FOOD SHORTAGE IN SMALL FRAGMENTS: EVIDENCE FROM AN AREA-SENSITIVE PASSERINE

Summary 1. Many songbird species are absent from small forest fragments. One possible explanation for this finding is that populations in small fragments decline to extinction because reproductive success is too low to compensate for adult mortality (λ < 1). I tested this hypothesis from 1995 to 1997 by measuring reproductive success, adult survival, and population viability for four banded populations of the Eastern yellow robin (Eopsaltria australis, White), an area‐sensitive songbird. The populations inhabited two small (55 ha, termed S1, S2) and two large (500–1000 ha, termed L1, L2) forest fragments set within an agricultural landscape. 2. Females produced more fledglings but a similar number of independent young per capita in the small compared to the large fragments. Local female survival did not vary with fragment size. 3. Population models were used to calculate the finite rate of increase (λ) from the number of young produced (female yearlings per female per year) and adult female survival rates. Combining data from all four populations, λ = 0·96, indicating that robins produced 4% fewer female young per year than were needed to compensate for adult losses. Combining data from the two small fragments, λ = 1·02, suggesting that populations in small fragments were viable on average. In contrast, λ = 0·85 for the two large fragments combined, indicating that, on average, 15% fewer young were produced than were needed to replace missing adults each year. Averaging across fragments within the same size class masked the variability in population dynamics that existed between the replicates. Considering each of the four fragments separately, there was one potential demographic source (S2, λ = 1·12; L2, λ = 1·23) and one demographic sink (S1, λ = 0·90; L1, λ = 0·46) within each of the fragment size categories. 4. Observed changes in population sizes were similar to those predicted by the population models. Combining all populations, recruitment replaced most but not all missing females, and the observed population size declined by 8% per year. In both the small and large fragments, observed recruitment was insufficient to replace missing females and population sizes declined 5% and 8% per year, respectively. Within each of the fragment size categories, there was one stable population (S2 and L2) and one population that declined in size (S1 and L1, respectively, declined by 14% and 20% per year). 5. Reproductive success, adult survival, and population viability did not vary with fragment size. To explain area‐sensitivity, I suggest that a threshold in fragment size exists below which populations are never viable, but above which populations may or may not be viable depending on factors unrelated to fragment size, such as nest predation rates.

Area sensitivity in songbirds is commonly attributed to increased nest predation in forest fragments. In 1995 and 1996, we tested whether the nest predators and nesting success of an area-sensitive forest bird, the Eastern Yellow Robin (Eopsaltria australis), varied with fragment size, and we also conducted an artificial nest experiment. The study occurred in two small (55 ha) and two large (>400 ha) forest fragments in a matrix of agricultural land in New South Wales, Australia. Predation accounted for 95% of all failures of 282 robin nests, and the survival of robin nests was negatively correlated with how frequently we observed avian nest predators near nests (i.e. nest-predator activity). Of 461 artificial nests, 84% were depredated, nearly all (99%) by birds. Thus, birds were important predators of nests. The abundance, species richness, and activity of avian nest predators were not related to fragment size. Survival of robin nests averaged 19%; nests in small fragments had a 22% chance of producing at least one fledgling compared with 15% in large fragments, but the difference was not significant. Survival of artificial nests averaged 12% in both small and large fragments. Nest-predator activity accounted for the most variation (68%) in the fate of robin nests, followed by the cumulative density of open-cup nesters (16%). The placement of robin nests had no influence on nest fate. We conclude that nest predation was not area dependent and propose food supply as an alternative hypothesis to explain area sensitivity. We suggest that, rather than being related to fragment size, nest predation increases with decreasing forest cover in a landscape. Increased nest predation in fragmented compared with contiguous landscapes may lower the population viability of songbirds in a region, and hence regional numbers. Therefore, the spatial scale at which fragmentation influences nest predation and songbird populations must be considered carefully.

Large habitat fragments are generally thought to host more species and to offer more diverse and/or better quality habitats than small fragments. However, the importance of small fragments for population dynamics in general and for reproductive performance in particular is highly controversial. Using an information-theoretic approach, we examined reproductive performance and probability of local recruitment of color-banded reed buntings Emberiza schoeniclus in relation to the size of 18 wetland fragments in northeastern Switzerland over 4 years. We also investigated if reproductive performance and recruitment probability were density-dependent. None of the four measures of reproductive performance (laying date, nest failure probability, fledgling production per territory, fledgling condition) nor recruitment probability were found to be related to wetland fragment size. In terms of fledgling production, however, fragment size interacted with year, indicating that small fragments were better reproductive grounds in some years than large fragments. Reproductive performance and recruitment probability were not density-dependent. Our results suggest that small fragments are equally suited as breeding grounds for the reed bunting as large fragments and should therefore be managed to provide a habitat for this and other specialists occurring in the same habitat. Moreover, large fragments may represent sinks in specific years because a substantial percentage of all breeding pairs in our study area breed in large fragments, and reproductive failure in these fragments due to the regularly occurring floods may have a much stronger impact on regional population dynamics than comparable events in small fragments.

Habitat fragmentation poses a significant threat to primate populations, necessitating behavioral adaptations in activity patterns and feeding strategies. This study investigated the impact of habitat fragmentation on the endangered San Martin titi monkey (Plecturocebus oenanthe) in the Moyobamba region of Peru. Activity budgets and feeding strategies were compared between groups inhabiting large (400 ha) and small (5 ha) forest fragments. Results revealed that resting was the predominant activity in both habitats, followed by feeding. Notably, individuals in large fragments allocated significantly more time to feeding (30.1%) than those in small fragments (23.0%) did. Social interactions were more prevalent in the small fragments (15.1%) than in the large fragments (9.1%). Fruit was the primary dietary component in both sites, accounting for 59.9% in the large fragment and 44.9% in the small fragment. However, individuals in small fragments exhibited a higher consumption of leaves (13.7%) and prey (14.6%) than those in large fragments (7.1% and 8.5%, respectively). The feeding time for specific food resources varied according to the fragment size and seasonal availability. These findings demonstrate the ecological plasticity of P. oenanthe in adapting to fragmented habitats by modifying their activity budgets and feeding strategies. However, continued habitat degradation may compromise these adaptive mechanisms, highlighting the need for conservation efforts focused on habitat restoration and connectivity to ensure long-term persistence of this endangered primate species.

Seed dispersal is a key process driving the structure, composition, and regeneration of tropical forests. Larger frugivores play a crucial role in community structuring by dispersing large seeds not dispersed by smaller frugivores. We assessed the hypothesis that brown howler monkeys (Alouatta guariba clamitans) provide seed dispersal services for a wide assemblage of plant species in both small and large Atlantic forest fragments. Although fruit availability often decreases in small fragments compared with large ones, we predicted that brown howlers are efficient seed dispersers in quantitative and qualitative terms in both forest types given their high dietary flexibility. After a 36-month study period and 2,962 sampling hours, we found that howlers swallowed and defecated intact the vast majority of seeds (96%-100%) they handled in all study sites. Overall, they defecated ca. 315,600 seeds belonging to 98 species distributed in eight growth forms. We estimated that each individual howler dispersed an average of 143 (SD = 49) seeds >2 mm per day or 52,052 (SD = 17,782) seeds per year. They dispersed seeds of 58% to 93% of the local assemblages of fleshy-fruit trees. In most cases, the richness and abundance of seed species dispersed was similar between small and large fragments. However, groups inhabiting small fragments tended to disperse a higher diversity of seeds from rarely consumed fruits than those living in large fragments. We conclude that brown howlers are legitimate seed dispersers for most fleshy-fruit species of the angiosperm assemblages of their habitats, and that they might favor the regeneration of Atlantic forest fragments with the plentiful amount of intact seeds that they disperse each year.

Primates display varying degrees of behavioral flexibility that allow them to adjust their diet to temporal changes in food availability. This trait might be critical for the survival of folivorous-frugivorous species inhabiting small forest fragments, where the availability of food resources tends to be lower than in large fragments and continuous forests. However, the scarcity of studies addressing this issue hampers our understanding of the adaptive behaviors that favor the survival of these primates in low-quality habitats. We conducted a 36-mo study testing the hypothesis that brown howler monkeys (Alouatta guariba clamitans) are able to adjust their diet in response to local and seasonal changes in resource availability. We compared the diet of six free-ranging groups inhabiting three small (<10 ha) and three large (>90 ha) Atlantic forest fragments in southern Brazil and estimated the temporal availability of their top food species (i.e., those species that together contribute ≥80% of total feeding records). We found that brown howlers exploited similarly rich diets in small (45, 54, and 57 plant species) and large (48, 51, and 56 species) fragments. However, intermonth diet similarity was higher for groups in small fragments, where howlers also fed on plant items from nine alien species. Fruits and leaves were the most consumed plant items in both small (42% and 49% of feeding records, respectively) and large (51% and 41%, respectively) fragments. The consumption of young leaves was higher in small than in large fragments, whereas the consumption of other plant items did not show a pattern related to fragment size. Regarding the contribution of growth forms as food sources, only the exploitation of palms showed a pattern related to fragment size. Palms contributed more to the diet of groups inhabiting large fragments. The availability of seasonal food items–ripe fruits and young leaves–influenced their consumption in both habitat types. Therefore, brown howlers cope with local and seasonal fluctuations in food availability by opportunistically exploiting resources. We believe that this feeding flexibility is a key component of the phenotypic plasticity that enables howlers to thrive in disturbed habitat patches, where periods of scarcity of preferred foods shall be more common.

The effect of the spatial scale of recruitment on tree diversity in Afromontane forest fragments

Food limitation may interact with nest predation and influence nesting patterns, such as breeding season length and renesting intervals. If so, reproductive effort should change with food availability. Thus, when food is limited, birds should have fewer attempts and shorter seasons than when food is not limiting. Here we experimentally test that increased food availability results in increased reproductive effort in a fragmented landscape in the Variable Antshrike (Thamnophilus caerulescens) in southern Brazil. We followed nesting pairs in five natural fragments (4, 23, 24, 112, 214 ha) in which food was supplemented for half of those pairs, beginning with the first nest. Nest success in the largest (214 ha) fragment was 59%, compared to 5% in the 112 ha fragment and no nest was successful in the smallest (24 ha) fragment. Birds were seen, but evidence of nesting was never found in the two smallest fragments. Pairs with supplemented food were more likely to increase clutch size from two to three eggs, tended to renest sooner (20 d on average) than control pairs. Also, fragment size interacted with breeding and pairs in the largest fragment had greater daily nest survival rates, and so nests tended to last longer, and so these pairs had fewer nesting attempts than those in the 112 ha fragment while more than those in the smallest fragment with nesting (24 ha). Clearly, pairs increased their reproductive effort when food was supplemented in comparison to control pairs and fragment size seems to influence both predation risk and food abundance.

This study characterized rainforest economic tree species composition and indices of diversity in Delta State, South-southern Nigeria: Implications for biodiversity conservation. The study area was stratified into 25 divisions based on the existing Local Government Areas (LGA). From each LGA, one larger fragment (&gt;15,000m²) and one smaller (≤15,000m²) fragment were selected, making a total of 50 fragments from which data were collected on the size of rainforest fragment, population of individual tree species, and the species of trees of the rainforest origin using standard approaches in quadrats of 15m×15m. Data analysis involved the descriptive and inferential statistics, as well as Simpson’s index of tree species diversity. Results revealed that larger fragments have higher population of trees than smaller fragments. The mean population values of tree species were 306.40 and 80.88 for the larger and smaller fragments; with paired mean, standard deviation, and standard error of mean values of 225.52, 103.89 and ±20.78 respectively. With P (0.000) &lt; 0.05, and t-value of 10.85, the observed mean differences in the population of trees between the larger and smaller rainforest fragments was significant at 0.05 level of confidence. The mean population density values of 0.013 and 0.006 for the larger and smaller fragments were observed; with paired mean, standard deviation, and standard error of mean values of 0.0069, 0.003 and ±0.0005 respectively. With P (0.000) &lt; 0.05, and t-value of 14.11, the observed mean differences in the population density of trees between the larger and smaller rainforest fragments was significant at 0.05 level of confidence. Piptadeniastrum africanum was the most dominant tree species in the study area. Effective biodiversity conservation is recommended to prevent the extinction of rainforest tree species of economic importance in the study area.

Agricultural intensification has been shown to reduce biodiversity through processes such as habitat degradation and fragmentation. We tested whether several small or single large habitat fragments (re-visiting the 'single large or several small' debate) support more species across a wide range of taxonomic groups (plants, leafhoppers, true bugs, snails). Our study comprised 14 small (<1 ha) and 14 large (1.5-8 ha) fragments of calcareous grassland in Central Germany along orthogonal gradients of landscape complexity and habitat connectivity. Each taxon was sampled on six plots per fragment. Across taxa, species richness did not differ between large and small fragments, whereas species-area accumulation curves showed that both overall and specialist species richness was much higher on several small fragments of calcareous grassland than on few large fragments. On average, 85% of the overall species richness was recorded on all small fragments taken together (4.6 ha), whereas the two largest ones (15.1 ha) only accounted for 37% of the species. This could be due to the greater geographic extent covered by many small fragments. However, community composition differed strongly between large and small fragments, and some of the rarest specialist species appeared to be confined to large fragments. The surrounding landscape did not show any consistent effects on species richness and community composition. Our results show that both single large and many small fragments are needed to promote landscape-wide biodiversity across taxa. We therefore question the focus on large fragments only and call for a new diversified habitat fragmentation strategy for biodiversity conservation.

Tropical forests are seriously threatened by fragmentation and habitat loss. The impact of fragment size and forest configuration on the composition of seed rain is insufficiently studied. For the present study, seed rain composition of small and large forest fragments (8-388 ha) was assessed in order to identify variations in seed abundance, species richness, seed size and dispersal mode. Seed rain was documented during a 1-year period in three large and four small Atlantic Forest fragments that are isolated by a sugarcane matrix. Total seed rain included 20,518 seeds of 149 species of trees, shrubs, palms, lianas and herbs. Most species and seeds were animal-dispersed. A significant difference in the proportion of seeds and species within different categories of seed size was found between small and large fragments. Small fragments received significantly more very small-sized seeds (<0.3 cm) and less large-seeded species (>1.5 cm) that were generally very rare, with only one species in small and eight in large fragments. We found a negative correlation between the inflow of small-sized seeds and the percentage of forest cover. Species richness was lower in small than in large fragments, but the difference was not very pronounced. Given our results, we propose changing plant species pools through logging, tree mortality and a high inflow of pioneer species and lianas, especially in small forest fragments and areas with low forest cover. Connecting forest fragments through corridors and reforestation with local large-seeded tree species may facilitate the maintenance of species diversity.

Abstract: For some species of forest‐breeding birds, productivity is much higher in large forest fragments than in small forest fragments and is higher than the level needed to replace individuals within the fragment (sources). Thus large forest fragments potentially provide excess individuals to neighboring small fragments where productivity is much lower and not adequate for replacement (sinks). We used occurrence data and distances between putative sources and sinks for four species of forest birds with this pattern of demography in large and small forest fragments to predict occupancy in small fragments. For the Ovenbird ( Seiurus aurocapilla), Wood Thrush ( Hylocichla mustelina), Veery (Catharus fuscescens), and Rose‐breasted Grosbeak (Pheucticus ludovicianus), distance from large woodlots was a significant predictor of occurrence in small woodlots. Distance from large woodlots did not predict occupancy of Red‐eyed Vireos ( Vireo olivaceus), a species with no apparent difference in productivity in relation to fragment size. Neither vegetation features nor area of the small woodlots adequately explained patterns of occupancy for any species. These results suggest that maintenance of large, productive forest fragments benefit regional occurrence of forest‐breeding birds by providing individuals to neighboring small woodlots. Limited reproduction in these small woodlots means that they may act as a reservoir for excess individuals in the case of catastrophic events that result in vacancies in source woodlots. Conservation and sound forest management of small forests should be directed particularly toward those closer to large regional forests.

The process of bone union after arthroscopic bony Bankart repair in younger athletes with a subcritical glenoid defect: An advantage of remained large bone fragment

Habitat loss in highly deforested landscapes such as the Brazilian Atlantic Forest has been severely affecting the diversity and survival of palm species. As some species are more sensitive than others, trait responses to the environment, as well as environmental effects on fecundity, growth, and mortality rates, may affect species demography. Considering this context, we studied functional and demographic responses of three palm species (Astrocaryum aculeatissimum, Euterpe edulis, and Geonoma schottiana) to habitat loss in the Atlantic Forest in southeastern Brazil by measuring morpho-physiological traits related to plant growth and light acquisition for photosynthesis. We also tested the response of population fitness to fragment size. Plant survival and growth was subsequently monitored in 2006 and 2007, and population dynamics were summarized in pool matrices for large and small forest fragments in the monitoring periods comprehending one full year between 2005–2006 and 2006–2007. The asymptotic growth rate of populations (defined here as population fitness, λ) in five forest fragments was then calculated. Diameter of individuals of the demography plots (from year 2005 to 2007) was used to calculate the relative diameter growth rate. Later, in 2015, we measured a set of morpho-physiological functional traits in palms in the same plots used in the demographic studies. While A. aculeatissimum populations were stable in both monitoring periods in small and large fragments, E. edulis populations were predicted to decline due to intense predation by monkeys in the large fragment, but were stable in the smaller fragments, and G. schottiana populations were stable in the large fragments in both monitoring periods, but populations in the smaller fragments were predicted to decline in the second period, i.e., with lower fitness in these fragments. In addition, the functional traits analyzed showed that G. schottiana is a forest interior species associated with the shade/understory environment response. E. edulis was also affected by the size of the fragment, but due to a disruptive interaction with a predator and showed intermediate functional traits values. On the other hand, A. aculeatissimum thrived in areas with higher and lower incidence of light and was not demographically affected by forest remnant size. This suggests that E. edulis and A. aculeatissimum are habitat generalists. We concluded that differences in the ecophysiological performance of palms due to distinct morpho-physiological functional traits related to leaf economic spectrum, such as LDMC or specific leaf area (SLA) and to photosynthetic responses to light environment as electron transport rate (ETR) and saturation irradiance (Ik) were linked to the demographic variation observed in forest remnants of different size.

To determine the minimum size requirements and influence of landscape context on reproductive success of forest-breeding songbirds, we monitored nesting success of five species of songbirds on 40 fragments (12–2350 ha in total woodlot size) and two continuous forest sites in south-central Ontario from 1994 through 1997. Woodlot size was the most important variable contributing to differences in reproductive success, with local forest cover (within a 10-km radius) having no significant additional effect on productivity for any species. For all species, except Red-eyed Vireo (Vireo olivaceous), adult female reproductive success was at or above replacement levels in large fragments (mean of 121 ha core area, 849 ha woodlot area) and continuous forest, and below replacement levels in small fragments (mean of 7.8 ha core area, 93 ha woodlot area). Red-eyed Vireo productivity was particularly low, with single-brooded females unable to maintain populations in any woodlots monitored, although populations were close to replacement levels in continuous forest. Only the very largest fragments acted as sources for Ovenbird (Seiurus aurocapillus) and Wood Thrush (Hylocichla mustelina) (i.e., >23 ha in core area, 225 ha in total woodlot area). Ovenbird and Wood Thrush were the most area-sensitive, and Rose-breasted Grosbeak (Pheucticus ludovicianus) and Veery (Catharus fuscens) the least, although all four species had significantly lower reproductive success in small as compared to large forest fragments. Reproductive success of Rose-breasted Grosbeaks and Veery were similar in large forest fragments and continuous forest and were close to maintenance levels in small fragments, despite published reports of significant declines that might be attributable to poor productivity. Nest predation was the major cause of nest failure, with brood parasitism by Brown-headed Cowbirds (Molothrus ater) further reducing the number of host young fledged from Ovenbird and Red-eyed Vireo nests. Parasitism rates on the remaining species were very low and not analyzed statistically. Local forest cover had little effect on predation rate or rate of parasitism by Brown-headed Cowbirds. Although local forest cover had little effect on reproductive success within fragmented landscapes, nest success was significantly higher in continuous forest than in all fragments combined for both Ovenbird and Red-eyed Vireo. Our data show a poor agreement with Breeding Bird Survey population trend estimates for Ontario between 1986 and 1996. As only 1% of the forest fragments locally available are large enough to function as source habitats, we strongly advocate their preservation. We recommend preservation of forest tracts at least 500 ha in size, particularly woodlots with >90 ha in core area, to function as sources that will help guard against population declines on a local scale.