Palaeohistology reveals a slow pace of life for the dwarfed Sicilian elephant.

The 1-m-tall dwarf elephant Palaeoloxodon falconeri from the Pleistocene of Sicily (Italy) is an extreme example of insular dwarfism and epitomizes the Island Rule. Based on scaling of life-history (LH) traits with body mass, P. falconeri is widely considered to be 'r-selected' by truncation of the growth period, associated with an early onset of reproduction and an abbreviated lifespan. These conjectures are, however, at odds with predictions from LH models for adaptive shifts in body size on islands. To settle the LH strategy of P. falconeri, we used bone, molar, and tusk histology to infer growth rates, age at first reproduction, and longevity. Our results from all approaches are congruent and provide evidence that the insular dwarf elephant grew at very slow rates over an extended period; attained maturity at the age of 15 years; and had a minimum lifespan of 68 years. This surpasses not only the values predicted from body mass but even those of both its giant sister taxon (P. antiquus) and its large mainland cousin (L. africana). The suite of LH traits of P. falconeri is consistent with the LH data hitherto inferred for other dwarfed insular mammals. P. falconeri, thus, not only epitomizes the Island Rule but it can also be viewed as a paradigm of evolutionary change towards a slow LH that accompanies the process of dwarfing in insular mammals.

Nontraditional systems in aging research: an update.

Research on the evolutionary and mechanistic aspects of aging and longevity has a reductionist nature, as the majority of knowledge originates from experiments on a relatively small number of systems and species. Good examples are the studies on the cellular, molecular, and genetic attributes of aging (senescence) that are primarily based on a narrow group of somatic cells, especially fibroblasts. Research on aging and/or longevity at the organismal level is dominated, in turn, by experiments on Drosophila melanogaster, worms (Caenorhabditis elegans), yeast (Saccharomyces cerevisiae), and higher organisms such as mice and humans. Other systems of aging, though numerous, constitute the minority. In this review, we collected and discussed a plethora of up-to-date findings about studies of aging, longevity, and sometimes even immortality in several valuable but less frequently used systems, including bacteria (Caulobacter crescentus, Escherichia coli), invertebrates (Turritopsis dohrnii, Hydra sp., Arctica islandica), fishes (Nothobranchius sp., Greenland shark), reptiles (giant tortoise), mammals (blind mole rats, naked mole rats, bats, elephants, killer whale), and even 3D organoids, to prove that they offer biogerontologists as much as the more conventional tools. At the same time, the diversified knowledge gained owing to research on those species may help to reconsider aging from a broader perspective, which should translate into a better understanding of this tremendously complex and clearly system-specific phenomenon.

Ivory Coast without ivory: Massive extinction of African forest elephants in Côte d'Ivoire.

In pre-colonial and colonial times Côte d'Ivoire probably hosted one of the largest elephant populations in West Africa, resulting in the country's name Côte d'Ivoire (in English Ivory Coast) by French settlers. Numbers declined and by the early 90s it was estimated that the total number of both savannah and forest elephants had reached 63 to 360 elephants in the entire country. Here we present updated information on the distribution and conservation status of forest elephant in Côte d'Ivoire based on multiple sources-dung counts on line transects, records of human-elephant conflict, media reports, sign and interview surveys-obtained during the period 2011-2017. We used Pearson correlation to determine the correlation between the presence of forest elephant and site variables (size of the forest, percentage of area converted into plantation, size of the forest left, size of human population inside the PA, poaching index, distance to the nearest road, population density in the Department, level of protection of the PA). To examine the effect of ecological traits on elephant extirpation, we used Principal Components Analysis (PCA) to check for multicollinearity among variables. Based on dung count elephant presence was confirmed in only 4 of the 25 protected areas surveyed. PAs with higher level of protection have higher probability to be home of elephant population. The viability of these populations is uncertain, since they have a small size and are isolated. Aggressive conservation actions including law enforcement for the protection of their remaining habitat and ranger patrolling are needed to protect the remaining forest elephant populations.

Development of motor control and behaviour in Asian elephants in the Kabini elephant population, southern India.

Although neonates of precocial mammals are capable of locomotory, sensory, nutritional, and thermoregulatory independence to some extent soon after birth, they attain their adult body mass more slowly than altricial mammals, allowing for an extended period of learning or perfecting skills to an adult-like degree. Asian elephants are precocial but are nutritionally dependent on the mother for at least two years and are long-lived and social. We wanted to examine the ontogeny of trunk motor control and various behaviours in Asian elephant calves and see whether the former develops faster than the latter since limb motor control is achieved soon after birth. We collected field data on trunk use, lateralisation, and behaviours from individually identified, free-ranging elephants in southern India and examined how they were affected by age and other factors. Unlike limb motor control, we found trunk motor skills and behaviours to develop gradually with age. Trunk lateralisation occurred very early on, was not highly dependent on trunk motor skills, and is probably not a developmental marker in Asian elephants. Adult-like behaviours that required low trunk usage emerged within 3 months, while some feeding behaviours emerged later. Calves spent less time resting and more time feeding as they grew, and their activity budgets resembled those of adults only after a year; hence, mother-offspring behavioural synchrony was low for young calves and increased with age. Behavioural development and trunk motor control in Asian elephants are both gradual processes, taking about a year to mature.

Demographic and reproductive associations with nematode infection in a long-lived mammal.

Infection by macroparasites, such as nematodes, varies within vertebrate host systems; elevated infection is commonly observed in juveniles and males, and, for females, with different reproductive states. However, while such patterns are widely recognized in short-lived model systems, how they apply to long-lived hosts is comparatively understudied. Here, we investigated how infection varies with host age, sex, and female reproduction in a semi-captive population of individually marked Asian elephants Elephas maximus. We carried out 1,977 faecal egg counts (FECs) across five years to estimate nematode loads for 324 hosts. Infection patterns followed an established age-infection curve, whereby calves (5 years) exhibited the highest FECs and adults (45 years) the lowest. However, males and females had similar FECs across their long lifespan, despite distinct differences in life-history strategy and clear sexual dimorphism. Additionally, although mothers invest two years in pregnancy and a further three to five years into lactation, nematode load did not vary with four different measures of female reproduction. Our results provide a much-needed insight into the host-parasite dynamics of a long-lived host; determining host-specific associations with infection in such systems is important for broadening our knowledge of parasite ecology and provides practical applications for wildlife medicine and management.

Anthropometric and blood data on a hand-reared captive Asian elephant (Elephas maximus) calf: A retrospective case report.

The anthropometric and blood data of an unsuccessfully hand-reared Asian elephant (Elephas maximus) calf were retrospectively compared with the data for calves raised by their real mothers or allomothers, to identify potential reasons for poor outcomes in the hand-reared case. The hand-reared calf grew normally in terms of body weight and withers height. However, blood biochemical data suggested reduced bone metabolism, low immune status, and malnutrition during its life. Blood bone markers were measured to determine whether a skeletal disorder was present in the Asian elephant calf, which was not clear from the anthropometric data. Monitoring these parameters in hand-reared Asian elephant calves, with the aim of keeping them within the normal range, may increase the success rate of hand-rearing of Asian elephant calves.

Weigh and see-Body mass recordings versus body condition scoring in European zoo elephants (Loxodonta africana and Elephas maximus).

Regular body mass (BM) monitoring plays a key role in preventative health care of zoo animals. In some species, including African (Loxodonta africana) and Asian elephants (Elephas maximus), the process of weighing can be challenging, and alternative methods such as visual body condition scoring (BCS) have been developed. We investigated the temporal development of both parameters regarding correlation patterns between them, and their suitability as monitoring measures in dependence of an elephant's life stage. While BM is more suitable in calves and juveniles under the age of 8 years, both BM and BCS are considered equally reliable in adult elephants. In elephants over the age of 40 years, BCS might be more suitable for assessing the physical status. Independent of species and sex, juvenile zoo elephants grow in BM nearly linearly with age, and reach a higher BM at an earlier age compared with conspecifics of free-ranging and semi-captive populations in the countries of origin. The BCS typically remains constant during this life stage, seemingly unaffected by growth. In adult animals, breeding females have a lower BM and BCS than nonbreeders, and BM and BCS typically indicate fluctuations in the same direction. In geriatric elephants (>40 years) a drop in BCS occurs commonly, while BM may even increase in this life stage. We recommend regular body mass recording in zoo elephants to enhance our knowledge of body mass development and allow the formulation of objective practical recommendations. BCS presents a valuable and simple tool for complementary monitoring of an elephant's condition, especially in adult and geriatric individuals.

Savanna elephant numbers are only a quarter of their expected values.

Savannas once constituted the range of many species that human encroachment has now reduced to a fraction of their former distribution. Many survive only in protected areas. Poaching reduces the savanna elephant, even where protected, likely to the detriment of savanna ecosystems. While resources go into estimating elephant populations, an ecological benchmark by which to assess counts is lacking. Knowing how many elephants there are and how many poachers kill is important, but on their own, such data lack context. We collated savanna elephant count data from 73 protected areas across the continent estimated to hold ~50% of Africa's elephants and extracted densities from 18 broadly stable population time series. We modeled these densities using primary productivity, water availability, and an index of poaching as predictors. We then used the model to predict stable densities given current conditions and poaching for all 73 populations. Next, to generate ecological benchmarks, we predicted such densities for a scenario of zero poaching. Where historical data are available, they corroborate or exceed benchmarks. According to recent counts, collectively, the 73 savanna elephant populations are at 75% of the size predicted based on current conditions and poaching levels. However, populations are at <25% of ecological benchmarks given a scenario of zero poaching (~967,000)-a total deficit of ~730,000 elephants. Populations in 30% of the 73 protected areas were <5% of their benchmarks, and the median current density as a percentage of ecological benchmark across protected areas was just 13%. The ecological context provided by these benchmark values, in conjunction with ongoing census projects, allow efficient targeting of conservation efforts.

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.

Trophy Hunting and Sustainability: Temporal Dynamics in Trophy Quality and Harvesting Patterns of Wild Herbivores in a Tropical Semi-Arid Savanna Ecosystem.

The selective nature of trophy hunting may cause changes in desirable phenotypic traits in harvested species. A decline in trophy size of preferred species may reduce hunting destination competitiveness thus compromising the sustainability of trophy hunting as a conservation tool. We explored the trophy quality and trends in harvesting patterns (i.e., 2004-2015) of Cape buffalo (Syncerus caffer), African elephant (Loxodonta africana), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in Matetsi Safari Area, northwest Zimbabwe. We used long-term data on horn and tusk size, age, quota size allocation and offtake levels of selected species. To analyse the effect of year, area and age on the trophy size, quota size and offtake levels, we used linear mixed models. One sample t-test was used to compare observed trophy size with Safari Club International (SCI) minimum score. Trophy sizes for Cape buffalo and African elephant were below the SCI minimum score. Greater kudu trophy sizes were within the minimum score threshold whereas sable trophy sizes were above the SCI minimum score between 2004 and 2015. Age at harvest for Cape buffalo, kudu and sable increased whilst that of elephant remained constant between 2004 and 2015. Quota size allocated for buffalo and the corresponding offtake levels declined over time. Offtake levels of African elephant and Greater kudu declined whilst the quota size did not change between 2004 and 2015. The quota size for sable increased whilst the offtake levels fluctuated without changing for the period 2004-2015. The trophy size and harvesting patterns in these species pose a conservation and management dilemma on the sustainability of trophy hunting in this area. We recommend: (1) temporal and spatial rotational resting of hunting areas to create refuge to improve trophy quality and maintenance of genetic diversity, and (2) introduction of variable trophy fee pricing system based on trophy size.

Conserving wildlife amongst the cotton fields. A third of a century of experience at the Nazinga Game Ranch, Burkina Faso.

Established in the early 1970 as a participatory wildlife production area, the Nazinga Game Ranch turned into an island of conservation surrounded by cultivation. We asked ourselves how long-term ungulate trends are affected in a context of continuous human pressure. To find out, we compiled and analysed the data of yearly line-transect counts of mammals carried out since 1985. Results showed that large species such as the elephant (Loxodonta africana) and large antelopes increased or showed stable populations. In contrast, medium and small ungulates showed continuously decreasing trends. During the same period, rainfall, water availability from artificial water points and the crop encroaching outside Nazinga Game Ranch increased. After an initial significant reduction, illegal human signs increased. However, we showed that human signs were positively correlated with the abundance of large ungulates but negatively correlated with the abundance of medium and small ones. In conclusion, this study showed that some isolated mammal populations could be restored and maintained in the long term, in spite of being surrounded by highly cultivated areas.

Distinguishing between determinate and indeterminate growth in a long-lived mammal.

BACKGROUND: The growth strategy of a species influences many key aspects of its life-history. Animals can either grow indeterminately (throughout life), or grow determinately, ceasing at maturity. In mammals, continued weight gain after maturity is clearly distinguishable from continued skeletal growth (indeterminate growth). Elephants represent an interesting candidate for studying growth because of their large size, long life and sexual dimorphism. Objective measures of their weight, height and age, however, are rare. RESULTS: We investigate evidence for indeterminate growth in the Asian elephant Elephas maximus using a longitudinal dataset from a semi-captive population. We fit growth curves to weight and height measurements, assess sex differences in growth, and test for indeterminate growth by comparing the asymptotes for height and weight curves. Our results show no evidence for indeterminate growth in the Asian elephant; neither sex increases in height throughout life, with the majority of height growth completed by the age of 15 years in females and 21 years in males. Females show a similar pattern with weight, whereas males continue to gain weight until over age 50. Neither sex shows any declines in weight with age. CONCLUSIONS: These results have implications for understanding mammalian life-history, which could include sex-specific differences in trade-offs between size and reproductive investment.

The progression of small-follicle reserves in the ovaries of wild African elephants (Loxodonta africana) from puberty to reproductive senescence.

This study aimed to determine whether the follicle reserve in the ovary of the African elephant declines progressively after puberty and whether its depletion constrains the fertility of older females. Elephant ovaries were fixed in 4% neutral buffered formalin and small-follicle counts made using stereological protocols. Excepting a slight rise in small-follicle numbers between 16 and 25 years of age, there was a trend for follicle numbers to fall from puberty to 70 years. Reproductive status did not impact significantly on small-follicle numbers (P=0.31). The number of early primary follicles, initially higher in number than true primary follicles, fell from post-puberty to nil at 45 years of age. Six of the seven oldest animals in the study showed signs of recent ovarian activity in the form of antral follicles, corpora lutea or large corpora nigra. The four oldest elephants (mean age 69 years) had a median small-follicle count of 11,113. In summary, it appears that the elephant ovary is capable of supplying oocytes for ovulation right up to the time of death at the age of maximum life expectancy, although the follicle reserve becomes depleted in some older elephants.

Enduring consequences of early experiences: 40 year effects on survival and success among African elephants (Loxodonta africana).

Growth from conception to reproductive onset in African elephants (Loxodonta africana) provides insights into phenotypic plasticity, individual adaptive plastic responses and facultative maternal investment. Using growth for 867 and life histories for 2652 elephants over 40 years, we demonstrate that maternal inexperience plus drought in early life result in reduced growth rates for sons and higher mortality for both sexes. Slow growth during early lactation was associated with smaller adult size, later age at first reproduction, reduced lifetime survival and consequently limited reproductive output. These enduring effects of trading slow early growth against immediate survival were apparent over the very long term; delayed downstream consequences were unexpected for a species with a maximum longevity of 70+ years and unpredictable environmental experiences.

Unmanned aerial survey of elephants.

The use of a UAS (Unmanned Aircraft System) was tested to survey large mammals in the Nazinga Game Ranch in the south of Burkina Faso. The Gatewing ×100™ equipped with a Ricoh GR III camera was used to test animal reaction as the UAS passed, and visibility on the images. No reaction was recorded as the UAS passed at a height of 100 m. Observations, made on a set of more than 7000 images, revealed that only elephants (Loxodonta africana) were easily visible while medium and small sized mammals were not. The easy observation of elephants allows experts to enumerate them on images acquired at a height of 100 m. We, therefore, implemented an aerial strip sample count along transects used for the annual wildlife foot count. A total of 34 elephants were recorded on 4 transects, each overflown twice. The elephant density was estimated at 2.47 elephants/km(2) with a coefficient of variation (CV%) of 36.10%. The main drawback of our UAS was its low autonomy (45 min). Increased endurance of small UAS is required to replace manned aircraft survey of large areas (about 1000 km of transect per day vs 40 km for our UAS). The monitoring strategy should be adapted according to the sampling plan. Also, the UAS is as expensive as a second-hand light aircraft. However the logistic and flight implementation are easier, the running costs are lower and its use is safer. Technological evolution will make civil UAS more efficient, allowing them to compete with light aircraft for aerial wildlife surveys.

Elephant transcriptome provides insights into the evolution of eutherian placentation.

The chorioallantoic placenta connects mother and fetus in eutherian pregnancies. In order to understand the evolution of the placenta and provide further understanding of placenta biology, we sequenced the transcriptome of a term placenta of an African elephant (Loxodonta africana) and compared these data with RNA sequence and microarray data from other eutherian placentas including human, mouse, and cow. We characterized the composition of 55,910 expressed sequence tag (i.e., cDNA) contigs using our custom annotation pipeline. A Markov algorithm was used to cluster orthologs of human, mouse, cow, and elephant placenta transcripts. We found 2,963 genes are commonly expressed in the placentas of these eutherian mammals. Gene ontology categories previously suggested to be important for placenta function (e.g., estrogen receptor signaling pathway, cell motion and migration, and adherens junctions) were significantly enriched in these eutherian placenta-expressed genes. Genes duplicated in different lineages and also specifically expressed in the placenta contribute to the great diversity observed in mammalian placenta anatomy. We identified 1,365 human lineage-specific, 1,235 mouse lineage-specific, 436 cow lineage-specific, and 904 elephant-specific placenta-expressed (PE) genes. The most enriched clusters of human-specific PE genes are signal/glycoprotein and immunoglobulin, and humans possess a deeply invasive human hemochorial placenta that comes into direct contact with maternal immune cells. Inference of phylogenetically conserved and derived transcripts demonstrates the power of comparative transcriptomics to trace placenta evolution and variation across mammals and identified candidate genes that may be important in the normal function of the human placenta, and their dysfunction may be related to human pregnancy complications.

How immunocontraception can contribute to elephant management in small, enclosed reserves: Munyawana population as a case study.

Immunocontraception has been widely used as a management tool to reduce population growth in captive as well as wild populations of various fauna. We model the use of an individual-based rotational immunocontraception plan on a wild elephant, Loxodonta africana, population and quantify the social and reproductive advantages of this method of implementation using adaptive management. The use of immunocontraception on an individual, rotational basis stretches the inter-calving interval for each individual female elephant to a management-determined interval, preventing exposing females to unlimited long-term immunocontraception use (which may have as yet undocumented negative effects). Such rotational immunocontraception can effectively lower population growth rates, age the population, and alter the age structure. Furthermore, such structured intervention can simulate natural process such as predation or episodic catastrophic events (e.g., drought), which regulates calf recruitment within an abnormally structured population. A rotational immunocontraception plan is a feasible and useful elephant population management tool, especially in a small, enclosed conservation area. Such approaches should be considered for other long-lived, social species in enclosed areas where the long-term consequences of consistent contraception may be unknown.

From flat foot to fat foot: structure, ontogeny, function, and evolution of elephant "sixth toes".

Several groups of tetrapods have expanded sesamoid (small, tendon-anchoring) bones into digit-like structures ("predigits"), such as pandas' "thumbs." Elephants similarly have expanded structures in the fat pads of their fore- and hindfeet, but for three centuries these have been overlooked as mere cartilaginous curiosities. We show that these are indeed massive sesamoids that employ a patchy mode of ossification of a massive cartilaginous precursor and that the predigits act functionally like digits. Further, we reveal clear osteological correlates of predigit joint articulation with the carpals/tarsals that are visible in fossils. Our survey shows that basal proboscideans were relatively "flat-footed" (plantigrade), whereas early elephantiforms evolved the more derived "tip-toed" (subunguligrade) morphology, including the predigits and fat pad, of extant elephants. Thus, elephants co-opted sesamoid bones into a role as false digits and used them for support as they changed their foot posture.

Age determination by back length for African savanna elephants: extending age assessment techniques for aerial-based surveys.

Determining the age of individuals in a population can lead to a better understanding of population dynamics through age structure analysis and estimation of age-specific fecundity and survival rates. Shoulder height has been used to accurately assign age to free-ranging African savanna elephants. However, back length may provide an analog measurable in aerial-based surveys. We assessed the relationship between back length and age for known-age elephants in Amboseli National Park, Kenya, and Addo Elephant National Park, South Africa. We also compared age- and sex-specific back lengths between these populations and compared adult female back lengths across 11 widely dispersed populations in five African countries. Sex-specific Von Bertalanffy growth curves provided a good fit to the back length data of known-age individuals. Based on back length, accurate ages could be assigned relatively precisely for females up to 23 years of age and males up to 17. The female back length curve allowed more precise age assignment to older females than the curve for shoulder height does, probably because of divergence between the respective growth curves. However, this did not appear to be the case for males, but the sample of known-age males was limited to ≤27 years. Age- and sex-specific back lengths were similar in Amboseli National Park and Addo Elephant National Park. Furthermore, while adult female back lengths in the three Zambian populations were generally shorter than in other populations, back lengths in the remaining eight populations did not differ significantly, in support of claims that growth patterns of African savanna elephants are similar over wide geographic regions. Thus, the growth curves presented here should allow researchers to use aerial-based surveys to assign ages to elephants with greater precision than previously possible and, therefore, to estimate population variables.