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Successional dynamics in Neotropical forests are as uncertain as they are predictable

Natalia Nordena, Héctor A. Angaritab, Frans Bongers, Miguel Martínez-Ramos, Iñigo Granzow-de la Cerda, Michiel van Breugelf, Edwin Lebrija-Trejosg, Jorge A. Meavei, John Vandermeer, G. Bruce Williamson,Bryan Finegan, Rita Mesquitam, and Robin L. Chazdon

Abstract


Although forest succession has traditionally been approached as a deterministic process, successional trajectories of vegetation change vary widely, even among nearby stands with similar environmental conditions and disturbance histories. Here, we provide the first attempt, to our knowledge, to quantify predictability and uncertainty during succession based on the most extensive long-term datasets ever assembled for Neotropical forests. We develop a novel approach that integrates deterministic and stochastic components into different candidate models describing the dynamical interactions among three widely used and interrelated forest attributes—stem density, basal area, and species density. Within each of the seven study sites, successional trajectories were highly idiosyncratic, even when controlling for prior land use, environment, and initial conditions in these attributes. Plot factors were far more important than stand age in explaining successional trajectories. For each site, the best-fit model was able to capture the complete set of time series in certain attributes only when both the deterministic and stochastic components were set to similar magnitudes. Surprisingly, predictability of stem density, basal area, and species density did not show consistent trends across attributes, study sites, or land use history, and was independent of plot size and time series length. The model developed here represents the best approach, to date, for characterizing autogenic successional dynamics and demonstrates the low predictability of successional trajectories. These high levels of uncertainty suggest that the impacts of allogenic factors on rates of change during tropical forest succession are far more pervasive than previously thought, challenging the way ecologists view and investigate forest regeneration.

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Ecological disturbance regimes caused by agricultural land uses and their effects on tropical forest regeneration

Isela Zermeño-Hernández, Moisés Méndez-Toribio, Christina Siebe, Julieta Benítez-Malvido & Miguel Martınez-Ramos Key

Abstract


Aims: Across tropical regions, large forest areas have been converted to different agricultural land uses. These uses impose ecological disturbances affecting forest regeneration potential after field abandonment. Finding ways to identify those agricultural land uses limiting forest regeneration is a critical issue for conserving biodiversity in human-modified landscapes. Here, we developed a fast and inexpensive index, useful for quantifying ecological disturbance regimes associated with agricultural land uses, and tested its power to predict forest regeneration potential.

Location: Municipality of Marques de Comillas, southeast Mexico.

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Biomass is the main driver of changes in ecosystem process rates during tropical forest succession

Madelon Lohbeck, Lourens Poorter, Miguel MartíNez-Ramos, And Frans Bongers

Abstract


ver half of the world’s forests are disturbed, and the rate at which ecosystem processes recover after disturbance is important for the services these forests can provide. We analyze the drivers’ underlying changes in rates of key ecosystem processes (biomass productivity, litter productivity, actual litter decomposition, and potential litter decomposition) during secondary succession after shifting cultivation in wet tropical forest of Mexico. We test the importance of three alternative drivers of ecosystem processes: vegetation biomass (vegetation quantity hypothesis), community-weighted trait mean (mass ratio hypothesis), and functional diversity (niche complementarity hypothesis) using structural equation modeling. This allows us to infer the relative importance of different mechanisms underlying ecosystem process recovery. Ecosystem process rates changed during succession, and the strongest driver was aboveground biomass for each of the processes. Productivity of aboveground stem biomass and leaf litter as well as actual litter decomposition increased with initial standing vegetation biomass, whereas potential litter decomposition decreased with standing biomass. Additionally, biomass productivity was positively affected by community-weighted mean of specific leaf area, and potential decomposition was positively affected by functional divergence, and negatively by community-weighted mean of leaf dry matter content.

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How do light and water acquisition strategies affect species selection during secondary succession in moist tropical forests?

Leonie Schönbeck, Madelon Lohbeck, Frans Bongers, Miguel Martínez Ramos and Frank Sterck 1

Abstract


Pioneer tree species have acquisitive leaf characteristics associated with high demand of light and water, and are expected to be shade and drought intolerant. Using leaf functional traits (specific leaf area, photosynthetic rate, relative water content and stomatal conductance) and tree performance (mortality rate) in the field, we assessed how shade and drought tolerance of leaves are related to the species’ positions along a successional gradient in moist tropical forest in Chiapas, Mexico. We quantified morphological and physiological leaf shade and drought tolerance indicators for 25 dominant species that characterize different successional stages. We found that light demand decreases with succession, confirming the importance of light availability for species filtering during early stages of succession. In addition, water transport levels in the leaves decreased with succession, but high water transport did not increase the leaf’s vulnerability to drought. In fact, late successional species showed higher mortality in dry years than early successional ones, against suggestions from leaf drought tolerance traits. It is likely that pioneer species have other drought-avoiding strategies, like deep rooting systems and water storage in roots and stems. More research on belowground plant physiology is needed to understand how plants adapt to changing environments, which is crucial to anticipate the effects of climate change on secondary forests.

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Functional trait strategies of trees in dry and wet tropical forests are similar but differ in their consequences for succession

Madelon Lohbeck, Edwin Lebrija-Trejos, Miguel Martínez-Ramos, Jorge A. Meave, Lourens Poorter and Frans Bongers

Abstract


Global plant trait studies have revealed fundamental trade-offs in plant resource economics. We evaluated such trait trade-offs during secondary succession in two species-rich tropical ecosystems that contrast in precipitation: dry deciduous and wet evergreen forests of Mexico. Species turnover with succession in dry forest largely relates to increasing water availability and in wet forest to decreasing light availability. We hypothesized that while functional trait trade-offs are similar in the two forest systems, the successful plant strategies in these communities will be different, as contrasting filters affect species turnover. Research was carried out in 15 dry secondary forest sites (5-63 years after abandonment) and in 17 wet secondary forest sites (<1-25 years after andonment). We used 11 functional traits measured on 132 species to make species-trait PCA biplots for dry and wet forest and compare trait trade-offs. We evaluated whether multivariate plant strategies changed during succession, by calculating a 'CommunityWeighted Mean' plant strategy, based on species scores on the first two PCA-axes. Trait spectra reflected two main trade-off axes that were similar for dry and wet forest species: acquisitive versus conservative species, and drought avoiding species versus evergreen species with large animal-dispersed seeds.

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Effects of grass clearing and soil tilling on establishment of planted tree seedlings in tropical riparian pastures

Paula Meli, José María Rey Benayas, Miguel Martínez Ramos and Julia Carabias

Abstract


Anthropogenic disturbances frequently exceed resilience of riparian forests. In small-scale restoration projects revegetation is a common technique, but its success depends on the plant species used and some environmental filters. We investigated whether grass competition and soil compaction together with seasonal drought and flooding limit the establishment of seven tree species experimentally planted in abandoned riparian pastures in Southern Mexico. We tested the effects of grass clearing and soil tilling and analyzed seasonal variation of vertical distance to water level, and changes in tree performance,microclimate, and vegetation biomass after planting. Seedling survival was low (19 ± 3 %), ranging between 3 (Brosimum alicastrum, Moraceae) and 38 % (Pachira aquatica, Bombacaceae). Survival was negatively correlated to vertical distance to water level, highlighting the importance of the short but severe dry season that may occur in the humid tropics, which reduced survival by [60 %. Flooding events also produced high seedling mortality (80 % after the two first event).

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Recovery of Amphibian and Reptile Assemblages During Old-Field Successionof Tropical Rain Forests

Omar Hernandez-Ordoñez, Nicolas Urbina-Cardona and Miguel Martınez-Ramos

Abstract


Conversion of tropical forests to agriculture affects vertebrate assemblages, but we do not know how fast or to what extent theseassemblages recover afterfield abandonment. We addressed this question by examining amphibians and reptiles in secondary forests insoutheastern Mexico. We used chronosequence data (12 secondary forests fallow for 1–23 yr and 3 old-growth forest sites) to analyzesuccessional trajectories and estimate recovery times of assemblage attributes for amphibians and reptiles. We conducted 6 surveys ateach site over 14 mo (1200 person-hours) and recorded 1552 individuals, including 25 species of amphibians and 36 of reptiles, repre-senting 96 and 74 percent of the expected regional number of species, respectively. Abundance, species richness, and species diversity ofamphibians increased rapidly with successional age, approaching old-growth forest values in

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Structure and diversity of phyllostomid bat assemblages on riparian corridors in a human-dominated tropical landscape

Erika de la Peña-Cuéllar, Julieta Benítez-Malvido, Luis Daniel Avila-Cabadilla, Miguel Martínez-Ramos and Alejandro Estrada

Abstract


Tropical forests around the world have been lost, mainly because of agricultural activities. Linear elements like riparian vegetation in fragmented tropical landscapes help maintain the native flora and fauna. Information about the role of riparian corridors as a reservoir of bat species, however, is scanty. We assessed the value of riparian corridors on the conservation of phyllostomid bat assemblage in an agricultural landscape of southern Mexico. For 2 years (2011–2013), mist-netting at ground level was carried out twice during the dry season (December to May) and twice during the wet season (June to November) in different habitats: (1) riparian corridors in mature forest, (2) riparian corridors in pasture, (3) continuous forest away from riparian vegetation, and (4) open pastures. Each habitat was replicated three times. To determine the influence of vegetation structure on bat assemblages, all trees (≥10 cm dbh) were sampled in all habitats. Overall, 1752 individuals belonging to 28 species of Phyllostomidae were captured with Sternodermatinae being the most rich and abundant subfamily. Riparian corridors in mature forest and pastures had the greatest species richness and shared 65% of all species. Open pastures had the lowest richness and abundance of bats with no Phyllostominae species recorded. Six of the 18 species recorded could be considered as habitat indicators. There was a positive relationship between bat species composition and tree basal area. Our findings suggest that contrary to our expectations, bats with generalist habits and naturally abundant could be useful detector taxa of habitat modification, rather than bats strongly associated with undisturbed forest. Also in human-dominated landscapes, the maintenance of habitat elements such as large trees in riparian corridors can serve as reservoirs for bat species, especially for those that are strongly associated with undisturbed forest.

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Land-use Change Dynamics, Soil Type and Species Forming Mono-dominant Patches: the Case of Pteridium aquilinum in a Neotropical Rain Forest Region

Ireri Suazo-Ortuno, Leonel Lopez-Toledo1, Javier Alvarado-Dıaz, and Miguel Martınez-Ramos

Abstract


Deforestation and agricultural land degradation in tropical regions can create conditions for growth of perennial plant species forming mono-dominated patches (MDP). Such species might limit forest regeneration, and their proliferation forces the abandonment of fields and subsequent deforestation to establish new fields. Therefore, identifying factors fostering MDP species is critical for biodiversity conservation in human-modified landscapes. Here, we propose a conceptual framework to identify such factors and apply it to the case of Pteridium aquilinum (bracken fern), a light-demanding species, tolerant of low soil fertility and fire. We hypothesize that bracken proliferation is promoted by land-use changes that increase light availability, especially in sites with low soil fertility and land uses involving fire. We assessed this idea using agricultural fields in southeastern Mexico with different land-use change histories and quantifying prevalence and cover of bracken. Five different land-use change histories resulted from transitions among forest, crop, pasture, and fallow field stages. Of the 133 fields sampled, 71 percent had P. aquilinum; regression tree analysis indicated that 65 percent of inter-field variation in prevalence and 90 percent in cover was explained by land-use change history and soil type. Maximum prevalence, cover, and rates of increase in bracken were found on fields with low fertility sandy/clay soils, which had been used for crops and pasture, were frequently burned, and had high levels of light. Fields on fertile alluvial soil never used for pasture were bracken-free. Agriculture promoting high light environments on less fertile soils is a major cause of bracken proliferation and likely that of other MDP species.

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Testing Chronosequences through Dynamic Approaches: Time and Site Effects on Tropical Dry Forest Succession

Francisco Mora, Miguel Martınez-Ramos, Guillermo Ibarra-Manrıquez, Alfredo Perez-Jiménez, Jenny Trilleras, and Patricia Balvanera1

Abstract


Chronosequences, commonly used to assess succession, have been questioned because of their failure to project successional trajectories.Here, we develop a simple analytical approach combining both chronosequence and dynamic data to test the power of age of abandonment and site factors to explain and predict succession. The approach proceeds by first fitting statistical models relating age to attribute values (the chronosequence model) and their observed changes (the dynamic model) to test explanatory power. Predictive power is then tested by bootstrapping the chronosequence model to derive confidence intervals for expected changes and comparing them with the dynamic model. Finally, residuals from both models are tested against site factors. The procedure was applied to six attributes (basal area, plant density, mean plant height, species richness, evenness, and composition) of the woody community (plants >1 cm dbh within 0.1-ha plots) in nine abandoned cattle pastures (0–12 yr) and three old growth tropical dry forests monitored over 6 yr. Age explained 60–97 percent of the variance in community attributes and only 32–57 percent in observed changes. It significantly overestimated basal area and mean height, while species richness and composition were highly predicted. Besides age, management history also explained successional dynamics. Our results suggest age is not necessarily a reliable predictor of short-term successional dynamics, and explanatory power is not indicative of predictive power. Because of this low reliability, caution is needed when applying chronosequences to evaluate ecosystem services’ recovery. The analytical approach developed here contributes to a better exploration of those possible limitations.

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Sustainable harvesting of non-timber forest products based on ecological and economic criteria

Juan C. Hernández-Barrios1, Niels P. R. Anten and Miguel Martınez-Ramos

Abstract


1. Harvesting of highly valuable non-timber forest products (NTFPs) has been considered a win-win strategy where local people profit while conserving forest biodiversity ecosystem services. Nevertheless the sustainability of NTFP harvesting has been debated as the nature of NTFPs harvesting regimes scale of commercialization are highly heterogeneous, few studies have evaluated the cumulative ecological economic effects of such regimes. Here we assessed the medium-term (10 years) sustainability of NTFP harvesting using Chamaedorea palm leaves a major NTFP from Mesoamerica that is highly valued in the international floral industry as a case study.

2. We used an experimental ecological study and an economic assessment to analyse the sustainability of leaf harvesting in C. ernesti-augustii. A 4-year leaf removal experiment was conducted to assess effects of increasing levels of defoliation (0%, 25%, 50%, 75%, 100% leaf removal, biannually) on palm survivorship, leaf production and leaf quality. Results of this experiment were combined with estimations of harvest economic value to make projections of the availability of leaves and profit per unit area. Finally, we determined harvesting regimes that maximize profit while maintaining medium-term viability of exploited populations.

 

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Experimental defaunation of terrestrial mammalian herbivores alters tropical rainforest understorey diversity

Angela A. Camargo-Sanabria, Eduardo Mendoza, Roger Guevara, Miguel Martínez-Ramos, Rodolfo Dirzo

Abstract


It has been suggested that tropical defaunation may unleash community-wide cascading effects, leading to reductions in plant diversity. However, experimental evidence establishing cause–effect relationships thereof is poor. Through a 5 year exclosure experiment, we tested the hypothesis that mammalian defaunation affects tree seedling/sapling community dynamics leading to reductions in understorey plant diversity. We established plot triplets (n = 25) representing three defaunation contexts: terrestrial-mammal exclosure (TE), medium/large mammal exclosure (PE) and open access controls (C). Seedlings/saplings 30–100 cm tall were marked and identified within each of these plots and re-censused three times to record survival and recruitment. In the periods 2010–2011 and 2011–2013, survival was greater in PE than in C plots and recruitment was higher in TE plots than in C plots. Overall, seedling density increased by 61% in TE plots and 23% in PE plots, whereas it decreased by 5% in C plots. Common species highly consumed by mammals (e.g. Brosimum alicastrum and Ampelocera hottlei) increased in their abundance in TE plots. Rarefaction curves showed that species diversity decreased in TE plots from 2008 to 2013, whereas it remained similar for C plots. Given the prevalence of tropical defaunation, we posit this is an anthropogenic effect threatening the maintenance of tropical forest diversity.

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Combining ecological, social and technical criteria to select species for forest restoration

Paula Meli, Miguel Martınez-Ramos, Jose Mar  ıa Rey-Benayas & Julia Carabias

Abstract


Question: How to evaluate and integrate relevant ecological, social and technical criteria to select species to be introduced in restoration projects of highly diverse ecosystems such as tropical riparian forests.

Location: Riparian forest, Marques de Comillas municipality, southeast Mexico (16°54′N, 92°05′W).

Methods: We proposed a ‘species selection index’ (SSI) using five independent criteria related to ecological, social and technical information. SSI targeted species that (1) are important in the reference forest; (2) are less likely to establish following disturbance; (3) are not specific to a particular habitat; (4) are socially accepted; and (5) their propagation requires a reasonable time and financial investment. SSI may range between zero and 50, with higher values meaning higher potential for restoration purposes.

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Restoration Enhances Wetland Biodiversity and Ecosystem Service Supply, but Results Are Context- Dependent: A Meta-Analysis

JPaula Meli, José María Rey Benayas, Patricia Balvanera, Miguel Martínez Ramos

Abstract


Wetlands are valuable ecosystems because they harbor a huge biodiversity and provide key services to societies. When natural or human factors degrade wetlands, ecological restoration is often carried out to recover biodiversity and ecosystem services (ES). Although such restorations are routinely performed, we lack systematic, evidence-based assessments of their effectiveness on the recovery of biodiversity and ES. Here we performed a meta-analysis of 70 experimental studies in order to assess the effectiveness of ecological restoration and identify what factors affect it. We compared selected ecosystem performance variables between degraded and restored wetlands and between restored and natural wetlands using response ratios and random-effects categorical modeling. We assessed how context factors such as ecosystem type, main agent of degradation, restoration action, experimental design, and restoration age influenced post-restoration biodiversity and ES. Biodiversity showed excellent recovery, though the precise recovery depended strongly on the type of organisms involved. Restored wetlands showed 36% higher levels of provisioning, regulating and supporting ES than did degraded wetlands. In fact, wetlands showed levels of provisioning and cultural ES similar to those of natural wetlands; however, their levels of supporting and regulating ES were, respectively, 16% and 22% lower than in natural wetlands. Recovery of biodiversity and of ES were positively correlated, indicating a win-win restoration outcome. The extent to which restoration increased biodiversity and ES in degraded wetlands depended primarily on the main agent of degradation, restoration actions, experimental design, and ecosystem type. In contrast, the choice of specific restoration actions alone explained most differences between restored and natural wetlands. These results highlight the importance of comprehensive, multi- factorial assessment to determine the ecological status of degraded, restored and natural wetlands and thereby evaluate the effectiveness of ecological restorations. Future research on wetland restoration should also seek to identify which restoration actions work best for specific habitats.

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Transplanting native tree seedlings to enrich tropical live fences: an ecological and socio-economic analysis

Beatriz D. Fuentealba and Miguel Martínez-Ramos

Abstract

The enrichment of live fences with native tree species has been proposed as a conservation strategy in agricultural landscapes; however, little research has explored ways to do this in tropical areas. This study examines selection of native tree species, effects of damage caused by mammals (mainly cattle) in performance (survival and growth) of transplanted seedlings, and cost-benefit balances as critical steps to enrich tropical live fences. Seven native tree species,with ecological and socio-economic importance, were selected in a Mexican agricultural landscape to grow as seedlings, and six of them were transplanted into live fences of cattle ranches with different levels of cattle activity (none/moderate/high). Costs associated with propagation and seedling protection in the field were calculated, and performance and damage in seedlings were measured over 2 years. We developed an index to identify species with the best performance and lowest costs in sites with cattle activity. Our results showed that damage, caused mainly by cattle, reduced the performance of transplanted seedlings. The effect of this damage varied depending on its severity (level and frequency) and the identity and life history of species. All selected species performed well in the site without cattle access.
Dendropanax arbo-reus was the best species at site with moderate cattle activity, and Trema micrantha and Saurauia scabrida at site with high cattle activity. These species are recommended for enriching live fences because of good cost-benefit balance. This approach could be an important quantitative method to select species useful not only in agroforestry but also in restoration projects, which normally remain under the pressure of domestic and wild animals.

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Defoliation effects on seed dispersal and seedling recruitment in a tropical rain forest understorey palm

Jeffrey van Lent, Juan C. Hernández-Barrios, Niels P. R. Anten and Miguel Martínez-Ramos

 Abstract

Assessing the demographic effects of leaf area losses in perennial plants is important to determine population resilience to natural and anthropogenic disturbances. Yet, while impacts of defoliation on vital rates of adult plants have been well documented, consequences for seed dispersal and seedling recruitment have been barely explored.
Here, we assessed the effects of defoliation on fruit production, fruit/seed size, seed dispersal and seedling recruitment in populations of Chamaedorea ernesti-augustii, a tropical rain forest, understorey palm from Mesoamerica, whose leaves are exploited as a highly valuable non-timber forest product (NTFP).
Fruit size and seed production were quantified in mature palms that were subjected to 0% (control), 25%, 50%, 75% and 100% experimental removal of standing leaves, applied every 6 months over 2 years. Seed dispersal by birds and gravity, and seed predation by small vertebrates on the ground were also quantified. Rates of seedling recruitment were recorded in non-defoliated and ‘sterile’ populations (75% sustained defoliation and periodical removal of all produced fruits). Finally, a stochastic model was used to quantify the overall effect of defoliation on seedlings recruited from locally (i.e. dispersed by gravity) and immigrant produced seeds (i.e. dispersed by birds). Increasing defoliation strongly reduced seed production. The probability of bird dispersal was positively correlated with fruit/seed size. Isolated (i.e. bird-dispersed) seeds endured lower predation rates than grouped (i.e. gravity-dispersed) ones. Modelling showed that seedling recruitment rate is severely reduced in highly defoliated populations due to a strong decline in the number of seedlings coming from local seed sources. Surrounding non-defoliated populations could partly compensate for this effect via seedlings coming from immigrant seeds.
Synthesis. Chronic and intense defoliation negatively affects seed production and dispersal, which reduces the probability of seedling recruitment. Such effects may have a profound impact on the dynamics and genetic variability of populations, which should be taken into account when considering the effects of natural defoliation and sustainability of leaf-harvesting regimes.

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Distribution and conservation status of amphibian and reptile species in the Lacandona rainforest, Mexico: an update after 20 years of research

Omar Hernández-Ordóñez, Miguel Martínez-Ramos, Víctor Arroyo-Rodríguez, Adriana González- Hernández, Arturo González-Zamora,Diego A. Zárate and Víctor Hugo Reynoso

Abstract

Mexico has one of the richest tropical forests, but is also one of the most deforested in Mesoamerica. Species lists updates and accurate information on the geographic distribution of species are necessary for baseline studies in ecology and conservation of these sites. Here, we present an updated list of the diversity of amphibians and reptiles in the Lacandona region, and actualized information on their distribution and conservation status.Although some studies have discussed the amphibians and reptiles of the Lacandona, most herpetological lists came from the northern part of the region, and there are no confirmed records for many of the species assumed to live in the region. After reviewing databases of scientific collections and published herpetological lists, and adding new information from our 2007 to 2013 inventories of the southeastern Lacandona rainforest, we recorded 124 species (89 reptiles and 35 amphibians) for the region. Nine amphibians (25.7%) and 2 (2.2%)reptiles are endemic to the Mayan forest (from the Lacandona, in Mexico, to the Mayan Mountains in Belize). Four amphibians and three reptiles appeared to be restricted to the Montes Azules Biosphere Reserve, suggesting that they are particularly vulnerable to habitat loss and degradation. On average, the region shares less than 60% of the species with neighboring Mexican tropical forests (e.g., Las Choapas region and Los Tuxtlas, Calakmul, and Sian-Ka’an Biosphere Reserves). Of the herpetofauna evaluated,the Mexican government threatened species list (NOM 059)indicates that seven species (20%) are under a risk category, while the IUCN indicates that only seven species (10.1%) are at risk. Our findings indicate that Lacandona is of great importance for the conservation of the Mesoamerican herpetofauna. Nevertheless, increasing deforestation levels add further uncertainties to the mainten ance of amphibians and reptiles among other vertebrates in the region.

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Effects of ENSO and Temporal Rainfall Variation on the Dynamics of Successional Communities in Old-Field Succession of a Tropical Dry Forest

Susana Maza-Villalobos, Lourens Poorter, Miguel Martínez-Ramos

Abstract

The effects of temporal variation of rainfall on secondary succession of tropical dry ecosystems are poorly understood. We studied effects of inter-seasonal and inter-year rainfall variation on the dynamics of regenerative successional communities of a tropical dry forest in Mexico. We emphasized the effects caused by the severe El Niño Southern Oscillation (ENSO) occurred in 2005. We established permanent plots in sites representing a chronosequence of Pasture (abandoned pastures,0–1 years fallow age), Early (3–5), Intermediate (8–12), and Old-Growth Forest categories (n = 3 per category). In total, 8210 shrubs and trees 10 to 100-cm height were identified, measured, and monitored over four years. Rates of plant recruitment, growth and mortality, and gain and loss of species were quantified per season (dry vs. rainy), year, and successional category, considering whole communities and separating seedlings from sprouts and shrubs from trees. Community rates changed with rainfall variation without almost any effect of successional stage. Mortality and species loss rates peaked during the ENSO year and the following year; however, after two rainy years mortality peaked in the rainy season. Such changes could result from the severe drought in the ENSO year, and of the outbreak of biotic agents during the following rainy years. Growth, recruitment and species gain rates were higher in the rainy season but they were significantly reduced after the ENSO year. Seedlings exhibited higher recruitment and mortality rate than sprouts, and shrubs showed higher recruitment than trees. ENSO strongly impacted both the dynamics and trajectory of succession, creating transient fluctuations in the abundance and species richness of the communities. Overall, there was a net decline in plant and species density in most successional stages along the years. Therefore, strong drought events have critical consequences for regeneration dynamics, delaying the successional process and modifying the resilience of these systems.

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