Behavioral responses of terrestrial mammals to COVID-19 lockdowns.

COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals' 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.

Mapping potential connections between Southern Africa's elephant populations.

Southern Africa spans nearly 7 million km2 and contains approximately 80% of the world's savannah elephants (Loxodonta africana) mostly living in isolated protected areas. Here we ask what are the prospects for improving the connections between these populations? We combine 1.2 million telemetry observations from 254 elephants with spatial data on environmental factors and human land use across eight southern African countries. Telemetry data show what natural features limit elephant movement and what human factors, including fencing, further prevent or restrict dispersal. The resulting intersection of geospatial data and elephant presences provides a map of suitable landscapes that are environmentally appropriate for elephants and where humans allow elephants to occupy. We explore the environmental and anthropogenic constraints in detail using five case studies. Lastly, we review all the major potential connections that may remain to connect a fragmented elephant metapopulation and document connections that are no longer feasible.

State-space models reveal a continuing elephant poaching problem in most of Africa.

The most comprehensive data on poaching of African elephants comes from the Monitoring the Illegal Killing of Elephants (MIKE) program, which reports numbers of illegally killed carcasses encountered by rangers. Recent studies utilizing MIKE data have reported that poaching of African elephants peaked in 2011 and has been decreasing through 2018. Closer examination of these studies, however, raises questions about the conclusion that poaching is decreasing throughout the continent. To provide more accurate information on trends in elephant poaching, we analyzed MIKE data using state-space models. State-space models account for missing data and the error inherent when sampling carcasses. Using the state-space model, for 2011-2018, we found no significant temporal trends in rates of illegal killing for Southern, Central and Western Africa. Only in Eastern Africa have poaching rates decreased substantially since 2011. For Africa as a whole, poaching did decline for 2011-2018, but the decline was entirely due to Eastern African sites. Our results suggest that poaching for ivory has not diminished across most of Africa since 2011. Continued vigilance and anti-poaching efforts will be necessary to combat poaching and to conserve African elephants.

Drone-based effective counting and ageing of hippopotamus (Hippopotamus amphibius) in the Okavango Delta in Botswana.

Accurately estimating hippopotamus (Hippopotamus amphibius) numbers is difficult due to their aggressive nature, amphibious lifestyle, and habit of diving and surfacing. Traditionally, hippos are counted using aerial surveys and land/boat surveys. We compared estimates of numbers of hippos in a lagoon in the Okavango Delta, counted from land to counts from video taken from a DJI Phantom 4TM drone, testing for effectiveness at three heights (40 m, 80 m, and 120 m) and four times of day (early morning, late morning, early afternoon, and late afternoon). In addition, we determined effectiveness for differentiating age classes (juvenile, subadult, and adult), based on visual assessment and measurements from drone images, at different times and heights. Estimates in the pool averaged 9.18 (± 0.25SE, range 1-14, n = 112 counts). Drone counts at 40 m produced the highest counts of hippos, 10.6% higher than land counts and drone counts at 80 m, and 17.6% higher than drone counts at 120 m. Fewer hippos were counted in the early morning, when the hippos were active and most likely submerged, compared to all other times of day, when they tended to rest in shallow water with their bodies exposed. We were able to assign age classes to similar numbers of hippos from land counts and counts at 40 m, although land counts were better at identifying juveniles and subadults. Early morning was the least effective time to age hippos given their active behaviour, increasingly problematic with increasing height. Use of a relatively low-cost drone provided a rigorous and repeatable method for estimating numbers and ages of hippos, other than in the early morning, compared to land counts, considered the most accurate method of counting hippos.

Evidence of a Growing Elephant Poaching Problem in Botswana.

Botswana holds roughly one-third of Africa's remaining savannah elephants (Loxodonta africana) [1, 2] and will play a key role in the future conservation of this species. To date, Botswana has been one of the safest countries for elephants, with little poaching reported [3]. Here, we present evidence of a new outbreak of elephant poaching for ivory in northern Botswana. Comparing results from 2014 and 2018 aerial surveys, we found that elephant populations were stable, but numbers of elephant carcasses have increased, especially for newer carcasses dead for less than roughly 1 year. Newer carcasses were clustered in five "hotspots" averaging 3,522 km2 in area. We compared elephant populations in hotspots to the immediately surrounding areas and found that since 2014, elephants have decreased by 16% in hotspots but increased by 10% in surrounding areas. Numbers of "old" carcasses, dead for more than 1 year, increased by 78% in hotspots between 2014 and 2018 but decreased by 3% in surrounding areas. To verify that poaching has been occurring, we used helicopters to visit 148 elephant carcasses and assess their cause of death. We confirmed poaching for all 72 newer carcasses assessed. We also confirmed poaching for 62 of 76 (82%) carcasses older than 1 year, primarily in one hotspot. Poached older carcasses were all males aged 30-60 and likely killed for their large tusks. This evidence suggests that ivory poaching on the scale of hundreds of elephants per year has been occurring in northern Botswana since 2017 or possibly earlier.

Partial migration in savanna elephant populations distributed across southern Africa.

Migration is an important, but threatened ecological process. Conserving migration requires the maintenance of functional connectivity across sufficiently large areas. Therefore, we need to know if, where and why species migrate. Elephants are highly mobile and can travel long distances but we do not know if they migrate. Here, we analysed the movement trajectories of 139 savanna elephants (Loxodonta africana) within eight clusters of protected areas across southern Africa to determine if elephants migrate, and if so, where, how and why they migrate. Only 25 of these elephants migrated. Elephants are a facultative partially migratory species, where only some individuals in a population migrate opportunistically, and not every year. Elephants migrated between distinct seasonal ranges corresponding to southern Africa's dry and wet seasons. The timing of wet season migrations was associated with the onset of rainfall and the subsequent greening up of forage. Conversely, the duration, distance, and the timing of dry season migrations varied idiosyncratically. The drivers of elephant migration are likely a complex interaction between individual traits, density, and the distribution and availability of resources. Despite most migrations crossing administrative boundaries, conservation networks provided functional space for elephants to migrate.

Poaching and human encroachment reverse recovery of African savannah elephants in south-east Angola despite 14 years of peace.

With populations of African savannah elephants (Loxodonta africana) declining across the continent, assessing the status of individual elephant populations is important for conservation. Angola's elephant population represents a key linkage between the larger populations of Namibia and Botswana. Elephants in Angola were decimated during the 1975-2002 Angolan civil war, but a 2005 survey showed that populations were recolonizing former habitats. Between 2005 and 2015, no research was permitted on elephants in Angola, but elsewhere in Africa many elephant populations experienced a poaching crisis. In 2015, we were able to resume elephant research in Angola. We used aerial surveys and satellite monitoring of collared elephants to determine the current status of elephant populations in Angola and to learn how human populations may be affecting elephant habitat usage. The aerial survey revealed a population of 3,395 ± SE of 797 elephants, but populations had declined 21% from the 2005 estimate. The high number of carcasses observed on the survey suggests that populations may have increased after the 2005 survey but were declining rapidly as of 2015. Satellite-collared elephants avoided areas <6 km from human indicators but preferred areas nearer humans at scales of 6-40 km, suggesting that humans may be displacing elephants from preferred habitats near rivers. Taken together, these results suggest that Angola's elephant population is experiencing intense poaching and may be losing habitat to human settlements. Without action to conserve their populations, Angola's elephants face an uncertain future.

Inactivity/sleep in two wild free-roaming African elephant matriarchs - Does large body size make elephants the shortest mammalian sleepers?

The current study provides details of sleep (or inactivity) in two wild, free-roaming African elephant matriarchs studied in their natural habitat with remote monitoring using an actiwatch subcutaneously implanted in the trunk, a standard elephant collar equipped with a GPS system and gyroscope, and a portable weather station. We found that these two elephants were polyphasic sleepers, had an average daily total sleep time of 2 h, mostly between 02:00 and 06:00, and displayed the shortest daily sleep time of any mammal recorded to date. Moreover, these two elephants exhibited both standing and recumbent sleep, but only exhibited recumbent sleep every third or fourth day, potentially limiting their ability to enter REM sleep on a daily basis. In addition, we observed on five occasions that the elephants went without sleep for up to 46 h and traversed around 30 km in 10 h, possibly due to disturbances such as potential predation or poaching events, or a bull elephant in musth. They exhibited no form of sleep rebound following a night without sleep. Environmental conditions, especially ambient air temperature and relative humidity, analysed as wet-bulb globe temperature, reliably predict sleep onset and offset times. The elephants selected novel sleep sites each night and the amount of activity between sleep periods did not affect the amount of sleep. A number of similarities and differences to studies of elephant sleep in captivity are noted, and specific factors shaping sleep architecture in elephants, on various temporal scales, are discussed.

Testing the Accuracy of Aerial Surveys for Large Mammals: An Experiment with African Savanna Elephants (Loxodonta africana).

Accurate counts of animals are critical for prioritizing conservation efforts. Past research, however, suggests that observers on aerial surveys may fail to detect all individuals of the target species present in the survey area. Such errors could bias population estimates low and confound trend estimation. We used two approaches to assess the accuracy of aerial surveys for African savanna elephants (Loxodonta africana) in northern Botswana. First, we used double-observer sampling, in which two observers make observations on the same herds, to estimate detectability of elephants and determine what variables affect it. Second, we compared total counts, a complete survey of the entire study area, against sample counts, in which only a portion of the study area is sampled. Total counts are often considered a complete census, so comparing total counts against sample counts can help to determine if sample counts are underestimating elephant numbers. We estimated that observers detected only 76% ± SE of 2% of elephant herds and 87 ± 1% of individual elephants present in survey strips. Detectability increased strongly with elephant herd size. Out of the four observers used in total, one observer had a lower detection probability than the other three, and detectability was higher in the rear row of seats than the front. The habitat immediately adjacent to animals also affected detectability, with detection more likely in more open habitats. Total counts were not statistically distinguishable from sample counts. Because, however, the double-observer samples revealed that observers missed 13% of elephants, we conclude that total counts may be undercounting elephants as well. These results suggest that elephant population estimates from both sample and total counts are biased low. Because factors such as observer and habitat affected detectability of elephants, comparisons of elephant populations across time or space may be confounded. We encourage survey teams to incorporate detectability analysis in all aerial surveys for mammals.

Continent-wide survey reveals massive decline in African savannah elephants.

African elephants (Loxodonta africana) are imperiled by poaching and habitat loss. Despite global attention to the plight of elephants, their population sizes and trends are uncertain or unknown over much of Africa. To conserve this iconic species, conservationists need timely, accurate data on elephant populations. Here, we report the results of the Great Elephant Census (GEC), the first continent-wide, standardized survey of African savannah elephants. We also provide the first quantitative model of elephant population trends across Africa. We estimated a population of 352,271 savannah elephants on study sites in 18 countries, representing approximately 93% of all savannah elephants in those countries. Elephant populations in survey areas with historical data decreased by an estimated 144,000 from 2007 to 2014, and populations are currently shrinking by 8% per year continent-wide, primarily due to poaching. Though 84% of elephants occurred in protected areas, many protected areas had carcass ratios that indicated high levels of elephant mortality. Results of the GEC show the necessity of action to end the African elephants' downward trajectory by preventing poaching and protecting habitat.

A Comparison of Walking Rates Between Wild and Zoo African Elephants.

With increased scrutiny surrounding the welfare of elephants in zoological institutions, it is important to have empirical evidence on their current welfare status. If elephants are not receiving adequate exercise, it could lead to obesity, which can lead to many issues including acyclicity and potentially heart disease. The goal of the current study was to compare the walking rates of elephants in the wild versus elephants in zoos to determine if elephants are walking similar distances relative to their wild counterparts. Eleven wild elephants throughout different habitats and locations in Botswana were compared to 8 elephants at the San Diego Zoo Safari Park. Direct comparisons revealed no significant difference in average walking rates of zoo elephants when compared with wild elephants. These results suggest that elephants at the San Diego Zoo Safari Park walk similar rates to those of wild elephants and may be meeting their exercise needs.