Conservation of tropical forest birds in countryside habitats

The pressing need to increase agricultural production often seems at odds with conserving biodiversity. We find that if managed properly, the tropical countryside may provide a substantial opportunity for tropical bird conservation. We detected 144 bird species from 29 families in agricultural areas outside of extensive native forest in southern Costa Rica. The majority of the species detected were observed foraging, often kilometres from extensive native forest. We estimate that 46% of those native to this region (excluding nocturnal species and waterfowl) are utilizing the countryside in some manner. Forecasts of biodiversity change under various land-use scenarios indicate that policies that affect habitat composition could greatly impact the persistence of these species in the countryside. In particular, if tall trees and edge habitats were removed from this landscape, we predict that bird richness in the countryside would decline by approximately 40%.     

Behaviour and ecology of birds in tropical rain forest canopies

The canopy of rain forests exhibits several distinct features that determine avian adaptations. Light intensity and colour, wind, and temperature are the most important components of the physical environment. Tree architecture poses many challenges for the evolution of avian morphological features, and resource availability varies greatly. The canopy is easily accessible for birds that are equipped to travel over considerable distances. Given these conditions, high behavioural flexibility, and adaptations in behaviour and morphology related to a nomadic life-style, should prevail among canopy birds. The easy access for certain groups of birds also has consequences for the historical processes that shape canopy bird communities. Birds may contribute greatly to forest dynamics and composition, mainly due to their role as seed dispersers. We elaborate on these points with behavioural, morphological, and ecological data collected at our Neotropical study site in southern Venezuela (Surumoni Canopy Crane Project). We discuss how behaviour and morphological features reflect the ecological demands in the canopy. We contrast the specific behavioural and ecological characteristics of canopy birds with those of species found in the other tiers of the rain forest. It is pointed out that much of our knowledge is biased towards the Neotropics, and that better knowledge of the larger canopy species is needed for conservation.     

Partial migration in birds: tests of three hypotheses in a tropical lekking frugivore

1. Partially migratory species provide opportunities to understand which ecological factors cause some animals to migrate when others remain resident year round. Partial migration in birds has been explained by the dominance, arrival-time, and body-size hypotheses. 2. Testing these hypotheses has proven difficult due to the similarities of the predictions they make in temperate-breeding long-distance migrants. In tropical altitudinal migrants, however, these hypotheses make different predictions regarding the sex, age, and condition of migrants and residents. 3. Among white-ruffed manakins in Costa Rica, young birds were not more likely to migrate (as predicted by the dominance hypothesis), nor were females more likely to migrate (as predicted by the arrival-time hypothesis). All condition-related variables interacted with sex, together explaining much of the variation in migratory behaviour. 4. I re-articulate the body-size hypothesis in the context of tropical altitudinal bird migration, focusing explicitly on how limited foraging opportunities and differences in individual condition affect fasting ability during torrential rains. Despite ample food, the smallest birds or those stressed by parasites or moult may risk starvation at breeding elevations due to a reduction in foraging time. These results highlight how intrinsic and extrinsic factors may interact to produce observed patterns of within- and among-species variation in migratory behaviour.     

Above and below ground impacts of terrestrial mammals and birds in a tropical forest

Understanding the impact of losing trophic diversity has global significance for managing ecosystems as well as important theoretical implications for community and ecosystem ecology. In several tropical forest ecosystems, habitat fragmentation has resulted in declines and local extinctions of mammalian and avian terrestrial insectivores. To assess the ability of a tropical rainforest community in Ivory Coast to resist perturbation from such loss of trophic diversity, I traced feedbacks in above and below ground communities and measured changes in nutrient levels and herbivory rates in response to an experimental exclosure of avian and mammalian terrestrial insectivores. I present evidence that loss of this functional group may result in increased tree seedling herbivory and altered nutrient regimes through changes in the abundance and guild structure of invertebrates. Exclusion of top predators of the forest floor resulted in increased seedling herbivory rates and macro-invertebrate (>5 mm) densities with strongest effects on herbivorous taxa, spiders and earthworms. Densities of microbivores including Collembola, Acarina and Sciaridae showed the opposite trend as did levels of inorganic phosphorus in the soil. Results were evaluated using path analysis which supported the presence of a top down trophic cascade in the detrital web which ultimately affected turnover of phosphorus, a limiting nutrient in tropical soils. Results illustrate the potential importance of vertebrate predators in both above and belowground food webs despite the biotic diversity and structural heterogeneity of the rainforest floor.     

Effects of an ENSO-related fire on birds of a lowland tropical forest in Sumatra

Comparisons of bird community composition in burned and unburned areas of a lowland tropical rainforest in Sumatra, Indonesia indicated the following during the first 5 years after burning: (1) original burn severity strongly affected bird community composition at both the genus and family levels; (2) bird community composition continued to change progressively away from immediate post-burn composition in medium and severely burned forest as well as adjacent unburned forest; and (3) the degree of impact was both taxon and guild specific, with understory insectivores most detrimentally affected. Although species richness may temporarily increase in burned areas, this study suggests that multiple wildfires will lead to a decline in diversity over a large scale as birds of open fields replace interior forest specialists.     

Lekking in birds: choice, competition and reproductive constraints

This paper discusses the importance of female choice and intrasexual competition in lek species. The importance of phenotypic differences between individuals is stressed. Reproductive constraints limit the possible behavioural strategies that can be used by males and females. I suggest that it is difficult to understand reproductive behaviour without a knowledge of these reproductive constraints.     

The distribution of forest passerine birds and Quaternary climatic change in tropical Africa

A cluster analysis of the distribution patterns of forest passerine birds in tropical Africa showed that they could be divided into three main distribution types: those of restricted distribution (144 species), those of disjunct distributions across West, Central and rarely to East Africa (65 species) and those of very wide distribution (81 species). Centres of species richness, endemism and disjunction coincide spatially, and are identified as forest refugia, where forest persisted throughout Quaternary climatic vicissitudes. These distribution patterns agree with modern interpretations of Quaternary palaeoclimatic changes, which show that glacial periods were arid and interglacials humid. Glacial periods were therefore too dry for montane forest to have spread into areas at present occupied by lowland forest, as advocated by Moreau and other supporters of ihe pluvial theory; montane forest is probably as extensive now as at any time during the Quaternary and there is no evidence of past connections between currently isolated montane forests. Bird distribution can be explained largely by slow dispersal outwards from refugia as climatic conditions allowed forest to spread, combined with a relaxing of the distinction between montane and lowland species under conditions of reduced interspecific competition.     

Trophic partitioning in tropical rain forest birds: insights from stable isotope analysis

Bird communities reach their highest taxonomic and trophic diversity in tropical rain forest, but the use of different foraging strategies to meet food requirements in such competitive environments is poorly understood. Conventional dietary analyses are poorly suited to investigate dietary patterns in complex systems. We used stable carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) isotope analysis of whole blood to examine avian trophic patterns and sources of diet in the tropical rain forest of Los Tuxtlas, Veracruz, Mexico. We used stable nitrogen isotope analysis to delineate trophic levels, and stable carbon isotope analysis to distinguish the relative contribution of C-3 and CAM/C-4 ultimate sources of proteins to diets. There was large inter- and intraspecific variation in whole blood ¹³C and ¹⁵N values in 23 species of birds. Stable nitrogen isotope analysis separated birds into several trophic levels, including species that obtained their dietary protein mostly from plants, insects or a combination of both food sources. Stable carbon isotope analysis showed that most birds fed on C3-based foods but Stub-tailed Spadebills (Platyrinchus cancrominus) included C-3- and C-4/CAM-specialist individuals. Our analyses provided insights into the nutritional contribution of plant and animal sources of protein and distinguish their photosynthetic origin over relatively long average time periods.     

Conservation value of secondary forest habitats for endemic birds,a perspective from two widely separated tropical ecosystems

Tropical secondary forests are increasingly widespread, but their potential for conserving endemic birds remains unclear. Previous studies report different results; however all have been restricted to geographically discreet locations. This is important as different ecosystems are influenced by different external factors, possibly influencing conservation potential. Here we use consistent survey methods to examine how endemic bird richness varies between primary and secondary forest habitats in two widely separated tropical ecosystems, providing a more global context for evaluating the conservation value of secondary forests. Research was completed in Lambusango Forest Reserve (LFR) on Buton Island, Sulawesi, and Cusuco National Park (CNP), a Honduran cloud forest reserve. Bird communities in both forests were surveyed using 50 m radius point counts. Vulnerability assessments based on ecological theory on avifaunal assemblages were then conducted, which suggested endemics in LFR to be more susceptible to disturbance than those in CNP. Contrary to the results from our vulnerability assessments, endemics in CNP were less tolerant of moderate habitat modification than those in LFR. Richness of Mesoamerican endemics per study site declined significantly between core zone forest (6.34 ± 0.81) and more degraded forest in the boundary zone (3.86 ± 0.69). Richness of Wallacean endemics was similar in primary (4.89 ± 1.68) and disturbed secondary forest (4.52 ± 1.62). We recommend considering local and regional biogeographical and ecological factors when determining the conservation value of secondary forests, and suggest examples of potential importance, including differential community richness, influence of figs and human settlement patterns.     

Species characteristics associated with extinction vulnerability and nestedness rankings of birds in tropical forest fragments

Following habitat fragmentation, species are predicted to go locally extinct from remnant patches in a predictable order due to differential extinction vulnerabilities. This selective species loss will result in nested distributions of species such that species found in depauperate patches will also tend to be found in larger, more speciose patches. Therefore, it should be possible to determine the relationship between species-specific characteristics and extinction vulnerability by comparing the order in which species are nested [i.e. nestedness ranking (NR)] with various natural history characteristics available from the literature and/or collected in the field. In this study, we investigate the relationship between the NRs of 41 resident forest-interior bird species inhabiting recently isolated landbridge islands in Lago Guri, Venezuela, with a large number of natural history characteristics collected from the literature (regional abundance, body length, habitat specificity, trophic guild, sensitivity to disturbance, range size) and from the field (local population density). In a comparison of the best regression models generated using just variables available through the literature (i.e. no local population density) with the best model generated using all possible variables, we found that the inclusion of field-based data significantly improved the amount of variation explained. The best overall model ( r ²=0.40, P <0.001) included body size, habitat specificity, zoogeographic distribution (a measure of range size) and local population density as predictors of NR. Understanding the factors that influence extinction vulnerability has important implications for conservation and could be used to help direct management efforts.     

Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland forests in East Africa

Burgess, N.D. & Mlingwa, C.O.F. 2000. Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland forests in East Africa. Ostrich 71 (1 & 2): 184–190.

In this paper we assess the evidence for altitudinal movements of forest birds from the montane forests of the Eastern Arc mountains of East Africa to nearby lowland forest patches. For 34 montane species, including all the Eastern Arc endemics except Banded Green Sunbird Anthreptes rubritorques there is no evidence that they undertake seasonal movements to lower altitudes. An additional 26 montane species, of somewhat wider distribution, have been recorded at low (<500 m) altitudes during the cold/dry season (June to September). Most records of these montane birds at lower altitudes are from sites adjoining montane forest areas, although a few records are from lowland coastal forests at 100–240 km distance from montane areas. Only five of the 26 species (White-chested Alethe Alethe fulleborni, White-starred Forest Robin Pogonocichla stellata, Orange Ground Thrush Zoothera gurneyi, Evergreen Forest Warbler Bradypterus mariae and Barred Long-tailed Cuckoo Cercococcyx montanus) are regularly and commonly reported in the lowlands. They are also found in the lowlands in small numbers during the warm/wet season (October to February), when they may breed. The abundance of at least four, and probably more, of the forest birds with a more widespread distribution in the lowland and montane forests of East Africa declines greatly at high altitudes from the onset of the cold/wet season (February) and only increases again at the start of the warm/wet season (September). It is not known how far these species move as they cannot be easily separated from resident populations in lowland forests, and there are no ringing recoveries in different forests. Altitudinal migration of a proportion of the Eastern Arc avifauna is the most likely explanation for available data, although source-sink metapopulation theories may be helpful to explain the distributions of some species. As the movement of forest birds from the Eastern Arc to the lowland forests does not involve the rare endemics, they are of lower conservation concern, but the presence of montane and lowland forest may be important for the long-term survival of some more widely distributed forest species.     

Nitrogen conservation in a tropical rain forest

Summary A series of plots in an Amazonian Rain Forest were trenched and treated with calcium to determine the effects of perturbation on numbers and activity of nitrifying bacteria. Although treatment resulted in between 2 and 26% of the nitrogen being lost from the humus layer, virtually all of it was in the NH ₄ ⁺ form. Numbers of nitrifying bacteria in the plots were relatively low. The low numbers and low activity of nitrifying bacteria is attributed to the low pH and high concentration of tannins in the root mat. It is hypothesized that the suppression of nitrifying bacteria results in nitrogen conservation in the rain forest.     

Non-structural carbohydrate pools in a tropical forest

The pool size of mobile, i.e. non-structural carbohydrates (NSC) in trees reflects the balance between net photosynthetic carbon uptake (source) and irreversible investments in structures or loss of carbon (sink). The seasonal variation of NSC concentration should reflect the sink/source relationship, provided all tissues from root to crown tops are considered. Using the Smithsonian canopy crane in Panama we studied NSC concentrations in a semi-deciduous tropical forest over 22 months. In the 9 most intensively studied species (out of the 17 investigated), we found higher NSC concentrations (starch, glucose, fructose, sucrose) across all species and organs in the dry season than in the wet season (NSC 7.2% vs 5.8% of dry matter in leaves, 8.8/6.0 in branches, 9.7/8.5 in stems, 8.3/6.4 in coarse and 3.9/2.2 in fine roots). Since this increase was due to starch only, we attribute this to drought-constrained growth (photosynthesis less affected by drought than sink activity). Species-specific phenological rhythms (leafing or fruiting) did not overturn these seasonal trends. Most of the stem volume (diameter at breast height around 40 cm) stores NSC. We present the first whole forest estimate of NSC pool size, assuming a 200 t ha^–1 forest biomass: 8% of this i.e. ca. 16 t ha^–1 is NSC, with ca. 13 t ha^–1 in stems and branches, ca. 0.5 and 2.8 t ha^–1 in leaves and roots. Starch alone (ca. 10.5 t ha^–1) accounts for far more C than would be needed to replace the total leaf canopy without additional photosynthesis. NSC never passed through a period of significant depletion. Leaf flushing did not draw heavily upon NSC pools. Overall, the data imply a high carbon supply status of this forest and that growth during the dry season is not carbon limited. Rather, water shortage seems to limit carbon investment (new tissue formation) directly, leaving little leeway for a direct CO₂ fertilization effects.     

Embryonic sex hormones in birds

Hormone activity of embryonic gonads in birds was demonstrated by grafting and culture experiments. Anti-Müllerian hormone responsible for the regression of the Müllerian ducts in the male is most probably a glycoprotein. Whether the testis also secretes testosterone has long been disputed, but most arguments are against this possibility. From early stages of development, the ovary secretes estrome and estradiol. However, it could not be demonstrated unambiguously whether estrogen is identical with the sex inducing substance in the female. The hypophysis seems to control ovarian estrogen secretion at 10-13 days of incubation in the chick embryo.     

Seedling interactions in a tropical forest in Panama

Competition is believed to be a central force limiting local diversity and controlling the structure of plant communities. However, it has been proposed that the stressed understory environment limits total understory plant density to such low levels that competitive exclusion cannot be an important factor limiting the local diversity of understory plants. To evaluate the importance of inter-seedling competition, we performed a seedling competition experiment with five shade-tolerant species in a tropical moist forest in Panama. Three-month-old seedlings were transplanted into the forest singly or with their roots intertwined with a single conspecific or heterospecific seedling in all pairwise species combinations. If competition is important, performance (survival, stem height, and number of leaves after one and six years) would be expected to be lowest with a conspecific neighbor and greatest without a neighbor. The experiment was replicated in five 0.24-m² plots at each of 20 sites in tall secondary forest. To test whether seedling performance differed among treatments we fitted linear mixed models (LMM) and generalized linear mixed models (GLMM), treating species identity and microsite (site and plot) as random effects. The five shade-tolerant study species all experienced good establishment with relatively high survival and growth rates. The neighbor treatment consistently affected seedling performance, but the effect was always very small, both in absolute terms and relative to the much stronger species and microsite effects. Seedlings with a conspecific neighbor consistently performed worse than seedlings with a heterospecific neighbor, but having no neighbor generally did not cause superior performance relative to the other treatments. We conclude that direct competitive interactions are relatively unimportant among understory plants in humid tropical forests.     

Traffic noise affects forest bird species in a protected tropical forest

The construction of roads near protected forest areas alters ecosystem function by creating habitat fragmentation and through several direct and indirect negative effects such as increased pollution, animal mortality through collisions, disturbance caused by excessive noise and wind turbulence. Noise in particular may have strong negative effects on animal groups such as frogs and birds, that rely on sound for communication as it can negatively interfere with vocalizations used for territorial defense or courtship. Thus, birds are expected to be less abundant close to the road where noise levels are high. In this study, we examined the effects of road traffic noise levels on forest bird species in a protected tropical forest in Costa Rica. Data collection was conducted in a forest segment of the Carara National Park adjacent to the Coastal Highway. We carried out 120 ten minute bird surveys and measured road noise levels 192 times from the 19th to the 23rd of April and from the 21st to the 28th of November, 2008. To maximize bird detection for the species richness estimates we operated six 12 m standard mist nets simultaneously with the surveys. The overall mist-netting effort was 240 net/h. In addition, we estimated traffic volumes by tallying the number of vehicles passing by the edge of the park using 24 one hour counts throughout the study. We found that the relative abundance of birds and bird species richness decreased significantly with the increasing traffic noise in the dry and wet season. Noise decreased significantly and in a logarithmic way with distance from the road in both seasons. However, noise levels at any given distance were significantly higher in the dry compared to the wet season. Our results suggest that noise might be an important factor influencing road bird avoidance as measured by species richness and relative abundance. Since the protected forest in question is located in a national park subjected to tourist visitation, these results have conservation as well as management implications. A decrease in bird species richness and bird abundance due to intrusive road noise could negatively affect the use of trails by visitors. Alternatives for noise attenuation in the affected forest area include the enforcement of speed limits and the planting of live barriers.     

Revised evidence for facultative sex ratio adjustment in birds: a correction

We provide a revision to the calculation of effect sizes and heterogeneity statistics in our original article, 'Facultative primary sex ratio variation: a lack of evidence in birds' (Ewen et al. 2004). Our revision shows that significant heterogeneity in sex ratio study effect sizes does indeed exist and that for a series of key traits the average effect sizes (while still weak) are in fact significantly different from zero.     

Silvicultural intensification for tropical forest conservation

Minimizing the deleterious environmental impacts of logging and other silvicultural treatments is the primary conservation goal in tropical forests managed for timber production. While it is always environmentally beneficial to minimize unnecessary damage, more intensive silviculture should not be discouraged in tropical forests in which regeneration and growth of commercially valuable timber species requires such treatments. Failing to regenerate commercial species may render forests more susceptible to conversion to other, more lucrative land uses. Increasing the intensity of silviculture may also decrease the total area of forest exploited for timber, thereby reducing the impacts of over-hunting, timber theft, wildfires, colonization, and conversion, which are facilitated by the increased accessibility of logged areas.