Driver ants ( Dorylus nigricans species group) are keystone species in forest and savanna ecosystems across tropical Africa. However, their study has been hampered by considerable taxonomic confusion. A multitude of worker forms has been described under the name Dorylus nigricans Illiger, 1802, even though the link between the name-bearing type specimens, two males from Sierra Leone, and the respective conspecific workers has never been established. Extensive surveys of driver ants were first conducted at numerous West African sites from Senegal to Ghana. These yielded a total of only three species recognisable by morphological differences. We then examined the two D. nigricans syntypes and found that they belong to different species. By analysing mitochondrial and nuclear DNA sequences of males and workers from Taï National Park (Ivory Coast), males with a phenotype similar to that of the larger syntype specimen were matched to workers originally described under the name Anomma arcens by Westwood (1847). This larger syntype male is designated as the lectotype of D. nigricans Illiger, 1802, so that Anomma arcens Westwood, 1847 is synonymised under D. nigricans . A worker neotype for the species originally described as Anomma burmeisteri Shuckard, 1840, whose type specimen was lost, is designated. A key to the large workers of the three driver ant species occurring in this region, D. burmeisteri , D. nigricans , and D. mayri , is provided. By clarifying the taxonomy of the regional driver ant fauna, we resolve a centuries-old puzzle and pave the way for a comprehensive revision of the entire species group.

Understanding the genetic basis of adaption is a central task in biology. Populations of the honey bee Apis mellifera that inhabit the mountain forests of East Africa differ in behavior and morphology from those inhabiting the surrounding lowland savannahs, which likely reflects adaptation to these habitats. We performed whole genome sequencing on 39 samples of highland and lowland bees from two pairs of populations to determine their evolutionary affinities and identify the genetic basis of these putative adaptations. We find that in general, levels of genetic differentiation between highland and lowland populations are very low, consistent with them being a single panmictic population. However, we identify two loci on chromosomes 7 and 9, each several hundred kilobases in length, which exhibit near fixation for different haplotypes between highland and lowland populations. The highland haplotypes at these loci are extremely rare in samples from the rest of the world. Patterns of segregation of genetic variants suggest that recombination between haplotypes at each locus is suppressed, indicating that they comprise independent structural variants. The haplotype on chromosome 7 harbors nearly all octopamine receptor genes in the honey bee genome. These have a role in learning and foraging behavior in honey bees and are strong candidates for adaptation to highland habitats. Molecular analysis of a putative breakpoint indicates that it may disrupt the coding sequence of one of these genes. Divergence between the highland and lowland haplotypes at both loci is extremely high suggesting that they are ancient balanced polymorphisms that greatly predate divergence between the extant honey bee subspecies.

Root herbivory incurs fitness costs, but herbivory by nematodes can additionally increase nutrient availability mediated by enhanced root exudation and soil microbial activity and change plant susceptibility to aboveground herbivores due to systemic changes in plant defence. We hypothesized that such positive indirect effects may outweigh the negative direct effects of root herbivory by nematodes on plant performance.

Cultural variation has been identified in a growing number of animal species ranging from primates to cetaceans. The principal method used to establish the presence of culture in wild populations is the method of exclusion. This method is problematic, since it cannot rule out the influence of genetics and ecology in geographically distant populations. A new approach to the study of culture compares neighbouring groups belonging to the same population. We applied this new approach by comparing ant-dipping tool length between two neighbouring communities of chimpanzees (Pan troglodytes schweinfurthii) in the Kalinzu Forest, Uganda. Ant-dipping tool length varies across chimpanzee study sites in relation to army ant species (Dorylus spp.) and dipping location (nest vs. trail). We compared the availability of army ant species and dipping tool length between the two communities. M-group tools were significantly longer than S-group tools, despite identical army ant target species availabilities. Moreover, tool length in S-group was shorter than at all other sites where chimpanzees prey on epigaeic ants at nests. Considering the lack of ecological differences between the two communities, the tool length difference appears to be cultural. Our findings highlight how cultural knowledge can generate small-scale cultural diversification in neighbouring chimpanzee communities.

Chimpanzees are renowned for their use of foraging tools in harvesting social insects and some populations use tools to prey on aggressive army ants ( Dorylus spp.). Tool use in army ant predation varies across chimpanzee study sites with differences in tool length, harvesting technique, and army ant species targeted. However, surprisingly little is known about the detailed ecology of army ant predation. We studied army ant predation by chimpanzees ( Pan troglodytes verus ) at the Seringbara study site in the Nimba Mountains, Guinea (West Africa), over 10 years (2003–2013). We investigated chimpanzee selectivity with regards to army ant prey species. We assessed the temporal variation in army ant‐feeding and examined whether army ant predation was related to rainfall or ripe fruit availability. Moreover, we examined whether chimpanzees showed selectivity regarding plant species used for tool manufacture, as well as the relationship between tool species preference and tool collection distance. Lastly, we measured tool properties and investigated the use of tool sets and composite tools in army ant predation. Seringbara chimpanzees preyed on one army ant species ( D. nigricans ) more often than expected based on encounter rates, which may be explained by the overlap in altitudinal distribution between chimpanzees and D. nigricans . Army ant predation was not related to rainfall or fruit availability. Chimpanzees were selective in their choice of tool materials and collected their preferred tool species ( Alchornea hirtella ) from greater distances than they did other species. Lastly, Seringbara chimpanzees used both tool sets and composite tools (tree perch) in army ant predation. Tool types (dig vs. dip) differed in width and strength, but not length. Tool composites were found at 40% of ant‐feeding sites. Our study sheds new light on the ecology of army ant predation and provides novel insights into chimpanzee selection of army ant prey and tool species. Am. J. Primatol. 77:319–329, 2015. © 2014 Wiley Periodicals, Inc.

Identifying the forces shaping intraspecific phenotypic and genotypic divergence are of key importance in evolutionary biology. Phenotypic divergence may result from local adaptation or, especially in species with strong gene flow, from pronounced phenotypic plasticity. Here, we examine morphological and genetic divergence among populations of the western honey bee Apis mellifera in the topographically heterogeneous East African region. The currently accepted “mountain refugia hypothesis” states that populations living in disjunct montane forests belong to a different lineage than those in savanna habitats surrounding these forests. We obtained microsatellite data, mitochondrial sequences, and morphometric data from worker honey bees collected from feral colonies in three montane forests and corresponding neighboring savanna regions in Kenya. Honey bee colonies from montane forests showed distinct worker morphology compared with colonies in savanna areas. Mitochondrial sequence data did not support the existence of the two currently accepted subspecies. Furthermore, analyses of the microsatellite data with a Bayesian clustering method did not support the existence of two source populations as it would be expected under the mountain refugia scenario. Our findings suggest that phenotypic plasticity rather than distinct ancestry is the leading cause behind the phenotypic divergence observed between montane forest and savanna honey bees. Our study thus corroborates the idea that high gene flow may select for increased plasticity.

Harmonic radar tracking was used to record the flights of scout bees during takeoff and initial flight path of two honeybee swarms. One swarm remained intact and performed a full flight to a destination beyond the range of the harmonic radar, while a second swarm disintegrated within the range of the radar and most of the bees returned to the queen. The initial stretch of the full flight is characterized by accelerating speed, whereas the disintegrating swarm flew steadily at low speed. The two scouts in the swarm displaying full flight performed characteristic flight maneuvers. They flew at high speed when traveling in the direction of their destination and slowed down or returned over short stretches at low speed. Scouts in the disintegrating swarm did not exhibit the same kind of characteristic flight performance. Our data support the streaker bee hypothesis proposing that scout bees guide the swarm by traveling at high speed in the direction of the new nest site for short stretches of flight and slowing down when reversing flight direction.

Deciphering the evolutionary processes driving nucleotide variation in multiallelic genes is limited by the number of genetic systems in which such genes occur. The complementary sex determiner (csd) gene in the honey bee Apis mellifera is an informative example for studying allelic diversity and the underlying evolutionary forces in a well-described model of balancing selection. Acting as the primary signal of sex determination, diploid individuals heterozygous for csd develop into females, whereas csd homozygotes are diploid males that have zero fitness. Examining 77 of the functional heterozygous csd allele pairs, we established a combinatorical criteria that provide insights into the minimum number of amino acid differences among those pairs. Given a data set of 244 csd sequences, we show that the total number of csd alleles found in A. mellifera ranges from 53 (locally) to 87 (worldwide), which is much higher than was previously reported (20). Using a coupon-collector model, we extrapolate the presence of in total 116-145 csd alleles worldwide. The hypervariable region (HVR) is of particular importance in determining csd allele specificity, and we provide for this region evidence of high evolutionary rate for length differences exceeding those of microsatellites. The proportion of amino acids driven by positive selection and the rate of nonsynonymous substitutions in the HVR-flanking regions reach values close to 1 but differ with respect to the HVR length. Using a model of csd coalescence, we identified the high originating rate of csd specificities as a major evolutionary force, leading to an origin of a novel csd allele every 400,000 years. The csd polymorphism frequencies in natural populations indicate an excess of new mutations, whereas signs of ancestral transspecies polymorphism can still be detected. This study provides a comprehensive view of the enormous diversity and the evolutionary forces shaping a multiallelic gene.

East African montane forests have been subjected to heavy logging, particularly of Ocotea usambarensis Engl., formerly one of the dominant tree species of moist mid-altitude forests. At Mt. Kenya, logging was suspended in 2000 after a conspicuous decline in population size, but the success of this conservation measure has not yet been evaluated. Given that a management scheme of O. usambarensis forests based on vegetative regeneration has been suggested, we hypothesized that natural regeneration mainly by root suckers would be sufficient for a recovery of this species. Demography and regeneration (both sexual and vegetative) of O. usambarensis were studied in 45 study plots between 1700 and 2500 m asl along a gradient of historical logging intensity, while taking altitude and light incidence into account as predictor variables. The diameter distribution showed a high percentage of old individuals and rather low recruitment in O. usambarensis. In heavily logged areas (removed basal area >25 m(2)ha(-1)), smaller trees (<50 cm DBH, >130 cm high) were completely absent. The number of seedlings was low and independent of logging intensity. It increased with higher light incidence. The number of root suckers was 5.6-fold the number of seedlings, underscoring the importance of vegetative reproduction. However, number of root suckers and logging intensity were negatively correlated. We conclude that regeneration of O. usambarensis at Mt. Kenya is generally low and negatively influenced by historical logging. Therefore, natural regeneration is inadequate for the recovery of this valuable timber species, and additional conservation measures such as enrichment planting should be considered. (C) 2012 Elsevier B.V. All rights reserved.

SUMMARY The ectoparasitic mite Varroa destructor and honey bee pathogenic viruses have been implicated in the recent demise of honey bee colonies. Several studies have shown that the combination of V. destructor and deformed wing virus (DWV) poses an especially serious threat to honey bee health. Mites transmitting virulent forms of DWV may cause fatal DWV infections in the developing bee, while pupae parasitised by mites not inducing or activating overt DWV infections may develop normally. Adult bees respond to brood diseases by removing affected brood. This hygienic behaviour is an essential part of the bees' immune response repertoire and is also shown towards mite-parasitised brood. However, it is still unclear whether the bees react towards the mite in the brood cell or rather towards the damage done to the brood. We hypothesised that the extent of mite-associated damage rather than the mere presence of parasitising mites triggers hygienic behaviour. Hygienic behaviour assays performed with mites differing in their potential to transmit overt DWV infections revealed that brood parasitised by ‘virulent’ mites (i.e. mites with a high potential to induce fatal DWV infections in parasitised pupae) were removed significantly more often than brood parasitised by ‘less virulent’ mites (i.e. mites with a very low potential to induce overt DWV infections) or non-parasitised brood. Chemical analyses of brood odour profiles suggested that the bees recognise severely affected brood by olfactory cues. Our results suggest that bees show selective, damage-dependent hygienic behaviour, which may be an economic way for colonies to cope with mite infestation.

Army ant colonies do not have permanent nests but frequently move to new patches. Local food depletion is considered the ultimate cause of this nomadic behaviour, but the proximate causes are not well understood. We tested if and how patch departure time of the aboveground-hunting army ant Dorylus molestus under field conditions is influenced by food availability and nest attacks by predators. In the first food supplement experiment, colonies receiving additional food throughout an entire nest stay did not reside in their nests for longer periods than control colonies. However, the distances travelled by colonies after nest stays during which colonies obtained food were shorter than those before these nest stays, indicating that colonies do assess food availability and avoid moving too far away from patches of high food availability. In the second food supplement experiment, in which colonies were given even larger amounts of food in the second half of their nest stay to mimic a rich unpredictable food source that these highly polyphagous predators are likely to encounter sometimes, patch departure times likewise did not differ between treated and control colonies. Either patch departure time is independent of food availability or there is another, as yet unappreciated proximate cause of colony movements in this species which we were unable to control for in our field experiments. One possibility is that encounters between neighbouring colonies influence patch departure time. In the experiment on the effect of predation, colonies responded to simulated nest attacks by mammals by leaving nests almost instantaneously and thus much earlier than control colonies. Rapid nest evacuation is likely a response to minimize the probability of repeat attacks by predators which cannot be repelled in other ways. Future studies will be necessary to definitively determine whether food availability influences patch departure times and to elucidate the consequences of colony encounters.

BACKGROUND:Hybridization can have complex effects on evolutionary dynamics in ants because of the combination of haplodiploid sex-determination and eusociality. While hybrid non-reproductive workers have been found in a range of species, examples of gene-flow via hybrid queens and males are rare. We studied hybridization in East African army ants (Dorylus subgenus Anomma) using morphology, mitochondrial DNA sequences, and nuclear microsatellites.RESULTS:While the mitochondrial phylogeny had a strong geographic signal, different species were not recovered as monophyletic. At our main study site at Kakamega Forest, a mitochondrial haplotype was shared between a "Dorylus molestus-like" and a "Dorylus wilverthi-like" form. This pattern is best explained by introgression following hybridization between D. molestus and D. wilverthi. Microsatellite data from workers showed that the two morphological forms correspond to two distinct genetic clusters, with a significant proportion of individuals being classified as hybrids.CONCLUSIONS:We conclude that hybridization and gene-flow between the two army ant species D. molestus and D. wilverthi has occurred, and that mating between the two forms continues to regularly produce hybrid workers. Hybridization is particularly surprising in army ants because workers have control over which males are allowed to mate with a young virgin queen inside the colony.

A trademark of Homo sapiens is the enormous variation in behavioural patterns across populations. Insight into the development of human cultures can be aided by studies of Pan communities across Africa, which display unique combinations of social behaviour and elementary technology. Only cross-population comparisons can reveal whether this diversity reflects differential genetics, environmental constraints, or arbitrary cultural patterns. However, the recently recognised and most endangered subspecies Pan troglodytes vellerosus remains completely unstudied in this respect. We report on the Nigerian chimpanzees at Gashaka. At this site, diet composition is highly varied and the apes have to cope with high concentrations of anti-feedant defenses of plants against consumption. It is not surprising therefore, that Gashaka chimpanzees use a varied tool-kit for extractive foraging. For example, they harvest insects throughout the year, employing digging sticks and probes to obtain honey from nests of stingless bees and honey bees, dipping wands to prey on army ants and fishing rods to eat arboreal ants. Tools appeared to be custom-made with a considerable degree of standardisation and preferential use of distal ends. Many of these expressions of subsistence technology seem to be environmentally constrained. Most notably, the absence of termite eating could reflect a low abundance of mounds. Other traits may represent arbitrary cultural variation. For example, two types of hard-shelled nuts found in the habitat are not opened with tools, unlike what is observed elsewhere in West Africa. The prevalence of elementary technology may indicate that the material culture of Gashaka chimpanzees is most closely related to core cultural tendencies of Central African populations of these apes.

References [1] Thakur, R. K., Bienefeld, K.; Keller, R.: Varroa defence behavior in Apis mellifera carnica. American Bee Journal 137, pp. 143-148, 1997. [2] Viola, P., Jones, M.: Rapid Object Detection using a Boosted Cascade of Simple Features. CVPR 2001. [3] Knauer, U. et al.: Application of an Adaptive Background Model for Monitoring Honeybees. VIIP 2005 . [4] Knauer, U. et al.: Evaluation based combining of classifiers. WACV 2009. Methods TPR FPR Rank 1 Template Matching 93,01% 4,98% 3 2 Haarlike Features 91,26% 4,01% 5 3 Contour Detection 95,97% 1,74% 1 4 Interest Point Density 95,87% 4% 2 5 Hough Transform 92,59% 5,23% 4 Methods Precision Recall Rank 1 Template Matching 69,40% 82,50% 5 2 Haarlike Features 91,17% 90,50% 2 3 Contour Detection 94,66% 91,20% 1 4 Interest Point Density 86,70% 88% 3 5 Hough Transform 79,41% 84,37% 4

Several populations of chimpanzees have been reported to prey upon Dorylus army ants. The most common tool‐using technique to gather these ants is with “dipping” probes, which vary in length with regard to aggressiveness and lifestyle of the prey species. We report the use of a tool set in army ant predation by chimpanzees in the Goualougo Triangle, Republic of Congo. We recovered 1,060 tools used in this context and collected 25 video recordings of chimpanzee tool‐using behavior at ant nests. Two different types of tools were distinguished based on their form and function. The chimpanzees use a woody sapling to perforate the ant nest, and then a herb stem as a dipping tool to harvest the ants. All of the species of ants preyed upon in Goualougo are present and consumed by chimpanzees at other sites, but there are no other reports of such a regular or widespread use of more than one type of tool to prey upon Dorylus ants. Furthermore, this tool set differs from other types of tool combinations used by chimpanzees at this site for preying upon termites or gathering honey. Therefore, we conclude that these chimpanzees have developed a specialized method for preying upon army ants, which involves the use of an additional tool for opening nests. Further research is needed to determine which specific ecological and social factors may have shaped the emergence and maintenance of this technology. Am. J. Primatol. 72:17–24, 2010. © 2009 Wiley‐Liss, Inc.

African driver ants are nomadic social mesopredators feeding on a highly diverse array of prey species at different trophic levels. Colonies of certain driver ant species have a biomass which can equal that of medium-sized mammalian carnivores and the ultimate cause of their nomadic life-style is thought to be local prey depletion. The impact of driver ant swarm raids is therefore expected to be strong but the degree to which they reduce prey populations has not been quantified and it is unknown whether these spectacular predators exert significant top-down effects. We examined the combined effect of driver ant (Dorylus molestus) and swarm-attending bird (Alethe poliocephala) predation on the population dynamics of earthworms, which constitute the ants' main prey type in the montane forest of Mount Kenya. Pre-raid earthworm biomass densities in the soil layer down to a depth of 8 cm varied by a factor of 31. The immediate effect of swarm raids was a reduction in earthworm numbers in this layer, but 8 days later earthworm numbers had recovered to pre-raid levels. When earthworm biomass densities were compared, no significant effect of swarm raids was detected. The estimated proportion of earthworm prey biomass extracted from 0 to 8 cm layer by driver ants and birds together was about 2.2%. Although colony distribution was overdispersed as expected based on knowledge of D. molestus migratory behaviour, predation events were highly localized. Predation frequency was low (once every 62 days on average) and highly variable. These results indicate that earthworm prey is highly abundant but at the same time so difficult to harvest that swarm raids exert only a marginal influence on earthworm populations. Longer-term studies would be required to determine whether earthworm populations are limited by swarm raids. The small impacts of individual raids and rapid recovery of earthworm prey populations likely underlie the low frequency of migrations and short distances travelled by migrating colonies of D. molestus.

Theory predicts that altruism is only evolutionarily stable if it is preferentially directed towards relatives, so that any such behaviour towards seemingly unrelated individuals requires scrutiny. Queenless army ant colonies, which have anecdotally been reported to fuse with queenright foreign colonies, are such an enigmatic case. Here we combine experimental queen removal with population genetics and cuticular chemistry analyses to show that colonies of the African army ant Dorylus molestus frequently merge with neighbouring colonies after queen loss. Merging colonies often have no direct co-ancestry, but are on average probably distantly related because of overall population viscosity. The alternative of male production by orphaned workers appears to be so inefficient that residual inclusive fitness of orphaned workers might be maximized by indiscriminately merging with neighbouring colonies to increase their reproductive success. We show that worker chemical recognition profiles remain similar after queen loss, but rapidly change into a mixed colony Gestalt odour after fusion, consistent with indiscriminate acceptance of alien workers that are no longer aggressive. We hypothesize that colony fusion after queen loss might be more widespread, especially in spatially structured populations of social insects where worker reproduction is not profitable.

Although swarm-raiding army ants are considered keystone predators in tropical forest ecosystems, information on the prey spectra of most species is based on anecdotal reports and not on systematic studies with extensive sampling. We analysed prey samples of 18 colonies of the two afrotropical species Dorylus (Anomma) molestus and Dorylus (Anomma) wilverthi (4289 prey items in total) to examine the prey composition variation within and between species and to determine the best methodology to obtain reliable prey spectrum estimates for a given species, site and season. Variation in prey composition was substantial for D. molestus even within a single site and season, with the biomass proportion of the most important prey type differing by a factor of 12. Conclusions from studies using small samples sizes may thus be misleading. We demonstrate that the method of assessing prey spectra in terms of relative prey item numbers often produces biased results, and therefore recommend relative prey biomass as the more useful parameter. The near absence of earthworms, which always constituted a substantial part of the D. molestus prey, in the diet of D. wilverthi is interpreted to result from subtle differences in swarm-raiding behaviour between these two species, but could alternatively also be due to low availability. Similar studies recording prey composition as biomass proportions and analysing large samples sizes from many colonies are needed to understand the effect of army ant swarm raids on invertebrate communities in afrotropical forests.