An assessment of the maximum sustainable yield of ivory from African elephant populations.

A general, logistic population model is used to explore the dynamics of harvested elephant populations. The model includes two features peculiar to elephant populations and the harvesting of ivory. First, because of the shape of the growth curve of tusks with age, the conversion factor that relates the number of elephants killed to the ivory yield in weight is not constant, but a function of the population size. Second, tusks from animals that die from natural causes can be retrieved and included in the total yield of ivory. The implications of the relationship between tusk size and age of an animal on the maximum sustainable yield in terms of ivory tonnage and in terms of the number of tusks are explored. The nonequilibrium implications of the tusk growth curve on the population dynamics under different harvesting strategies are also investigated. Results indicate that the maximum sustainable yield is achieved at very low harvest rates with population levels close to the pristine equilibrium. When tusks from animals that die of natural causes are included in the harvest, the maximum yield may, depending on the mortality and recruitment parameters, occur when there is no direct harvest.     

Sexual Dimorphism and Inter-Generic Variation in Proboscidean Tusks: Multivariate Assessment of American Mastodons (Mammut americanum) and Extant African Elephants

Characteristics of social structure, mating strategies, and parental investment can be inferred for mammalian species based on degree of sexual dimorphism, especially when males are substantially larger than females. American mastodons (Mammut americanum) and extant African elephants (Loxodonta africana, Loxodonta cyclotis) both exhibit marked dimorphism in tusk size. To evaluate the hypothesis that this dimorphism might be indicative of similar, and possibly conserved, behavioral patterns in each lineage, we undertook a detailed evaluation of the pattern of tusk growth in these two genera. Separate discriminant function analyses (DFA) of 21 adult mastodon tusks of inferred sex and 48 adult elephant tusks of known sex show that patterns of ontogenetic change in tusk circumference, regardless of genus, effectively discriminate between sexes. Canonical variates analysis (CVA) of tusks from male and female mastodons and male and female elephants shows that male tusks in both genera are larger than female tusks across all measurements, especially for maximum tusk circumference and pulp cavity depth. CVA’s emphasis of inter-group differences in tusk morphology also shows that mastodon tusks are more robust than elephant tusks. Overall, this study illustrates that there is a characteristic male and a characteristic female tusk form shared by elephants and mastodons. This shows that elephants and mastodons display a shared syndrome of traits beyond sex-linked differences in tusk size, supporting the hypothesis that mastodons exhibited behaviors similar to those we observe today in African elephants.     

THE COLLECTION OF FOUND IVORY IN MURCHISON FALLS NATIONAL PARK, UGANDA

The paper concerns the circumstances surrounding the collection of ivory from dead elephants, with particular reference to Murchison Falls National Park. The characteristics of the interval between death and complete disintegration of an elephant are described. These, combined with observations of known age skeletons, comprised the criteria used in classifying skeletons found from the air into three relative age classes. Average annual mortality is estimated for the population north of the Nile (MFPN) at 147 animals yielding 1945 kg of ivory, and for that south of the Nile (MFPS) at 474 animals yielding 7497 kg ivory. Park-found ivory records are analyzed for the 11 y 1959–69. The expected age distribution of deaths is compared with the observed. For MFPN a bias in favour of large (male) tusks is present, explicable by the concentration of ranger search effort in areas of known high male density. For MFPS a bias towards small tusks is thought to be caused by elephants wounded outside the park dying inside it. The National Park recovers an average of 27.6 % of its available ivory per annum, with large annual fluctuations probably correlated with the incidence of wounding outside the park. High losses to poachers are evident. An aerial search for ivory showed a tendency for elephants to die near watercourses. A finding rate of one carcass every 4.3 km of watercourse was obtained. As only 5 % of carcasses still had tusks the aerial searching was prematurely terminated. The results indicated a finding efficiency of 26.4% of the available current year carcasses. Comparative costing suggests that ground searching would be a more efficient method of finding ivory than aerial searching. The high value of the available ivory in Murchison and other areas justifies intensive searching. The low collection rates prevailing in East Africa are largely attributable to the absence of appropriate search efforts.     

Body scarring as an indicator of social function of dugong (Dugong dugon) tusks

The fully aquatic lifestyle of dugongs means that direct observation of social tusk use is not usually possible. This study used body scarring as an indicator of tusk function by males. Tusk rake scars on 298 live wild dugongs, of both sexes and all sizes, were categorized and counted in over 1,000 photographs, and examined in relation to maturity and reproductive activity over seasons. All dugongs had tusk scars, but adults were the main recipients. Sexually active adults acquired the greatest number of fresh tusk wounds during the mating season. Subadults received fresh rakes at similar numbers year-round. Adult males had more scars on the mid and posterior dorsum, indicating that males direct combative force to these regions of the male body when competing for females. Adult females had heaviest scarring and more tusk puncture wounds on the anterior-mid dorsum and head, suggesting that male dugongs use tusks in sexual coercion. Heavy scarring sustained by solitary calves compared to dependent ones, suggests that mothers afford some protection. Body scarring caused by tusks may serve as an indicator of reproductive contribution of the recipients, providing that successful males are involved in more reproductive competitions, and successful females in more mating events.     

Trend in Pacific walrus (Odobenus rosmarus divergens) tusk asymmetry, 1990–2014

We used the basal circumference of Pacific walrus (Odobenus rosmarus divergens) tusks (upper canine teeth, n = 21,068 pairs) to estimate fluctuating asymmetry (FA1 index) from 1990 to 2014. The mean difference in circumference between paired tusks was ?0.006 (SEM = 0.002) cm and approximately normally distributed. Measurement error was 0.6 (0.02)%, similar between biologists and lay persons (P = 0.83), and ≤15% of FA1. Tusk FA1 was greatest in 1990 then declined by 56% (P = 0.0001) through 2014. Male and female trends differed (P = 0.0001) and male FA1 was 40% greater (P = 0.0001) and the rate of decline 28% steeper (P = 0.3) than females. A quartic polynomial model (r² = 0.66, w_i = 0.685) fit the trend for female data better than simpler forms, whereas a linear model (r² = 0.55, w_i = 0.693) was a better fit for male data. Walrus tusk FA1 reflected periods when the population was stressed due to food limitations and then recovered, and perhaps when females began to experience the loss of preferred sea ice habitat in summer and FA1 is an easily monitored indicator. More work is needed to confirm the link between FA1, individual fitness, and adaptive potential.     

西双版纳三岔河地区野生亚洲象的个体识别、种群数量和活动特点

2004 年10 月至2005 年10 月,在云南西双版纳三岔河野象谷用直接观察、拍照、摄像等方法直接记录野生亚洲象的形态学特征,通过耳、门齿、背、尾、疤痕、面部骨骼等特征对经过本地区的象群和独象进行个体识别并建立个体信息库。直接观测记录野象524 头次,通过照片资料和访问间接记录289 头次,识别为46 头个体,由7 个家族(庭)群共38 头和8 头单独活动的雄象组成;发现象群和独象的活动具有季节性;不同象群在野象谷的活动频次差异显著;象群大小和其在本地区的活动频率之间呈明显的正相关。     

When does selective hunting select,how can we tell,and what should we do about it?

Potential evolutionary consequences of selective hunting of mammals are controversial because of limited evidence and important socio‐economic impacts. Several ecological and management variables facilitate evolutionary responses to selection for horn, tusk or antler size, including strong selective hunting pressure; harvest of males with large horns, tusks or antlers before they can breed; unavailable or ineffective sources of unselected immigrants; and age‐dependent relationships between horn, tusk or antler size and male mating success. Plastic responses of male horns, tusks and antlers to environment are probably more common than evolutionary changes. Evidence for evolutionary effects of selective hunting is strong for large mammals where biological characteristics and hunting regulations combine to favour them.     

Growth of the African elephant (Loxodonta africana)

The three-stage desk calculation of the von Bertalanffy equation to describe growth in height and weight with age in the elephant is compared with a new approach to calculating the three coefficients in the function by a computer. The two methods give different results with respect to the weight/age calculations. Theoretical von Bertalanffy equations calculated by both methods to describe growth in height and weight with age in the African elephant in Zambia are compared with previously published equations for the elephant in East Africa. Details are given of growth in height in two known-age African elephants, a female ‘Diksie’ and a male ‘Kartoum’. Theoretical growth in height curves for the female African and Asiatic elephant are compared. The coefficients K and / for growth in height are not transferable to the growth in weight equations. Inherent inaccuracies in the calculation of the coefficients in the von Bertalanffy equation are discussed, and it is concluded that in animals which have a long life-span such as the elephant, the equation serves as a purely empirical representation of weight-at-age data and that there is little biological significance in the parameters it contains. The computer-calculated curves give the best fit to the data. The regression of log age on log shoulder height from 2–20 years of age provides a more realistic approach to comparative growth studies. The increase in adrenal weight with age is linear. Tusk growth in relation to age and sex in Zambia is compared with East Africa. It is concluded that the tusks in Zambia are smaller and are more difficult to sex correctly than their East African counterparts, possibly a consequence of the Zambian elephant having a greater degree of tusk wear. Allometric growth is described with emphasis on the estimation of body weight from shoulder height. The most reliable estimates are obtained from a purely empirical representation of the data, a semilog plot of log body weight on shoulder height.     

The mechanical design of the tusk of the narwhal (Monodon nonoceros: Cetacea)

The mechanical properties of the tissues of the tusk of a narwhal Monodon monoceros , their histological arrangement, and the gross anatomy of the tusk were examined. The histology of the tusk shows various toughening mechanisms. The tusk tissues are of low stiffness, but are very tough, suiting the tusks for loading on impact, but not for longitudinal loading. These mechanical properties suggest the possible functions of the tusk, namely to be long in order to be sexually attractive, while being tough in order to withstand 'testing' blows from other males.     

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.     

Devastating Decline of Forest Elephants in Central Africa

African forest elephants– taxonomically and functionally unique–are being poached at accelerating rates, but we lack range-wide information on the repercussions. Analysis of the largest survey dataset ever assembled for forest elephants (80 foot-surveys; covering 13,000 km; 91,600 person-days of fieldwork) revealed that population size declined by ca. 62% between 2002–2011, and the taxon lost 30% of its geographical range. The population is now less than 10% of its potential size, occupying less than 25% of its potential range. High human population density, hunting intensity, absence of law enforcement, poor governance, and proximity to expanding infrastructure are the strongest predictors of decline. To save the remaining African forest elephants, illegal poaching for ivory and encroachment into core elephant habitat must be stopped. In addition, the international demand for ivory, which fuels illegal trade, must be dramatically reduced.     

African elephants show high levels of interest in the skulls and ivory of their own species

An important area of biology involves investigating the origins in animals of traits that are thought of as uniquely human. One way that humans appear unique is in the importance they attach to the dead bodies of other humans, particularly those of their close kin, and the rituals that they have developed for burying them. In contrast, most animals appear to show only limited interest in the carcasses or associated remains of dead individuals of their own species. African elephants (Loxodonta africana) are unusual in that they not only give dramatic reactions to the dead bodies of other elephants, but are also reported to systematically investigate elephant bones and tusks that they encounter, and it has sometimes been suggested that they visit the bones of relatives. Here, we use systematic presentations of object arrays to demonstrate that African elephants show higher levels of interest in elephant skulls and ivory than in natural objects or the skulls of other large terrestrial mammals. However, they do not appear to specifically select the skulls of their own relatives for investigation so that visits to dead relatives probably result from a more general attraction to elephant remains.     

Structure of population,grouping patterns and density of forest elephants in north-west Congo

The structure of the population of forest elephants visiting a clearing in north-west Congo was studied over 8 months. Out of 3314 sightings, 629 elephants were identified (including 64%, of adults). The sex ratio was about 1:1. The number of offspring per female was 1.12. Solitary elephants (91.4%, of males) made up 35% of the population. Of females, 93.6% were grouped. Groups (mean size 3.5) included females and offspring (3.4), males and females (5.0), or males only (2.3). The return rate of elephants at the clearing suggested that up to 1900 individuals have visited the clearing. This confirms that elephant densities are especially high in north Congo.     

The African savannah elephant population in Kidepo Valley National Park, Uganda: changes in size and structure from 1967 to 2000

The age and sex structure of the elephant population in Kidepo Valley National Park were studied using recognition files. Moreover, population trends were reviewed using past studies. From 1967 to 2000, the elephant population varied between 200 and 500 individuals. Of the minimum number of 374 elephants found to use the Park, 352 were individually identified. Seventy‐nine per cent of these were recorded in 29 families ranging from three to 22 animals with a mean group size of ten. Young animals dominated the population (45% aged 0–9.9 years), while 11% were 10–14.9 years of age and only 18% over 25 years of age. Overall, males constituted 45% of the population, but only 32% of the individuals in the 10–14.9‐year age class were males. Individuals ≥15 years of age displayed a skewed sex ratio of 1 : 1.4 in favour of females. The extent of skew was greatest among mature animals (≥25 years of age), which had four times as many females as males. The skewed age structure in the year 2000 caused by poaching and drought, led to a significant difference between the age distribution in 1970 and 2000. Recent counts suggest that the Kidepo elephant population may be increasing.     

The influence of increased line-fishing mortality on the sex ratio and age of sex reversal of the venus tusk fish

The age of sex reversal of the venus tusk fish Choerodon venustus , caught by line fishing at various locations on the southern Great Barrier Reef, indicated that C. venustus is capable of modifying its life cycle in response to increased mortality. The evidence suggests Masthead Reef fish, which experience the highest mortality, underwent sex reversal at a smaller size and younger age than at the other sites. The largest female fish, sexually transitional fish and males were smaller at Masthead Reef than at the Swains Reefs or One Tree Reef at Masthead Reef. There was also considerable overlap in the size of males and females within the exploited populations indicating that sex reversal is not initiated at a particular length but may have a social cause. The sex ratio of fish was essentially the same for fish fully susceptible to line fishing in the Swains and Masthead samples. Circumstantial evidence suggested that the absence of large males in a population may initiate sex reversal, indicating the maintenance of a constant sex ratio may have a social basis.     

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.     

Employing participatory surveys to monitor the illegal killing of elephants across diverse land uses in Laikipia–Samburu,Kenya

Levels and trends of illegal killing of elephants are measured by the Convention on International Trade in Endangered Species (CITES) Monitoring the Illegal Killing of Elephants (MIKE) programme in sites across Africa and Asia. In the mostly unprotected Laikipia–Samburu MIKE site in northern Kenya, elephant mortality data were collected using both standard law enforcement monitoring procedures, relying on patrolling, and participatory methods involving local communities. Qualitatively, traditional patrolling techniques were more successful in protected areas whereas participatory approaches provided more information outside protected areas, where elephant were most at risk from ivory poachers. A minimum of 35% of the 389 verified carcasses during 2001–2003 were illegally killed. In this baseline study, land uses ranked from highest to lowest by the proportion of illegally killed elephants (PIKE) were community conservation areas, government trust lands, forest reserves, private ranches, settlement areas and national reserves. PIKE trends derived from traditional and participatory data sources were similar across years and indicate elephants were at greater risk in regions outside government or privately patrolled areas. We suggest that PIKE is a useful index for comparing levels and trends in illegal killing of elephants, and that carcass ratios and presence/absence of tusks are useful proxy indicators of mortality and its causes.     

AGE CRITERIA FOR THE AFRICAN ELEPHANT

1. The paper is based on a collection of 385 lower jaws of the African elephant (Loxodonta a. africana Blumenbach) from western Uganda. 2. Thirty age groups are described and illustrated, which are related to the progress of eruption and wear of the six teeth in each side of the lower jaw. 3. Correct identification of individual teeth in the series is essential and several checks are described. Thus, when length is plotted against width for the six teeth the points fall into six well - defined groups. Frequency distributions of laminary indices and lamella numbers support the belief that teeth have been correctly identified, but overlapping, distributions preclude identification on these characters alone. 4. The presence of abnormal seventh molars in four jaws is described. 5. Chronological ages have been assigned to the relative age groups. Information on age and growth of captive animals has been considered and the general form of the growth curve established. Arbitrary estimates of the intervals between successive age groups were made and subsequently checked against growth and seasonal ridges on the roots. These indicate an upper age limit of about 60 years, which is compatible with fuller information on Indian elephants. 6. Growth curves support the validity of the ages assigned. Asymptotic heights at shoulder (∞) are respectively 317 cm and 298 cm for males from Murchison Falls and Queen Elizabeth National Parks. For females no distinction is made (because the sample from the latter area is small) and ∞ is 272 cm. 7. Growth in weight has also been established but shows no significant differences between the two populations. The mean maximum weight of females is 2766 kg (6100 lb), and of males 5450 kg (12000 lb). Growth apparently continues throughout life. 8. Tusk growth is analysed. There is a linear increase in weight of female tusks from eruption at 1–3 years up to the oldest group, with an indicated mean combined weight of 17.7 kg (39 lb) at 60 years. Male tusks show an increasing rate of growth throughout life to a mean combined weight of about 109 kg (240 lb) at 60 years. Big tusks are generally the result of prolonged growth; extremely big tusks probably result from prolonged and above average rate of growth. 9. Seasonal and annual incremental layers on the root 3 of the teeth are briefly described; these give an objective estimate of the chronological intervals between the relative age groups. 10. Field age criteria are presented which are derived from these growth curves. 11. The age at puberty in the female elephant is discussed and compared with earlier conclusions. There is evidence of a retardation in recent years in the Murchison Falls National Park (South bank) population and of a lengthening of the mean calving interval. This considerable depression of the reproductive rate, which is almost certainly density dependent, agrees with the observed lower recruitment in this population. 12. Survivorship curves constructed from material representing 325 natural deaths are presented for the two populations. If estimates of the age at puberty and the reproductive rate are taken into account, the expected differences in recruitment are found. 13. The calculated mean expectation of life is less than 15 years. 14. Mean individual weight is estimated at about 3800 lb.     

Demography of captive Asian elephants (Elephas maximus) in southern India

Historically, the Asian elephant has never bred well in captivity. We have carried out demographic analyses of elephants captured in the wild or born in captivity and kept in forest timber camps in southern India during the past century. The average fecundity during this period was 0.095/adult female/year. During 1969–89, however, the fecundity was higher at 0.155/adult female/year, which compares favorably with wild populations. There was seasonality in births with a peak in January. The sex ratio of 129 male to 109 female calves at birth is not significantly different from equality, although the excess of male calves born mainly to mothers 20–40 years old may have biological significance. Mortality rates were higher in females than in males up to age 10, but much lower in females than in males above age 10 years. The population growth rate, based on the lower secundity over the century, was 0.5% per year, and based on the higher secundity during 1969–89, was 1.8% per year. The analyses thus showed that timber camp elephants in southern India could potentially maintain a stationary or increasing population without resorting to captures from the wild. Breeding efforts for elephants in zoos can thus profitably learn from the experience of traditional management systems in part of Asia. Zoo Biol 16:263–272, 1997. © 1997 Wiley-Liss, Inc.     

Resource wars and conflict ivory: the impact of civil conflict on elephants in the Democratic Republic of Congo--the case of the Okapi Reserve

Human conflict generally has substantial negative impacts on wildlife and conservation. The recent civil war (1995-2006) in the Democratic Republic of Congo (DRC) resulted in a significant loss of wildlife, including elephants, due to institutional collapse, lawlessness and unbridled exploitation of natural resources such as minerals, wood, ivory and bushmeat. We used data from distance sampling surveys conducted before and after the war in a protected forest, the Okapi Faunal Reserve, to document changes in elephant abundance and distribution. We employed Generalized Additive Models to relate changes in elephant distribution to human and environmental factors. Populations declined by nearly fifty percent coinciding with a major increase in elephant poaching as indicated by reports of ivory trade during the war. Our results suggest that humans influenced elephant distribution far more than habitat, both before and after the war, but post-war models explained more of the variation. Elephant abundance declined more, closer to the park boundary and to areas of intense human activity. After the war, elephant densities were relatively higher in the centre of the park where they were better protected, suggesting that this area may have acted as a refuge. In other sites in Eastern DRC, where no protection was provided, elephants were even more decimated. Post-war dynamics, such as weakened institutions, human movements and availability of weapons, continue to affect elephants. Survival of remaining populations and recovery will be determined by these persistent factors and by new threats associated with growing human populations and exploitation of natural resources. Prioritizing wildlife protection, curbing illegal trade in ivory and bushmeat, and strengthening national institutions and organizations in charge of conservation will be crucial to counter these threats.