Research Priorities for Neotropical Dry Forests1

Our understanding of the human and biophysical dimensions of tropical dry forest change and its cumulative effects is still in the early stages of academic discovery. The papers in this special section on Neotropical dry forests cover a wide range of sites and problems ranging from the use of multispectral and hyperspectral remote sensing platforms to the impact of hurricanes on tropical dry forest regeneration. Here, we present to the scientific community the results of a workshop on which research priorities for tropical dry forests were discussed. This discussion focuses on the need to develop linkages between remote sensing, ecological, and social science research. The incorporation of social sciences into ecological research could contribute dramatically to our understandings of tropical dry forests by providing important contextual information to ecologists, and by helping to develop an important science–policy–public nexus on which environmental management can succeed.     

Regional‐Scale Variation in Litter Production and Seasonality in Tropical Dry Forests of Southern Mexico1

Highly seasonal rainfall creates a pulse of litterfall in the southern Yucatan peninsula region, with cascading effects on the timing of essential nutrient fluxes, microbial dynamics, and vegetation growth. I investigated whether forest age or a regional environmental gradient related to rainfall has a greater effect on patterns of litterfall in this increasingly human‐dominated landscape. Litterfall was sampled in 10–13 stands in each of three locations spanning a rainfall gradient of ca 900–1400 mm/yr. Litter was collected monthly from November 1998 through January 2000 in mature forests and in secondary forests aged 2–25 yr. Despite a substantial precipitation gradient, age was the only significant predictor of annual litter mass. Two‐ to five‐yr‐old forests produced significantly less litter than 12–25‐yr‐old secondary forests (4.6 vs. 6.2 Mg/ha/yr), but the difference between older secondary forests and mature forests (9 percent) was not significant. Litter production increased with rainfall, but not significantly so. The pattern of litterfall was similar across locations and age classes, with a peak during late March or early April. However, litterfall seasonality was most pronounced in the old secondary and mature forests. Litterfall was more evenly distributed throughout the year in forests under 10 yr old. Seasonality of litterfall was also less pronounced at the wettest site, with less disparity between peak litterfall and off‐peak months. Seasonality was not related to soil texture. Forest age and rainfall are important drivers of litterfall dynamics; however, both litter mass and degree of seasonality depended more strongly on forest age. Thus, the impact of land‐use change on litter nutrient cycling is as great, if not greater, than the constraint imposed by the major natural environmental factor affecting tropical dry forests.     

Phyllostomid Bat Community Structure and Abundance in Two Contrasting Tropical Dry Forests1

Although tropical wet forests are generally more diverse than dry forests for many faunal groups, few studies have compared bat diversity among dry forests. I compared ground level phyllostomid bat community structure between two tropical dry forests with different precipitation regimes. Parque National Palo Verde in northwestern Costa Rica represents one of the wettest tropical dry forests (rainfall 1.5 m/yr), whereas the Chamela‐Cuixmala Biosphere Reserve on the Pacific coast of central Mexico represents one of the driest (750 mm/yr). Mist net sampling was conducted at the two study sites to compare changes in ground level phyllostomid bat community structure between regions and seasons. Palo Verde was more diverse than Chamela and phyllostomid species showed low similarity between sites (Classic Jaccard = 0.263). The distinct phyllostomid communities observed at these two dry forest sites demonstrates that variants of tropical dry forest can be sufficiently different in structure and composition to affect phyllostomid communities. At both dry forest sites, abundance of the two most common foraging guilds (frugivores and nectarivores) differed between seasons, with greatest numbers of individuals captured coinciding with highest chiropterophilic resource abundance.     

The Utility of Spectral Indices from Landsat ETM+ for Measuring the Structure and Composition of Tropical Dry Forests1

There is a growing emphasis on developing methods for quantifying the structure and composition of tropical forests that can be applied over large landscapes, especially for tropical dry forests that are severely fragmented and have a high conservation priority. This study investigates the relationships between various measures of forest structure (annual woody increment, canopy closure, stand density, stand basal area) and composition (tree species diversity, tree community composition) measured in semi‐deciduous tropical dry forests on islands in Lago Guri, Venezuela and three spectral indices derived from Landsat ETM+: Normalized Difference Vegetation Index (NDVI), Infrared Index (IRI), and Mid‐Infrared Index (MIRI). Even though there were significant autocorrelations among spectral indices, there were significant differences in the relationships between spectral indices and forest attributes. IRI was not significantly correlated with any of the structural variables while MIRI was correlated with canopy closure and NDVI values were correlated with canopy closure as well as annual woody increment. NDVI and MIRI were both related to relative tree diversity and all three indices were associated with aspects of tree species composition. Based on the results of this study, it appears that spectral indices, and in particular NDVI, may be useful indicators of forest attributes in tropical dry forest habitats. Further research needs to be undertaken to identify if the results of this study can be applied to other tropical dry forests at a global spatial scale.     

Effects of Season and Successional Stage on Leaf Area Index and Spectral Vegetation Indices in Three Mesoamerican Tropical Dry Forests1

We compared plant area index (PAI) and canopy openness for different successional stages in three tropical dry forest sites: Chamela, Mexico; Santa Rosa, Costa Rica; and Palo Verde, Costa Rica, in the wet and dry seasons. We also compared leaf area index (LAI) for the Costa Rican sites during the wet and dry seasons. In addition, we examined differences in canopy structure to ascertain the most influential factors on PAI/LAI. Subsequently, we explored relationships between spectral vegetation indices derived from Landsat 7 ETM+ satellite imagery and PAI/LAI to create maps of PAI/LAI for the wet season for the three sites. Specific forest structure characteristics with the greatest influence on PAI/LAI varied among the sites and were linked to climatic differences. The differences in PAI/LAI and canopy openness among the sites were explained by both the past land‐use history and forest management practices. For all sites, the best‐fit regression model between the spectral vegetation indices and PAI/LAI was a Lorentzian Cumulative Function. Overall, this study aimed to further research linkages between PAI/LAI and remotely sensed data while exploring unique challenges posed by this ecosystem.     

Vegetation Structure,Composition, and Species Richness Across a 56‐year Chronosequence of Dry Tropical Forest on Providencia Island,Colombia1

We compared vegetation structure and species richness across a 56‐yr chronosequence of six replicated age classes of dry tropical forest on the island of Providencia, Colombia, in the Southwest Caribbean. Stand age classes were determined using sequential, orthorectified panchromatic aerial photos acquired between 1944 and 1996 and Landsat 7 ETM + satellite imagery from 2000. Along the chronosequence we established 59 plots of 2 × 50 m (0.01 ha) to document changes in species richness, basal area, tree height, stem density, and sprouting ability. All woody trees and shrubs >2.5 cm diameter at breast height (DBH) were censused and measured. Although woody species density reached a peak in stands from 32 to 56 yr old, rarefaction analysis showed that species richness increased linearly with stand age and was highest in stands 56 yr old or greater. Nonparametric, abundance‐based estimators of species richness also showed positive and linear associations with age. Basal area and mean tree height were positively associated with age since abandonment, while sprouting ability showed a negative relationship. Our results indicate rapid recovery of woody species richness and structural characteristics along this tropical dry forest chronosequence.     

Recovery of a Subtropical Dry Forest After Abandonment of Different Land Uses1

We studied the ecological characteristics of 45–50‐yr‐old subtropical dry forest stands in Puerto Rico that were growing on sites that had been deforested and used intensively for up to 128 yr. The study took place in the Guánica Commonwealth Forest. Our objective was to assess the long‐term effects of previous land use on this forest—i.e., its species composition, structure, and functioning. Previous land‐use types included houses, farmlands, and charcoal pits. Stands with these land uses were compared with a nearby mature forest stand. The speed and path of forest recovery after deforestation and land‐use abandonment depended on the conditions of the land. Study areas where land uses had removed the forest canopy and altered soil conditions (houses and farmlands) required a longer time to recover and had a different species composition than study areas where land uses retained a forest canopy (charcoal pits). Different forest attributes recovered at different rates. Crown area index, stem density, and litterfall rate recovered faster than stemwood and root, biomass, tree height, and basal area. Where previous land uses removed the canopy, Leucaena leucocephala, a naturalized alien pioneer species, dominated the regrowth. Native species dominated abandoned charcoal pits and mature forest. The change in species composition, including the invasion of alien species, appears to be the most significant long‐term effect of human use and modification of the landscape.     

Linking the Conservation of Culture and Nature: A Case Study of Sacred Forests in Zimbabwe

This paper examines the role of traditional religious beliefs and traditional leaders in conserving remnant patches of a unique type of dry forest in the Zambezi Valley of northern Zimbabwe. We examined aerial photographs spanning more than three decades, interviewed and surveyed local residents, and met with communities to learn about the environmental history of the forests and the factors that have affected land use in the area. Our results show that forest loss is dramatically less in forests that are now considered sacred, or were in the past connected to sacred forests. This supports our hypothesis that traditional spiritual values have influenced human behavior affecting the forests, and have played a role in protecting them until now. We also found that rates of forest loss have been much higher in an area where traditional leaders are relatively disempowered within the post-independence political system compared to an area where traditional leaders have more power. These findings lead us to conclude that a strategy that links the conservation of culture and nature is likely to be more effective in conserving forests than a strategy that ignores traditional beliefs, values, and institutions.     

Quantifying Tropical Dry Forest Type and Succession: Substantial Improvement with LiDAR

Improved technologies are needed to advance our knowledge of the biophysical and human factors influencing tropical dry forests, one of the world's most threatened ecosystems. We evaluated the use of light detection and ranging (LiDAR) data to address two major needs in remote sensing of tropical dry forests, i.e., classification of forest types and delineation of forest successional status. We evaluated LiDAR‐derived measures of three‐dimensional canopy structure and subcanopy topography using classification‐tree techniques to separate different dry forest types and successional stages in the Guánica Biosphere Reserve in Puerto Rico. We compared the LiDAR‐based results with classifications made from commonly used remote sensing data, including Landsat satellite imagery and radar‐based topographic data. The accuracy of the LiDAR‐based forest type classification (including native‐ and exotic‐dominated forest classes) was substantially higher than those from previously available data (kappa = 0.90 and 0.63, respectively). The best result was obtained when combining LiDAR‐derived metrics of canopy structure and topography, and adding Landsat spectral data did not improve the classification. For the second objective, we observed that LiDAR‐derived variables of vegetation structure were better predictors of forest successional status (i.e., mid‐secondary, late‐secondary, and primary forests) than was spectral information from Landsat. Importantly, the key LiDAR predictors identified within each classification‐tree model agreed with previous ecological knowledge of these forests. Our study highlights the value of LiDAR remote sensing for assessing tropical dry forests, reinforcing the potential for this novel technology to advance research and management of tropical forests in general.     

Litter Nutrient Dynamics During Succession in Dry Tropical Forests of the Yucatan: Regional and Seasonal Effects

Land-use change in the tropics is creating secondary forest at an unprecedented rate. In the tropical Americas, mature dry tropical forest is rapidly being converted to secondary forest during the fallow period of shifting cultivation. We investigated litter phosphorus (P) and nitrogen (N) dynamics in forests recovering from shifting cultivation of maize (corn) in three regions of the Southern Yucatan Peninsula, Mexico. Our goal was to understand how nutrient and water availability affect forest recovery following conversion of mature forest to agricultural land. To investigate such changes at a regional scale, newly fallen litter was collected monthly along a seasonal, a successional, and a precipitation gradient. Reflecting possible P limitation, litter P concentration declined with forest age, while litter N concentration did not differ between age classes. Average litter P concentration from the southern, wettest region was 0.87 mg/g, almost twice the litter P concentration in the drier central and northern regions (0.44 and 0.45 mg/g, respectively). Average N concentrations of litter from the three regions ranged from 1.1% to 1.2%, with no regional differences. However, minima in both P and N concentration from all regions were pronouncedly timed with peak litterfall, suggesting nutrient retranslocation during periods of water stress. Additionally, successional differences in litter P were clearest during wetter months. P nutrient-use efficiency was lowest in the southern region and highest in the central and northern study regions. N nutrient-use efficiency was up to 40 times lower than P nutrient-use efficiency and showed no regional differences. Overall, our results suggest that litter nutrient dynamics in secondary dry tropical forests of the Southern Yucatan are strongly influenced by water and nutrient availability, especially P, as well as land-use history.     

Genetic Diversity of a Tropical Tree Species,Shorea leprosula Miq. (Dipterocarpaceae), in Malaysia: Implications for Conservation of Genetic Resources and Tree Improvement1

Genetic diversity and population genetic structure of Shorea leprosula was investigated using seven natural populations distributed throughout Peninsular Malaysia and one natural population from Borneo. The mean population and species level genetic diversity were exceptionally high (H_e= 0.369 ± 0.025 and 0.406 ± 0.070, respectively). Heterozygosity varied among populations, ranging from 0.326 to 0.400, with the highest values found in the populations from central Peninsular Malaysia. Correlations among ecological factors (longitude, latitude, and annual rainfall) were not significant (P > 0.05), indicating that these ecological variables were not responsible for the observed genetic differences among populations. The Bangi adult population exhibited a higher level of observed heterozygosity but lower fixation indices in comparison to its seedling population. All other seedling populations also showed positive fixation indices (f), indicating a general excess of homozygotes. This also may suggest selection against homozygotes between the seedling and adult stages. A low level of population differentiation was detected (G_ST= 0.117 with the Lambir population and GST= 0.085 without the Lambir population). Furthermore, gene flow (N_m) between populations was not significantly correlated with geographical distances for the populations within Peninsular Malaysia. Cluster analysis also did not reflect geographical proximity and gave little insight into the genetic relatedness of the populations. This may indicate that the populations sampled are part of a continuous population with fragmentation having occurred in the recent past.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

Restoration ecology in the Anthropocene: learning from responses of tropical forests to extreme disturbance events

Extreme disturbance events denote another aspect of global environmental changes archetypal of the Anthropocene. These events of climatic or anthropic origin are challenging our perceived understanding about how forests respond to disturbance. I present a general framework of tropical forest responses to extreme disturbance events with specific examples from tropical dry forests. The linkage between level of disturbance severity and dominant mechanism of vegetation recovery is reflected on a variety of initial trajectories of forest succession. Accordingly, more realistic and cost‐effective restoration goals in many tropical forests likely consist in maintaining a mosaic of different successional trajectories while promoting landscape connectivity, rather than encouraging full‐ecosystem recovery to pre‐disturbance conditions. Incorporating extreme disturbance events into the global restoration ecology agenda will be essential to design well‐informed ecosystem management strategies in the coming decades.     

Diversity and Coexistence of Ectoparasites in Small Rodents in a Tropical Dry Forest

To understand the mechanisms driving species diversity is central to community ecology. Here, we explored if habitat partitioning is associated with a species‐rich ectoparasite community in small rodents from a tropical dry forest in western Mexico. We trapped 199 mice in three 0.5 ha‐plots from eight small rodent species for every two months, from July 2011 to April 2012, and collected their ectoparasites. We identified 17 species of mites, two sucking lice species, two phoretic species, and one commensal species. The most abundant ectoparasite species was Steptolaelaps liomydis, representing 42 percent of all ectoparasites collected; seven ectoparasite species had < 10 individuals. Eighteen ectoparasite species (of 22 species) were collected from the most abundant rodent Liomys pictus. C‐score and the number of checkerboard species pairs were significantly higher against a random expectation. Ectoparasite species in L. pictus mice showed host microhabitat partitioning; Fahrenholzia ehrlichi and Fahrenholzia texana were found only in the anterior dorsal area, Ornithonysus sp. occurred along the dorsal part, Ixodes species were restricted to the ears, and Steptolaelaps liomydis was found throughout the body. We also identified ectoparasite communities with distinct species composition in two rodent species that use contrasting macrohabitats (L. pictus, strictly terrestrial; Peromyscus perfulvus, mostly arboreal). The remaining and low abundant rodent species showed a species‐poor ectoparasite community composition. We conclude that habitat partitioning at both macro and microhabitat scales appeared to characterize the species‐rich ectoparasite community. Conversely, most rodent host species with low abundances showed a species‐poor ectoparasite community.     

Tropical forest and savanna ecosystems show differential impact of N and P additions on soil organic matter and aggregate structure

Landscape transformation and atmospheric nutrient depositions, important global change drivers, are affecting the vegetation and soil properties of natural dry tropical forest and derived savanna ecosystems in India. This study assessed the effect of continuous N and P additions for 6 years on the size distribution and properties of soil aggregates in forest–ecotone–savanna gradient. Addition of N significantly increased the proportion of macroaggregates in forest and ecotone, whereas the same input significantly decreased their proportion in the savanna. Consequently, the stability of soil aggregates increased significantly in forest and ecotone, whereas it decreased significantly in the savanna. The effect of P addition on soil aggregate stability was marginal. N addition also altered the biological and chemical qualities of soil aggregates. It caused increase in microbial biomass C (MBC) associated with macroaggregates in forest and ecotone; however, in savanna, MBC increased in the microaggregates. P addition did not affect the amount of MBC in both types of soil aggregates. Because of rapid accumulation of applied N and P in the microbial biomass, the ratios of MBC to microbial biomass nitrogen (MBN) as well as microbial biomass phosphorous (MBP) were decreased in both aggregates. Overall, the effect of N addition was more marked than that of P addition, suggesting that N is more limiting than P in these dry tropical ecosystems. In the current scenario of N loading, continued soil N loading in forest may lead to increased macroaggregates with associated MBC and MBN and greater aggregate stability. In contrast, the extensively distributed savannas may show the reverse trend leading to a decrease in soil fertility.     

Land use change and the global carbon cycle: the role of tropical soils

Millions of hectares of tropical forest are cleared annually for agriculture, pasture, shifting cultivation and timber. One result of these changes in land use is the release of CO₂ from the cleared vegetation and soils. Although there is uncertainty as to the size of this release, it appears to be a major source of atmospheric CO₂, second only to the release from the combustion of fossil fuels. This study estimates the release of CO₂ from tropical soils using a computer model that simulates land use change in the tropics and data on (1) the carbon content of forest soils before clearing; (2) the changes in the carbon content under the various types of land use; and (3) the area of forest converted to each use. It appears that the clearing and use of tropical soils affects their carbon content to a depth of about 40 cm. Soils of tropical closed forests contain approximately 6.7 kg C · m^-2; soils of tropical open forests contain approximately 5.2 kg C · m^-2 to this depth. The cultivation of tropical soils reduces their carbon content by 40% 5 yr after clearing; the use of these soils for pasture reduces it by about 20%. Logging in tropical forests appears to have little effect on soil carbon. The carbon content of soils used by shifting cultivators returns to the level found under primary forest about 35 yr after abandonment. The estimated net release of carbon from tropical soils due to land use change was 0.11–0.26 × 10¹⁵ g in 1980.     

Structural diversity and representativeness of forest vegetation in a protected area of Kumaun Himalaya, India: implications for conservation

Forest vegetation of a Protected Area (Askot Wildlife Sanctuary) in Kumaun (west Himalaya) was analysed for structure, composition and representativeness. A high percentage of non-natives was noticed in herbaceous flora of all representative forest types. Floristic representativeness in all growth forms (tree, shrub and herbs) increased significantly (tree, p<0.05; shrub and herbs, p<0.01) with altitude. The population structures of trees suggest, (i) expansion of riverine and Pinus roxburghii forests; (ii) compositional changes in Quercus leucotrichophora and Quercus lanuginosa forests, largely owing to preferred extraction demand of inhabitants; and (iii) infrequent regeneration and declining population of Quercus semecarpifolia and Abies pindrow forests. The possible pathways of non-native introduction in the study area were examined. Considering the existing status of forest vegetation and future trends, proliferation of non-native species in most forest types was discussed. It is suggested that the compositional changes vis-a`-vis proliferation of non-native species need priority attention while initiating conservation activities in the reserve.     

Multiple Methods in the Study of Driving Forces of Land Use and Land Cover Change: A Case Study of SE Kajiado District,Kenya

This landscape-scale study combines analysis of multitemporal satellite imagery spanning 30 years and information from field studies extending over 25 years to assess the extent and causes of land use and land cover change in the Loitokitok area, southeast Kajiado District, Kenya. Rain fed and irrigated agriculture, livestock herding, and wildlife and tourism have all experienced rapid change in their structure, extent, and interactions over the past 30 years in response to a variety of economic, cultural, political, institutional, and demographic processes. Land use patterns and processes are explored through a complementary application of interpretation of satellite imagery and case study analysis that explicitly addresses the local–national spatial scale over a time frame appropriate to the identification of fundamental causal processes. The results illustrate that this combination provides an effective basis for describing and explaining patterns of land use and land cover change and their root causes.