Heard but not seen: an acoustic survey of the African forest elephant population at Kakum Conservation Area, Ghana

This study, designed to survey forest elephants ( Loxodonta africana cyclotis ) at Kakum Conservation Area, Ghana, is the first to apply acoustic methods to elephant abundance estimation and to compare results with independent survey estimates. Nine acoustic sensors gathered sound continuously for 38 days. Low-frequency calling rates have been established as useful elephant abundance indices at a Namibian watering hole and a central African forest clearing. In this study, we estimated elephant population size by applying an abundance index model and detection function developed in central Africa to data from simultaneous sampling periods on Kakum sensors. The sensor array recorded an average of 1.81 calls per 20-min sampling period from an effective detection area averaging 10.27 km². The resulting estimate of 294 elephants (95% CI: 259–329) falls within confidence bounds of recent dung-based surveys. An extended acoustic model, estimating the frequency with which elephants are silent when present, yields an estimate of 350 elephants (95% CI: 315–384). Acoustic survey confidence intervals are at least half as wide as those from dung-based surveys. This study demonstrates that acoustic surveying is a valuable tool for estimating elephant abundance, as well as for detecting other vocal species and anthropogenic noises that may be associated with poaching.     

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.     

Small-scale dung survey reveals high forest elephant density and preference for mixed species forest in an intact protected area

Understanding the ecological factors influencing African forest elephant (Loxodonta cyclotis) abundance and distribution is crucial for their conservation management in Central Africa. Dung surveys have been conducted at the landscape scale and confirmed the overwhelming impact of anthropogenic activities on forest elephants. We present results from a small-scale survey in a pristine protected area without anthropogenic activities to elucidate the ecological factors influencing forest elephant density. We conducted a line transect dung survey in a small study area (110 km²) around Mbeli Bai, a natural forest clearing in the Nouabalé-Ndoki National Park, Republic of Congo, and compare results with a landscape survey conducted during the same period. We used habitat specific dung decay data collected on site to estimate elephant density using distance sampling. We fitted Generalized Additive Models to elephant dung encounter rate using explanatory variables collected during the transect survey and from geospatial data. The small-scale survey revealed a precise estimate of forest elephant density that was twice as high as the result from a landscape survey with higher density in mixed species forest for the small-scale survey. We could not find an impact of the proximity of forest clearings and proximity to rivers at the small scale. Fine-scale habitat features, e.g. degree of canopy and understorey closure, had little explanatory power for elephant dung encounter rate. Small-scale dung surveys are a useful method to reveal spatio-temporal variation in forest elephant density and distribution which can inform conservation practitioners in a timely manner. Combining monitoring methods at various spatial scales improves our knowledge and conservation efforts of forest elephants. Nouabalé-Ndoki National Park is a stronghold for forest elephants and of global importance for their conservation.

     

A survey of elephants (Loxodonta africana) in the Kahuzi-Biega National Park lowland sector and adjacent forest in eastern Zaire

There has been considerable uncertainty about the abundance estimation of Loxodonta africana within tropical lowland moist forests in Zaire. We surveyed a 15,570 km² area within the forests of eastern Zaire using transect sampling methods and estimated the elephant population to be 3720 (range 2300–5000) individuals. Dung pile densities were significantly different between adjacent settlement, deep forest, and deep forest core strata, with the most remote area harbouring the highest density. Evidence of elephant poaching was encountered throughout the survey area suggesting that elephant populations continue to be at severe risk.     

Opening a larger window onto forest elephant ecology

African forest elephants (Loxodonta cyclotis) can be observed reliably in forest clearings but are difficult to observe directly in their otherwise forested habitat. To obtain data on population demography and activity cycles, conservation programmes commit considerable human and financial resources to observations at clearings. However, individual elephants spend only a tiny proportion of their annual activity budget in a particular clearing, and there may be demographic differences between daytime and nighttime populations. Using acoustic monitoring tools to assess elephant numbers continuously for long time periods at multiple clearings, we show that daytime‐only observations are likely to be biased. In this study, (i) 79% of all elephant visitation occurred at night, (ii) sometimes large changes in nocturnal elephant numbers occurred without proportional changes in daytime elephant numbers, and (iii) there were indications that a different demographic of the population visits clearings at night compared with the day. These results suggest that acoustic monitoring should be incorporated into forest elephant monitoring programmes to augment direct observation and that more study is needed to explicitly identify any biases inherent in daytime‐only observation.     

A case study of apparent conflict between molecular phylogenies: the interrelationships of African elephants

Recent molecular phylogenies of the African elephants suggest that there is an evolutionary structure within Loxodonta africana. Some nuclear results ( Roca et al., 2001 ) support the separation of the forest African elephant subspecies L. a. cyclotis as a species distinct from the savannah elephant L. a. africana, on the basis of the recognition of both forming highly divergent (reciprocally monophyletic) clades. Conversely, a mitochondrial survey ( Eggert et al., 2002 ), while admitting a geographic partitioning of the genetic structure within African elephants, suggests retaining the status quo. They recognize three diagnosible entities (western, central and south‐eastern Africa) with non‐overlapping ranges within L. africana sensu lato. In order to address these conflicting views (historical fragmentation and speciation or isolation by distance, respectively), we have sequenced two datasets of 1961 bp (for 50 elephants) and about 3700 bp, respectively (for 20 elephants) of the mitochondrial DNA for both forms of elephants (cyclotis and africana). They span the cytochrome b gene, the control region and several RNAs. When compared with former mtDNA data, they provide the most comprehensive view of the African elephant phylogeny (78 mtDNA haplotypes, of which 44 are new) and provide the first insight into populations from the Democratic Republic of Congo. The genetic diversity of mtDNA was appraised and the stability of alternative phylogenetic trees was investigated. Our results are inconsistent with both those prior studies. They revealed two highly divergent molecular clades referred to as F and S, that do not conform to the morphological delineations of cyclotis and africana. A non‐negligible proportion of specimens of L. a. africana display haplotypes prevailing in forest elephant populations (clade F). The geographic distribution of clades and areas of their co‐occurrence support the hypothesis of incomplete isolation between forest and savannah African elephant populations, followed by recurrent interbreeding between the two forms. We state that the conclusions of prior studies resulted from insufficient character and/or geographic sampling. We conclude that there is no satisfying argument which can recognize two or more species of African elephants. We briefly comment on the meaning of such an attitude in a conservation viewpoint. © The Willi Hennig Society 2005.     

Group size, density and biomass of large mammals in the Réserve de Faune du Petit Loango, Gabon

Group size, density and biomass of large‐bodied diurnal mammal species in the Réserve de Faune du Petit Loango, Gabon (now Parc National de Loango) was determined over a 12‐month period using standard line‐transect methods. Petit Loango encompasses a range of distinct habitat types, including coastal scrub, savanna, swamps and disturbed and mature forest. Such intact coastal habitats are increasingly rare on the Central‐West African coastline. Faecal censusing indicated highest forest elephant (Loxodonta africana cyclotis) and buffalo (Syncerus caffer nanus) ecological densities at the extreme coast (2.48 and 1.29 km⁻² respectively), probably reflecting high intensity of use of this habitat. Ape density was comparable with that at other Central African study sites at 1.01 individuals km⁻². Mean total biomass of diurnal primates, elephants and other ungulates over the 20 km² site was 3290 kg km⁻². Forest elephants and red river hogs (Potamochoerus porcus) constituted the bulk of the biomass, at 67% and 14% respectively. Primates made up 5% of the biomass. This is the first estimation of mammal density and biomass over an annual cycle at a Central African coastal site, and provides baseline data for long‐term studies in such habitats and to aid habitat and wildlife management decisions.     

Forest elephant crisis in the Congo Basin

Debate over repealing the ivory trade ban dominates conferences of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Resolving this controversy requires accurate estimates of elephant population trends and rates of illegal killing. Most African savannah elephant populations are well known; however, the status of forest elephants, perhaps a distinct species, in the vast Congo Basin is unclear. We assessed population status and incidence of poaching from line-transect and reconnaissance surveys conducted on foot in sites throughout the Congo Basin. Results indicate that the abundance and range of forest elephants are threatened from poaching that is most intense close to roads. The probability of elephant presence increased with distance to roads, whereas that of human signs declined. At all distances from roads, the probability of elephant occurrence was always higher inside, compared to outside, protected areas, whereas that of humans was always lower. Inside protected areas, forest elephant density was correlated with the size of remote forest core, but not with size of protected area. Forest elephants must be prioritised in elephant management planning at the continental scale.     

Quantifying past and present connectivity illuminates a rapidly changing landscape for the African elephant

There is widespread concern about impacts of land‐use change on connectivity among animal and plant populations, but those impacts are difficult to quantify. Moreover, lack of knowledge regarding ecosystems before fragmentation may obscure appropriate conservation targets. We use occurrence and population genetic data to contrast connectivity for a long‐lived mega‐herbivore over historical and contemporary time frames. We test whether (i) historical gene flow is predicted by persistent landscape features rather than human settlement, (ii) contemporary connectivity is most affected by human settlement and (iii) recent gene flow estimates show the effects of both factors. We used 16 microsatellite loci to estimate historical and recent gene flow among African elephant (Loxodonta africana) populations in seven protected areas in Tanzania, East Africa. We used historical gene flow (F_ST and G'_ST) to test and optimize models of historical landscape resistance to movement. We inferred contemporary landscape resistance from elephant resource selection, assessed via walking surveys across ~15 400 km² of protected and unprotected lands. We used assignment‐based recent gene flow estimates to optimize and test the contemporary resistance model, and to test a combined historical and contemporary model. We detected striking changes in connectivity. Historical connectivity among elephant populations was strongly influenced by slope but not human settlement, whereas contemporary connectivity was influenced most by human settlement. Recent gene flow was strongly influenced by slope but was also correlated with contemporary resistance. Inferences across multiple timescales can better inform conservation efforts on large and complex landscapes, while mitigating the fundamental problem of shifting baselines in conservation.     

Focusing Conservation Efforts for the Critically Endangered Brown-headed Spider Monkey (<Emphasis Type="Italic">Ateles fusciceps</Emphasis>) Using Remote Sensing,Modeling, and Playback Survey Methods

Brown-headed spider monkeys (Ateles fusciceps), endemic to the Choco-Darien forests and lower Andean forests of NW Ecuador, are considered critically endangered. Unfortunately, scientific data regarding the actual status of populations is lacking. We combined satellite image analysis, species-specific habitat assessment, and a field survey technique using playback to focus conservation efforts for this species. First, we identified remaining forest via a LANDSAT mosaic and then applied species-specific criteria to delineate remaining forest with potential to hold populations. By combining this with the historical distribution from ecological niche modeling and predicted hunting intensity we generated a species-specific landscape map. Within our study area, forest capable of sustaining Ateles fusciceps covers 5872 km², of which 2172 km² (40%) is protected. Unprotected forest considered suitable for Ateles fusciceps extends to 3700 km² but within this only 989 km² (23%) is under low hunting pressure and likely to maintain healthy populations of Ateles fusciceps. To overcome problems of sampling at low primate density and in difficult mountain terrain we developed a field survey technique to determine presence and estimate abundance using acoustic sampling. For sites under low hunting pressure density of primates varied with altitude. Densities decreased from 7.49 individuals/km² at 332 masl to 0.9 individuals/km² at 1570 masl. Based on combining data sets in a gap analysis, we recommend conservation action focus on unprotected lowland forest to the south and west of the Cotacachi-Cayapas Ecological Reserve where hunting pressure is low and population densities of Ateles fusciceps are greatest.     

Estimating elephant density using motion-sensitive cameras: challenges,opportunities, and parameters for consideration

With extinction rates far exceeding the natural background rate, reliable monitoring of wildlife populations has become crucial for adaptive management and conservation. Robust monitoring is often labor intensive with high economic costs, particularly in the case of those species that are subject to illegal poaching, such as elephants, which require frequent and accurate population estimates over large spatial scales. Dung counting methods are commonly employed to estimate the density of elephants; however, in the absence of a full survey calibration, these can be unreliable in heterogeneous habitats where dung decay rates may be highly variable. We explored whether motion-sensitive cameras offer a simple, lower cost, and reliable alternative for monitoring in challenging forest environments. We estimated the density of African savanna elephants (Loxodanta africana) in a montane forest using the random encounter model and assessed the importance of surveying parameters for future survey design. We deployed motion-sensitive cameras in 65 locations in the Aberdare Conservation Area in Kenya during June to August in 2015 to 2017, for a survey effort of 967 days, and a mean encounter rate of 0.09 ± 0.29 (SD) images/day. Elephants were captured in 16 locations. Density estimates varied between vegetation types, with estimates ranging from 6.27/km² in shrub, 1.1/km² in forest, 0.53/km² in bamboo (Yushania alpine), and 0.44/km² in the moorlands. The average speed of animal movement and the camera detection zone had the strongest linear associations with density estimates (R = −0.97). The random encounter model has the potential to offer an alternative, or complementary method within the active management framework for monitoring elephant populations in forests at a relatively low cost.     

Estimating population sizes for elusive animals: the forest elephants of Kakum National Park,Ghana

African forest elephants are difficult to observe in the dense vegetation, and previous studies have relied upon indirect methods to estimate population sizes. Using multilocus genotyping of noninvasively collected samples, we performed a genetic survey of the forest elephant population at Kakum National Park, Ghana. We estimated population size, sex ratio and genetic variability from our data, then combined this information with field observations to divide the population into age groups. Our population size estimate was very close to that obtained using dung counts, the most commonly used indirect method of estimating the population sizes of forest elephant populations. As their habitat is fragmented by expanding human populations, management will be increasingly important to the persistence of forest elephant populations. The data that can be obtained from noninvasively collected samples will help managers plan for the conservation of this keystone species.     

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.     

Changes in Elephant Abundance Affect Forest Composition or Regeneration?

While overall numbers of African elephant have declined dramatically in recent times, some populations are now confined to protected areas and are locally overabundant—an undesirable situation for both biodiversity conservation and elephants. In forested protected areas, options to manage elephants are limited because it is difficult to safely approach animals, yet it is vital that these populations are managed because browsing by elephants can dramatically alter forest ecosystems. Using data collected over 50 yr in Kibale National Park, Uganda, we examine the prediction that increasing elephant numbers and associated changes in their foraging behavior have caused a shift in tree community composition. Although the relative abundance of elephants increased significantly between 1996 and 2010, the population structure of their preferred tree food species did not change, nor did tree community composition change in favor of species able to re‐sprout after elephant damage. Furthermore, over the last 50 yr Kibale elephants have not become more selective foragers, as would be expected if more nutritious tree species were declining. However, elephants are more abundant in disturbed areas dominated by shrubs and grasses and appear to have arrested forest succession in these areas. At their current abundance, elephants have not selectively altered the composition of intact old growth forest, but they do inhibit the regeneration of disturbed areas.     

Elephants caught in the middle: impacts of war, fences and people on elephant distribution and abundance in the Caprivi Strip, Namibia

We conducted wet [26 March–4 April 2003 (Apr03)] and dry [1–8 November 2005 (Nov05)] season aerial surveys of African elephants ( Loxodonta africana Blumenbach) in the Caprivi Strip, Namibia to provide an updated status report on elephant numbers and distribution and assist with a historical analysis of elephant distribution and abundance in the Caprivi Strip. During the wet season when water was available in seasonal pans, elephants were widely distributed throughout the survey area. In contrast, during the dry season, a majority of elephant herds occurred within 30 km of the perennial Kwando, Linyanti and Okavango rivers and few herds occurred within the West Caprivi Game Reserve where water in the seasonal pans was limited. We estimated 5318 elephants for the 7731-km² survey area (0.71 elephants km⁻²) for the Apr03 wet season survey and 6474 elephants for the 8597-km² survey area (0.75 elephants km⁻²) for the Nov05 dry season survey. Based on our aerial surveys and reports of elephant numbers and distribution from historical aerial surveys and telemetry studies, civil war, veterinary fences and human activities appear to have effected changes in African elephant abundance, distribution and movements in the Caprivi Strip, Namibia since 1988 when the first comprehensive aerial surveys were conducted.     

Assessing the effects of elephant foraging on the structure and diversity of an Afrotropical forest

African forest elephants (Loxodonta cyclotis) are ecosystem engineers that browse and damage large quantities of vegetation during their foraging and movement. Though elephant trail networks and clearings are conspicuous features of many African forests, the consequences of elephant foraging for forest structure and diversity are poorly documented. In this study in northeastern Gabon, we compare stem size, stem density, proportional damage, species diversity, and species relative abundance of seedlings and saplings in the vicinity of seven tree species that produce elephant-preferred fruits (“elephant trees”) relative to control trees that do not. Across 34 survey trees, with a combined census area of 2.04 ha, we recorded data on 26,128 woody stems in three sizes classes. Compared with control trees, the area around elephant trees had the following: (a) a significantly greater proportion of damaged seedlings and a marginally greater proportion of damaged saplings (with 82% and 24% greater odds of damage, respectively); (b) no significant difference in stem density or species diversity; and (c) a significantly greater relative abundance of seedlings of elephant tree species. Increasing distance away from focal elephant trees was associated with significantly reduced sapling stem damage, significantly increased sapling stem density, and significantly increased sapling species diversity. Considered in sum, our results suggest that elephants can affect the structure and diversity of Afrotropical forests through their foraging activities, with some variation based on location and plant size class. Developing a more complete understanding of elephants’ ecological effects will require continued research, ideally with manipulative experiments.     

Evaluating monitoring methods to guide adaptive management of a threatened amphibian (Litoria aurea)

Prompt detection of declines in abundance or distribution of populations is critical when managing threatened species that have high population turnover. Population monitoring programs provide the tools necessary to identify and detect decreases in abundance that will threaten the persistence of key populations and should occur in an adaptive management framework which designs monitoring to maximize detection and minimize effort. We monitored a population of Litoria aurea at Sydney Olympic Park over 5 years using mark–recapture, capture encounter, noncapture encounter, auditory, tadpole trapping, and dip‐net surveys. The methods differed in the cost, time, and ability to detect changes in the population. Only capture encounter surveys were able to simultaneously detect a decline in the occupancy, relative abundance, and recruitment of frogs during the surveys. The relative abundance of L. aurea during encounter surveys correlated with the population size obtained from mark–recapture surveys, and the methods were therefore useful for detecting a change in the population. Tadpole trapping and auditory surveys did not predict overall abundance and were therefore not useful in detecting declines. Monitoring regimes should determine optimal survey times to identify periods where populations have the highest detectability. Once this has been achieved, capture encounter surveys provide a cost‐effective method of effectively monitoring trends in occupancy, changes in relative abundance, and detecting recruitment in populations.     

Large‐scale changes in abundance of terricolous bryophytes and macrolichens in Finland

Abstract. Major changes in forest floor vegetation were identified on the basis of three nationwide surveys conducted as part of national forest inventories in 1951–1953, 1985–1986 and 1995. These surveys provided objectively selected, statistically representative samples of all forested land in Finland. The 1951–1953 data consist of over 10000 sample plots, while the later surveys were conducted on ca. 3000 permanent plots. Changes in relative abundance of dominant species (i.e. in the proportions of species of the total cover of forest floor vegetation) were analysed across biogeographical provinces. Spatial correlation, systematic sampling, partial re‐measurement and multiple testing were taken into account in assessment of the statistical significance of the observed changes. The most notable changes in forest floor vegetation were a decrease in the relative abundance of Hylocomium splendens and an increase in Dicranum polysetum. In N Finland, where forests are grazed by semi‐domestic reindeer, we observed a decline in the abundance of Cladina lichens and an increase in Dicranum mosses. Peltigera aphthosa declined throughout the country. Polytrichum juniperinum, Pohlia nutans, and Brachythecium species, which occupy disturbed sites or grow on litter, increased in abundance. The relative abundance of Sphagnum species decreased, particularly in W Finland, where Pleurozium schreberi was favoured. A major decline in S. fuscum was also recorded in C and E Finland. Many of the changes detected in this study are apparently related to intensified forest management; but solely on the basis of this study, its effects cannot be distinguished from those of other large‐scale environmental changes.     

Response of native ungulates to drought in semi‐arid Kenyan rangeland

The distribution and abundance of native ungulates were measured on commercially managed, semi‐arid rangeland in central Kenya over a 3‐year period that encompassed severe drought and above‐average rainfall. Native ungulate biomass density averaged 5282 kg km^?2 over the study and was dominated by elephant (Loxodonta africana), impala (Aepyceros melampus) and dik‐dik (Madoqua kirkii). Biomass density of domestic cattle (Bos taurus) averaged 2280 kg km^?2 during the study. Responses of native ungulates to severe drought were variable. Impala densities were similar to or greater than densities for similar habitat in protected areas, and varied from 12 to 16 km^?2 during and following the drought to 24–29 km^?2 following above‐average rainfall. Dik‐dik densities were also greater than densities reported for protected areas and were surprisingly stable throughout the study despite the wide annual fluctuations in rainfall. Elephant migrated out of the region during drought but were present at high densities (2.9–5.2 km^?2) during wet seasons, consistent with telemetry studies emphasizing the importance of Acacia bushland habitat on commercial rangelands for the migratory portion of the Laikipia–Samburu elephant population. Results show that substantial densities of native browsing and mixed‐feeding ungulates can occur on rangeland managed for commercial beef production and suggest that the capacity for ungulates to move over large spatial scales (>100 km²) and to shift distributions in response to locally variable thunderstorms may be important for sustaining these populations.     

The use of a non‐invasive tool for capture–recapture studies on a seahorse Hippocampus guttulatus population

In this study, the spot pattern in Hippocampus guttulatus was analysed using a computer programme algorithm that allowed individual comparison. This methodology was first tested in a controlled environment using 51 adult and 55 juvenile H. guttulatus. Positive matches were obtained in 86·3 and 83·6% of the adults and juveniles, respectively. In a second experiment, monthly surveys were carried out in five selected locations in the Ria Formosa Lagoon, south Portugal, over the course of a year and a total of 980 photographs were analysed. Photographed H. guttulatus were re‐sighted one to nine times during the course of the survey period with an overall re‐sight record of over 30%. Photo‐identification was therefore shown to be a useful tool for non‐invasive mark–recapture studies that can be successfully used to survey the population abundance of H. guttulatus aged 6 months or older in consecutive years. This could be of great value when considering the assessment of H. guttulatus populations and understanding changes over time.