Community structure and diversity of tropical forest mammals: data from a global camera trap network

Terrestrial mammals are a key component of tropical forest communities as indicators of ecosystem health and providers of important ecosystem services. However, there is little quantitative information about how they change with local, regional and global threats. In this paper, the first standardized pantropical forest terrestrial mammal community study, we examine several aspects of terrestrial mammal species and community diversity (species richness, species diversity, evenness, dominance, functional diversity and community structure) at seven sites around the globe using a single standardized camera trapping methodology approach. The sites-located in Uganda, Tanzania, Indonesia, Lao PDR, Suriname, Brazil and Costa Rica-are surrounded by different landscape configurations, from continuous forests to highly fragmented forests. We obtained more than 51 000 images and detected 105 species of mammals with a total sampling effort of 12 687 camera trap days. We find that mammal communities from highly fragmented sites have lower species richness, species diversity, functional diversity and higher dominance when compared with sites in partially fragmented and continuous forest. We emphasize the importance of standardized camera trapping approaches for obtaining baselines for monitoring forest mammal communities so as to adequately understand the effect of global, regional and local threats and appropriately inform conservation actions.     

Camera trapping mammals in the scrubland’s of the Cape Floristic Kingdom—the importance of effort,spacing and trap placement

As a non-invasive monitoring method camera traps are noted as being an effective, accurate and rapid means of compiling species richness estimates of medium to large terrestrial mammals. However, crucial elements of camera trap survey design are rarely empirically addressed, which has raised the need for both a standardised and optimised camera trapping protocol. Our study confirms that an appropriate camera placement buffer and targeting areas of animal activity, contributes to more complete species richness estimates as well as significantly reducing the rate of false trigger events. However, attaining the required survey effort in terms of camera days was the most important factor in providing accurate species richness estimates. Our results suggest that reliable estimates of species richness can be achieved in open scrubland when cameras are spaced 1 × 1 km apart and left in the targeted area until a survey effort of a 1000 camera days is realised.     

Optimising sampling methods for small mammal communities in Neotropical rainforests

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Seasonal Change of Species Diversity Patterns of Non‐volant Small Mammals along Three Subtropical Elevational Gradients

Our understanding of geographic patterns of species diversity and the underlying mechanisms is increasing rapidly, whereas the temporal variation in these patterns remains poorly understood. We examined the seasonal species richness and species turnover patterns of non‐volant small mammals along three subtropical elevational gradients in southwest China. Small mammal diversity was surveyed in two seasons (early wet season and late wet season) using a standardized sampling protocol. The comparison of species richness patterns between two seasons indicated a temporal component in magnitude and shape, with species richness at high elevations clearly increased during the late wet season. Species richness demonstrated weak correlations with modelled temperature and precipitation. The elevational pattern of species turnover measured by Chao‐Sørenson similarity index also changed seasonally, even though the temporal pattern varied with scale. Species turnover between neighboring elevations at high elevations was slower in the late wet season. Meanwhile, there was an acceleration of species turnover along the whole range of the gradient. The seasonal change in species diversity patterns may be due to population‐level increases in abundance and elevational migration, whereas seasonal variation in factors other than temperature and precipitation may play a greater role in driving seasonal diversity patterns. Our study strongly supports the seasonality in elevational patterns of small mammal diversity in subtropical montane forests. Thus it is recommended that subsequent field surveys consider temporal sampling replicate for elevational diversity studies.     

Estimating Species Richness and Modelling Habitat Preferences of Tropical Forest Mammals from Camera Trap Data

Medium-to-large mammals within tropical forests represent a rich and functionally diversified component of this biome; however, they continue to be threatened by hunting and habitat loss. Assessing these communities implies studying species’ richness and composition, and determining a state variable of species abundance in order to infer changes in species distribution and habitat associations. The Tropical Ecology, Assessment and Monitoring (TEAM) network fills a chronic gap in standardized data collection by implementing a systematic monitoring framework of biodiversity, including mammal communities, across several sites. In this study, we used TEAM camera trap data collected in the Udzungwa Mountains of Tanzania, an area of exceptional importance for mammal diversity, to propose an example of a baseline assessment of species’ occupancy. We used 60 camera trap locations and cumulated 1,818 camera days in 2009. Sampling yielded 10,647 images of 26 species of mammals. We estimated that a minimum of 32 species are in fact present, matching available knowledge from other sources. Estimated species richness at camera sites did not vary with a suite of habitat covariates derived from remote sensing, however the detection probability varied with functional guilds, with herbivores being more detectable than other guilds. Species-specific occupancy modelling revealed novel ecological knowledge for the 11 most detected species, highlighting patterns such as ‘montane forest dwellers’, e.g. the endemic Sanje mangabey (Cercocebus sanjei), and ‘lowland forest dwellers’, e.g. suni antelope (Neotragus moschatus). Our results show that the analysis of camera trap data with account for imperfect detection can provide a solid ecological assessment of mammal communities that can be systematically replicated across sites.     

Camera trapping surveys of forest mammal communities in the Eastern Arc Mountains reveal generalized habitat and human disturbance responses

Large-bodied mammals are a rich and diversified faunal group in tropical rainforests. However, knowledge on community size and composition, and on species’ distribution and ecology remains often scant and inadequate against their chronic status of threats. We used camera trapping to detect mammals in the forests of the Eastern Arc Mountains (EAM) of Tanzania, a world renowned region for biodiversity comprised by a series of distinct and ancient mountain ranges partially covered in moist montane forest. We conducted surveys from 2003 to 2011 in eight of the 12 mountain blocks in Tanzania, and, through an overall sampling effort of 11,500 camera days, we detected 43 species. We normalized species richness and species’ detection events by effort, and used these metrics to assess the effect of habitat and human disturbance variables. We found that rarefied richness is positively affected by forest area at the block level, and that richness at forest patch level is also affected by forest area as well as surrounding human density (negative effect). For a subset of 17 species, we found consistent patterns of avoidance or tolerance of human disturbance and forest edges, and increased occurrence in areas at higher elevation, matching the historical forest loss that in most mountains occurred at lower elevation. Our study provides ecological insights that are novel for most species and sites, and reveals a general trend of negative impact of human disturbance on both community size and species’ relative abundance. Increased protection of the EAM forests in Tanzania is of urgent importance for the persistence of diversified mammal communities.     

The influence of trap density and sampling duration on the detection of small mammal species richness

Assessing species richness of small mammal communities is an important research objective for many live-trapping studies designed to assess or monitor biological diversity. We tested the effectiveness and efficiency of various trap densities for determining estimates and counts of small mammal species richness. Trapping was conducted in grassland habitats in northeastern Kansas during spring and fall of 2002 and 2003. Estimates and counts of species richness were higher at increased trap densities. This effect appeared to be primarily due to the higher number of individuals sampled at higher trap densities. At least 3 nights duration was needed to produce a stable estimate of species richness for the range of trap densities tested (9–144 trap stations/ha). Higher trap densities generally reached stable richness estimates in fewer nights than low density trapping arrangements. Given that counts and estimates of species richness were influenced by trap density and sampling duration, it is critical that these parameters are selected to most effectively meet research objectives.     

The efficacy of performing field surveys during the dry and rainy seasons to estimate species richness and composition of local bat assemblages

Here we assessed the efficacy of performing field surveys during both dry and rainy seasons to estimate species richness and composition of phyllostomid bats in 12 sites of the Brazilian Atlantic Forest. Our goal was to elucidate two opposite scenarios. If bat species compositions differ between the dry and rainy seasons, then field surveys in both seasons will be considered complementary for estimating local bat assemblages. In contrast, if bat species compositions in the dry season are a nested subset of the species present in the rainy season, then field surveys in both seasons will be considered redundant. Our results showed that field surveys performed only during the rainy season exhibited the same efficacy in recording bat species as field surveys considering both dry and rainy seasons combined. Consequently, the rare species sampled in the rainy season contributed to the uniqueness of communities in terms of species composition. Considering that time, funding, and the number of field personnel available for work are important factors when determining the sampling effort in highly biodiverse regions like the tropics, our results can be useful in establishing protocols for research projects with limited resources aiming to create a comprehensive list of species.     

Effects of peat swamp logging and agricultural expansion on species richness of native mammals in Peninsular Malaysia

The biodiversity inhabiting tropical peat swamp forests in Southeast Asia is currently threatened by commercial logging and agricultural expansion. The occurrence of mammals in such forests is often poorly known and the factors influencing their occurrence in these ecosystems have rarely been quantified. We aim to determine the key habitat and landscape drivers of mammal species richness in fragmented peat swamp reserves. We conducted camera trap surveys in the North Selangor Peat Swamp Forest (NSPSF), the last remaining area of peat swamp forest on the west coast of Peninsular Malaysia. We also measured vegetation structure and landscape metrics to investigate the relationship between these factors and mammal richness. We recorded a total of 16 mammal species from 45 sampling sites using camera traps located in peat swamp forest reserves. Mammal species richness increased with the abundance of large trees and distance away from roads. Species richness decreased significantly with canopy cover and height, the abundance of fallen trees, the abundance of forest palms and saplings, distance away from rivers, and a measure of landscape compositional heterogeneity. Our findings underscore the high conservation value of logged peat swamp forests and the urgent need to halt further deforestation. We recommend: (1) protecting riparian habitat; (2) avoiding further forest conversion particularly areas supporting large trees into oil palm plantations; and (3) limiting road development within and around the NSPSF.     

Revealing kleptoparasitic and predatory tendencies in an African mammal community using camera traps: a comparison of spatiotemporal approaches

Camera trap data are increasingly being used to characterise relationships between the spatiotemporal activity patterns of sympatric mammal species, often with a view to inferring inter‐specific interactions. In this context, we attempted to characterise the kleptoparasitic and predatory tendencies of spotted hyaenas Crocuta crocuta and lions Panthera leo from photographic data collected across 54 camera trap stations and two dry seasons in Tanzania's Ruaha National Park. We applied four different methods of quantifying spatiotemporal associations, including one strictly temporal approach (activity pattern overlap), one strictly spatial approach (co‐occupancy modelling), and two spatiotemporal approaches (co‐detection modelling and temporal spacing at shared camera trap sites). We expected a kleptoparasitic relationship between spotted hyaenas and lions to result in a positive spatiotemporal association, and further hypothesised that the association between lions and their favourite prey in Ruaha, the giraffe Giraffa camelopardalis and the zebra Equus quagga, would be stronger than those observed with non‐preferred prey species (the impala Aepyceros melampus and the dikdik Madoqua kirkii). Only approaches incorporating both the temporal and spatial components of camera trap data resulted in significant associative patterns. The latter were particularly sensitive to the temporal resolution chosen to define species detections (i.e. occasion length), and only revealed a significant positive association between lion and spotted hyaena detections, as well as a tendency for both species to follow each other at camera trap sites, during the dry season of 2013, but not that of 2014. In both seasons, observed spatiotemporal associations between lions and each of the four herbivore species considered provided no convincing or consistent indications of any predatory preferences. Our study suggests that, when making inferences on inter‐specific interactions from camera trap data, due regards should be given to the potential behavioural and methodological processes underlying observed spatiotemporal patterns.     

基于红外相机技术对广东车八岭国家级自然保护区大中型兽类与雉类的编目清查与评估

近年来, 红外相机技术已被广泛应用于国内外自然保护地内地栖鸟兽的物种编目和动态评估。本文以广东车八岭国家级自然保护区为例, 探讨基于红外相机技术如何进行保护区全境大中型兽类和雉鸡类的物种编目清查与评估。通过对车八岭保护区全境为期1年的调查, 共记录兽类和雉鸡类18种, 其中兽类15种, 鸡形目鸟类3种。基于物种累计曲线, 采用全年数据所需的最小调查网格数、最少调查相机日均要少于雨季或旱季, 而旱季调查需要的最小调查网格数和最少调查相机日比雨季更少。通过红外相机图像数据获得了车八岭保护区的大中型兽类和地栖雉鸡类物种名录、物种丰富度、每个物种的相对多度、分布图和凭证标本等重要内容。     

Species reordering,not changes in richness,drives long‐term dynamics in grassland communities

Determining how ecological communities will respond to global environmental change remains a challenging research problem. Recent meta‐analyses concluded that most communities are undergoing compositional change despite no net change in local species richness. We explored how species richness and composition of co‐occurring plant, grasshopper, breeding bird and small mammal communities in arid and mesic grasslands changed in response to increasing aridity and fire frequency. In the arid system, grassland and shrubland plant and breeding bird communities were undergoing directional change, whereas grasshopper and small mammal communities were stable. In the mesic system, all communities were undergoing directional change regardless of fire frequency. Despite directional change in composition in some communities, species richness of all communities did not change because compositional change resulted more from reordering of species abundances than turnover in species composition. Thus, species reordering, not changes in richness, explains long‐term dynamics in these grass and shrub dominated communities.     

Temporal Patterns of Species Accumulation in a Survey of Lepidoptera in a Beech-Maple Forest

One of the most significant challenges to insect conservation is lack of information concerning species diversity and distribution. Because a complete inventory of all species in an area is virtually impossible, interest has turned to developing statistical techniques to guide sampling design and to estimate total species richness within a site. We used two such techniques, diversity partitioning and non-parametric richness estimation, to determine how variation in sampling effort over time affected species accumulation for a survey of Lepidoptera in an old-growth beech-maple forest. Temporal scaling of sampling effort had significant effects on two measures of species diversity. Increases in species richness were primarily driven by changes in species occurrences with season, while Shannon diversity was largely determined at the scale of individual sampling units (i.e. by spatial effects). Variation in sampling effort affected the values of the two most widely regarded richness estimators (ICE and Chao 2); neither diversity estimator achieved stable values across a range of sampling efforts. Even after 52 trap-nights and accounting for seasonality, rare species (singletons and uniques) remained a significant component of the moth community. To the extent that moth communities in other forest systems are similarly comprised of many rare species, non-parametric richness estimators should be expected to yield variable estimates with increased effort and should only be used to provide a minimum benchmark for predicting the number of species remaining to be sampled. Our results suggest the best strategy for a short-term survey of forest Lepidoptera should emphasize spreading sampling intervals throughout a given year rather than focusing on intensive sampling during a short time period or prolonged sampling over many years.     

Carrion fly‐derived DNA metabarcoding is an effective tool for mammal surveys: Evidence from a known tropical mammal community

Metabarcoding of vertebrate DNA derived from carrion flies has been proposed as a promising tool for biodiversity monitoring. To evaluate its efficacy, we conducted metabarcoding surveys of carrion flies on Barro Colorado Island (BCI), Panama, which has a well‐known mammal community, and compared our results against diurnal transect counts and camera trapping. We collected 1,084 flies in 29 sampling days, conducted metabarcoding with mammal‐specific (16S) and vertebrate‐specific (12S) primers, and sequenced amplicons on Illumina MiSeq. For taxonomic assignment, we compared blast with the new program protax , and we found that protax improved species identifications. We detected 20 mammal, four bird, and one lizard species from carrion fly metabarcoding, all but one of which are known from BCI. Fly metabarcoding detected more mammal species than concurrent transect counts (29 sampling days, 13 species) and concurrent camera trapping (84 sampling days, 17 species), and detected 67% of the number of mammal species documented by 8 years of transect counts and camera trapping combined, although fly metabarcoding missed several abundant species. This study demonstrates that carrion fly metabarcoding is a powerful tool for mammal biodiversity surveys and has the potential to detect a broader range of species than more commonly used methods.     

Temporal Dynamics of Rich Moth Ensembles in the Montane Forest Zone in Southern Ecuador1

We studied temporal dynamics of diverse moth ensembles (Arctiidae and Geometridae) in early and late succession stages of forest recovery in the montane zone of southern Ecuador. Moths were sampled using weak light sources (2 × 15 W tubes per trap) during three sampling periods (March–April 2002, wet season; October–November 2002, and August–October 2003, both “dry” seasons). Arctiid moth abundance hardly varied between sampling periods. Estimates of local diversity were lower in 2002 (wet and dry season) than in the dry season 2003, and ensemble composition was more strongly affected by sampling period rather than habitat differences. Geometridae ensembles revealed stronger temporal patterns. Geometrid abundance increased about twofold from the wet to the dry season, and temporal effects on species composition were far more pronounced that in arctiids. These differences might hint to variation in the dependence of geometrid versus arctiid larvae on ephemeral plant resources. Despite these significant temporal dynamics, in both families only a few of the more common species analyzed individually showed strong temporal changes in abundance. Almost all common species occurred as adults during all sampling periods. Therefore, even though temporal dynamics of moth ensembles are not negligible, both moth families are suitable “indicators” of community diversity and change along the succession gradient. Samples must be large enough, however, and preferably should be collated over various times of the year and in parallel, to allow for valid statements about moth diversity and species compositions in relation to habitat differences. These recommendations undermine the validity of the concept of “rapid biodiversity inventories” for speciose tropical insect communities.     

A preliminary investigation of temporal patterns in semiarid ant communities: Variation with habitat type

Abstract This study reports on preliminary findings of habitat‐contingent temporal variability in ant assemblages in Purnululu National Park in northern Australia's semiarid tropics, by sampling at the end of the dry season (October 2004) and the end of the wet season (April 2005). Six grids of 15 pitfall traps were established in each of the spinifex, sandplain and gorge habitats. Community composition was dominated by behaviourally dominant ants (Iridomyrmex spp.) and climate specialists (Melophorus and Meranoplus spp.). Ant activity was higher in the wet season sampling period, with greater species richness and abundance. Interestingly, temporal variation in ant assemblage richness, abundance and composition varied markedly with habitat type. While there were large differences between sampling periods for the spinifex and sandplain habitat, this was not the case in the gorges. These temporal changes in ant assemblages are postulated to be linked with major environmental differences between the two sampling periods, driven by seasonal climatic conditions. It is likely that these changes influenced the ant assemblages through species differences in physiological tolerance levels, ecological requirements and competitive ability. This study demonstrates the need, in highly seasonal environments, to consider the temporal context of studies in relation to habitat type, particularly when undertaking biodiversity surveys and monitoring.     

Effects of alien pine plantations on small mammal community structure in a southern African biodiversity hotspot

Commercial plantations and alien tree invasions often have substantial negative impacts on local biodiversity. The effect of plantations on faunal communities in the fire‐adapted fynbos vegetation of the Cape Floristic Region biodiversity hotspot is not yet well quantified. We studied small mammal community structure in alien Pinus radiata plantations and adjacent fynbos regenerating after clear‐felling of plantations on the Cape Peninsula, South Africa. Small mammal sampling over 1,800 trap‐nights resulted in 480 captures of 345 individuals (excluding recaptures) representing six species. Significantly more species, individuals (12 X) and biomasses (29 X) of small mammals occurred on recovering fynbos sites compared to plantations. This was commensurate with a higher diversity of plant growth forms, vegetation densities and live vegetation biomass. Only one small mammal species, the pygmy mouse (Mus minutoides), was consistently trapped within plantations. Fynbos sites were dominated by three small mammal species that are ecological generalists and early successional pioneer species, rendering the recovering fynbos slightly depauperate in terms of species richness and evenness relative to other studies done in pristine fynbos. We make three recommendations for forestry that would facilitate the restoration of more diverse natural plant communities and progressively more diverse and dynamic small mammal assemblages in a key biodiversity hotspot.     

Effects of sample standardization on mean species detectabilities and estimates of relative differences in species richness among assemblages

Ecological surveys provide the basic information needed to estimate differences in species richness among assemblages. Comparable estimates of the differences in richness between assemblages require equal mean species detectabilities across assemblages. However, mean species detectabilities are often unknown, typically low, and potentially different from one assemblage to another. As a result, inferences regarding differences in species richness among assemblages can be biased. We evaluated how well three methods used to produce comparable estimates of species richness achieved equal mean species detectabilities across diverse assemblages: rarefaction, statistical estimators, and standardization of sampling effort on mean taxonomic similarity among replicate samples (MRS). We used simulated assemblages to mimic a wide range of species-occurrence distributions and species richness to compare the performance of these three methods. Inferences regarding differences in species richness based on rarefaction were highly biased when richness estimates were compared among assemblages with distinctly different species-occurrence distributions. Statistical estimators only marginally reduced this bias. Standardization on MRS yielded the most comparable estimates of differences in species richness. These findings have important implications for our understanding of species-richness patterns, inferences drawn from biological monitoring data, and planning for biodiversity conservation.     

Tiger-Moths in Savannas in Eastern Amazon: First Assessment of Diversity and Seasonal Aspects

Biodiversity knowledge on insects is urgently needed due to the ever growing demand for food and the consequent deforestation process and loss of natural habitats in many understudied tropical regions. In this paper, we describe the outcome of a biodiversity research on tiger moths performed for the first time in a poorly studied Amazonian landscape—the savanna. We sampled tiger moths monthly with UV automatic light traps for 12 consecutive months in two sampling points in an area of savanna in eastern Amazon, and we compared our results to previously available data for eastern Amazon. We found a total of 91 species of which 80 were identified to species level. The most species-rich subtribes were Phaegopterina and Euchromiina with 32 species each. Species richness and abundance did not differ among sampling sites, but in general the species richness was higher during the dry season while abundance was higher during the wet season. This seasonal diversity pattern differs from the most common patterns recorded for savannas in other parts of the world. The species composition also changed in wet and dry seasons and correlated significantly with temperature and relative humidity. Our results suggest that the alpha diversity of the Amazonian savannas in our sampling area is lower than that in nearby rain forests and similar to that in agriculturally disturbed areas surrounded by rain forests. However, the species composition differed considerably from natural and disturbed areas. These results highlight the need of basic biodiversity surveys of insects in Amazonian savannas.