Insectivory of savanna chimpanzees (Pan troglodytes verus) at Fongoli, Senegal

Little is known about the behavior of chimpanzees living in savanna-woodlands, although they are of particular interest to anthropologists for the insight they can provide regarding the ecological pressures affecting early hominins living in similar habitats. Fongoli, Senegal, is the first site where savanna chimpanzees have been habituated for observational data collection and is the hottest and driest site where such observation of chimpanzees occurs today. Previously, indirect evidence suggested these chimpanzees consumed termites throughout the year, an unusual occurrence for western and eastern chimpanzees. Although meat eating by chimpanzees continues to receive much attention, their use of invertebrate prey has received less emphasis in scenarios of hominin evolution. Here, we further examine the invertebrate diet of Fongoli chimpanzees using direct observational methods and accounting for potential environmental influences. Termite feeding positively correlated with high temperatures. Fongoli chimpanzees spend more time obtaining termites than any other chimpanzee population studied, and this extensive insectivory contributes to the list of distinctive behaviors they display relative to chimpanzees living in more forested habitats. We suggest that savanna chimpanzees at Fongoli differ significantly from chimpanzees elsewhere as a result of the selective pressures characterizing their harsh environment, and this contrast provides an example of a viable referential model for better understanding human evolution. Specifically, our results support the hypotheses that invertebrate prey may have figured more prominently into the diet of early hominins in similar habitats, especially given that invertebrates are an important source of protein and other essential nutrients in a highly seasonal environment.     

Arboreal nesting as anti-predator adaptation by savanna chimpanzees (Pan troglodytes verus) in southeastern Senegal

Chimpanzees (Pan troglodytes) make nests for resting and sleeping, which is unusual for anthropoid primates but common to all great apes. Arboreal nesting has been linked to predation pressure, but few studies have tested the adaptive nature of this behavior. We collected data at two chimpanzee study sites in southeastern Senegal that differed in predator presence to test the hypothesis that elevated sleeping platforms are adaptations for predator defense. At Assirik in the Parc National du Niokolo-Koba, chimpanzees face four species of large carnivore, whereas at Fongoli, outside national park boundaries, humans have exterminated almost all natural predators. We quantified the availability of vegetation at the two sites to test the alternative hypothesis that differences in nesting reflect differences in habitat structure. We also examined possible sex differences in nesting behavior, community demographic differences, seasonality and nest age differences as variables also potentially affecting nest characteristics and nesting behavior between the two sites. Chimpanzees at Fongoli nested at lower heights and farther apart than did chimpanzees at Assirik and sometimes made nests on the ground. The absence of predators outside of the national park may account for the differences in nest characteristics at the two sites, given the similarities in habitat structure between Fongoli and Assirik. However, Fongoli chimpanzees regularly build arboreal nests for sleeping, even under minimal predation pressure, and this requires explanation.     

Termite fishing laterality in the fongoli savanna chimpanzees (Pan troglodytes verus): Further evidence of a left hand preference

Whether nonhuman primates show population‐level handedness is a topic of much scientific debate. A previous study of handedness for termite fishing reported population‐level left handedness in the chimpanzees from Gombe National Park, Tanzania. In the current study, we examined whether similar hand preferences were evident in a savanna‐dwelling chimpanzee population with regards to termite fishing. Hand preference data were collected for 27 chimpanzees from February 2007 through July 2008 and November 2011 through January 2012 in southeastern Senegal. Overall, the Fongoli chimpanzees demonstrate a trend toward population‐level handedness, though the results did not reach conventional levels of statistical significance likely due to the limited sample size. Fongoli chimpanzees showed the same pattern of left hand preference as reported at Gombe and the two populations did not differ significantly. When the data were combined across all studies, wild chimpanzees showed a population‐level left hand preference for termite fishing. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Ecological context of savanna chimpanzee (Pan troglodytes verus) termite fishing at Fongoli, Senegal

Chimpanzees (Pan troglodytes verus) at Fongoli, Senegal, consume termites year-round. Understanding the ecological context behind this behavior is especially important in light of the environmental conditions at Fongoli. This mosaic savanna habitat is one of the hottest and driest sites where chimpanzees have been studied. Two genera and four species of termites were found in association with tools used by chimpanzees in a sample of 124 termite mounds that were monitored. The chimpanzees of Fongoli termite fish predominantly in woodland and forest habitat types, and, although woodland accounts for the majority of the chimpanzees' home range, forest habitat types comprise only about 4% of their range. Thus, habitat type has an influence on the Fongoli chimpanzees' termite fishing. Termite consumption to the degree seen at Fongoli may have particular significance for hominid evolution, given the expansion of Pliocene hominids into increasingly open, hot, and dry habitats.     

集约化生产对农田土壤碳氮含量及δ13C和δ15N同位素丰度的影响

集约化生产下农田土壤碳、氮含量变化是衡量土壤肥力持久性的重要指标.对常规水稻-蚕豆轮作地、露地蔬菜地、3年塑料大棚地和10年以上塑料大棚地的土壤pH、电导率(EC)、土壤有机碳(SOC)和总氮(TN)含量及δ13C和δ15N同位素丰度进行测定,研究了集约化生产程度对土壤特性的影响.结果表明:与水稻-蚕豆轮作地相比,露地蔬菜地、3年塑料大棚地和10年以上塑料大棚地0 ～20 cm耕层土壤pH分别降低1.1、0.8和0.7,而土壤EC分别是水稻-蚕豆轮作地的4.2、4.9和5.2倍;土壤碳、氮含量随塑料大棚地生产年限的增加总体上呈先增大后减小的趋势.与水稻-蚕豆轮作地相比,10年以上塑料大棚地0～20、20～40、40 ～60、60 ～ 80、80 ～ 100 cm土层的土壤SOC含量分别下降了54％、46％、60％、63％和59％,土壤TN含量分别下降了53％、53％、71％、82％和85％.农田集约化生产程度显著影响土壤SOC、TN含量和δ13C、δ15N丰度,土壤δ13C丰度与SOC含量呈显著负相关.土壤δ13C丰度可作为评价农田土壤碳循环受人为干扰强度的指标.     

Elemental variation in the termite fishing of wild chimpanzees (Pan troglodytes)

Chimpanzee tool behaviours vary dramatically in their complexity and extent of geographical distribution. The use of tool sets with specific design features to gather termites extends across a large portion of central Africa. Detailed examination of the composition and uniformity of such complex tool tasks has the potential to advance our understanding of the cognitive capabilities of tool users and processes underlying the maintenance of technological skills. In this study, we examined variation in chimpanzee tool use in termite gathering from video-recorded sequences that were scored to the level of functionally distinct behavioural elements. Overall, we found a high degree of similarity in tool-using techniques exhibited by individuals in this population. The number of elements in each individual's repertoire often exceeded that necessary to accomplish the task, with consistent differences in repertoire sizes between age classes. Adults and subadults had the largest repertoires and more consistently exhibited element strings than younger individuals. Larger repertoires were typically associated with incorporation of rare variants, some of which indicate flexibility and intelligence. These tool using apes aid us in understanding the evolution of technology, including that of our human ancestors, which showed a high degree of uniformity over large spatial scales.     

Diet of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik,Senegal: I. Composition

The composition of the diet of a savanna-living population of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik in Senegal is presented. The study site had a prolonged dry season, high temperatures, and vegetation dominated by grasses. Data came from direct observation, fecal specimens, and feeding traces; thus, strict criteria for acceptance of the indirect data were specified. Composition of diet was given in terms of species and family of prey, parts eaten, life-form, type of habitat, and criteria for inclusion. Forty-three species of plants with 60 parts were eaten; mostly fruits, from trees, in woodland. Nine species of animal prey were eaten, mostly social insects. An additional 41 species of plants with 53 parts were classed as likely to be eaten by chimpanzees, mostly on the grounds of their being eaten by sympatric anthropoids. Overall, the diet of the apes at Mt. Assirik resembles that of this species elsewhere in Africa, but the size of the dietary repertoire seems small and the proportion of low-quality foodstuffs high. The latter are mostly time-consuming to collect or tedious to obtain or process, and include underground storage organs.     

流溪河水库颗粒有机物及浮游动物碳、氮稳定同位素特征

为了解影响流溪河水库颗粒有机物(POM)碳和氮稳定同位素(δ¹³C和δ¹⁵N)变化的主要因素,及其与浮游动物δ¹³C和δ¹⁵N之间的关系,于2008年5月至12月份对POM及浮游动物的δ¹³C和δ¹⁵N进行了研究。颗粒有机物碳稳定同位素(δ¹³C_POM)和氮稳定同位素(δ¹⁵N_POM)的季节性变化幅度分别为5.1‰和2.2‰,5月和7月份δ¹³C_POM较高,而在10月和12月份降低,这主要与降雨将大量外源有机物带入水库而引起的外源及内源有机物在POM组成上发生变化有关。δ¹⁵N_POM总体呈上升趋势,可能是由降雨引起的外源负荷、初级生产力、生物固氮等因素共同作用的结果。浮游动物的δ¹³C及δ¹⁵N总的变化趋势与POM的相似,也具有明显的季节性变化,食物来源的季节变化可能是造成其变化的主要原因。在5月份,浮游动物的食物来源为POM中δ¹³C较高的部分,也就是外源有机物,而在10月及12月份,其食物则可能主要为浮游植物。     

Stable isotope signatures confirm carbon and nitrogen gain through ectomycorrhizas in the ghost orchid Epipogium aphyllum Swartz

Epipogium aphyllum is a rare Eurasian achlorophyllous forest orchid known to associate with fungi that form ectomycorrhizas, while closely related orchids of warm humid climates depend on wood- or litter-decomposer fungi. We conducted (13) C and (15) N stable isotope natural abundance analyses to identify the organic nutrient source of E. aphyllum from Central Norway. These data for orchid shoot tissues, in comparison to accompanying autotrophic plants, document C and N flow from ectomycorrhizal fungi to the orchid. DNA data from fungal pelotons in the orchid root cortex confirm the presence of Inocybe and Hebeloma, which are both fungi that form ectomycorrhizas. The enrichment factors for (13) C and (15) N of E. aphyllum are used to calculate a new overall average enrichment factor for mycoheterotrophic plants living in association with ectomycorrhizal fungi (ε(13) C ± 1 SD of 7.2 ± 1.6 ‰ and ε(15) N ± 1 SD of 12.8 ± 3.9 ‰). These can be used to estimate the fungal contribution to organic nutrient uptake by partially mycoheterotrophic plants where fully mycoheterotrophic plants are lacking. N concentrations in orchid tissue were unusually high and significantly higher than in accompanying autotrophic leaf samples. This may be caused by N gain of E. aphyllum from obligate ectomycorrhizal fungi. We show that E. aphyllum is an epiparasitic mycoheterotrophic orchid that depends on ectomycorrhizal Inocybe and Hebeloma to obtain C and N through a tripartite system linking mycoheterotrophic plants through fungi with forest trees.     

Feeding ecology of the highly threatened common bottlenose dolphin of the Gulf of Ambracia,Greece, through stable isotope analysis

The Gulf of Ambracia, in northwestern Greece, hosts a highly threatened community of about 150 common bottlenose dolphins (Tursiops truncatus). Until now, information on their feeding habits was derived exclusively from fish scale samples collected during surface-feeding events targeting small schooling epipelagic fish. The aim of this study was to determine the diet of bottlenose dolphins living in the Gulf of Ambracia through the application of Bayesian isotopic mixing models. Skin biopsy samples of 16 dolphins were analyzed and no difference related to sex or age-class was found in δ¹³C and δ¹⁵N values. Results suggested that the dolphin diet was mainly based on Trachurus trachurus, species belonging to the family Sparidae: Diplodus annularis, Lithognathus mormyrus, and Sepia officinalis, which represented together about 42% ± 15% of the biomass ingested, followed by species belonging to the order Clupeiformes (Engraulis encrasicolus, Sardinella aurita, and Sardina pilchardus) and the genus Gobius (37% ± 17%). A better understanding of the feeding habits of these dolphins sheds light on the feeding ecology of this highly threatened population by, for instance, evidencing interactions with artisanal fisheries sharing the same target species, and is key for identifying adequate management measures consistent with an ecosystem-based approach.     

Stable nitrogen isotope analysis of dentine serial sections elucidate sex differences in weaning patterns of wild chimpanzees (Pan troglodytes)

Offspring provisioning is one of the most energetically demanding aspects of reproduction for female mammals. Variation in lactation length and weaning strategies between chimpanzees (Pan troglodytes), our closest living relative, and modern human societies have been reported. When and why these changes occurred is frequently debated. Our study used stable nitrogen isotope data of tooth root dentine from wild Western chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d'Ivoire, to quantify weaning in these chimpanzees and explore if infant sex plays a role in maternal investment. We analyzed serial sections of deciduous lateral incisor root dentine from four Taï chimpanzees to establish the δ¹⁵N signal of nursing infants; we then analyzed serial sections of first permanent mandibular molar root dentine from 12 Taï chimpanzees to provide quantitative δ¹⁵N data on weaning in this population. Up to 2 years of age both sexes exhibited dentine δ¹⁵N values ≈2–3‰ higher than adult female Taï chimpanzees, consistent with a nursing signal. Thereafter a steady decrease in δ¹⁵N values consistent with the onset, and progression, of weaning, was visible. Sex differences were also evident, where male δ¹⁵N values decreased at a significantly slower rate compared to females. Confirmation of sex differences in maternal investment among Taï chimpanzees, demonstrates the viability of using isotope analysis to investigate weaning in non‐human primates. Additionally, assuming that behaviors observed in the Taï chimpanzees are illustrative of the ancestral pattern, our results provide a platform to enable the trajectory of weaning in human evolution to be further explored. Am J Phys Anthropol 153:635–642, 2014. © 2013 Wiley Periodicals, Inc.     

Social influences on ranging patterns among chimpanzees (Pan troglodytes verus) in the Tai National Park, Cote d'Ivoire

Group size is expected to be an important factor to predicthome-range (HR) size in social animals. In chimpanzees adultmales play an important role in defending the HR against neighbors,and therefore it has been suggested that HR size depends onthe number of adult males. In this long-term study on wild WestAfrican chimpanzees, we analyzed the relative importance ofcommunity size and composition on ranging patterns over a 10-yearperiod, using multivariate statistics. Because community sizedecreased from 51 individuals with 6 adult males in 1992 to22 individuals with only 1 adult male in 2001, we expected adecrease in HR size, which should be better predicted by thenumber of males than by community size. We further investigatedthe effect of fruit availability on monthly HRs over a 4-yearperiod. As predicted, HR size decreased during the first 7 yearsof our study but increased during the last 3 years. Overall,the number of adult males was the best predictor of HR size,whereas fruit availability did not correlate with HR size. HRuse remained stable over the entire study period, with a constantproportion of about 35% of the HR used as core area. High HRand core-area overlap values between different years indicatedstrong site-fidelity. Although the number of males within thecommunity explained the decrease in HR size, the recent increasein size remains unexplained. This finding suggests that otherfactors such as relative fighting power of males affect HR size.     

Stable isotope analysis of larval mosquito diets in agricultural wetlands in the coastal plain of Georgia,U.S.A.

Previous studies have used C and N isotope ratios to investigate the use of different food resources such as plant and animal detritus by container‐breeding mosquitoes. This study is the first to report on the potential food resources assimilated by larval mosquitoes in agricultural and reference wetlands. Larval mosquitoes (Diptera: Culcidae) were sampled, along with their potential food resources, from agricultural and reference wetland habitats throughout a seasonal hydroperiod. IsoSource mixing model results indicated that food resources had greater δ¹⁵N isotope values in agricultural wetlands compared with cypress‐gum swamps. In February, Aedes vexans (Meigen) and Culex territans Walker larvae fed primarily on lower quality food resources (coarse particulate organic matter and sediment) based on C:N. In contrast, higher quality food resources (fine particulate organic matter) were utilized by Anopheles spp. throughout the study and by Psorophora columbiae (Dyer and Knab) in May. This research contributes to a more comprehensive understanding of the food resources available and assimilated by larval mosquitoes in agricultural wetlands.     

Animal foods in the diets of wild chimpanzees (Pan troglodytes): Why cross-cultural variation?

The common chimpanzee (Pan troglodytes) in nature is an omnivore, i.e. it eats animals as well as plants. Six long-term field studies reveal differences in diet amongst populations of chimpanzees in east, central and west Africa. Specific comparisons can be made for social insect (ants, bees, termites) and mammalian (primates, ungulates) prey. Most of the differences can be explained in terms of environmental influences: presence or absence of prey species, abundance and distribution of prey, range of potential prey secies, competing predators, characteristics of habitat, human interference. However, other differences appear to reflect true social customs, independent of the bio-physical environment, and therefore can be termed cross-cultural.     

A longitudinal study of urinary dipstick parameters in wild chimpanzees (Pan troglodytes verus) in Côte d'Ivoire

We performed 796 dip‐stick tests on urine from 100 wild West African chimpanzees (Pan troglodytes verus) from 4 habituated groups in the tropical rain forest of Taï National Park, Cote d'Ivoire, to establish reference values for health monitoring. Specific gravity was also measured on 359 urine samples from 62 chimpanzees. The effect of age, sex, group, month, estrus, pregnancy, meat consumption, and acute respiratory disease on pH, leucocytes, protein, blood, hemoglobin, and glucose was examined using ordinal logistic regression. The presence of nitrite, ketones, bilirubin, and urobilinogen in urine was also recorded. Outbreak of acute respiratory disease did not influence any of the urinary parameters. Thirty‐seven percent of the samples had a pH <7 and the whole range of pH was found through the year, in all age groups, and in both sexes. Meat consumption lowered the urinary pH. Our results show that all pH levels must be considered normal for the West African chimpanzee subspecies P. troglodytes verus living in the rainforest. We also found a cluster of glucose‐positive samples at a specific point in time which was not attributed to diabetes mellitus. These findings highlight that there are differences in normal physiological parameters among wild chimpanzees living in different habitats. Am. J. Primatol. 72:689–698, 2010. © 2010 Wiley‐Liss, Inc.     

Variation in performance of two co‐occurring mosquito species across diverse resource environments: insights from nutrient and stable isotope analyses

1. Inputs of animal and plant detritus are the main energy sources for food webs in a number of isolated container systems, including discarded automobile tyres and tree holes. Containers are dominated by mosquitoes in the genera Culex and Aedes, which among other differences often engage in different foraging behaviours. We hypothesised that because Aedes feed more by browsing surfaces, whereas Culex often filter the water column, these mosquitoes would show variation in performance and differentially affect detritus. Effects of different ratios of animal and plant detritus on survival, mass, and development time for two common container mosquito species, Culex restuans L. and Aedes albopictus Skuse, were examined. We also quantified detrital contribution to biomass via isotopic and nutrient analysis and the effect of larvae on detrital decay. 2. Adult male and female mass of both species was highest with some animal detritus and lowest in only leaf detritus. Aedes albopictus survival was higher than C. restuans across most detritus ratios. 3. Aedes albopictus had higher values of ¹⁵N and in some cases ¹³C across all detritus ratios compared with C. restuans; A. albopictus had lower nitrogen in tissue. Aedes albopictus appeared to be more efficient at obtaining potentially limiting nutrients and had a greater overall effect on detrital decay – a possible consequence of greater foraging effort. 4. Findings further support the view that mosquito performance can be influenced by detritus type, and provide a more precise hypothesis (i.e. lower need for nitrogen) that may explain the superior competitive ability of A. albopictus over other container mosquitoes.     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.