Identifying multiple wildlife species-crop interactions using network analysis

Managing crop damage by wildlife is a complex challenge in agrarian-wildlife landscapes. Losses to farmers and their negative attitude towards crop-raiding wildlife, compromise wildlife conservation and farmer-wildlife coexistence. Priority crop types, crop-raiding wildlife and socio-ecological factors are highly integrated. Here we use cross-sectional network analysis as a conservation planning tool to simultaneously identify the crop types used by crop-raiding wildlife, and evaluate the importance of these crop-wildlife interactions relative to the socio-ecological factors that affect these interactions. The most vulnerable crops were maize, millet and fruit trees, while the most problematic crop-raiding wildlife were African elephant, bushpig, hippo, warthog and red-billed quelea. Crop damage declined with countermeasure effectiveness, farm size, and distance from river to farm, but increased with richness of attractive crops. Undamaged crops included cash crops, such as chili, onion, ginger, lemon grass and garlic. Applying network analysis to different crops and damage scenarios enables identification of the most important and influential crops, crop-raiding wildlife and socio-ecological factors needed to develop effective crop protection strategies.     

Beehive fences as effective deterrents for crop‐raiding elephants: field trials in northern Kenya

Increasing elephant populations in Kenya since 1989 have been widely praised as a conservation success story. However, where elephants and agricultural land overlap, incidents of human–elephant conflict are on the increase. Wildlife managers and farmers are now trying different farm‐based deterrents to keep elephants out of crops. Here, we present data on the effectiveness of a novel beehive fence deployed in a Turkana community of 62 communally run farms in Kenya. Specifically, 1700 m of beehive fences semi‐surrounded the outer boundaries of seventeen farms, and we compared elephant farm invasion events with these and to seventeen neighbouring farms whose boundaries were ‘protected’ only by thorn bush barriers. We present data from 45 farm invasions, or attempted invasions, recorded over 2 years. Thirteen groups of elephants approached the beehive fences and turned away. Of the 32 successful farm invasions, only one bull elephant broke through the beehive fences. These results demonstrate that beehive fences are more effective than thorn bush barriers at deterring elephants and may have a role to play in alleviating farmer–elephant conflict. Additionally, the harvesting of 106 kg of honey during the trial period suggests that beehive fences may also improve crop production and enhance rural livelihoods through honey sales.     

Quantitative Interpretation of Tracks for Determination of Body Mass

To better understand the biology of extinct animals, experimentation with extant animals and innovative numerical approaches have grown in recent years. This research project uses principles of soil mechanics and a neoichnological field experiment with an African elephant to derive a novel concept for calculating the mass (i.e., the weight) of an animal from its footprints. We used the elephant''s footprint geometry (i.e., vertical displacements, diameter) in combination with soil mechanical analyses (i.e., soil classification, soil parameter determination in the laboratory, Finite Element Analysis (FEA) and gait analysis) for the back analysis of the elephant''s weight from a single footprint. In doing so we validated the first component of a methodology for calculating the weight of extinct dinosaurs. The field experiment was conducted under known boundary conditions at the Zoological Gardens Wuppertal with a female African elephant. The weight of the elephant was measured and the walking area was prepared with sediment in advance. Then the elephant was walked across the test area, leaving a trackway behind. Footprint geometry was obtained by laser scanning. To estimate the dynamic component involved in footprint formation, the velocity the foot reaches when touching the subsoil was determined by the Digital Image Correlation (DIC) technique. Soil parameters were identified by performing experiments on the soil in the laboratory. FEA was then used for the backcalculation of the elephant''s weight. With this study, we demonstrate the adaptability of using footprint geometry in combination with theoretical considerations of loading of the subsoil during a walk and soil mechanical methods for prediction of trackmakers weight.     

Effects of diet,habitat, and phylogeny on the fecal microbiome of wild African savanna (Loxodonta africana) and forest elephants (L. cyclotis)

The gut microbiome, or the community of microorganisms inhabiting the digestive tract, is often unique to its symbiont and, in many animal taxa, is highly influenced by host phylogeny and diet. In this study, we characterized the gut microbiome of the African savanna elephant (Loxodonta africana) and the African forest elephant (Loxodonta cyclotis), sister taxa separated by 2.6–5.6 million years of independent evolution. We examined the effect of host phylogeny on microbiome composition. Additionally, we examined the influence of habitat types (forest versus savanna) and diet types (crop‐raiding versus noncrop‐raiding) on the microbiome within L. africana. We found 58 bacterial orders, representing 16 phyla, across all African elephant samples. The most common phyla were Firmicutes, Proteobacteria, and Bacteroidetes. The microbiome of L. africana was dominated by Firmicutes, similar to other hindgut fermenters, while the microbiome of L. cyclotis was dominated by Proteobacteria, similar to more frugivorous species. Alpha diversity did not differ across species, habitat type, or diet, but beta diversity indicated that microbial communities differed significantly among species, diet types, and habitat types. Based on predicted KEGG metabolic pathways, we also found significant differences between species, but not habitat or diet, in amino acid metabolism, energy metabolism, and metabolism of terpenoids and polyketides. Understanding the digestive capabilities of these elephant species could aid in their captive management and ultimately their conservation.     

Exploring the relationship between NDVI and African elephant population density in protected areas

Assessing the extent to which populations are limited by bottom‐up processes driven by food limitation is crucial to our understanding of how ecosystems should be managed. Using satellite‐derived Normalised Difference Vegetation Index (NDVI) as an index of resource availability, we investigated the relationships between greenness levels and African elephant (Loxodonta africana) population densities. Our results unveiled a positive relationship between NDVI and elephant densities in nonforest populations, but failed to capture any significant effects of survey area and type, number of years after the continental poaching ban and the IUCN category of the protected area in which the survey was undertaken. The number of forest elephant populations for which density estimates were available was very low, and no significant relationship between NDVI and forest elephant density could be established. Altogether, our study suggests that NDVI can successfully be linked to megaherbivore abundance across Africa and further highlights that a continental approach to energy–abundance relationships can be conducted in a relatively low‐cost manner and short timescale, supporting managers' efforts to identify future suitable areas for elephant populations.     

Genomic DNA sequences from mastodon and woolly mammoth reveal deep speciation of forest and savanna elephants

To elucidate the history of living and extinct elephantids, we generated 39,763 bp of aligned nuclear DNA sequence across 375 loci for African savanna elephant, African forest elephant, Asian elephant, the extinct American mastodon, and the woolly mammoth. Our data establish that the Asian elephant is the closest living relative of the extinct mammoth in the nuclear genome, extending previous findings from mitochondrial DNA analyses. We also find that savanna and forest elephants, which some have argued are the same species, are as or more divergent in the nuclear genome as mammoths and Asian elephants, which are considered to be distinct genera, thus resolving a long-standing debate about the appropriate taxonomic classification of the African elephants. Finally, we document a much larger effective population size in forest elephants compared with the other elephantid taxa, likely reflecting species differences in ancient geographic structure and range and differences in life history traits such as variance in male reproductive success.     

Crop damage by African elephants assessment in kaftasheraro national park,Ethiopia

This study was conducted in Kafta‐Sheraro wereda in Northwest Tigray region of Ethiopia to assess community take on crop damage by the African Elephant, to identify visiting time of Elephants to agricultural plots and identify any other animals causing crop damage. A total of 240 households were selected from twelve villages using a stratified random sampling procedure. Half of the villages represented samples closer to the park and the other half, represented samples farther away. Whether the farmer owned or rented plots on which crops were grown as well as the distance to the park from the plots were the primary factors significantly (P < 0.05) influencing crop damage by elephants in the area. Time of the elephants visiting the agricultural plots was significantly (P < 0.05) higher at night. In a ranking exercise, among seven species considered, Elephants were ranked second in importance after Warthogs for crop destruction in villages close to the park but last in villages farther away. In the latter area, the most problematic animal was the Warthog followed by the Crested Porcupine. In the former areas where elephants' damage to crops was more frequent, farmers ranked fire as the most effective way of controlling the invasion. The study concludes that bringing the community to the discussion on elephant crop damage, increasing the time of keeping vigil during the night; improving the extension services and training in awareness creation and additional research's should be undertaken to evaluate in monetary terms the value of crop damage.     

Wild Asian elephants distinguish aggressive tiger and leopard growls according to perceived danger

Prey species exhibit antipredator behaviours such as alertness, aggression and flight, among others, in response to predators. The nature of this response is variable, with animals reacting more strongly in situations of increased vulnerability. Our research described here is the first formal study to investigate night-time antipredator behaviour in any species of elephants, Asian or African. We examined the provocative effects of elephant-triggered tiger and leopard growls while elephants attempted to crop-raid. Tigers opportunistically prey on elephant calves, whereas leopards pose no threat; therefore, we predicted that the elephant response would be reflective of this difference. Elephants reacted similarly cautiously to the simulated presence of felids of both species by eventually moving away, but differed markedly in their more immediate behavioural responses. Elephants retreated silently to tiger-growl playbacks, whereas they responded with aggressive vocalizations, such as trumpets and grunts, to leopard-growl playbacks. Elephants also lingered in the area and displayed alert or investigative behaviours in response to leopard growls when compared with tiger growls. We anticipate that the methods outlined here will promote further study of elephant antipredator behaviour in a naturalistic context, with applications for conservation efforts as well.     

Molecular Characterization of Pro-Inflammatory Cytokines Interleukin-1β and Interleukin-8 in Asian Elephant (Elephas maximus)

Interleukin (IL)-1β and IL-8 are pro-inflammatory cytokines produced primarily by monocytes and macrophages in response to a variety of microbial and nonmicrobial agents. As yet, no molecular data have been reported for IL-1β and IL-8 of the Asian elephant. In the present study, we have cloned and sequenced the cDNA encoding IL-1β and IL-8 of the Asian elephant. The open reading frame (ORF) of Asian elephant IL-1β is 789 bp in length, encoded a propeptide of 263 amino acid polypeptide. The predicted protein revealed the presence of IL-1 family signature motif and an ICE cut site. Whereas, IL-8 contained 321 bp of open reading frame. Interestingly, the predicted protein sequence of 106 aa, contains an ELR motif immediately upstream of the CQC residues, common in all vertebrate IL-8 molecules. Identity levels of the nucleic acid and deduced amino acid sequences of Asian elephant IL-1β ranged from 68.48 (Squirrel monkey) to 98.57% (African elephant), and 57.78 (Sheep) to 98.47% (African elephant), respectively, whereas that of IL-8 ranged from 72.9% (Human) to 87.8% (African elephant), and 63.2 (human, gorilla, chimpanzee) to 74.5% (African elephant, buffalo), respectively. The phylogenetic analysis based on deduced amino acid sequenced showed that the Asian elephant IL-1β and IL-8 were most closely related to African elephant. Molecular characterization of these two cytokines, IL-1β and IL-8, in Asian elephant provides fundamental information necessary to progress the study of functional immune responses in this animal and gives the potential to use them to manipulate the immune response as recombinant proteins.     

太行山华北豹袭击家畜的时空特点与管理建议: 以山西省和顺县为例

人兽冲突是全球野生动物保护面临的共同挑战。袭击家畜是大型食肉动物引发人兽冲突的主要原因之一, 如果管理不当还会导致针对大型食肉动物的报复性猎杀等严重威胁。冲突的缓解需要深入了解冲突发生的规律与原因, 评估当地社区与居民的态度与看法, 以采取因地制宜的管理措施。太行山是华北豹(Panthera pardus japonensis)的核心分布区, 华北豹袭击家牛的问题是该区域野生华北豹保护中面临的一大挑战。本研究以山西省和顺县为研究区域, 在多方联合发起的“和顺县华北豹袭击家牛肇事补偿项目”实施过程中, 系统收集了2015-2019年期间华北豹袭击家畜事件的记录, 结合同期的红外相机监测数据, 探究了袭击事件的时空分布格局, 并从生境和猎物的角度分析了背后的原因。本文还评估了豹肇事造成的经济损失, 以及受损农户对华北豹的态度和对补偿的满意度。2015-2019年, 项目区域内共记录到华北豹袭击家牛事件195起, 向116户受损农户提供补偿资金共27.05万元。分析结果显示, 华北豹袭击家牛事件在夏季最多, 时间上与家牛上山散放的季节重叠; 袭击风险随狍(Capreolus pygargus)相对多度的上升而下降, 随家牛相对多度和距村庄距离的增加而上升, 但保护区内外和华北豹相对多度对袭击风险的影响不显著。对107名受损农户的访谈结果显示, 华北豹袭击家牛事件虽然对当地畜牧业所造成的整体损失较小, 但对受损农户而言其家庭收入损失严重, 因此导致受损农户对华北豹持负面态度, 且认为现有的补偿金额偏低。基于本研究的结果, 我们提出如下管理建议以期缓解未来该地区华北豹带来的人兽冲突: (1)加强放牧管理、改进管理方式, 特别是限制牛群离开村庄的距离, 以及在家牛散放期间将牛群每晚赶回牛圈以降低家畜被袭击的风险; (2)开展禁牧试点, 严格划分允许放牧与禁止放牧的区域; (3)继续开展华北豹袭击家牛的定损与补偿工作, 降低当地农户的经济损失, 同时开展社区走访、自然教育, 提升当地居民对华北豹的容忍度; (4)加强野生动物种群的监测与保护, 促进人与野生动物的和谐共存。     

Molecular and morphological evidence on the phylogeny of the Elephantidae

The African and Asian elephants and the mammoth diverged ca. 4-6 million years ago and their phylogenetic relationship has been controversial. Morphological studies have suggested a mammoth Asian elephant relationship, while molecular studies have produced conflicting results. We obtained cytochrome b sequences of up to 545 base pairs from five mammoths, 14 Asian and eight African elephants. A high degree of polymorphism is detected within species. With a dugong sequence used as the outgroup, parsimony and maximum-likelihood analyses support a mammoth African elephant clade. As the dugong is a very distant outgroup, we employ likelihood analysis to root the tree with a molecular clock, and use bootstrap and Bayesian analyses to quantify the relative support for different topologies. The analyses support the mammoth African elephant relationship, although other trees cannot be rejected. Ancestral polymorphisms may have resulted in gene trees differing from the species phylogeny Examination of morphological data, especially from primitive fossil members, indicates that some supposed synapomorphies between the mammoth and Asian elephant are variable, others convergent or autapomorphous. A mammoth African elephant relationship is not excluded. Our results highlight the need, in both morphological and molecular phylogenetics, for multiple markers and close attention to within-taxon variation and outgroup selection.     

The problem of precision and trend detection posed by small elephant populations in West Africa

The precision of elephant estimates from aerial sample surveys and dung counts is inversely proportional to abundance. West African elephant populations are already small, and the power of a monitoring programme to detect changes in abundance diminishes as the population shrinks in size. Thus it will be difficult to evaluate the effects on elephant numbers of new management policies in West Africa. The same will be true of monitoring schemes for antelope and primate populations that are hunted for bushmeat. Elephant estimates from dung counts are more precise than those from aerial sample surveys, and changes in elephant numbers are more likely to be detected in the subregion by dung counts than by aerial sample surveys.     

Heat transfer in elephants: thermal partitioning based on skin temperature profiles

The elephant with its low surface-to-volume ratio presents an interesting problem concerning heat dissipation. To understand how such large mammals remain in thermal balance, we determined the major avenues of heat loss for an adult African elephant and an immature Indian elephant. Because conventional physiological measurements are difficult for these animals, the present study used a non-invasive technique, infrared thermography, to measure skin temperatures of each elephant. Detailed surface temperature profiles and surface area measurements of each elephant were used in standard equations for convective, conductive and radiant heat transfer. Results demonstrated that heat transfer by free convection and radiation accounted for 86% of the total heat loss for the elephants at T _a= 12·6 °C. Heat transfer across the ears, an important thermal window at high ambient temperatures, represented less than 8% of the total heat loss. Surface area of the animals, and metabolic heat production calculated from total heat loss of the African elephant, scaled predictably with body mass. In contrast, the thermal conductance of the elephants (71·6 W /°C, African; 84·5 W /°C, Indian) was three to five times higher than predicted from an allometric relationship for smaller mammals. The high thermal conductance of elephants is attributed to the absence of fur and appears to counteract reduced heat transfer associated with a low surface-to-volume ratio.     

Assessing the cost of living with elephants (Loxodonta africana) in areas adjacent to Meru National Park,Kenya

Crop raiding by the elephants is a serious and recurring management problem around protected areas in Kenya such as Meru National Park. Crop-raiding menace is one of the most significant of human–elephant conflicts in Meru National Park. The distribution, impact, and conservation implications of the increased elephant crop raiding in areas adjacent to Meru National Park is attributed to the changes in land use systems within these areas. Crop raiding by African elephants (Loxodonta africana) was monitored in the area adjacent to Meru National Park between August 2010 and July 2011. From the study, 144 farms were raided and farmers lost crops amounting to USD 120,308.60. Crop raiding was higher during the month of August 2010 (KES 2,714,295 or USD 33,928). The study suggests capacity building for communities in order to safeguard their crops against elephant raiding. Other probable measures put forward to mitigate this include the development of alternative water sources and the need to implement electric fence around the remaining section of the Park boundary which is affected by elephant.     

Economics of carnivore depredation: A case study from the northern periphery of Corbett Tiger Reserve,Uttarakhand, India

Corbett Tiger Reserve (CTR) supports several important species such as the tiger (Panthera tigris), leopard (Panthera pardus), elephant (Elephas maximus), and sambar (Cervus unicolor). Some of the species are involved in crop damage, livestock depredation, and human injuries or causalities, causing economic and social losses. Tigers and leopards are perceived as the major threat, causing economic losses to the people living around the protected areas (PAs), because of livestock depredation. Alternatively, the local community also draws benefits from the forest, which are seldom considered part of their economy. Here, we assessed the economic losses and benefits to the people at the northern periphery of the CTR based on semi-structured questionnaire survey of 301 respondents in 26 villages, between January and June 2018. We also determined the socio-economic factors associated with forest dependency using Ordinary Least Square regression (OLS). Most respondents (98%) reported reliance on the forest for fuelwood, fodder, and cattle grazing. Income from forest resources (US$ 1102.2 ± 63.6/hh/yr)² contributed significantly to the livelihood of the people (26.5 ± 1.3%). The income from fuelwood (US$ 188.1 ± 2.2/hh/yr), fodder (US$ 335.0 ± 10.0/hh/yr) and livestock grazing (US$ 618.1 ± 59.9/hh/yr) contributed 5.6 ± 0.3%, 9.3 ± 0.6%, and 11.6 ± 1%, respectively, to the household's total income. Reliance on forest resources was positively associated with the number of livestock and female participation in resource extraction, and negatively with the employment, age, education of respondents, and use of modern source of energy -Liquid Petroleum Gas. People incurred a cost of US$ 110.5 ± 4.7/hh/yr due to livestock depredation. It was estimated that benefits drawn from the forest exceed the losses incurred due to livestock depredation. Therefore, we recommend that a cost-benefit analysis is essential before designing conflict mitigation strategies. Since the compensatory schemes in India have largely struggled to achieve their objectives, it is, therefore, emphasised to find other measures, such as payment of ecosystem services (PES), to reduce conflict for maintaining a peaceful co-existence of human and carnivores.     

Feeding patterns of elephants at the Atherstone Collaborative Nature Reserve

The African savannah elephant (Loxodonta africana Blumenbach) is a generalist herbivore that relies on widely distributed resources. Vegetation decline, aggravated by these elephants, can compromise local conservation efforts. Thus it imperative to understand the factors that drive them to consume specific plant species and plant parts. The objective of our study was to investigate the feeding patterns of African savannah elephants in the enclosed bushveld savannah at the Atherstone Collaborative Nature Reserve in South Africa. For 1 year, we examined elephant selection of woody versus herbaceous vegetation, and identified which plant species and parts were preferentially consumed. We accomplished this by directly observing feeding elephants, and by utilizing data collected on elephant footprints, dung piles, stripped bark and broken branches. We further conducted vegetation surveys to determine selection frequency relative to species abundance. Elephants showed a preference for different plant parts consumption in the feeding plots. In total, leaves, branches and bark contributed mostly to their diet. Seasonal selection patterns showed an increasing proportion of bark and branch consumption during the dry season.     

Characterization of tetranucleotide microsatellite loci in the African Savannah Elephant (Loxodonta africana africana)

Most African elephant (Loxodonta africana africana) populations are isolated and thus threatened by a loss of genetic diversity. As a consequence, genetic analysis of African elephant populations will play an increasing role in their conservation, and microsatellite loci will be an important tool in these analyses. Previously published sets of polymorphic microsatellites developed for African elephants are all dinucleotide repeats, which are prone to typing error. Here, we characterize 11 tetranucleotide microsatellite loci in the African elephant. All loci were polymorphic in 32 faecal samples and two tissue samples from 33 individual African savannah elephants.     

The influence of lunar cycles on crop‐raiding elephants; evidence for risk avoidance

Long‐term solutions to crop raiding by elephants (Loxodonta africana) should be based on an understanding of their behaviour and ecology. The real and perceived risks from humans have been shown to affect elephant behaviour. This is evidenced by elephants predominantly raiding crops at night, avoiding the height of human activity. If such human avoidance behaviours are apparent, it might also be expected that elephants avoid risks associated with higher visibility and increased human activity as may occur during the full moon. However, elephant nocturnal crop‐raiding behaviour in relation to lunar cycles has largely been a neglected factor in studies of human–elephant interactions. In this study around Mikumi National Park, Tanzania, we apply circular statistics in this context for the first time to show a significant decrease in crop raiding during the full moon and apply this method retrospectively to data from another site in West Africa with similar results. Additionally, a greater proportion of farms raided was guarded during the full moon than any other moon phase. Our results indicate that variations in crop raiding with lunar phase could be a general feature of elephant behaviour and thus could be used to design and time mitigation efforts.     

Understanding the effect of landscape fragmentation and vegetation productivity on elephant habitat utilization in Amboseli ecosystem,Kenya

Understanding factors affecting the distribution of the African elephant is important for its conservation in increasingly human‐dominated savannah landscapes. However, understanding how landscape fragmentation and vegetation productivity affect elephant habitat utilization remains poorly understood. In this study, we tested whether landscape fragmentation and vegetation productivity explain elephant habitat utilization in the Amboseli ecosystem in Kenya. We used GPS (Global Positioning System) telemetry data from five elephants to quantify elephant habitat utilization. Habitat utilization was determined by calculating the time elephants spent within a unit area. We then used generalized additive models (GAMs) to model the relationship between time density and landscape fragmentation, as well as vegetation productivity. Results show that landscape fragmentation and vegetation productivity significantly (P < 0.05) explain elephant habitat utilization. A significant (P < 0.05) unimodal relationship between vegetation productivity and habitat utilization was observed. Results suggest that elephants spend much of their time in less fragmented landscapes of intermediate productivity.     

Reconciling apparent conflicts between mitochondrial and nuclear phylogenies in African elephants

Conservation strategies for African elephants would be advanced by resolution of conflicting claims that they comprise one, two, three or four taxonomic groups, and by development of genetic markers that establish more incisively the provenance of confiscated ivory. We addressed these related issues by genotyping 555 elephants from across Africa with microsatellite markers, developing a method to identify those loci most effective at geographic assignment of elephants (or their ivory), and conducting novel analyses of continent-wide datasets of mitochondrial DNA. Results showed that nuclear genetic diversity was partitioned into two clusters, corresponding to African forest elephants (99.5% Cluster-1) and African savanna elephants (99.4% Cluster-2). Hybrid individuals were rare. In a comparison of basal forest "F" and savanna "S" mtDNA clade distributions to nuclear DNA partitions, forest elephant nuclear genotypes occurred only in populations in which S clade mtDNA was absent, suggesting that nuclear partitioning corresponds to the presence or absence of S clade mtDNA. We reanalyzed African elephant mtDNA sequences from 81 locales spanning the continent and discovered that S clade mtDNA was completely absent among elephants at all 30 sampled tropical forest locales. The distribution of savanna nuclear DNA and S clade mtDNA corresponded closely to range boundaries traditionally ascribed to the savanna elephant species based on habitat and morphology. Further, a reanalysis of nuclear genetic assignment results suggested that West African elephants do not comprise a distinct third species. Finally, we show that some DNA markers will be more useful than others for determining the geographic origins of illegal ivory. These findings resolve the apparent incongruence between mtDNA and nuclear genetic patterns that has confounded the taxonomy of African elephants, affirm the limitations of using mtDNA patterns to infer elephant systematics or population structure, and strongly support the existence of two elephant species in Africa.