Age- and tactic-related paternity success in male African elephants

Information on age- and tactic-related paternity success isessential for understanding the lifetime reproductive strategyof males and constitutes an important component of the fitnesstrade-offs that shape the life-history traits of a species.The degree of reproductive skew impacts the genetic structureof a population and should be considered when developing conservationstrategies for threatened species. The behavior and geneticstructure of species with large reproductive skew may be disproportionatelyimpacted by anthropogenic actions affecting reproductively dominantindividuals. Our results on age- and tactic-specific paternitysuccess in male African elephants are the first from a free-rangingpopulation and demonstrate that paternity success increasesdramatically with age, with the small number of older bullsin the competitive state of musth being the most successfulsires. However, nonmusth males sired 20% of genotyped calves,and 60% of mature bulls (>20 years old) were estimated tohave sired offspring during the 5-year study period. The 3 mostsuccessful males sired less than 20% of the genotyped offspring.Hence, contrary to prediction from behavior and life-historytraits, reproduction was not heavily skewed compared with manyother mammalian systems with a similar breeding system. Nevertheless,these results indicate that trophy hunting and ivory poaching,both of which target older bulls, may have substantial behavioraland genetic effects on elephant populations. In addition, theseresults are critical to the current debate on methods for managingand controlling increasing populations of this species.     

Unusually extensive networks of vocal recognition in African elephants

Research on acoustic communication has often focused on signalling between territorial individuals or static neighbouring groups. Under these circumstances, receivers have the opportunity to learn to recognize the signals only of the limited number of conspecifics with which they are in auditory contact. In some mammals, however, social units move freely with respect to one another and range widely, providing individuals with opportunities to learn to recognize the signals of a wide range of conspecifics in addition to those of their immediate neighbours. We conducted playback experiments on African elephants, Loxodonta africana, in Amboseli National Park, Kenya, to determine the extent to which adult female elephants, which have a highly fluid social system, can recognize others in the population through infrasonic contact calls. Female elephants could distinguish the calls of female family and bond group members from those of females outside of these categories; moreover, they could also discriminate between the calls of family units further removed than bond group members, on the basis of how frequently they encountered them. We estimated that subjects would have to be familiar with the contact calls of a mean of 14 families in the population (containing around 100 adult females in total), in order to perform these discriminations. Female elephants thus appear to have unusually extensive networks of vocal recognition, which may prove to be typical of long-lived species that have both fluid social systems and the means for long-distance vocal communication. Copyright 2000 The Association for the Study of Animal Behaviour.     

Physiological stress and refuge behavior by African elephants

Physiological stress responses allow individuals to adapt to changes in their status or surroundings, but chronic exposure to stressors could have detrimental effects. Increased stress hormone secretion leads to short-term escape behavior; however, no studies have assessed the potential of longer-term escape behavior, when individuals are in a chronic physiological state. Such refuge behavior is likely to take two forms, where an individual or population restricts its space use patterns spatially (spatial refuge hypothesis), or alters its use of space temporally (temporal refuge hypothesis). We tested the spatial and temporal refuge hypotheses by comparing space use patterns among three African elephant populations maintaining different fecal glucocorticoid metabolite (FGM) concentrations. In support of the spatial refuge hypothesis, the elephant population that maintained elevated FGM concentrations (iSimangaliso) used 20% less of its reserve than did an elephant population with lower FGM concentrations (Pilanesberg) in a reserve of similar size, and 43% less than elephants in the smaller Phinda reserve. We found mixed support for the temporal refuge hypothesis; home range sizes in the iSimangaliso population did not differ by day compared to nighttime, but elephants used areas within their home ranges differently between day and night. Elephants in all three reserves generally selected forest and woodland habitats over grasslands, but elephants in iSimangaliso selected exotic forest plantations over native habitat types. Our findings suggest that chronic stress is associated with restricted space use and altered habitat preferences that resemble a facultative refuge behavioral response. Elephants can maintain elevated FGM levels for ≥ 6 years following translocation, during which they exhibit refuge behavior that is likely a result of human disturbance and habitat conditions. Wildlife managers planning to translocate animals, or to initiate other management activities that could result in chronic stress responses, should consider the potential for, and consequences of, refuge behavior.     

Nocturnal behavior in a group of female African elephants

The nocturnal behavior of a stable group of female, African elephants (Loxodonta africana africana) was studied to: (1) examine their behavior as a function of hour of night; (2) qualitatively compare the elephants' activity budgets to those reported in a previous study; and (3) document the presence of aggressive and stereotypic behaviors that might necessitate a change in their management. The elephants were systematically observed at least five times per week for 10 weeks between 17:00 and 08:00. Instantaneous focal samples of behavior, location, and proximity were taken every minute on a rotating basis, and all observed occurrences of social behavior were recorded. The hour of night affected elephant activity: significant relationships were found between hour of night and percent of time they spent feeding, lying, and standing. The overall activity budgets of the elephants were similar to the activity budgets reported in a previous study, although differences were evident in lying, stereotypic, and social behaviors. These differences might be a function of age. Affiliative behaviors accounted for 57% of the elephants' social behaviors, and agonistic behaviors among the elephants occurred infrequently and caused no injuries. Additionally, the elephants used all areas to which they had access. These findings provide compelling evidence that unrestricted social access during the night is the appropriate management strategy for these elephants. The results from the present study also highlight the importance of replicating existing studies and using multiple behavioral measures to make decisions regarding the welfare and management of stable groups of captive elephants. Zoo Biol 25:173–186, 2006. © 2006 Wiley‐Liss, Inc.     

Serum testosterone and musth in captive male African and Asian elephants

Testosterone concentrations in serum samples collected weekly over a 5-year period from a young adult male Asian elephant (Elephas maximus) and a young adult male African forest elephant (Loxodonta africana cyclotis) were measured by radioimmunoassay. Testosterone profiles during this maturational period were compared between the two species and related to the occurrence of musth, a recurring physiological and behavioral condition exhibited by most mature Asian, and some African, bull elephants. Musth is characterized by secretion from the bull's temporal glands, dribbling urine, and increased aggression. Serum testosterone concentrations in the Asian bull were elevated substantially between April and September each year, coincident with the presence of temporal gland secretion, urine dribbling, and aggressive behavior. Testosterone levels from April through September averaged (± SEM) 41.2 ± 2.8 ng/ml, compared to 7.6 ± 1.0 ng/ml during the rest of the year. In contrast, the testosterone profile of the African bull showed greater variation and lower levels overall, the only pattern being a tendency for levels to be lowest from November to February (avg. 6.8 ± 1.5 vs. 10.3 ± 0.8 ng/ml during the rest of the year). Temporal gland secretion and other signs of musth were first observed in this bull in 1988, at age 17. While his testosterone profile did not show a pattern comparable to that in the Asian bull, average testosterone values were significantly greater in 1988 compared to previous years. The Asian bull showed sexual attention to preovulatory (estrous) cows whether in musth or not, and exposure to estrous cows did not appear to alter the highly consistent, annual pattern of musth as evidenced in temporal gland flow.     

Seasonal variation of feeding patterns and food selection by crop-raiding elephants in Zimbabwe

Elephants and humans are increasingly coming into conflict because of the conversion of elephant habitat into agricultural areas. In order to identify trends that influence raiding behaviour, the nutritional makeup of food items consumed by crop‐raiding elephants over a 2‐year period were analysed and a trigger for crop raiding was identified. The point at which the quality of wild grasses declines below the quality of crop species corresponded to the movement of bull elephants out of a protected area and into fields. This finding may have wider implications for developing predictive models of elephant/human interactions.     

Population structure and behaviour of crop-raiding elephants in Kibale National Park, Uganda

We examined patterns of crop raiding by elephants across gender and age classes in relation to elephant life history and sociobiology and estimated the quantitative contribution of crops to elephant diet in Kibale National Park (KNP). Elephant dung‐boli sizes were used to estimate age and sex, while the presence of crop remains in the dung of crop‐raiding elephants was used as evidence of repeated raiding. To estimate the expected proportion of elephants raiding per age class, the age distribution of raiders was compared with the age distribution of all KNP elephants. Elephants raiding crops were predominantly males. They began raiding in expected proportions at 10–14 years while a higher than expected proportion raided crops at 20–24 years. These results suggest that crop raiding is initiated at an age when male elephants leave their families and a large proportion of elephants raid when they are approaching reproductive competition. Evidence from dung of crop raiders, suggests that repeated raiding increases with age. Crop raiders derived 38% of their daily forage from the short time spent raiding, consistent with expectations of foraging theory. Males may be more likely to learn crop raiding because they are socially more independent and experience intense mating competition than females.     

Nocturnal behavior in a group of unchained female African elephants

The chaining of elephants at night is a common management strategy in zoos, yet the costs and benefits of such a strategy are relatively unknown. A study of three unchained female African elephants was undertaken to document their nocturnal behavior. The subjects were observed between the hours of 1800 and 0800 for 10 weeks in the summer of 1992 (total of 172 hr) and 14 weeks in the summer of 1994 (total of 153 hr). Scan data were collected every 5 min to gather information on activity budgets, social proximity, and space utilization. All‐occurrence data were collected on social and non‐social behaviors. In each year of the study, the subjects spent equivalent amounts of time eating, lying, standing, and walking. Additionally, subjects spent half of their time within one body length of another animal and utilized all three available enclosures. Social and non‐social behaviors were frequent, and these data plus the activity profiles reveal the animals generally were most active between the hours of 1800 and 2400 and 0600 and 0700. The findings suggest that the use of no restraints is currently an effective strategy for this elephant group. The high activity levels observed during many of the early evening hours suggest that zoos could permit increased activity and social interactions by extending the hours when the elephants are unchained. Zoo Biol 18:101–109, 1999. © 1999 Wiley‐Liss, Inc.     

Plasticity in male mating behavior modulates female life history in fruit flies

In many species, intense male-male competition for the opportunity to sire offspring has led to the evolution of selfish reproductive traits that are harmful to the females they mate with. In the fruit fly, Drosophila melanogaster, males modulate their reproductive behavior based on the perceived intensity of competition in their premating environment. Specifically, males housed with other males subsequently transfer a larger ejaculate during a longer mating compared to males housed alone. Although the potential fitness benefits to males from such plasticity are clear, its effects on females are mostly unknown. Hence, we tested the long-term consequences to females from mating with males with distinct social experiences. First, we verified that competitive experience influences male mating behavior and found that males housed with rivals subsequently have shorter mating latencies and longer mating durations. Then, we exposed females every other day for 20 days to males that were either housed alone or with rivals, and subsequently measured their fitness. We found that females mated to males housed with rivals produce more offspring early in life but fewer offspring later in life and have shorter lifespans but similar intrinsic population growth rates. These results indicate that plasticity in male mating behavior can influence female life histories by altering females’ relative allocation to early versus late investment in reproduction and survival.     

Predictors of testosterone in zoo-managed male African elephants (Loxodonta Africana)

Reproductive complications for both male and female zoo-managed African elephants (Loxodonta africana) contribute to the rapidly declining population. In zoo-managed bull elephants, few studies have explored the potential physiological, physical, social, and environmental factors that influence bull fertility, particularly, androgen production. Testosterone is the essential steroid hormone for male sexual maturation and inadequate concentrations can be detrimental for spermatogenesis. Testosterone, fecal glucocorticoid metabolites, leptin, glucose, insulin, and triglycerides were analyzed from weekly fecal and blood serum samples taken over 6 months from six zoo-managed African elephant bulls (10–19 years of age). Testosterone levels were compared to endocrine factors, weekly social and environmental variables, daily musth signs, and body condition scores (BCS). The glucose-to-insulin ratio (G:I) was the only physiological biomarker found to be positively associated with testosterone. Predictive physical variables included Musth Score (+) and Moderate Exercise (+). Bulls with BCS signifying overweight (BCS 4) had lower testosterone (36.6 ± 1.6 ng/g fecal extraction [FE]) than bulls with healthy BCS 3; 51.2 ± 4.9 ng/g FE). Numerous social variables influenced testosterone concentrations, including Total Contact Day (+), Female Interaction Day (+), Indirect Contact Day (+), Indirect Contact Night (+) and Total No Contact (−). Both percentage of Time Outdoor and Time Mixed positively influenced testosterone, whereas testosterone decreased for percentage of Time Indoors. Each additional daily browse opportunity increased testosterone by approximately 7 ng/g FE. In managed care, the emphasis should be placed on optimizing these predictors of testosterone production to promote bull reproductive health.     

Complex phylogeographic history of central African forest elephants and its implications for taxonomy

Background

The mitochondrial genomes of snakes are characterized by an overall evolutionary rate that appears to be one of the most accelerated among vertebrates. They also possess other unusual features, including short tRNAs and other genes, and a duplicated control region that has been stably maintained since it originated more than 70 million years ago. Here, we provide a detailed analysis of evolutionary dynamics in snake mitochondrial genomes to better understand the basis of these extreme characteristics, and to explore the relationship between mitochondrial genome molecular evolution, genome architecture, and molecular function. We sequenced complete mitochondrial genomes from Slowinski's corn snake (Pantherophis slowinskii) and two cottonmouths (Agkistrodon piscivorus) to complement previously existing mitochondrial genomes, and to provide an improved comparative view of how genome architecture affects molecular evolution at contrasting levels of divergence.

Results

We present a Bayesian genetic approach that suggests that the duplicated control region can function as an additional origin of heavy strand replication. The two control regions also appear to have different intra-specific versus inter-specific evolutionary dynamics that may be associated with complex modes of concerted evolution. We find that different genomic regions have experienced substantial accelerated evolution along early branches in snakes, with different genes having experienced dramatic accelerations along specific branches. Some of these accelerations appear to coincide with, or subsequent to, the shortening of various mitochondrial genes and the duplication of the control region and flanking tRNAs.

Conclusion

Fluctuations in the strength and pattern of selection during snake evolution have had widely varying gene-specific effects on substitution rates, and these rate accelerations may have been functionally related to unusual changes in genomic architecture. The among-lineage and among-gene variation in rate dynamics observed in snakes is the most extreme thus far observed in animal genomes, and provides an important study system for further evaluating the biochemical and physiological basis of evolutionary pressures in vertebrate mitochondria.     

Social factors influence ovarian acyclicity in captive African elephants (Loxodonta africana)

Nearly one-third of reproductive age African elephants in North America that are hormonally monitored fail to exhibit estrous cycle activity, which exacerbates the nonsustainability of the captive population. Three surveys were distributed to facilities housing female African elephants to determine how social and environmental variables contribute to cyclicity problems. Forty-six facilities returned all three surveys providing information on 90% of the SSP population and 106 elephants (64 cycling, 27 noncycling and 15 undetermined). Logistic analyses found that some physiological and social history variables were related to ovarian acyclicity. Females more likely to be acyclic had a larger body mass index and had resided longer at a facility with the same herdmates. Results suggest that controlling the weight of an elephant might be a first step to helping mitigate estrous cycle problems. Data further show that transferring females among facilities has no major impact on ovarian activity. Last, social status appears to impact cyclicity status; at 19 of 21 facilities that housed both cycling and noncycling elephants, the dominant female was acyclic. Further studies on how social and environmental dynamics affect hormone levels in free-living, cycling elephants are needed to determine whether acyclicity is strictly a captivity-related phenomenon. Zoo Biol 28:1–15, 2009. © 2008 Wiley-Liss, Inc.     

The influence of Salix leaf abscission on leaf-miner survival and life history

Abstract. 1. Early abscission of mined leaves was an important mortality factor of a Phyllonorycter species (Lepidoptera: Gracillariidae) on Salix lasiolepis Benth. (Salicaceae). A larger percentage of mined leaves abscised early (34.4% in 1990; 24.5% in 1991), and Phyllonorycter survival was greatly reduced in these abscised leaves.
2. Leaf-mining by Phyllonorycter was associated with increased early leaf abscission. An egg removal experiment demonstrated that leaf mining induced this increase in leaf abscission.
3. The induction of early leaf abscission was dependent upon the timing of herbivory and simulated herbivory (mechanical damage). Early mechanical damage induced leaf abscission, late mechanical damage did not. Mines which expanded early were more likely to induce leaf abscission than mines which expanded more slowly.     

The influence of piscivory on life history traits of brown trout

Brown trout in Lake Femund migrated from the nursery streams mainly at 2 years old, but ranging between 1 and 8 years. Brown trout switched to piscivory from 3 years onwards, and a body length of 17·5 cm, according to back calculation from scales. Fast growers switched to piscivory at a younger age and smaller size than slow growers. The most slow-growing trout switched to fish feeding at 9 years old and a mean body length of 36 cm. The growth of the invertebrate feeders was almost rectilinear to c. 45 cm and 11 years of age. Switching to a fish diet induced increased growth rates. Age at sexual maturity increased with the age at which the fish became piscivorous. The invertebrate feeders matured at an age similar to that of the most fast growing piscivorous trout. The mortality rate of sub-adults and adults did not differ significantly between fish and invertebrate feeding trout. Longevity of piscivorous trout was estimated at 11 years and of invertebrate feeders at 10 years.     

Artificial evolution of life history and behavior

We present an individual-based model that uses artificial evolution to predict fit behavior and life-history traits on the basis of environmental data and organism physiology. Our main purpose is to investigate whether artificial evolution is a suitable tool for studying life history and behavior of real biological organisms. The evolutionary adaptation is founded on a genetic algorithm that searches for improved solutions to the traits under scrutiny. From the genetic algorithm's "genetic code," behavior is determined using an artificial neural network. The marine planktivorous fish Müller's pearlside (Maurolicus muelleri) is used as the model organism because of the broad knowledge of its behavior and life history, by which the model's performance is evaluated. The model adapts three traits: habitat choice, energy allocation, and spawning strategy. We present one simulation with, and one without, stochastic juvenile survival. Spawning pattern, longevity, and energy allocation are the life-history traits most affected by stochastic juvenile survival. Predicted behavior is in good agreement with field observations and with previous modeling results, validating the usefulness of the presented model in particular and artificial evolution in ecological modeling in general. The advantages, possibilities, and limitations of this modeling approach are further discussed.     

The sexually active states of free-ranging male African elephants (Loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behavior

Musth in male African elephants, Loxodonta africana, is associated with increased aggressive behavior, continuous discharge of urine, copious secretions from the swollen temporal glands, and elevated androgen levels. During musth, bulls actively seek out and are preferred by estrous females although sexual activity is not restricted to the musth condition. The present study combines recently established methods of fecal hormone analysis with long-term observations on male-female associations as well as the presence and intensity of physical signals to provide a more detailed picture about the physical, physiological, and behavioral characteristics of different states of sexual activity in free-ranging African elephants. Based on quantitative shifts in individual bull association patterns, the presence of different physical signals, and significant differences in androgen levels, a total of three potential sub-categories for sexually active bulls could be established. The results demonstrate that elevations in androgen levels are only observed in sexually active animals showing temporal gland secretion and/or urine dribbling, but are not related to the age of the individual. Further, none of the sexually active states showed elevated glucocorticoid output indicating that musth does not represent an HPA-mediated stress condition. On the basis of these results, we suggest that the term "musth" should be exclusively used for the competitive state in sexually active male elephants and that the presence of urine dribbling should be the physical signal used for defining this state.     

Heritability and Y-chromosome influence in the jack male life history of chinook salmon (Oncorhynchus tshawytscha)

Jacking in chinook salmon (Oncorhynchus tshawytscha) is an alternative reproductive strategy in which males sexually mature at least 1 year before other members of their year class. We characterize the genetic component of this reproductive strategy using two approaches; hormonal phenotypic sex manipulation, and a half-sib breeding experiment. We 'masculinized' chinook salmon larvae with testosterone, reared them to first maturation, identified jacks and immature males based on phenotype, and genotyped all fish as male ('XY') or female ('XX') using PCR-based Y-chromosome markers. The XY males had a much higher incidence of jacking than the XX males (30.8% vs 9.9%). There was no difference in body weight, gonad weight, and plasma concentrations of testosterone and 17beta-estradiol between the two jack genotypes, although XY jacks did have a higher gonadosomatic index (GSI) than XX jacks. In the second experiment, we bred chinook salmon in two modified half-sib mating designs, and scored the number of jacks and immature fish at first maturation. Heritability of jacking was estimated using two ANOVA models: dams nested within sires, and sires nested within dams with one-half of the half-sib families common to the two models. The sire component of the additive genetic variance yielded a high heritability estimate and was significantly higher than the dam component (h(2)(sire) = 0.62 +/- 0.21; h(2)(dam) = -0.14 +/- 0.12). Our experiments both indicated a strong sex-linked component (Y-chromosome) to jacking in chinook salmon, although evidence for at least some autosomal contribution was also observed.