, especially for old-growth forests that contain high species richness but are declining rapidly at a global scale (Nakamura et al., 2017;Watson et al., 2018). ...
... Canopy traits such as leaf biomass, branch density and structural complexity are important drivers of associated arthropod diversity (Nakamura et al., 2017, Sallé et al., 2021. Canopy biomass and vegetation density define the available resource amount and habitat space (Neves et al., 2013), supporting canopy arthropod abundance and diversity (Müller et al., 2018, Knuff et al., 2020. ...
... via nutrient quality (Tobisch et al., 2023). Disentangling such tree species effects from general structural dependencies is thus a major challenge for biodiversity research in canopies (Nakamura et al., 2017). Moreover, it remains unclear how mixing tree species alters canopy structural complexity and its habitat suitability for arthropods. ...
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Forest canopies host an abundant but often neglected diversity of arthropods, which requires careful attention in times of ongoing biodiversity loss. Yet, how tree species composition interacts with canopy structure in shaping arthropod communities remains largely unknown. Here, by combining mobile laser scanning and insecticidal fogging with a machine learning algorithm, we studied which canopy architectural properties affect canopy arthropod communities in monospecific and mixed stands of broadleaved European beech and the coniferous Norway spruce and non-native Douglas fir in Germany. Evaluating the abundances and ecological guild diversity of ~ 90,000 arthropods and 27 partly novel high-resolution structural variables, we identified vegetation volume and tree species identity as weak predictors of arthropod abundance and ecological guild diversity. In contrast, structural heterogeneity, i.e. structural complexity, vertical layering and variability of canopy gaps-which were highest in coniferous stands-were strong positive drivers. Despite this, arthropod ecological guild diversity was lower in non-native Douglas fir. Mixed stands had intermediate arthropod abundance and ecological guild diversity. Our study shows that habitat heterogeneity and tree species-identity are closely interlinked in shaping associated canopy arthropod communities. Positive effects of habitat heterogeneity on arthropod ecological guild diversity were often uncoupled from resource availability, and the key role of our novel intra-canopy gap indices suggests that they should be considered as indicators in future research on forest heterogeneity-diversity relationships. Broadleaf-conifer mixtures may be suitable to mediate negative tree-species identity effects when adapting forests to global change.
... A unique feature of forest ecosystems is their pronounced vertical structure. Forest canopies are the main place of forests-atmosphere interactions including carbon fixation (Nakamura et al., 2017). Canopy cover and structure determine microclimatic conditions in forests and thus play an important role for buffering effects of climate and weather Zellweger et al., 2020). ...
... Forest canopies remain one of the last ecological frontiers to be explored and understood by science (Nakamura et al. 2017). There is generally much more information about the ecology of ground and understory animals than about canopy specialists (Nakamura et al. 2017). ...
... Forest canopies remain one of the last ecological frontiers to be explored and understood by science (Nakamura et al. 2017). There is generally much more information about the ecology of ground and understory animals than about canopy specialists (Nakamura et al. 2017). This situation is partially related not only to their elusiveness but also due to the challenges of accessing and studying animal ecology in canopies, which in some forests can reach over 40 m (Lang et al. 2022;Nakamura et al. 2017;Santana et al. 2017). ...
... There is generally much more information about the ecology of ground and understory animals than about canopy specialists (Nakamura et al. 2017). This situation is partially related not only to their elusiveness but also due to the challenges of accessing and studying animal ecology in canopies, which in some forests can reach over 40 m (Lang et al. 2022;Nakamura et al. 2017;Santana et al. 2017). However, different mammals previously regarded as primarily arboreal have also occasionally been recorded on the ground (Mourthé et al. 2007;Pauli et al. 2014). ...
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The brown-eared woolly opossum (Caluromys lanatus) is an elusive frugivorous Neotropical canopy specialist marsupial considered primarily arboreal, but here we report the first records of individuals from this species captured on the ground in the Cerrado. We hypothesize two main reasons to explain this behavior: first related to canopy openness, which would force the animals to the ground to keep moving across the landscape; and the second related with search for food on the ground during periods of food scarcity due to increased canopy gaps. Our findings can have important implications for the species conservation and understanding forest ecological dynamics.
... El dosel del bosque tropical representa uno de los ambientes más fascinantes para desarrollar estudios de las comunidades de artrópodos (Erwin, 1982(Erwin, , 2013Lowman, 1992Lowman, , 1993Lowman, , 2012Basset et al., 2003Basset et al., , 2012Basset et al., , 2015Nakamura et al., 2017). Si bien es un hecho que la mayor diversidad de especies ocurre en los trópicos (Erwin, 2013), se estima que muchas interacciones en el bosque tropical se realizan en su dosel, debido al gran número de especies que en él habitan y a la gran cantidad de biomasa que se genera en este estrato del bosque (Erwin y Scott, 1980;Erwin, 1983;Stork, 1988;Nadkarni, 1994;Blüthgen et al., 2000;Ozanne et al., 2003;Basset et al., 2003, 2015, Nakamura et al., 2017. ...
... El dosel del bosque tropical representa uno de los ambientes más fascinantes para desarrollar estudios de las comunidades de artrópodos (Erwin, 1982(Erwin, , 2013Lowman, 1992Lowman, , 1993Lowman, , 2012Basset et al., 2003Basset et al., , 2012Basset et al., , 2015Nakamura et al., 2017). Si bien es un hecho que la mayor diversidad de especies ocurre en los trópicos (Erwin, 2013), se estima que muchas interacciones en el bosque tropical se realizan en su dosel, debido al gran número de especies que en él habitan y a la gran cantidad de biomasa que se genera en este estrato del bosque (Erwin y Scott, 1980;Erwin, 1983;Stork, 1988;Nadkarni, 1994;Blüthgen et al., 2000;Ozanne et al., 2003;Basset et al., 2003, 2015, Nakamura et al., 2017. ...
... El dosel del bosque tropical presenta características tales como: iluminación, temperatura, velocidad del viento, fluctuación de la humedad relativa, concentración de ozono y condensación del agua, radicalmente mayores a los de otros estratos del bosque (Blanc, 1990;Parker, 1995;Lamaud et al., 2002;Nakamura et al., 2017). Adicionalmente, el tipo y la cantidad de defensas químicas y físicas que contienen las hojas de los árboles son mayores en el dosel que en los otros ambientes del bosque (Barone y Coley, 1996;Dominy et al., 2003). ...
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Capítulo 5. Composición de la comunidad de artrópodos que habitan en el dosel en un bosque tropical de tierras bajas de Panamá Resumen Se estudió la composición de la comunidad de artrópodos asociados a 9 especies de árboles en el dosel del Parque Natural Metropolitano en Panamá. Durante 60 semanas se realizaron recolectas de artrópodos para lo cual se uti-lizaron trampas Malaise modificadas. Para llegar al dosel del bosque se empleó una grúa de construcción instalada en el parque. Se recolectaron 37 225 individuos, pertene-cientes a 162 familias. Los resultados indican que Diptera es el orden mejor representado. Cecidomyiidae, Chiro-nomidae, Psychodidae, Mycetophilidae, Ceratopogonidae, Agaonidae, Torymidae, Formicidae, y Cicadellidae fueron las familias más abundantes. Enterolobium cyclocarpum y Ana-cardium excelsum son las especies con mayor número de ar-trópodos asociados, tanto en número de familias como de
... In addition, the characteristics of the vegetation structure, e.g., life forms, stages of plant development and tolerance to shade, also change (Canham et al. 1994). Changes in microclimate create complex microhabitats, which consequently influence biodiversity (Nakamura et al. 2017). ...
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Measurements of biodiversity can be used to assess the scale of anthropogenic impact and predict species loss. A great number of diversity measures exist involving species richness and relative abundance that differ in how they are calculated. The choice of the adequate biodiversity measure to compare biological communities poses a challenge. The aim of this study was to evaluate the ability of 26 diversity measures to differentiate land cover types in the eastern Brazilian Amazon. Diversity was inventoried and quantified in three vertical strata in six locally dominant land cover types: primary and secondary forests, agroforestry systems, oil palm plantations, annual croplands and pastures. No diversity measure was able to differentiate all land cover types in every stratum. Fisher’s alpha was able to differentiate cover types in the middle and upper strata, while Smith-Wilson and Jentsch’s mixture quotient were able to differentiate in the lower and upper strata, and heterogeneity indices only differentiated in the upper stratum. The distinction and ranking of plant diversity among land covers depended on the diversity measure chosen. The number and types of land cover and vertical strata were a key factor in the ability of the diversity measure to differentiate among them.
... According to existing research, the highest caterpillar species richness, diversity, and density and abundance is found in the midstory where there are larger amounts of foliage, community overlap (Hirao et al. 2009) and structural complexity from overlapping host plant species which leads to increased niche availability (Basset 2001). Conversely, the upper-canopy has the lowest species richness and diversity of caterpillars, as there are harsh weather conditions (Ulyshen 2011;Nakamura et al. 2017) and reduced foliage quality (Coley and Barone 1996;Murakami et al. 2005). Additionally, in tropical adult lepidopteran assemblages, neighbouring strata have the highest similarity likely due to the overlap in biotic (e.g. ...
Preprint
Full-text available
Insect herbivores and their parasitoids play a crucial role in terrestrial trophic interactions in tropical forests. These interactions occur across the entire vertical gradient of the forest. This study compares how caterpillar communities, and their parasitism rates, vary across vertical strata and between caterpillar defensive strategies in a semi deciduous tropical forest in Nditam, Cameroon. Within a 0.1 ha plot, all trees with a diameter at breast height (DBH) ≥ 5 cm were felled and systematically searched for caterpillars. We divided the entire vertical gradient of the forest into eight, five-meter strata. All caterpillars were assigned to a stratum based on their collection height, reared, identified, and classified into one of three defensive traits: aposematic, cryptic and shelter-building. Caterpillar species richness and diversity showed a midstory peak, whereas density followed the opposite pattern, decreasing in the midstory and then increasing towards the highest strata. This trend was driven by some highly dense shelter-building caterpillars in the upper-canopy. Specialisation indices indicated decreasing levels of caterpillar generality with increasing height, a midstory peak in vulnerability, and increasing connectance towards the upper-canopy, although the latter was likely driven by decreasing network size. Both aposematic and shelter-building caterpillars had significantly higher parasitism rates than cryptic caterpillars. Our results highlight nuanced changes in caterpillar communities across forest strata and provide evidence that defences strategies are important indicators of parasitism rates in caterpillars and that both aposematic and shelter-building caterpillars could be considered a “ safe haven” for parasitoids.