Why elephant have trunks and giraffe long tongues: how plants shape large herbivore mouth morphology

We investigated whether mass and morphological spatial patterns in plants possibly induced the development of enlarged soft mouth parts in especially megaherbivores. We used power functions and geometric principles to explore allometric relationships of both morphological and foraging characteristics of mammalian herbivores in the South African savannah, covering a body size range of more than three orders magnitude. Our results show that, although intradental mouth volume scaled to a power slightly less than one to body mass, actual bite volume, as measured in the field, scaled to body mass with a factor closer to 1.75. However, when including the volume added to intradental mouth volume by soft mouth parts, such as tongue and lips (or trunks in elephant), mouth volume scaled linearly with actual bite volume and in a similar fashion as actual bite volume to body size. Bite mass and bite leaf mass scaled linearly with body size. We conclude that these scaling relationships indicate that large herbivores use their enlarged soft mouth parts to not only increase bite volume and thereby bite mass, but also select soft plant parts and thereby increase the leaf mass fraction per bite.     

The ranging behaviour of a large sexually dimorphic herbivore in response to seasonal and annual environmental variation

The allometric relationships of body size play a principle role in determining how large herbivores respond to the marked spatial and temporal heterogeneity of the savanna biome. Using location data collected over an 8‐year period from five distinct study sites, we investigated the influence of environmental variation (using phenological and rainfall data) on the ranging behaviour of the African elephant (Loxodonta africana), a species that exhibits pronounced sexual dimorphism. Both sexes expanded their annual ranges during years of high rainfall and contracted their ranges during periods of resource scarcity, concurring with the hypothesis that abiotic factors dictate the distribution of large generalist herbivores at the landscape scale. However, female elephant did not exhibit the same consistent response to rainfall at the seasonal scale. Furthermore, male elephant demonstrated a reduction in their daily displacement distances during the dry winter season, and altered their movement rates on the basis of seasonal rainfall. These results suggest that male elephant are able to consistently adapt their movement behaviour according to forage quality and abundance. Smaller‐bodied female elephant on the other hand, are unlikely to exhibit the same flexibility in their ranging behaviour because of their higher relative nutritional demands, lower tolerance to fibrous forage and the social and energetic constraints of group living with juveniles. Our study highlights the major role that body size and sociality plays in the decision making of sexually dimorphic herbivores. These differences can have important implications for effective conservation and management, particularly with regard to demographic (e.g. survival) and ecological (e.g. habitat use) factors.     

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.     

Intestinal length of three California pinniped species

Forty-eight intestinal tracts, extracted from both sexes of California sea lions, Harbour seals and Northern elephant seals, were measured. The majority of intestinal tracts were removed from stranded animals that died from various causes. The sea lions and elephant seals, approximately equal in size, were larger than the Harbour seals. All species possess a small intestine which is significantly longer than even the entire gastrointestinal tract of herbivores of comparable size. Elephant seal small intestines, averaging approximately 25 times the seal's body length, were considerably longer than the small intestines of either sea lions (averaged more than 18 times the body length) or Harbour seals (averaged nearly 16 times the body length). However, the large intestines of elephant seals were shorter than either of the other two species. Among the sea lions and Harbour seals the large intestines were approximately equal in length. Sea lions and Harbour seals also showed a close correlation between standard length and total intestinal length. Among elephant seals these two parameters showed greater variability. The functional significance of the extremely long small intestine remains unclear. Certainly, the large body mass and high energy requirements of these animals has contributed to the development of a long intestinal tract. It also appears likely that diet and the high motility rate of digesta influenced the intestinal development. Comparatively, the significantly shorter large intestine of elephant seals probably relates to this species' remarkable capabilities in water conservation and metabolic water retention.     

Observations on the length of the intestinal tract of African <Emphasis Type="Italic">Loxodonta africana</Emphasis> (Blumenbach 1797) and Asian elephants <Emphasis Type="Italic">Elephas maximus</Emphasis> (Linné 1735)

The digestive tract of elephants is surprisingly short compared to other herbivorous mammals. However, measurements relating the length of the intestine to the body mass of the respective individual are rare. In this study, we report such data for an African elephant and an Asian elephant. Our data support the hypothesis that Asian elephants have a longer intestinal tract than their African counterparts. These findings are in accord with the observation of longer retention times and higher digestion coefficients in Asian as compared to African elephants. This difference between the species could be the reflection of slightly different ecological niches, with Asian elephants adapted to a natural diet with a higher proportion of grass.     

Explaining the scaling of transport costs: the role of stride frequency and stride length

The mechanisms which enable large animals to transport a unit of body mass through a unit distance at a lower metabolic cost than smaller animals have been the subject of numerous studies. Recent investigations have concluded that stride frequency is a main determinant. We examine the role of both stride frequency and stride length in determining the scaling of the cost of transport.
Slopes for regressions between stride frequency and speed and stride length and speed were determined in four species of rodents. These data were pooled with literature values for the slopes of stride frequency, stride length and cost of locomotion (all vs. speed) for a total of 17 species ranging in size from 30 g to 250 kg. Interspecific equations were calculated for each of these slopes versus body mass, and residuals from these allometric lines were calculated. Residuals were compared to see if variation in the rate of cost increase at a given size is related to variation in the rates of stride frequency and/or stride length increase.
The residual analysis revealed that the variation in transport cost is explicable only in terms of the interaction of stride frequency and stride length slopes. The product of the scaling exponents for stride frequency slope and stride length slope is not significantly different from the scaling exponent for the cost of transport. A model seeking to explain the scaling of the cost of transport must therefore consider the influence of both stride length and stride frequency.
We propose that absolutely longer limbs allow large animals to minimize the rate of increase of stride frequency and stride length with speed, and that this allows utilization of muscles with lower intrinsic rates of contraction, which in turn results in a lower mass-specific cost of transport.     

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

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

Estimating daily walking distance of captive African elephants using an accelerometer

Two central concerns for elephant husbandry and management are whether zoological enclosures are appropriately sized and the degree to which naturalistic exercise and activity are observed in such enclosures. In order to address these issues, accurate data on the daily walking distance of elephants both in situ and ex situ are necessary. We used an accelerometer, a pedometer that measures step count and activity level, to estimate walking distance in African elephants (Loxodonta africana) at the San Diego Zoo's Wild Animal Park. The accelerometer was worn simultaneously with a GPS unit that recorded actual walking distance. Estimates of walking distance were extrapolated from the accelerometer and compared with actual distances determined by GPS data. The accelerometer was found to overestimate step count, and subsequently walking distance, by including false counts of steps. Extrapolating walking distance based upon stride length measurements did not match actual GPS walking distance. However, activity level output from the accelerometer significantly correlated with actual GPS walking distance. In addition, we report that the rate of movement is comparable to that reported in other zoological settings. We provide a linear regression equation that can be utilized by other institutions to estimate daily walking distance of elephants in their collection who are outfitted with accelerometers.     

Morphological adaptation of the digestive tract in relation to feeding ecology of raptors

The study examines some aspects of gross morphology in Falconiformes and Strigiformes. It is hypothesized that, in predatory birds, hunting strategy might influence the relative size of skeletal musculature and length of the digestive tract. Falconiform species were categorized as either 'attackers' or 'searchers' depending on the degree to which active, powered pursuit is required for prey capture. Attacking species feed predominantly on avian prey, requiring extreme agility, speed and acceleration for prey capture. Searchers feed largely on relatively slow-moving mammals and carrion. Comparisons between species of attackers and searchers showed that the former had heavier pectoral muscle mass, larger areas for flight muscle attachment and higher linearized wing loadings. Strigiformes had a pectoral muscle mass only half the size of that of attacking Falconiformes and had a correspondingly smaller sternum area. A skeletal body-size measure was determined to enable calculation of intestine length independent of body-size and shape differences. Attacking species have a small intestine which is 20–30% shorter than would be predicted on the basis of body-size and 50% shorter than found in searchers of equivalent body-size. Strigiformes that locate prey by active flight also have intestinal tracts shorter than expected. The likely effects of intestine length on digestive efficiency and food utilization are discussed and it is suggested that, in predatory birds, some species have evolved alimentary tracts that are shorter than necessary for maximum digestive efficiency in order to enhance prey capture.     

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.     

中国人群的体部指数

本文通过对63452例18～97岁的中国人体部12项指数值的统计分析,得出目前该年龄段中国人的体部形态特征。研究发现,中国人总体体型为长躯干型、中肩型、中骨盆型、中腿型。男性为中胸型,女性为宽胸型。随着年龄增长,上半身会显得更短一些,上半身与下半身比例更小一些,胸部更显宽厚一些,躯干下部显得更宽一些,腿显得更长一些。与南方族群相比,蒙古语族群、突厥语族群身体更壮实一些,胸部更显得宽厚一些,上身更高一些,上肢显得短一些,躯干的上部(肩部)相对窄一些,躯干的下部(骨盆)相对更宽一些,上肢长和下肢长度比例更小一些。研究还发现,同等身高的男性和女性相比,男性的上肢长度、下肢长度一般都小于女性,而女性比男性有一个更大的坐高值。从躯干长度来比较,女性确实比男性腿短一些。下身长相等的中国男性、女性之间相比,女性的坐高大于男性。同等身高情况下,中国人的坐高比欧亚人种、非洲人种的坐高要大,即有较高的上半身高度。     

Lumbar ontogenetic allometry and dimorphism in humans. A case for comparison between interspecific and intraspecific scaling

The ontogenetic allometry of the lumbar region of 1913 humans (1228 females and 685 males), ranging from newborn to 21-year-old individuals, was studied by means of length, width, projected surface area and bone mineral density of the segment L2 - L4, obtained by dual X-ray absorptiometry (DXA). All these parameters were regressed to body mass and height of the individuals, considered alternatively as the independent variable. Firstly, we addressed the comparison between the results obtained on both sexes in order to elucidate whether ontogenetic differences existed. Length of the segments increased significantly faster in females than in males, independently whether the regression was made against body mass or height, while in both types of regression width scaled in males faster than in females. Regarding bone mineral density, although males increased bone mineral density faster than females, slope differences were not significant. However, y-interception was significantly higher in females than in males when bone mineral density was regressed to body mass. Results on length and width are compared with others from previous research on allometry. Finally, global results are discussed as regards the slope predictions for interspecific scaling.     

The affects of artificial water availability on large herbivore ranging patterns in savanna habitats: a new approach based on modelling elephant path distributions

Aim  Artificial water points are often used in protected savanna ecosystems to maintain populations of large herbivores. However, these interventions lead to increased ranging and foraging pressure and can negatively impact important habitats and species. This study investigated the influence of artificial water provision on the foraging and movement paths of an African elephant population and modelled the impact of changing water availability on sensitive habitat types.
Location  Tembe Elephant Park (TEP), KwaZulu-Natal, South Africa.
Methods  We mapped and classified 414 km of elephant movement and foraging paths in a 300-km² fenced protected area. The data were analysed to determine the relationship between path size, distribution and distance to water. We also used a logistic modelling approach to explore the predicted effects of removing artificial water points on path distribution.
Results  Elephant paths were unevenly distributed throughout the habitats of TEP and the most established and heavily used paths were found closest to water. We also discovered a number of elephant 'rest areas' along the paths, which were distinct clearings that tended to be close to water and in sand forest habitat. Our model predicted that the removal of artificial water points would reduce the area crossed by elephant paths by 79%, leading to an 89% reduction in the presence of elephant paths in sand forest.
Main conclusions  Our study provides further evidence that manipulating surface water availability can be a useful tool for managing large herbivore impacts on vegetation and acts as the basis for further research on the trade-offs between conservation objectives.     

Morphometric relationships of take-off speed in anuran amphibians

Locomotory speed correlates with muscle mass (determining force and stride rate), limb length (stride rate and distance), and laterally compressed body trunk (force and stride distance). To delineate generalization of the locomotory-morphometric relationships specifically in anuran amphibians, we investigated take-off speed and the three morphological variables from seven species, Rana nigromaculata, R. rugosa, and Bombina orientalis, Eleuthrodectilus fitzingeri, E. diastema, Bufo typhonius, Colostethus flotator and Physalaemus pustulosus. The fastest jumper E. fitzingeri (3.41 m s(-1)) showed 2.49-fold greater speed than the slowest B. typhonius. Take-off speed correlated well with both thigh muscle mass relative to body mass and hindlimb length relative to snout-vent length (HL/SVL), but poorly correlated with the inter-ilial width relative to SVL. The best morphological predictor was HL/SVL (speed=-3.28+3.916 HL/SVL, r=0.968, P<0.0001), suggesting that anuran take-off speed is portrayed well with high gear and acceleration distance characterized by hindlimbs.     

An empirical evaluation of the African elephant as a focal species for connectivity planning in East Africa

Aim Large, charismatic and wide‐ranging animals are often employed as focal species for prioritizing landscape linkages in threatened ecosystems (i.e. ‘connectivity conservation’), but there have been few efforts to assess empirically whether focal species co‐occur with other species of conservation interest within potential linkages. We evaluated whether the African elephant (Loxodonta africana), a world‐recognized flagship species, would serve as an appropriate focal species for other large mammals in a potential linkage between two major protected area complexes. Location A 15,400 km² area between the Ruaha and Selous ecosystems in central Tanzania, East Africa. Methods We used walking transects to assess habitat, human activity and co‐occurrence of elephants and 48 other large mammal species (> 1 kg) at 63 sites using animal sign and direct sightings. We repeated a subset of transects to estimate species detectability using occupancy modelling. We used logistic regression and AIC model selection to characterize patterns of elephant occurrence and assessed correlation of elephant presence with richness of large mammals and subgroups. We considered other possible focal species, compared habitat‐based linear regression models of large mammal richness and used circuit theory to examine potential connectivity spatially. Results Elephants were detected in many locations across the potential linkage. Elephant presence was highly positively correlated with the richness of large mammals, as well as ungulates, carnivores, large carnivores and species > 45 kg in body mass (‘megafauna’). Outside of protected areas, both mammal richness and elephant presence were negatively correlated with human population density and distance from water. Only one other potential focal species was more strongly correlated with species richness than elephants, but detectability was highest for elephants. Main conclusions Although African elephants have dispersal abilities that exceed most other terrestrial mammals, conserving elephant movement corridors may effectively preserve habitat and potential landscape linkages for other large mammal species among Tanzanian reserves.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Ethnic differences in triceps surae muscle-tendon complex and walking economy

The purposes of this study were to (a) determine whether structural differences in triceps surae muscle-tendon complex and walking economy exist between 14 African American and 19 Caucasian sedentary women and (b) determine whether muscle-tendon parameters are associated with walking economy. African American and Caucasian subjects were matched on body weight, height, and body composition. Muscle-tendon parameters were determined by magnetic resonance imaging and walking economy was evaluated at 4.8 km.h(-1). Medial gastrocnemius and total triceps surae muscle shape were different across ethnicity despite no ethnic differences in plantar flexion strength or in maximal cross-sectional area for any triceps surae muscles. African American women had shorter gastrocnemius muscles and longer tendons and performed walking more economically. Tendon length was the only variable related to walking economy. No ethnic differences were observed in walking economy after adjusting for tendon length. Data show gastrocnemius tendon length is related to level walking and longer gastrocnemius tendons may partly explain more economical walking in African American women. These preliminary findings indicate the structure of the muscle-tendon complex could be a factor partially accounting for reported ethnic differences in certain types of athletic-related performance.     

The hard lives of trees in African savanna—Even without elephants

The ongoing loss of large trees and densification of shrubs are two prevalent processes that take place in African savannas, with profound consequences for their structure and function. We evaluated herbivore impacts on savanna woody communities using a long-term exclosure experiment in the Kruger National Park, South Africa, with three treatments: the exclusion of large mammals only (i.e. elephant and giraffe), exclusion of all herbivores larger than a hare, and areas open to all herbivores. We asked three questions: (1) How did variable exclusion of herbivores affect woody density and structure across the catena (i.e. riparian, sodic and crest vegetation)? (2) Did the exclusion of herbivores result in unique woody species composition? (3) Did herbivore exclusion result in a higher proportion of palatable species? After 17 years, we found that herbivores mainly affected the heights and densities of existing species, rather than leading to turnover of woody species assemblages. Although densities of individuals increased in the full exclosure (350 ha⁻¹), the change was more moderate than expected. By contrast, mixed mega-and meso-herbivores decreased the number of trees and shrubs (decreases of 780 ha⁻¹) via a variety of physical impacts. Meso-herbivores alone, on the other hand, had less impact on individual density (i.e. no change), but limited average height growth and canopy dimensions in certain habitat types. Where elephants are present, they are effective at reducing the density of woody stems to the point of counteracting woody encroachment, but at the same time are actively preventing the persistence of large trees (>5 m) as well as preventing trees from recruiting to larger size classes. However, the lack of massive recruitment and woody cover increases with elephant exclusion, especially for more preferred species, suggests that factors beyond elephants, such as dispersal limitation, seed predation, and drought, are also acting upon species.     

Fire frequency drives habitat selection by a diverse herbivore guild impacting top–down control of plant communities in an African savanna

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.     

象科动物对当地森林生态系统的影响

现存的象科动物(Elephantidae)分为非洲草原象(Loxodonta africana)、非洲森林象(Loxodonta cyclotis)和亚洲象(Elephas maximus)3 种,作为森林生态系统的关键物种,它们对当地森林生态系统的影响非常复杂,在一定环境条件下,既可能是积极的作用,也可能是消极的作用。积极的作用包括:帮助植物传播种子;促进种子萌发;创造断层,维持群落多样性;为其他动物增加食物资源;为其他动物创造栖息地。消极的作用包括:使一些物种的种群数量减小;使森林变成灌木丛和草原等。而由于活动受限导致的种群密度过高是象科动物对森林生态系统产生消极作用的主要原因。当前象科动物的3 个物种均面临种群数量锐减和生存空间不断缩小的危机,为此迫切需要针对其对生物多样性和生态平衡的影响开展深入和全面的研究,并且应根据实际情况因地制宜地制定管理措施。