Influence of Forest Fragmentation on Community Structure of Frogs and Lizards in Northeastern Costa Rica

Abstract:  To better understand responses of reptiles and amphibians to forest fragmentation in the lowland Neotropics, we examined community and population structure of frogs and lizards in the fragmented landscape surrounding La Selva Biological Station in the Sarapiquí region of northeastern Costa Rica. We used diurnal quadrats and nocturnal transects to sample frogs and lizards in nine forest fragments (1–7 ha each) and La Selva (1100 ha). Species richness in all fragments combined was 85% of that found in La Selva with comparable sampling effort. Richness varied from 10 to 24 species among forest fragments, compared with 36 species at La Selva. Lizard density was higher and frog density was lower in forest fragments than in La Selva. Community composition varied among sites and by fragment size class, and species occurrence was nested with respect to fragment area. Isolation and habitat variables did not significantly affect species richness, composition, or nestedness. We classified 34% of species as fragmentation sensitive because they were absent or occurred at low densities in fragments. Nevertheless, the relatively high diversity observed in the entire set of fragments indicates that preserving a network of small forest patches may be of considerable conservation value to the amphibians and reptiles of this region.     

Avian Community Response to Lowland Tropical Rainforest Isolation: 40 Years of Change at La Selva Biological Station, Costa Rica

Abstract:  Since 1960, most of the forest surrounding the La Selva Biological Station, an intensively studied tropical research facility in Costa Rica, has been converted to agricultural uses. We used quantitative censuses and analysis of previously published categorical abundances to assess changes in the bird community, and we evaluated potential causes of species-specific changes by assessing their association with habitat, diet, participation in mixed-species flocks, and nest type. Approximately the same percentage of species increased as decreased in abundance from 1960 to 1999 (10–20% of all species, depending on method of assessment). Diet was the single most important trait associated with declining species. At least 50% of the species that declined have insectivorous diets. Use of forest habitat and participation in mixed-species flocks were also significant factors associated with declines, but nest type was unrelated to change in abundance. The species that increased in abundance tended to occur in open habitats and have omnivorous diets. These results reinforce the importance of several population risk factors associated with tropical understory insectivory and mixed-species flocking: patchy spatial distribution, low population density, large home range, and dietary specialization. La Selva's protected area (1611 ha), despite a forested connection on one boundary with a higher elevation national park, is apparently too small to maintain at least one major guild (understory insectivores). This first quantitative assessment of bird community change at La Selva highlights the need to intensify study of the mechanisms and consequences of biological diversity change in tropical forest fragments.     

Seedling growth responses to soil resources in the understory of a wet tropical forest

Plant growth responses to resources may be an important mechanism that influences species' distributions, coexistence, and community structure. Irradiance is considered the most important resource for seedling growth in the understory of wet tropical forests, but multiple soil nutrients and species have yet to be examined simultaneously with irradiance under field conditions. To identify potentially limiting resources, we modeled tree seedling growth as a function of irradiance and soil nutrients across five sites, spanning a soil fertility gradient in old-growth, wet tropical forests at La Selva Biological Station, Costa Rica. We measured an array of soil nutrients including total nitrogen (total N), inorganic N (nitrate [NO3-] and ammonium [NH4+]), phosphate (PO4-), and sum of base cations (SBC; potassium, magnesium, and calcium). Shade in the forest understory did not preclude seedling growth correlations with soil nutrients. Irradiance was a significant predictor of growth in 52% of the species, inorganic N in 54% (NO3- in 32%; NH4+ in 34%), total N in 47%, SBC in 39%, and PO4- in 29%. Overall, growth was correlated with both irradiance and soil nutrients in 45% of species and with soil nutrients only in an additional 48%; rarely was irradiance alone correlated with growth. Contrary to expectations, the magnitudes of growth effects, assessed as the maximum growth response to significant resources for each species, were similar for irradiance and most soil nutrients. Among species whose growth correlated with soil nutrients, the rank importance of nutrient effects was SBC, followed by N (total N, NO3-, and/or NH4+) and PO4-. Species' growth responsiveness (i.e., magnitudes of effect) to irradiance and soil nutrients was negatively correlated with species' shade tolerance (survival under 1% full sun). In this broad survey of species and resources, the nearly ubiquitous effects of soil nutrients on seedling growth challenge the idea that soil nutrients are less important than irradiance in the light-limited understory of wet tropical forests.     

Controls over foliar N:P ratios in tropical rain forests

Correlations between foliar nutrient concentrations and soil nutrient availability have been found in multiple ecosystems. These relationships have led to the use of foliar nutrients as an index of nutrient status and to the prediction of broadscale patterns in ecosystem processes. More recently, a growing interest in ecological stoichiometry has fueled multiple analyses of foliar nitrogen:phosphorus (N:P) ratios within and across ecosystems. These studies have observed that N:P values are generally elevated in tropical forests when compared to higher latitude ecosystems, adding weight to a common belief that tropical forests are generally N rich and P poor. However, while these broad generalizations may have merit, their simplicity masks the enormous environmental heterogeneity that exists within the tropics; such variation includes large ranges in soil fertility and climate, as well as the highest plant species diversity of any biome. Here we present original data on foliar N and P concentrations from 150 mature canopy tree species in Costa Rica and Brazil, and combine those data with a comprehensive new literature synthesis to explore the major sources of variation in foliar N:P values within the tropics. We found no relationship between N:P ratios and either latitude or mean annual precipitation within the tropics alone. There is, however, evidence of seasonal controls; in our Costa Rica sites, foliar N:P values differed by 25% between wet and dry seasons. The N:P ratios do vary with soil P availability and/or soil order, but there is substantial overlap across coarse divisions in soil type, and perhaps the most striking feature of the data set is variation at the species level. Taken as a whole, our results imply that the dominant influence on foliar N:P ratios in the tropics is species variability and that, unlike marine systems and perhaps many other terrestrial biomes, the N:P stoichiometry of tropical forests is not well constrained. Thus any use of N:P ratios in the tropics to infer larger-scale ecosystem processes must comprehensively account for the diversity of any given site and recognize the broad range in nutrient requirements, even at the local scale.     

Tolerance of soil pathogens co-varies with shade tolerance across species of tropical tree seedlings

A negative feedback between local abundance and natural enemies could contribute to maintaining tree species diversity by constraining population growth of common species. Soil pathogens could be an important mechanism of such noncompetitive distance and density-dependent (NCDD) mortality, but susceptibility to local pathogens may be ameliorated by a life history strategy that favors survivorship. In a shade-house experiment (1% full sun), we tested seedling life span, growth, and mass allocation responses to microbial extract filtered from conspecific-cultured soil in 21 tree species that varied in abundance and shade tolerance in a wet tropical forest (La Selva Biological Station, Costa Rica). Forty-three percent of the species had significant reductions, and 10% of the species had significant increases in life span, growth, root length, or root surface area when inoculated with microbial extract; 10% of the species experienced opposing reductions and increases in these characteristics. Contrary to expectation, species' local abundance was not related to species-specific responses to microbial extracts from cultured soils. Across species, seedling shade tolerance (survival at 1% full sun) was negatively correlated with susceptibility to the microbial. treatment for both survival and total mass accumulation, thereby exaggerating shade tolerance differences among species. Thus, soil pathogens may contribute to species coexistence through heightening niche differentiation rather than through negative density dependence in common species.     

Multigenerational genetic analysis of tropical secondary regeneration in a canopy palm

Iriartea deltoidea (Arecaceae) is an abundant canopy palm with a wide geographic distribution in Neotropical wet forests. We analyzed the genetic profile across three generations of Iriartea within a 43-ha area encompassing two areas of second-growth and adjoining old-growth forest at La Selva Biological Field Station in northeastern Costa Rica. A total of 311 reproductively mature trees, 99 large saplings, 207 small saplings, and 601 seedlings were genotyped using 141 AFLP loci. Parentage analysis revealed high dispersal distances, both for seed (over 2.3 km) and pollen (over 3.8 km), indicating a large genetic neighborhood within La Selva Biological Station. In a 20-ha area of second growth, the founding palm population was dominated by a small number of parental trees located in the adjacent old-growth forest; two old-growth trees contributed 48% of the second-growth genes. The genetic diversity of reproductively mature trees in this second-growth forest was significantly reduced compared to adjacent old-growth forest. Within 400 m of the border with old-growth forest, we observed a similar reduction of genetic diversity in saplings, and an even greater loss of genetic diversity in the second generation of seedlings. Nearly half of these seedlings were offspring of local parents. In contrast, in the distant portion of second-growth forest (400-800 m from the old-growth border), parentage analysis showed that 40% of seedlings originated from outside the study area and only 10% were offspring of local parents. These high levels of gene flow maintained genetic diversity in saplings and seedlings similar to levels observed in old-growth forest. Our findings highlight the importance of gene flow from diverse seed and pollen sources for sustaining levels of genetic diversity of tree populations in second-growth forests.     

Prevalence and Intensity of Intestinal Parasites in Mantled Howling Monkeys (Alouatta palliata) in Northeastern Costa Rica: Implications for Conservation Biology

I examined the prevalence and intensity of intestinal parasitic infections in a population of mantled howling monkeys ( Alouatta palliata ) at La Selva Biological Reserve, a rainforest in northeastern Costa Rica. I collected fresh fecal samples from individual bowler monkeys from two focal troops from October 1991 through October 1992. The presence of parasite ova and larvae in samples was determined by using a formalin-ethyl acetate sedimentation technique. Three types of endoparasites were found: (1) a roundworm, Parabronema sp., (2) a physalopterid roundworm, species undetermined, and (3) a fluke, species undetermined. No differences were found in the prevalence of parasitic infections between sexes or troops. No statistical differences were found between the sexes for the intensity of infection. The troop that lived along the river showed a statistically significant higher intensity of nematode infection than the troop that lived mostly in primary forest. Parabronema sp. was found only from individuals in the river group. The prevalence of parasitic infection observed in howlers at La Selva (100%) is higher than has been reported for bowlers in a dry deciduous forest at La Pacifica, Costa Rica (47%). Microclimatic factors, ranging patterns, and home-range size are identified as important variables that may affect parasitic infections of howler populations. Data from this study suggest that narrow corridor designs may be inappropriate for conservation of primates and possibly other species of arboreal animals.     

Population and seasonal variation in response to prey calls by an eavesdropping bat

The fringe-lipped bat, Trachops cirrhosus, is an eavesdropping predator that hunts frogs and katydids by approaching these preys' sexual advertisement calls. In captivity, bats can rapidly learn to associate novel acoustic stimuli with food rewards. It is unknown how this learning ability is related to foraging behavior in the wild where prey and the calls that identify them vary over space and time. In two bat populations that differ in available prey species (Soberanía, Panama, and La Selva, Costa Rica), we presented wild-caught bats with frog calls, katydid calls, and control stimuli. Bats in Soberanía were significantly more responsive to complex calls and choruses of the túngara frog, Physalaemus pustulosus, than were bats in La Selva. La Selva bats were significantly more responsive to katydid calls (Steirodon sp.) than Soberanía bats. We also examined seasonal variation in bat response to prey cues. Bats were captured in Soberanía in dry and wet seasons and presented with the calls of a dry season breeding frog (Smilisca sila), a wet season breeding frog (P. pustulosus), and four katydid species. Bats captured in the dry season were significantly more responsive to the calls of S. sila than bats captured in the wet season, but there were no seasonal differences in response to the calls of P. pustulosus or the katydid calls. We demonstrate plasticity in the foraging behavior of this eavesdropping predator but also show that response to prey cues is not predicted solely by prey availability.     

Nutrient regulation of organic matter decomposition in a tropical rain forest

Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.     

Mammal Diversity and Conservation in the Selva Lacandona, Chiapas, Mexico

The Selva Lacandona region of Chiapas, Mexico, has high biodiversity, represents the last large portion of tropical rainforest in México, and faces imminent destruction. Through fieldwork and literature searches I found a total of 112 mammal species (including 17 Middle-American endemics) on the Lacandona's 331,200 ha Montes Azules Biosphere Reserve. This inventory plus those from eight additional Neotropical localities show local mammal species richness in the Neotropics to be in the range of 70–116 species. Richness is primarily correlated with the amount of annual rainfall, but within rainforest areas species richness is probably not correlated with rainfall, latitude, altitude, or area size. Rather, above a certain limit of rainfall and below a critical level of latitude and altitude, mammal species richness seem to reach an asymptotic maximum. Areas covered with tropical rainforest and with a well-known, relatively intact fauna have about 112–116 mammal species. In the Montes Azules Reserve most mammals (57%) are bats. The most heavily occupied feeding guilds are frugivores/herbivores and insectivores/omnivores. Large species and carnivores appear most vulnerable to local extinction. Montes Azules (and thus the Selva Lacandona) is especially noteworthy because (1) it contains a greater proportion of species facing conservation problems than expected from a random draw of Mexican mammals; (2) it is probably the most diverse ecosystem in Mexico; (3) many species sustain their only Mexican populations in this area; and (4) it is the largest remnant of tropical rainforest in Mexico and is part of the largest expanse of that vegetation type in Central America.     

Effects of Recent Land-Use Practices on Soil Nutrients and Succession under Tropical Wet Forest in Costa Rica

The effects of a variety of agricultural land uses were studied using soil nutrients, forest structure, and species assemblages as indicators. We compared soil properties and successional forests between abandoned cacao ( Theobroma cacao ) and abandoned palm ( Bactris gasipaes ) orchards, abandoned pasture, and mature forest. These sites co-occupy an alluvial terrace soil ( Andic Dystropept ) at La Selva Biological Station, Costa Rica. The agricultural sites were originally cleared of most or all forest vegetation approximately 30 years ago and went into succession approximately 7 years ago. Forest structure, species composition, soil nitrogen and phosphorus pools, and nitrogen-mineralization and nitrification rates were measured for each site. The abandoned palm orchard had lower basal area and stem density than other secondary forests of the same age. It also had significantly smaller nitrate (NaOH-extractable) and organic phosphorus pools and significantly lower net rates of nitrogen-mineralization and nitrification. It is evident that preserving tree cover does not necessarily maintain soil fertility. We found species richness and diversity in the secondary forests to be positively correlated with basal area at the time of abandonment.     

Background values and distribution trends of Cu and Zn in soils of humid Mediterranean environment

Soil is the foundation of the entire biosphere. Knowledge of its heavy metal content and regional variability is essential to assess the environmental quality of soil and the extent of any contamination. In this study, 250 soil samples (within 125 soil profiles) were collected in undisturbed soils of the La Rioja region (an ideal place of the humid Mediterranean environment of Spain). The ‘total and bioavailable’ contents of copper (Cu) and zinc (Zn) were measured and their spatial variability assessed. The results indicate that Zn and Cu values were found to be very close to national and global averages. The spatial distribution of those elements was related to the nature of the bedrock and, to a lesser extent, was of anthropogenic origin. The variation of vertical distributions can be related, firstly, to natural sources – mainly the bedrock – and, secondly, to soil processes. From an ecological perspective, the ecosystem has not been affected by pollution.     

Application of Extinction and Conservation Theories for Forest Birds in Nicaragua

Abstract: An increasing number of empirical studies have been done on the effects of tropical forest fragmentation on avian communities, but few researchers have applied these theories to assess the vulnerability of birds in poorly researched countries such as Nicaragua. I used a logistic regression to determine which natural-history characteristics were most important in predicting a list of threatened birds known to occur in Nicaragua. The best model included five macroecological variables ( body weight, habitat specificity, trophic group, forest preference, and biogeography within Nicaragua). I used this model to generate predicted probabilities of extinction for all forest birds in Nicaragua. The predicted probability of extinction from the best model ranked 63% of the extinction-prone birds from La Selva, Costa Rica, and 59% of the extinction-prone birds from Barro Colorado, Panama, in the first quartile of all forest birds recorded in Nicaragua. This method provides a first-order approximation of which species deserve global and national priorities for conservation. The central and Atlantic regions of Nicaragua deserve high priority for conservation at a global scale, whereas the Atlantic region deserves the highest priority for conservation at a national scale. The Nicaraguan Ministry of Natural Resources and the Environment has done an adequate job of identifying areas for conservation based on the proportion of decreed nature reserves in each biogeographic region and the distribution of forest birds with a high predicted probability of extinction. Forest birds in central Nicaragua, however, may currently be the most vulnerable to local extinction because of low forest cover within decreed reserves.     

The lichen-rich coastal heath vegetation on the isle of Anholt, Denmark — conservation and management

The high conservational value of the lichen-rich vegetation and landscape of the marine foreland Ørkenen on the isle of Anholt is treated from a Danish as well as a European perspective. The sensitivity of the lichen-rich vegetation to physical disturbance is emphasized. The impact of invasive species such asPinus mugo as well as the effect of atmospheric deposition of nutrients on the heaths at Anholt is described. Considerations related to the development of a management plan for Ørkenen are presented.     

Historical metal concentrations in lacustrine food webs revealed using fossil ephippia from Daphnia.

Metal contamination of freshwater ecosystems is increasingly prevalent due to anthropogenic activities such as metal smelting and fossil fuel combustion. While toxicological studies focus on aqueous metal concentrations that result in lethal or sublethal responses, currently the only method for reconstructing a lake's metal contamination history is through an examination of the sedimentary deposits. In this paper, we suggest that cladoceran diapausing eggs (ephippia), which are abundant in nature and accumulate maternally derived metals, can be used to measure historical variations in biologically relevant metals that derive from the water column (water, diet). Linear regressions of total metal content against ephippia density or mass were strong (R2 > 0.80, P < 0.04) and revealed that metals were incorporated into ephippia with little contamination from the sediment matrix. Comparison of metal concentrations in ephippia and bulk sediments from three lakes demonstrated that some metals associated with urban sources (Cd, Cr, Mo) were preferentially concentrated in ephippia, whereas concentrations of other metals indicating landscape erosion (Al, Ca, Fe, Mn) exhibited greater concentrations in bulk sediments than in diapausing eggs. Because historical changes in metals within fossils and bulk sediments were uncorrelated in most instances, past variation in the metal content of ephippia provided a unique history of food web exposure to metals in the water column.     

Resource availability and the abundance of an N-based defense in Australian tropical rain forests

Plant defense theories predict that relatively resource-rich environments (those with more fertile soil) will support a greater abundance of plants with nitrogen-based chemical defense, but this has yet to be adequately tested. We tested this prediction by measuring the diversity and contribution to total biomass of cyanogenic plants (those that release hydrogen cyanide from endogenous cyanide-containing compounds) in the Australian tropical rain forest. We examined 401 species in thirty 200-m2 plots, six at each of five sites, for cyanogenesis. In upland/highland rain forest, two pairs of sites similar in rainfall and altitude, but differing in soil nutrients, were selected, as well as one site in lowland rain forest. Sites differed markedly in species composition and foliar N was positively related to soil fertility. Holding altitude constant, we did not detect significant differences in the proportion of cyanogenic species with soil fertility, nor did we consistently detect significant increases in the contribution of cyanogenic species to total biomass on higher nutrient sites. Thus we found no clear evidence that soil fertility affects the distribution and prevalence of species investing in a constitutive N-based defense at the community level.     

Epiphytic cyanobacteria maintain shifts to macroalgal dominance on coral reefs following ENSO disturbance

Macroalgal dominance of some tropical reef communities in the Eastern Pacific after coral mortality during the 1997-1998 El Ni?o Southern Oscillation (ENSO) was facilitated by protection from herbivory by epiphytic cyanobacteria. Our results do not support that reduction in number of herbivores was a necessary precursor to coral reef decline and shifts to algal reefs in this system. Rather, macroalgae dominated the community for several years after this pulse disturbance with no concurrent change in herbivore populations. While results of microcosm experiments identified the importance of nutrients, especially phosphorus, in stimulating macroalgal growth, nutrient supply alone could not sustain macroalgal dominance as nutrient-stimulated growth rates in our in situ experiments never exceeded consumption rates of unprotected thalli. In addition, thalli with nutrient-enriched tissue were preferentially consumed, possibly negating the positive effects of nutrients on growth. These tropical reefs may be ideal systems to conduct experimental tests distinguishing phase shifts from alternative stable states. Shifts were initiated by a large-scale disturbance with no evidence of a changing environment except, perhaps, dilution in herbivory pressure due to increased algal cover. Community establishment was most likely stochastic, and the community was likely maintained by strongly positive interaction between macroalgal hosts and cyanobacterial epiphytes that uncoupled consumer control of community structure.     

Nutrient fluxes at the landscape level and the R* rule

Nutrient cycling in terrestrial ecosystems involves not only the vertical recycling of nutrients at specific locations in space, but also biologically driven horizontal fluxes between different areas of the landscape. This latter process can result in net accumulation of nutrients in some places and net losses in others. We examined the effects of such nutrient-concentrating fluxes on the R* rule, which predicts that the species that can survive in steady state at the lowest level of limiting resource, R*, can exclude all competing species. To study the R* rule in this context, we used a literature model of plant growth and nutrient cycling in which both nutrients and light may limit growth, with plants allocating carbon and nutrients between foliage and roots according to different strategies. We incorporated the assumption that biological processes may concentrate nutrients in some parts of the landscape. We assumed further that these processes draw nutrients from outside the zone of local recycling at a rate proportional to the local biomass density. Analysis showed that at sites where there is a sufficient biomass-dependent accumulation of nutrients, the plant species with the highest biomass production rates (roughly corresponding to the best competitors) do not reduce locally available nutrients to a minimum concentration level (that is, minimum R*), as expected from the R* rule, but instead maximize local nutrient concentration. These new results require broadening of our understanding of the relationships between nutrients and vegetation competition on the landscape level. The R* rule is replaced by a more complex criterion that varies across a landscape and reduces to the R* rule only under certain limiting conditions.     

Nocturnal accumulation of CO2 underneath a tropical forest canopy along a topographical gradient

Flux measurements of carbon dioxide and water vapor above tropical rain forests are often difficult to interpret because the terrain is usually complex. This complexity induces heterogeneity in the surface but also affects lateral movement of carbon dioxide (CO2) not readily detected by the eddy covariance systems. This study describes such variability using measurements of CO2 along vertical profiles and along a toposequence in a tropical rain forest near Manaus, Brazil. Seasonal and diurnal variation was recorded, with atmospheric CO2 concentration maxima around dawn, generally higher CO2 build-up in the dry season and stronger daytime CO2 drawdown in the wet season. This variation was reflected all along the toposequence, but the slope and valley bottom accumulated clearly more CO2 than the plateaus, depending on atmospheric stability. Particularly during stable nights, accumulation was along lines of equal altitude, suggesting that large amounts of CO2 are stored in the valleys of the landscape. Flushing of this store only occurs during mid-morning, when stored CO2 may well be partly transported back to the plateaus. It is clear that, for proper interpretation of tower fluxes in such complex and actively respiring terrain, the horizontal variability of storage needs to be taken into account not only during the night but also during the mornings.     

镧积累对红壤有效养分的影响

通过室内培养和盆栽试验 ,研究了镧积累对红壤有效养分的影响。结果表明 ,镧积累降低了土壤有效氮、有效磷含量 ,有效氮、有效磷含量与镧浓度呈显著的负相关 ,镧对土壤有效钾的影响不明显。