Correction: Serodiagnosis of Tuberculosis in Asian Elephants (Elephas maximus) in Southern India: A Latent Class Analysis.

[This corrects the article DOI: 10.1371/journal.pone.0049548.].

Animal movement ecology in India: insights from 2011-2021 and prospective for the future.

The field of animal movement ecology has advanced by leaps and bounds in the past few decades with the advent of sophisticated technology, advanced analytical tools, and multiple frameworks and paradigms to address key ecological problems. Unlike the longer history and faster growth of the field in North America, Europe, and Africa, movement ecology in Asia has only recently been gaining momentum. Here, we provide a review of the field from studies based in India over the last 11 years (2011-2021) curated from the database, Scopus, and search engine, Google Scholar. We identify current directions in the research objectives, taxa studied, tracking technology and the biogeographic regions in which animals were tracked, considering the years since the last systematic review of movement ecology research in the country. As an indication of the growing interest in this field, there has been a rapid increase in the number of publications over the last decade. Class Mammalia continues to dominate the taxa tracked, with tiger and leopard being the most common species studied across publications. Invertebrates and other small and medium-sized animals, as well as aquatic animals, in comparison, are understudied and remain among the important target taxa for tracking in future studies. As in the previous three decades, researchers have focussed on characterising home ranges and habitat use of animals. There is, however, a notable shift to examine the movement decision of animals in human-modified landscapes, although efforts to use movement ecology to understand impacts of climate change remain missing. Given the biogeographic and taxonomic diversity of India, and the fact that the interface between anthropogenic activity and wildlife interactions is increasing, we suggest ways in which the field of movement ecology can be expanded to facilitate ecological insights and conservation efforts. With the advancement of affordable technologies and the availability of analytical tools, the potential to expand the field of movement ecology, shift research foci, and gain new insights is now prime.

Viewing the rare through public lenses: insights into dead calf carrying and other thanatological responses in Asian elephants using YouTube videos.

Documenting the behavioural repertoire of an animal species is important for understanding that species' natural history. Many behaviours such as mating, parturition and death may be observed only rarely in the wild due to the low frequency of occurrence, short duration and the species' elusiveness. Opportunistic documentation of rare behaviours is therefore valuable for deciphering the behavioural complexity in a species. In this context, digital platforms may serve as useful data sources for studying rare behaviours in animals. Using videos uploaded on YouTube, we document and construct a tentative repertoire of thanatological responses (death-related behaviours) in Asian elephants (Elephas maximus). The most frequently observed thanatological responses included postural changes, guarding/keeping vigil, touching, investigating the carcass, epimeletic behaviours and vocalizations. We also describe some infrequently observed behaviours, including carrying dead calves by adult females, re-assurance-like behaviours and attempts to support dying or dead conspecifics, some of which were only known anecdotally in Asian elephants. Our observations indicate the significance of open-source video data on digital platforms for gaining insights into rarely observed behaviours and support the accumulating evidence for higher cognitive abilities of Asian elephants in the context of comparative thanatology.

Distribution of biomass dynamics in relation to tree size in forests across the world.

Tree size shapes forest carbon dynamics and determines how trees interact with their environment, including a changing climate. Here, we conduct the first global analysis of among-site differences in how aboveground biomass stocks and fluxes are distributed with tree size. We analyzed repeat tree censuses from 25 large-scale (4-52 ha) forest plots spanning a broad climatic range over five continents to characterize how aboveground biomass, woody productivity, and woody mortality vary with tree diameter. We examined how the median, dispersion, and skewness of these size-related distributions vary with mean annual temperature and precipitation. In warmer forests, aboveground biomass, woody productivity, and woody mortality were more broadly distributed with respect to tree size. In warmer and wetter forests, aboveground biomass and woody productivity were more right skewed, with a long tail towards large trees. Small trees (1-10 cm diameter) contributed more to productivity and mortality than to biomass, highlighting the importance of including these trees in analyses of forest dynamics. Our findings provide an improved characterization of climate-driven forest differences in the size structure of aboveground biomass and dynamics of that biomass, as well as refined benchmarks for capturing climate influences in vegetation demographic models.

Demographic composition, not demographic diversity, predicts biomass and turnover across temperate and tropical forests.

The growth and survival of individual trees determine the physical structure of a forest with important consequences for forest function. However, given the diversity of tree species and forest biomes, quantifying the multitude of demographic strategies within and across forests and the way that they translate into forest structure and function remains a significant challenge. Here, we quantify the demographic rates of 1961 tree species from temperate and tropical forests and evaluate how demographic diversity (DD) and demographic composition (DC) differ across forests, and how these differences in demography relate to species richness, aboveground biomass (AGB), and carbon residence time. We find wide variation in DD and DC across forest plots, patterns that are not explained by species richness or climate variables alone. There is no evidence that DD has an effect on either AGB or carbon residence time. Rather, the DC of forests, specifically the relative abundance of large statured species, predicted both biomass and carbon residence time. Our results demonstrate the distinct DCs of globally distributed forests, reflecting biogeography, recent history, and current plot conditions. Linking the DC of forests to resilience or vulnerability to climate change, will improve the precision and accuracy of predictions of future forest composition, structure, and function.

Plant dynamics in a tropical dry forest are strongly associated with climate and fire and weakly associated with stabilizing neighborhood effects.

Studies comparing the relative strengths of multiple key drivers of forest dynamics are rare, but can inform both our fundamental understanding of plant communities as well as community-ecology theory. We studied the dynamics of a woody plant community in a southern Indian seasonally-dry tropical forest (SDTF) in relation to environmental factors (precipitation, temperature, fire, soil nutrients, and topography) and conspecific and heterospecific plant neighborhoods to identify which of these best predicted recruitment, survival and growth of dominant species over a 24-year study period. We also assessed the relative prevalence of density-independent and density-dependent responses in the community. Climate and fire were more important than plant neighborhoods and topographic and edaphic variables in explaining variation in plant performance. Recruitment, survival and growth were lower during periods of low precipitation and immediately following fires. Recruitment increased, and growth and survival largely decreased, with increasing temperatures. Smaller-sized individuals were disproportionately strongly affected by the vagaries of climate and fire. Conspecific negative density-dependence, a population-fluctuation stabilizing process, was relatively unimportant. Density-dependent effects decayed rapidly with distance from the focal plant (growth, survival) or quadrat (recruitment); positive density-dependence was frequently found in recruitment, possibly resulting from limited dispersal and/or facilitation. Woody plant dynamics in this SDTF appear to be responding largely to fluctuations in environmental conditions, particularly precipitation, temperature, and fire. In contrast to wetter forests, population-fluctuation stabilizing processes in this ecosystem appear to be relatively weak. Changes in climatic or fire regimes are likely to result in large compositional shifts in this SDTF.

Interactions between all pairs of neighboring trees in 16 forests worldwide reveal details of unique ecological processes in each forest, and provide windows into their evolutionary histories.

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands' physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees' distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions.

Improving prediction and assessment of global fires using multilayer neural networks.

Fires determine vegetation patterns, impact human societies, and are a part of complex feedbacks into the global climate system. Empirical and process-based models differ in their scale and mechanistic assumptions, giving divergent predictions of fire drivers and extent. Although humans have historically used and managed fires, the current role of anthropogenic drivers of fires remains less quantified. Whereas patterns in fire-climate interactions are consistent across the globe, fire-human-vegetation relationships vary strongly by region. Taking a data-driven approach, we use an artificial neural network to learn region-specific relationships between fire and its socio-environmental drivers across the globe. As a result, our models achieve higher predictability as compared to many state-of-the-art fire models, with global spatial correlation of 0.92, monthly temporal correlation of 0.76, interannual correlation of 0.69, and grid-cell level correlation of 0.60, between predicted and observed burned area. Given the current socio-anthropogenic conditions, Equatorial Asia, southern Africa, and Australia show a strong sensitivity of burned area to temperature whereas northern Africa shows a strong negative sensitivity. Overall, forests and shrublands show a stronger sensitivity of burned area to temperature compared to savannas, potentially weakening their status as carbon sinks under future climate-change scenarios.

The tail-tale of stress: an exploratory analysis of cortisol levels in the tail-hair of captive Asian elephants.

BACKGROUND: Assessment of physiological states by measuring biomarkers, such as cortisol, has significantly contributed to the monitoring of health, welfare and management of animals. Immunoreactive cortisol in hair (hC) has been used widely for deciphering 'stressful' past-events in various wild and captive animals. However, no such studies have been done in long-lived mammals. METHODS: In this first exploratory study in elephants, we assessed (i) tail-hair growth rate (TGR) and (ii) hC levels in tail-hair samples from six captive Asian elephants from two zoos in Japan for comparing hC levels with zoo-keepers' records of distinct biological events over a c.0.5-2.0-year period. Tail-hair samples were cut into segments (based on monthly growth rate), pulverized or minced and a validated cortisol enzyme-immunoassay employed to measure hC levels. RESULTS: When the hC levels of all individuals were compared with the keepers' records, a posteriori, most of the high hC levels were found to be associated with 'stressful' or distinct behavioural events such as pathological (anaemia, colic infection, skin infection, oral sores), psychosocial (reluctance in entering the enclosure, presence of a calf) and husbandry practice-related (contact trials/ space sharing) conditions, indicating that tail-hair indeed can be a potential 'retrospective' calendar of physiological health of an animal. CONCLUSIONS: Our observations open up the possibility of using the tail-hair as an alternative matrix to reconstruct the physiological history of elephants.

Behavioural responses of free-ranging Asian elephants (Elephas maximus) towards dying and dead conspecifics.

Reactions to dying and dead conspecifics have been observed in many non-human animals. Elephants, particularly African elephants, are thought to have an awareness of the death of their conspecifics, as they show compassionate behaviour towards others in distress. However, there is a paucity of scientific documentation on thanatological responses displayed by Asian elephants. Here, we report three detailed, directly observed cases of free-ranging Asian elephants (Elephas maximus) responding to dying and dead conspecifics. Behavioural responses were recorded opportunistically and described as pre-, peri- and post-mortem phases based on the status of the individual before, near or after its death. In all three observations, elephants showed approach and exploratory (sniffing and inspecting) behaviours, and epimeletic or helping (physically supporting dying calves) in pre- and peri-mortem phases. We also recorded high-frequency vocalizations (trumpets) by an adult female in the presence of a dying calf. Our observations indicate that, like their African counterparts, Asian elephants might experience distress in response to the death of conspecifics, and may have some awareness of death. This information furthers our understanding of the emotional and cognitive complexities of highly social elephants, and contributes to the growing field of elephant thanatology.

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient.

Among the local processes that determine species diversity in ecological communities, fluctuation-dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.

All-Male Groups in Asian Elephants: A Novel, Adaptive Social Strategy in Increasingly Anthropogenic Landscapes of Southern India.

Male Asian elephants are known to adopt a high-risk high-gain foraging strategy by venturing into agricultural areas and feeding on nutritious crops in order to improve their reproductive fitness. We hypothesised that the high risks to survival posed by increasingly urbanising and often unpredictable production landscapes may necessitate the emergence of behavioural strategies that allow male elephants to persist in such landscapes. Using 1445 photographic records of 248 uniquely identified male Asian elephants over a 23-month period, we show that male Asian elephants display striking emergent behaviour, particularly the formation of stable, long-term all-male groups, typically in non-forested or human-modified and highly fragmented areas. They remained solitary or associated in mixed-sex groups, however, within forested habitats. These novel, large all-male associations, may constitute a unique life history strategy for male elephants in the high-risk but resource-rich production landscapes of southern India. This may be especially true for the adolescent males, which seemed to effectively improve their body condition by increasingly exploiting anthropogenic resources when in all-male groups. This observation further supports our hypothesis that such emergent behaviours are likely to constitute an adaptive strategy for male Asian elephants that may be forced to increasingly confront anthropogenically intrusive environments.

Linking Termite Feeding Preferences and Soil Physical Functioning in Southern-Indian Woodlands.

Termites are undoubtedly amongst the most important soil macroinvertebrate decomposers in semi-arid environments in India. However, in this specific type of environment, the influence of termite foraging activity on soil functioning remains unexplored. Therefore, this study examines the link between the quality of litter and the functional impact of termite feeding preferences on soil properties and soil hydraulic conductivity in a deciduous forest in southern India. Different organic resources (elephant dung: "ED", elephant grass: "EG", acacia leaves: "AL" and layers of cardboard: "CB") were applied on repacked soil cores. ED appeared to be the most attractive resource to Odontotermes obesus, leading to a larger amount of soil sheeting (i.e., the soil used by termites for covering the litter they consume), more numerous and larger holes in the ground and a lower soil bulk density. As a consequence, ED increased the soil hydraulic conductivity (4-fold) compared with the control soil. Thus, this study highlights that the more O. obesus prefers a substrate, the more this species impacts soil dynamics and water infiltration in the soil. This study also shows that ED can be used as an efficient substrate for accelerating the infiltration of water in southern-Indian soils, mainly through the production of galleries that are open on the soil surface, offering new perspectives on termite management in this environment.

Assessment of season-dependent body condition scores in relation to faecal glucocorticoid metabolites in free-ranging Asian elephants.

We studied seasonal and annual changes in visual body condition scores (BCSs), and assessed how these scores were related to levels of faecal glucocorticoid metabolites (fGCMs) in free-ranging Asian elephants (Elephas maximus) in the seasonally dry tropical forests of the Mysore and Nilgiri Elephant Reserves in southern India. We assessed the animals' BCS visually on a scale of 1 to 5; where 1 represents a very thin and 5 represents a very fat elephant. To understand the influence of seasonality on BCS, we sampled the population during dry (n = 398) and wet seasons (n = 255) of 2013 and 2015 while, for annual changes in BCS, we sampled nine free-ranging adult females from different family groups that had been repeatedly sighted over seven years. To evaluate the influence of body condition on fGCM, 307 faecal samples were collected from 261 different elephants and were analysed. As a parameter of adrenocortical activity, and thus stress, fGCM was measured (µg/g) in the ethanol-extracted samples using a group-specific 11-oxoaetiocholanolone EIA (antibody raised against 11-oxoaetiocholanolone-17-CMO:BSA and biotinylated-11-oxoaetiocholanolone as a label). Effect of age and season on BCS in relation to fGCM was also studied. A seasonal shift in BCS was observed as expected, i.e. individuals with low BCS were more frequent during the dry season when compared with the wet season. Concentrations of fGCM were highest in individuals with lowest BCS (BCS 1) and then significantly declined till BCS 3. fGCM levels were almost comparable for BCS 3, 4 and 5. This pattern was more conspicuous in female than in male elephants. Season-dependent BCS, hence, reflect the stress status as measured by fGCM, especially in female Asian elephants. This could be used as an important non-invasive approach to monitor the physiological health of free-ranging elephant populations.

Multidimensional tree niches in a tropical dry forest.

The extent to which interspecific niche differences structure plant communities is highly debated, with extreme viewpoints ranging from fine-scaled niche partitioning, where every species in the community is specialized to a distinct niche, to neutrality, where species have no niche or fitness differences. However, there exists a default position wherein niches of species in a community are determined by their evolutionary and biogeographic histories, irrespective of other species within the community. According to this viewpoint, a broad range of pair-wise niche overlaps-from completely overlapping to completely distinct-are expected in any community without the need to invoke interspecific interactions. We develop a method that can test for both habitat associations and niche differences along an arbitrary number of spatial and temporal niche dimensions and apply it to a 24-yr data set of the eight dominant woody-plant species (representing 84% and 76% of total community abundance and basal area, respectively) from a 50-ha permanent plot in a southern Indian tropical dry forest, using edaphic, topographic, and precipitation variables as niche axes. Species separated into two broad groups in niche space-one consisting of three canopy species and the other of a canopy species and four understory species-along axes that corresponded mainly to variation in soil P, Al and a topographic index of wetness. Species within groups tended to have significantly greater niche overlap than expected by chance. Community-wide niche overlap in spatial and temporal niche axes was never smaller than expected by chance. Species-habitat associations were neither necessary nor sufficient preconditions for niche differences to be present. Our results suggest that this tropical dry-forest community consists of several tree species with broadly overlapping niches, and where significant niche differences do exist, they are not readily interpretable as evidence for niche differentiation. We argue, based on a survey of the literature, that many of the observed niche differences in tropical forests are more parsimoniously viewed as autecological differences between species that exist independently of interspecific interactions.

Functional nonredundancy of elephants in a disturbed tropical forest.

Conservation efforts are often motivated by the threat of global extinction. Yet if conservationists had more information suggesting that extirpation of individual species could lead to undesirable ecological effects, they might more frequently attempt to protect or restore such species across their ranges even if they were not globally endangered. Scientists have seldom measured or quantitatively predicted the functional consequences of species loss, even for large, extinction-prone species that theory suggests should be functionally unique. We measured the contribution of Asian elephants (Elephas maximus) to the dispersal of 3 large-fruited species in a disturbed tropical moist forest and predicted the extent to which alternative dispersers could compensate for elephants in their absence. We created an empirical probability model with data on frugivory and seed dispersal from Buxa Tiger Reserve, India. These data were used to estimate the proportion of seeds consumed by elephants and other frugivores that survive handling and density-dependent processes (Janzen-Connell effects and conspecific intradung competition) and germinate. Without compensation, the number of seeds dispersed and surviving density-dependent effects decreased 26% (Artocarpus chaplasha), 42% (Careya arborea), and 72% (Dillenia indica) when elephants were absent from the ecosystem. Compensatory fruit removal by other animals substantially ameliorated these losses. For instance, reductions in successful dispersal of D. indica were as low as 23% when gaur (Bos gaurus) persisted, but median dispersal distance still declined from 30% (C. arborea) to 90% (A. chaplasha) without elephants. Our results support the theory that the largest animal species in an ecosystem have nonredundant ecological functionality and that their extirpation is likely to lead to the deterioration of ecosystem processes such as seed dispersal. This effect is likely accentuated by the overall defaunation of many tropical systems.

Fires in Seasonally Dry Tropical Forest: Testing the Varying Constraints Hypothesis across a Regional Rainfall Gradient.

The "varying constraints hypothesis" of fire in natural ecosystems postulates that the extent of fire in an ecosystem would differ according to the relative contribution of fuel load and fuel moisture available, factors that vary globally along a spatial gradient of climatic conditions. We examined if the globally widespread seasonally dry tropical forests (SDTFs) can be placed as a single entity in this framework by analyzing environmental influences on fire extent in a structurally diverse SDTF landscape in the Western Ghats of southern India, representative of similar forests in monsoonal south and southeast Asia. We used logistic regression to model fire extent with factors that represent fuel load and fuel moisture at two levels-the overall landscape and within four defined moisture regimes (between 700 and1700 mm yr-1)-using a dataset of area burnt and seasonal rainfall from 1990 to 2010. The landscape scale model showed that the extent of fire in a given year within this SDTF is dependent on the combined interaction of seasonal rainfall and extent burnt the previous year. Within individual moisture regimes the relative contribution of these factors to the annual extent burnt varied-early dry season rainfall (i.e., fuel moisture) was the predominant factor in the wettest regime, while wet season rainfall (i.e., fuel load) had a large influence on fire extent in the driest regime. Thus, the diverse structural vegetation types associated with SDTFs across a wide range of rainfall regimes would have to be examined at finer regional or local scales to understand the specific environmental drivers of fire. Our results could be extended to investigating fire-climate relationships in STDFs of monsoonal Asia.

Controls of Soil Spatial Variability in a Dry Tropical Forest.

We examined the roles of lithology, topography, vegetation and fire in generating local-scale (<1 km2) soil spatial variability in a seasonally dry tropical forest (SDTF) in southern India. For this, we mapped soil (available nutrients, Al, total C, pH, moisture and texture in the top 10 cm), rock outcrops, topography, all native woody plants ≥1 cm diameter at breast height (DBH), and spatial variation in fire frequency (times burnt during the 17 years preceding soil sampling) in a permanent 50-ha plot. Unlike classic catenas, lower elevation soils had lesser moisture, plant-available Ca, Cu, Mn, Mg, Zn, B, clay and total C. The distribution of plant-available Ca, Cu, Mn and Mg appeared to largely be determined by the whole-rock chemical composition differences between amphibolites and hornblende-biotite gneisses. Amphibolites were associated with summit positions, while gneisses dominated lower elevations, an observation that concurs with other studies in the region which suggest that hillslope-scale topography has been shaped by differential weathering of lithologies. Neither NO3(-)-N nor NH4(+)-N was explained by the basal area of trees belonging to Fabaceae, a family associated with N-fixing species, and no long-term effects of fire on soil parameters were detected. Local-scale lithological variation is an important first-order control over soil variability at the hillslope scale in this SDTF, by both direct influence on nutrient stocks and indirect influence via control of local relief.

Comparative sequence analyses of genome and transcriptome reveal novel transcripts and variants in the Asian elephant Elephas maximus.

The Asian elephant Elephas maximus and the African elephant Loxodonta africana that diverged 5-7 million years ago exhibit differences in their physiology, behaviour and morphology. A comparative genomics approach would be useful and necessary for evolutionary and functional genetic studies of elephants. We performed sequencing of E. maximus and map to L. africana at ~15X coverage. Through comparative sequence analyses, we have identified Asian elephant specific homozygous, non-synonymous single nucleotide variants (SNVs) that map to 1514 protein coding genes, many of which are involved in olfaction. We also present the first report of a high-coverage transcriptome sequence in E. maximus from peripheral blood lymphocytes. We have identified 103 novel protein coding transcripts and 66-long non-coding (lnc)RNAs. We also report the presence of 181 protein domains unique to elephants when compared to other Afrotheria species. Each of these findings can be further investigated to gain a better understanding of functional differences unique to elephant species, as well as those unique to elephantids in comparison with other mammals. This work therefore provides a valuable resource to explore the immense research potential of comparative analyses of transcriptome and genome sequences in the Asian elephant.

In the elephant's seed shadow: the prospects of domestic bovids as replacement dispersers of three tropical Asian trees.

As populations of the world's largest animal species decline, it is unclear how ecosystems will react to their local extirpation. Due to the unique ecological characteristics of megaherbivores such as elephants, seed dispersal is one ecosystem process that may be affected as populations of large animals are decimated. In typically disturbed South Asian ecosystems, domestic bovids (cattle, Bosprimigenius, and buffalo, Bubalus bubalis) may often be the species most available to replace Asian elephants (Elephas maximus) as endozoochorous dispersers of large-fruited mammal-dispersed species. We use feeding trials, germination trials, and movement data from the tropical moist forests of Buxa Tiger Reserve (India) to examine whether domestic bovids are viable replacements for elephants in the dispersal of three large- fruited species: Dillenia indica, Artocarpus chaplasha, and Careya arborea. We find that (1) once consumed, seeds are between 2.5 (C. arborea) and 26.5 (D. indica) times more likely to pass undigested into elephant dung than domestic bovid dung; and (2) seeds from elephant dung germinated as well as or better than seeds taken from bovid dung for all plant species, with D. indica seeds from elephant dung 1.5 times more likely to germinate. Furthermore, since wild elephants have less constrained movements than even free-roaming domestic bovids, we calculate that maximum dispersal by elephants is between 9.5 and 11.2 times farther than that of domestic bovids, with about 20% of elephant-dispersed seeds being moved farther than the maximum distance seeds are moved by bovids. Our findings suggest that, while bovids are able to disperse substantial numbers of seeds over moderate distances for two of the three study species, domestic bovids will be unable to routinely emulate the reliable, long-distance dispersal of seeds executed by elephants in this tropical moist forest. Thus while domestic bovids can attenuate the effects of losing elephants as dispersers, they may not be able to prevent the decline of various mammal-dispersed fruiting species in the face of overhunting, habitat fragmentation, and climate change.