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Although insect herbivorous communities in tropical forests are known to exhibit strong seasonality, few studies have systematically assessed temporal patterns of variation in community structure and plant-herbivore interactions in early successional arboreal communities. We assessed seasonal and interannual variation of the diversity and composition of herbivorous beetles and the tree-herbivore network in a recently established polyculture forest plantation, during the dry and the rainy seasons of 2012 and of 2013. Species richness was similar between years, while the ecological diversity was higher in 2012. Comparing seasons, no differences were found in 2012, whereas in 2013, the species richness and ecological diversity were higher during the dry season. The species composition differed radically across years and seasons. Moreover, a quantitative nested pattern was consistently found across both temporal scales, more influenced by species densities. We found temporal changes in the species strength, whereas connectance and interaction evenness remained stable. Rapid temporal changes in the structural complexity of recently established polyculture plantations and the availability and quality of the trophic resources they offer may act as drivers of beetle diversity patterns, promoting rapid variation in herbivore composition and some interacting attributes. Nonetheless, network structure, connectance, and interaction evenness remained similar, suggesting that reorganizations in the distribution of species may determine the maintenance of the patterns of interaction. Further work assessing long-term temporal dynamics of herbivore beetle assemblages are needed to more robustly relate diversity and interaction patterns to biotic and abiotic factors and their implications in management programs.

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Although commercial forest plantations have experienced a major growth in the tropics over the past decades, little attention has been paid to their role in the conservation of epigeal arthropod communities. We studied diversity patterns of the epigeal beetle community in monoculture and polyculture forest plantations with big-leaf mahogany (Swietenia macrophylla). Likewise, we explored the existence of indicator species of each plantation type. Our findings highlight that each plantation type promotes multiple impacts on diversity patterns. We found that monocultures positively influenced overall beetle species richness and ecological diversity. When broken down by guild, both predator and decomposer species richness were similar between monoculture and polyculture, whereas for beetle diversity we found contrasting responses by guild: decomposer diversity was greater in monoculture whereas predator diversity was higher in polyculture. In addition, species composition differed between monoculture and polyculture, except for the predator guild. Species turnover was the main component explaining beta diversity patterns at all levels, indicating that each plantation type promotes biologically distinct epigeal assemblages. Few superabundant heliophile species dominated the beetle community structure; moreover, monocultures had a composition skewed towards heliophile species whereas polyculture favored umbrophile species. These patterns could be attributed to differences in habitat complexity between plot types, namely differences in tree cover. Additionally, indicator species only were identified in polycultures, reflecting their higher spatial complexity. Monoculture and polyculture plantations with big-leaf mahogany are complementary agroecosystems for preserving diverse epigeal beetle communities and should be considered valuable tools for conservation purposes in the tropics.

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ABSTRACT Elevational patterns in flowering phenology have been reported for trees, shrubs and herbs. However, for vascular epiphytes that rely on atmospheric sources for humidity and nutrients, and depend on phorophyte microhabitat, elevational patterns of variation are unknown. In this study, we described the flowering phenology of Tillandsia carlos-hankii, an epiphytic bromeliad, along an elevational gradient in Capulálpam de Méndez, Oaxaca. We analyzed the onset, seasonality and duration of flowering along and within different elevation zones, and we evaluated the effect of phorophyte features (tree height, DBH and canopy diameter) on flowering start date and duration. From June 2016 to May 2017, we periodically recorded phenological data from six populations along three elevation zones ("low": 2151 to 2283 m. a. s. l., "medium": 2284 to 2416 m. a. s. l. and "high": 2417 to 2548 m. a. s. l.), monitoring two population per zone. Start of flowering occurred between December and January, beginning six to 16 days earlier at low elevations than in the other zones, although this difference was not statistically significant. We observed marked flowering seasonality at all the elevation zones, with differences between zones (W≥18.49, p≤0.0001) and between the populations at medium and high elevations (W≥8.57, p≤0.05). Flowering duration spanned from December to May. Phorophyte features were not related to the onset or duration of flowering (t≤-1.47, p≥0.14, in all cases). Our results suggest that vascular epiphytes follow the same elevational patterns in phenology as other life forms, and that populations in the same elevation range can vary. The causes of such intra-elevational variation merit further investigation.

RESUMEN Se han reportado los patrones altitudinales en la fenología de floración para árboles, arbustos y hierbas. Sin embargo, para epífitas vasculares, que dependen de fuentes atmosféricas para la humedad y los nutrientes, y del microhábitat del forofito, los patrones de variación en gradientes de elevación se desconocen. Se describe la fenología de floración de Tillandsia carlos-hankii, una bromelia epífita, en un gradiente de elevación en Capulálpam de Méndez, Oaxaca. Se analiza el inicio, estacionalidad y duración de su floración entre y dentro de diferentes elevaciones y se evalúa también el efecto del forofito (altura, DAP y diámetro de copa) sobre el inicio y la duración de la floración. De junio de 2016 a mayo de 2017, se monitorearon individuos localizados en seis poblaciones en tres elevaciones (bajo: 2151-2283 m. s. n. m., medio: 2284-2416 m. s. n. m., alto: 2417-2548 m. s. n. m.), dos poblaciones por zona. El inicio de la floración ocurrió entre diciembre y enero, empezando de seis a 16 días antes en la elevación baja que en las otras, aunque estas diferencias no fueron significativas. Hubo una marcada estacionalidad en todas las zonas de elevación, con diferencias entre elevaciones (W≥18,49, p≤0,0001) y entre poblaciones de media y alta elevación (W≥8,57, p≤0,05). La floración duró de diciembre a mayo. Las características del forofito no estuvieron relacionadas con el inicio ni la duración de la floración (t≤-1,47, p≥0,14, en todos los casos). Los resultados sugieren que las epífitas vasculares siguen el mismo patrón fenológico altitudinal de otras formas de vida, y que las poblaciones en un misma elevación altitudinal pueden variar. Las causas de tal variación intra-altitudinal merecen mayor investigación.

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Tree diversity exerts a strong influence on consumer communities, but most work has involved single time point measurements over short time periods. Describing temporal variation associated with diversity effects over longer time periods is necessary to fully understand the effects of tree diversity on ecological function. We conducted a year-long study in an experimental system in southern Mexico assessing the effects of tree diversity on the abundance and diversity of foraging birds. To this end, we recorded bird visitation patterns in 32 tree plots (21 × 21 m; 12 tree species monocultures, 20 four-species polycultures) every 45 days (n = 8 surveys) and for each plot estimated bird abundance, richness, functional diversity (FD) and phylogenetic diversity (PD). In each case, we reported temporal (intra-annual) variation in the magnitude of tree diversity effects, and calculated the temporal stability of these bird responses. Across surveys, tree diversity noticeably affected bird responses, demonstrated by significantly higher abundance (43%), richness (32%), PD (25%) and FD (25%) of birds visiting polyculture plots compared to monoculture plots, as well as a distinct species composition between plot types. We also found intra-annual variation in tree diversity effects on these response variables, ranging from surveys for which the diversity effect was not significant to surveys where a significant 80% increase (e.g. for bird FD and PD) was observed in polyculture relative to monoculture plots. Notably, tree diversity increased the stability of all bird responses, with polycultures having a greater stability abundance (18%), richness (38%), PD (32%), and FD (35%) of birds visiting tree species polycultures compared to monocultures. These results show that tree diversity not only increases bird visitation to plots, but also stabilizes bird habitat usage over time in ways that could implicate insurance-related mechanisms. Such findings are highly relevant for understanding the long-term effects of plant diversity on vertebrates and the persistence of bird-related ecosystem functions. More work is needed to unveil the ecological mechanisms behind temporal variation in vertebrate responses to tree diversity and their consequences for community structure and function.

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The interactions between pairs of native and alien plants via shared use of pollinators have been widely studied. Community level studies however, are necessary in order to fully understand the factors and mechanisms that facilitate successful plant invasion, but these are still scarce. Specifically, few community level studies have considered how differences in invasion level (alien flower abundance), and degree of floral trait similarity between native and invasive species, mediate effects on native plant-pollinator communities. Here, we evaluated the role of alien species on overall plant-floral visitor network structure, and on species-level network parameters, across nine invaded coastal communities distributed along 205 km in Yucatán, México that vary in alien species richness and flower abundance. We further assessed the potential the role of alien plant species on plant-floral visitor network structure and robustness via computational simulation of native and invasive plant extinction scenarios. We did not find significant differences between native and alien species in their functional floral phenotypes or in their visitation rate and pollinator community composition in these invaded sites. Variation in the proportion of alien plant species and flower abundance across sites did not influence plant-pollinator network structure. Species-level network parameters (i.e., normalized degree and nestedness contribution) did not differ between native and alien species. Furthermore, our simulation analyses revealed that alien species are functionally equivalent to native species and contribute equally to network structure and robustness. Overall, our results suggest that high levels of floral trait similarity and pollinator use overlap may help facilitate the integration of alien species into native plant-pollinator networks. As a result, alien species may also play a similar role than that of natives in the structure and stability of native plant and pollinator communities in the studied coastal sand dune ecosystem.

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PREMISE: The occurrence and amount of herbivory are shaped by bottom-up forces, primarily plant traits (e.g., defenses), and by abiotic factors. Addressing these concurrent effects in a spatial context has been useful in efforts to understand the mechanisms governing variation in plant-herbivore interactions. Still, few studies have evaluated the simultaneous influence of multiple sources of bottom-up variation on spatial variation in herbivory. METHODS: We tested to what extent chemical (phenolics, production of terpenoid glands) and physical (pubescence) defensive plant traits and climatic factors are associated with variation in herbivory by leaf-chewing insects across populations of wild cotton (Gossypium hirsutum). RESULTS: We found substantial population variation in cotton leaf defenses and insect leaf herbivory. Leaf pubescence, but not gossypol gland density or phenolic content, was significantly negatively associated with herbivory by leaf-chewing insects. In addition, there were direct effects of climate on defenses and herbivory, with leaf pubescence increasing toward drier conditions and leaf damage increasing toward wetter and cooler conditions. There was no evidence, however, of indirect effects (via plant defenses) of climate on herbivory. CONCLUSIONS: These results suggest that spatial variation in insect herbivory on wild G. hirsutum is predominantly driven by concurrent and independent influences of population variation in leaf pubescence and climatic factors.

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Biodiversity affects the structure of ecological communities, but little is known about the interactive effects of diversity across multiple trophic levels. We used a large-scale forest diversity experiment to investigate the effects of tropical tree species richness on insectivorous birds, and the subsequent indirect effect on predation rates by birds. Diverse plots (four tree species) had higher bird abundance (61%), phylogenetic diversity (61%), and functional diversity (55%) than predicted based on single-species monocultures, which corresponded to higher attack rates on artificial caterpillars (65%). Tree diversity effects on attack rate were driven by complementarity among tree species, with increases in attack rate observed on all tree species in polycultures. Attack rates on artificial caterpillars were higher in plots with higher bird abundance and diversity, but the indirect effect of tree species richness was mediated by bird diversity, providing evidence that diversity can interact across trophic levels with consequences tied to ecosystem services and function.

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While plant intra-specific variation in the stoichiometry of nutrients and carbon is well documented, clines for such traits have been less studied, despite their potential to reveal the mechanisms underlying such variation. Here we analyze latitudinal variation in the concentration of leaf nitrogen (N), phosphorus (P), carbon (C) and their ratios across 30 populations of the perennial herb Ruellia nudiflora. In addition, we further determined whether climatic and soil variables underlie any such latitudinal clines in leaf traits. The sampled transect spanned 5° latitude (ca. 900 km) and exhibited a four-fold precipitation gradient and 2 °C variation in mean annual temperature. We found that leaf P concentration increased with precipitation towards lower latitudes, whereas N and C did not exhibit latitudinal clines. In addition, N:P and C:P decreased towards lower latitudes and latitudinal variation in the former was weakly associated with soil conditions (clay content and cation exchange capacity); C:N did not exhibit a latitudinal gradient. Overall, these results emphasize the importance of addressing and disentangling the simultaneous effects of abiotic factors associated with intra-specific clines in plant stoichiometric traits, and highlight the previously underappreciated influence of abiotic factors on plant nutrients operating under sharp abiotic gradients over smaller spatial scales.

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PREMISE OF STUDY: The factors driving variation in species interactions are often unknown, and few studies have made a link between changes in interactions and the strength of selection. METHODS: We report on spatial variation in functional responses by a seed predator (SP) and its parasitic wasps associated with the herb Ruellia nudiflora. We assessed the influence of plant density on consumer responses and determined whether density effects and spatial variation in functional responses altered natural selection by these consumers on the plant. We established common gardens at two sites in Yucatan, Mexico, and planted R. nudiflora at two densities in each garden. We recorded fruit output and SP and parasitoid attack; calculated relative fitness (seed number) under scenarios of three trophic levels (accounting for SP and parasitoid effects), two trophic levels (accounting for SP but not parasitoid effects), and one trophic level (no consumer effects); and compared selection strength on fruit number under these scenarios across sites and densities. KEY RESULTS: There was spatial variation in SP recruitment, whereby the SP functional response was negatively density-dependent at one site but density-independent at the other; parasitoid responses were density-independent and invariant across sites. Site variation in SP attack led, in turn, to differences in SP selection on fruit output, and parasitoids did not alter SP selection. There were no significant effects of density at either site. CONCLUSIONS: Our results provide a link between consumer functional responses and consumer selection on plants, which deepens our understanding of geographic variation in the evolutionary outcomes of multitrophic interactions.

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Plant diversity can influence predators and omnivores and such effects may in turn influence herbivores and plants. However, evidence for these ecological feedbacks is rare. We evaluated if the effects of tree species (SD) and genotypic diversity (GD) on the abundance of different guilds of insect herbivores associated with big-leaf mahogany (Swietenia macrophylla) were contingent upon the protective effects of ants tending extra-floral nectaries of this species. This study was conducted within a larger experiment consisting of mahogany monocultures and species polycultures of four species and -within each of these two plot types- mahogany was represented by either one or four maternal families. We selected 24 plots spanning these treatment combinations, 10 mahogany plants/plot, and within each plot experimentally reduced ant abundance on half of the selected plants, and surveyed ant and herbivore abundance. There were positive effects of SD on generalist leaf-chewers and sap-feeders, but for the latter group this effect depended on the ant reduction treatment: SD positively influenced sap-feeders under ambient ant abundance but had no effect when ant abundance was reduced; at the same time, ants had negative effects on sap feeders in monoculture but no effect in polyculture. In contrast, SD did not influence specialist stem-borers or leaf-miners and this effect was not contingent upon ant reduction. Finally, GD did not influence any of the herbivore guilds studied, and such effects did not depend on the ant treatment. Overall, we show that tree species diversity influenced interactions between a focal plant species (mahogany) and ants, and that such effects in turn mediated plant diversity effects on some (sap-feeders) but not all the herbivores guilds studied. Our results suggest that the observed patterns are dependent on the combined effects of herbivore identity, diet breadth, and the source of plant diversity.

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BACKGROUND AND AIMS: Studies that have evaluated the effects of heterospecific pollen (HP) receipt on plant reproductive success have generally overlooked the variability of the natural abiotic environment in which plants grow. Variability in abiotic conditions, such as light and water availability, has the potential to affect pollen-stigma interactions (i.e. conspecific pollen germination and performance), which will probably influence the effects of HP receipt. Thus, a more complete understanding of the extent, strength and consequences of plant-plant interactions via HP transfer requires better consideration of the range of abiotic conditions in which these interactions occur. This study addresses this issue by evaluating the effects of two HP donors (Tamonea curassavica and Angelonia angustifolia) on the reproductive success of Cuphea gaumeri, an endemic species of the Yucatan Peninsula. METHODS: Mixed (conspecific pollen and HP) and pure (conspecific pollen only) hand-pollinations were conducted under varying conditions of water and light availability in a full factorial design. Reproductive success was measured as the number of pollen tubes that reached the bottom of the style. KEY RESULTS: Only one of the two HP donors had a significant effect on C. gaumeri reproductive success, but this effect was dependent on water and light availability. Specifically, HP receipt caused a decrease in pollen tube growth, but only when the availability of water, light or both was low, and not when the availability of both resources was high. CONCLUSIONS: The results show that the outcome of interspecific post-pollination interactions via HP transfer can be context-dependent and vary with abiotic conditions, thus suggesting that abiotic effects in natural populations may be under-estimated. Such context-dependency could lead to spatial and temporal mosaics in the ecological and evolutionary consequences of post-pollination interactions.

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Seasonal dynamics of the capitula infested by Dictyotrypeta sp. and Rhynencina spilogaster (Steyskal) (Diptera: Tephritidae) was evaluated throughout the flowering cycle of their host plant the sunflower, Smallanthus maculatus (Cavanilles) Robinson (Asterales: Asteraceae). In central Veracruz, Mexico, along 16 consecutive weeks, a total of 1,017 mature capitula were collected, recording the presence and abundance of immature stages (larvae and pupae) and their related parasitoids. Both fly species were present throughout the entire season, with overall infestation of 51.5% of the capitula examined. However, Dictyotrypeta sp. infested 11.3%, representing about one-fifth of them, and R. spilogaster was most abundant infesting four times as many capitula (42.9%), whereas both species were found together in only 2.6% of the capitula examined. Based on the temporal occurrence of larvae and pupae into flower heads as well as their associated parasitoids and times of emergence, Dictyotrypeta sp. had two yearly generations, and it seems that the second generation could enter a seasonal diapause; in contrast, R. spilogaster was a univoltine species that entered diapause that lasted until the next year.

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Mexican oregano (Lippia graveolens) is an important aromatic plant, mainly used as flavoring and usually harvested from non-cultivated populations. Mexican oregano essential oil showed important variation in the essential-oil yield and composition. The composition of the essential oils extracted by hydrodistillation from 14 wild populations of L. graveolens growing along an edaphoclimatic gradient was evaluated. Characterization of the oils by GC-FID and GC/MS analyses allowed the identification of 70 components, which accounted for 89-99% of the total oil composition. Principal component and hierarchical cluster analyses divided the essential oils into three distinct groups with contrasting oil compositions, viz., two phenolic chemotypes, with either carvacrol (C) or thymol (T) as dominant compounds (contents >75% of the total oil composition), and a non-phenolic chemotype (S) dominated by oxygenated sesquiterpenes. While Chemotype C was associated with semi-arid climate and shallower and rockier soils, Chemotype T was found for plants growing under less arid conditions and in deeper soils. The plants showing Chemotype S were more abundant in subhumid climate. High-oil-yield individuals (>3%) were identified, which additionally presented high percentages of either carvacrol or thymol; these individuals are of interest, as they could be used as parental material for scientific and commercial breeding programs.

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Dimorphic cleistogamy is a specialized form of mixed mating system where a single plant produces both open, potentially outcrossed chasmogamous (CH) and closed, obligately self-pollinated cleistogamous (CL) flowers. Typically, CH flowers and seeds are bigger and energetically more costly than those of CL. Although the effects of inbreeding and floral dimorphism are critical to understanding the evolution and maintenance of cleistogamy, these effects have been repeatedly confounded. In an attempt to separate these effects, we compared the performance of progeny derived from the two floral morphs while controlling for the source of pollen. That is, flower type and pollen source effects were assessed by comparing the performance of progeny derived from selfed CH vs. CL and outcrossed CH vs. selfed CH flowers, respectively. The experiment was carried out with the herb Ruellia nudiflora under two contrasting light environments. Outcrossed progeny generally performed better than selfed progeny. However, inbreeding depression ranges from low (1%) to moderate (36%), with the greatest value detected under shaded conditions when cumulative fitness was used. Although flower type generally had less of an effect on progeny performance than pollen source did, the progeny derived from selfed CH flowers largely outperformed the progeny from CL flowers, but only under shaded conditions and when cumulative fitness was taken into account. On the other hand, the source of pollen and flower type influenced seed predation, with selfed CH progeny the most heavily attacked by predators. Therefore, the effects of pollen source and flower type are environment-dependant and seed predators may increase the genetic differences between progeny derived from CH and CL flowers. Inbreeding depression alone cannot account for the maintenance of a mixed mating system in R. nudiflora and other unidentified mechanisms must thus be involved.

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Few studies have simultaneously addressed the effects of biotic and abiotic factors on pre-dispersal seed predation (PSP). Plant-seed predator interactions may be influenced by natural enemies and pollinators (the latter through changes in fruit or seed traits), and the activity of pre-dispersal seed predators and their natural enemies may both be affected by the abiotic environment. Additionally, in the case of cleistogamous plants with fruit dimorphism, PSP may be biased towards larger and more seeded chasmogamous (CH) fruits [relative to the smaller cleistogamous (CL) fruits], and the effects of biotic and abiotic factors may be contingent upon this fruit dimorphism. We studied PSP in the cleistogamous Ruellia nudiflora using a split-plot experimental design and asked the following: (1) is PSP biased towards CH fruits and is there an effect of pollen load on PSP? (2) Do parasitoids influence PSP and is their effect influenced by pollen load or fruit type? And (3) do light and water availability modify PSP and parasitoid effects? PSP was higher for CH relative to CL fruits, and under low water availability it was lower for pollen-supplemented CH fruits relative to open-pollinated CH fruits. Parasitoids were not influenced by abiotic conditions, but their negative effect on PSP was stronger for pollen-supplemented CH fruits. Overall, we show that fruit dimorphism, abiotic factors and natural enemies affect PSP, and that these effects can be non-additive.

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BACKGROUND AND AIMS: Mixed reproductive strategies may have evolved as a response of plants to cope with environmental variation. One example of a mixed reproductive strategy is dimorphic cleistogamy, where a single plant produces closed, obligately self-pollinated (CL) flowers and open, potentially outcrossed (CH) flowers. Frequently, optimal environmental conditions favour production of more costly CH structures whilst economical and reliable CL structures are produced under less favourable conditions. In this study we explore (1) the effect of light and water on the reproductive phenology and (2) the effect of pollen supplementation on resource allocation to seeds in the cleistogamous weed Ruellia nudiflora. METHODS: Split-plot field experiments were carried out to assess the effect of shade (two levels: ambient light vs. a reduction of 50 %) and watering (two levels: non-watered vs. watered) on the onset, end and duration of the production of three reproductive structures: CH flowers, CH fruit and CL fruit. We also looked at the effect of these environmental factors on biomass allocation to seeds (seed weight) from obligately self-pollinated flowers (CL), open-pollinated CH flowers and pollen-supplemented CH flowers. KEY RESULTS: CH structures were produced for a briefer period and ended earlier under shaded conditions. These conditions also resulted in an earlier production of CL fruit. Shaded conditions also produced greater biomass allocation to CH seeds receiving extra pollen. CONCLUSIONS: Sub-optimal (shaded) conditions resulted in a briefer production period of CH structures whilst these same conditions resulted in an earlier production of CL structures. However, under sub-optimal conditions, plants also allocated more resources to seeds sired from CH flowers receiving large pollen loads. Earlier production of reproductive structures and relatively larger seed might improve subsequent success of CL and pollen-supplemented CH seeds, respectively.

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The main goal of this work was to test for plant genetic variation in the phenotypic plasticity response of the weed Ruellia nudiflora to arbuscular mycorrhizal (AM) fungi inoculation. We collected plants in the field, kept them under homogeneous conditions inside a nursery, and then collected seeds from these parent plants to generate five inbred lines (i.e., genetic families). Half of the plants of each inbred line were inoculated with AM fungi while the other half were not (controls); a fully crossed experimental design was then used to test for the effects of treatment (with or without AM fungi inoculation) and inbred line (genetic family). For each plant, we recorded the number of leaves produced and the number of days it survived during a 2-month period. Results showed a strong positive treatment effect (plastic response to AM fungi inoculation) for leaf production and survival. Moreover, in terms of survival, the treatment effect differed between genetic families (significant genetic family by treatment interaction). These findings indicate that the positive effect of AM fungi on plant survival (and potentially also growth) differs across plant genotypes and that such condition may contribute to R. nudiflora's capacity to colonize new environments.

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Beetles were collected from April 2002 to July 2003 from the following sites in the centre of the state of Veracruz located every 200 m along an altitudinal gradient (1000 - 1400 m asl): three fragments of cloud forest, three shaded coffee plantations, an open canopy coffee plantation, secondary forest, and a pasture. A total of 9,982 specimens were captured, belonging to the families Scarabaeidae and Trogidae, and representing 21 genera and 50 species. The genera Ataenius Harold, Onthophagus Latreille and Aphodius Illiger represented 48% of the species of Scarabaeidae. Species richness was found to decrease with increasing altitude; there were 36 species at 1000 m asl, 27 species between 1200 and 1300 m asl, and 26 species at 1400 m asl. Abundance along the altitudinal gradient follows a pattern of few abundant species and many species with few specimens. Beetle activity is related to precipitation. In the nine communities studied, species richness was observed to increase when precipitation was greater than 100 mm at the beginning of the rainy season. On the landscape scale, there were 44 species during the rainy season, 22 during the windy nortes season, and 24 during the dry season. In terms of trophic guild, 40% of the beetles captured were saprophagous that feed on decomposing organic material from plants and 60% were saprophagous that feed on decomposing organic material from animals (30% necrophagous, 26% coprophagous, 4% telio-necrophagous).

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Durante 16 meses de colectas entre abril 2002 y julio 2003, en el centro del estado de Veracruz, en tres fragmentos de bosque mesófilo de montaña, tres cafetales bajo sombra, un cafetal sin sombra, un acahual y un pastizal, establecidos en un gradiente altitudinal (1000-1400 m), cada 200 m, se obtuvieron 9982 especimenes de Scarabaeidae y Trogidae que representan a 21 géneros y 50 especies. Los géneros Ataenius Harold, Onthophagus Latreilley Aphodius Illiger reúnen al 48 por ciento de las especies de Scarabaeidae. En el gradiente altitudinal se obtuvo que a mayor altitud menor riqueza específica, a los 1000 m coexisten 36 especies, entre los 1200-1300 m 27 especies y a 1400 m 26 especies. La abundancia general en el gradiente altitudinal sigue un patrón de pocas especies abundantes y muchas especies con pocos especimenes. La actividad de los escarabajos está relacionada con la precipitación, observándose con precipitaciones mayores a 100 mm al inicio de la época de lluvias un incremento en la riqueza específica en las nueve comunidades. Durante la época de lluvias coexisten 44 especies, en época de nortes 22 especies y en época de secas 24 especies. Los gremios tróficos están representados por el 40 por ciento de los saprófagos que se alimentan de materia orgánica en descomposición de origen vegetal, y por el 60 por ciento los saprófagos que se alimentan de materia orgánica en descomposición de origen animal (necrófagos 30 por ciento, coprófagos 26 por ciento, telio-necrófagos 4 por ciento).

Beetles were collected from April 2002 to July 2003 from the following sites in the centre of the state of Veracruz located every 200 m along an altitudinal gradient (1000 - 1400 m asl): three fragments of cloud forest, three shaded coffee plantations, an open canopy coffee plantation, secondary forest, and a pasture. A total of 9,982 specimens were captured, belonging to the families Scarabaeidae and Trogidae, and representing 21 genera and 50 species. The genera Ataenius Harold, Onthophagus Latreille and Aphodius Illiger represented 48 percent of the species of Scarabaeidae. Species richness was found to decrease with increasing altitude; there were 36 species at 1000 m asl, 27 species between 1200 and 1300 m asl, and 26 species at 1400 m asl. Abundance along the altitudinal gradient follows a pattern of few abundant species and many species with few specimens. Beetle activity is related to precipitation. In the nine communities studied, species richness was observed to increase when precipitation was greater than 100 mm at the beginning of the rainy season. On the landscape scale, there were 44 species during the rainy season, 22 during the windy nortes season, and 24 during the dry season. In terms of trophic guild, 40 percent of the beetles captured were saprophagous that feed on decomposing organic material from plants and 60 percent were saprophagous that feed on decomposing organic material from animals (30 percent necrophagous, 26 percent coprophagous, 4 percent telio-necrophagous).

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