Foraging at the Edge of Chaos: Internal Clock versus External Forcing

Activity rhythms in animal groups arise both from external changes in the environment, as well as from internal group dynamics. These cycles are reminiscent of physical and chemical systems with quasiperiodic and even chaotic behavior resulting from “autocatalytic” mechanisms. We use nonlinear differential equations to model how the coupling between the self-excitatory interactions of individuals and external forcing can produce four different types of activity rhythms: quasiperiodic, chaotic, phase locked, and displaying over or under shooting. At the transition between quasiperiodic and chaotic regimes, activity cycles are asymmetrical, with rapid activity increases and slower decreases and a phase shift between external forcing and activity. We find similar activity patterns in ant colonies in response to varying temperature during the day. Thus foraging ants operate in a region of quasiperiodicity close to a cascade of transitions leading to chaos. The model suggests that a wide range of temporal structures and irregularities seen in the activity of animal and human groups might be accounted for by the coupling between collectively generated internal clocks and external forcings.

Epigenetics and behavioural plasticity: drosophila euchromatin histone metiltransferase and foraging

A thesis submitted in fulfillment of the requirements for the degree of Masters in Molecular Genetics and Biomedicine

Observations on dynamics of foraging hole construction of two leaf-feeding, soil-inhabiting Syntermes species (insecta: Isoptera) in an Amazonian Rainforest, Brazil

The intensity of construction of foraging access holes by two leaf-litter feeding, soil- inhabiting termite species, Syntermes molestus and Syntermes spinosus, in a Central Amazonian rain forest, was observed on consecutive nights for two weeks. Between 11 and 48 nest entrances per m2 were counted. Interaction between the two species was intense; some entrance holes were overtaken by the larger species during the observations; however, both species coexist in the area. A calculated minimum of 35 entrances/m2 is built every year by both species, emphasizing the importance of soil-burrowing termites for soil structure, aeration and water regime.

Water depth selection during foraging and efficiency in prey capture by the egrets Casmerodius albus and Egretta thula (Aves, Ardeidae) in an urban lagoon in Rio de Janeiro State, Brazil

This study aimed to evaluate the water depth selection during foraging, the efficiency in prey capture, and the food items captured by Casmerodius albus (Linnaeus, 1758) and Egretta thula (Molina, 1782). The work was conducted at an urban lagoon, Lagoa Rodrigo de Freitas, Rio de Janeiro. Four transects were made each month (two in the morning and two in the afternoon) for six months. When the birds were detected foraging, the water depth and the types of prey captured were recorded. There was no significant relationship between the foraging efficiencies of the two species. However, they differed in relation to the water depth when foraging, and also in the food items captured. Casmerodius albus captured mainly fishes while Egretta thula captured mainly invertebrates. The results suggest that the differences in water depth when foraging and the food items captured allow a differential use of the food resources available by C. albus and E. thula at Lagoa Rodrigo de Freitas.

Trophic ecology and foraging behavior of Tropidurus hispidus and Tropidurus semitaeniatus (Squamata, Tropiduridae) in a caatinga area of northeastern Brazil

This study aimed to analyze the seasonal variation in diet composition and foraging behavior of Tropidurus hispidus (Spix, 1825) and T. semitaeniatus (Spix, 1825), as well as measurement of the foraging intensity (number of moves, time spent stationary, distance traveled and number of attacks on prey items) in a caatinga patch on the state of Rio Grande do Norte, Brazil. Hymenoptera/Formicidae and Isoptera predominated in the diet of both species during the dry season. Opportunistic predation on lepidopteran larvae, coleopteran larvae and adults, and orthopteran nymphs and adults occurred in the wet season; however, hymenopterans/Formicidae were the most important prey items. The number of food items was similar between lizard species in both seasons; however the overlap for number of prey was smaller in the wet season. Preys ingested by T. hispidus during the wet season were also larger than those consumed by T. semitaeniatus. Seasonal comparisons of foraging intensity between the two species differed, mainly in the wet season, when T. hispidus exhibited less movement and fewer attacks on prey, and more time spent stationary if compared to T. semitaeniatus. Although both lizards are sit-and-wait foragers, T. semitaeniatus is more active than T. hispidus. The diet and foraging behavior of T. hispidus and T. semitaeniatus overlap under limiting conditions during the dry season, and are segregative factors that may contribute to the coexistence of these species in the wet season.

Foraging behavioral of Phylloscartes ventralis (Aves, Tyrannidae) in native and planted forests of southern Brazil

Few studies have related the effects of silviculture practices to the behavior of bird species in the Neotropics. The present study examined the foraging behavior of Phylloscartes ventralis (Temminck, 1824) in a native forest and in silviculture areas of Pinus elliotti and Araucaria angustifolia with different structures and ages. We tested two general hypotheses: (1) areas of commercial forest plantation change the foraging behavior of P. ventralis in relation to native forest, and (2) the foraging behavior of P. ventralis in silviculture areas with understories (complex structures) is different from its behavior in areas without understory. The results showed that P. ventralis changed its foraging behavior depending on the type of forest, and on the presence of an understory in silviculture areas. Main changes involved the height and angle of substrate where the prey was captured. Phylloscartes ventralis showed the same set of attack maneuvers, with more maneuvers type in young Pinus planted without understory. The frequency of use of attack maneuvers was more similar in areas of silviculture with understory and in the native forest. The results highlight the importance of an understory structure and the utilization of native plant species in silviculture practices, to the foraging behavior of native bird species.

The locust foraging gene.

Our knowledge of how genes act on the nervous system in response to the environment to generate behavioral plasticity is limited. A number of recent advancements in this area concern food-related behaviors and a specific gene family called foraging (for), which encodes a cGMP-dependent protein kinase (PKG). The desert locust (Schistocerca gregaria) is notorious for its destructive feeding and long-term migratory behavior. Locust phase polyphenism is an extreme example of environmentally induced behavioral plasticity. In response to changes in population density, locusts dramatically alter their behavior, from solitary and relatively sedentary behavior to active aggregation and swarming. Very little is known about the molecular and genetic basis of this striking behavioral phenomenon. Here we initiated studies into the locust for gene by identifying, cloning, and studying expression of the gene in the locust brain. We determined the phylogenetic relationships between the locust PKG and other known PKG proteins in insects. FOR expression was found to be confined to neurons of the anterior midline of the brain, the pars intercerebralis. Our results suggest that differences in PKG enzyme activity are correlated to well-established phase-related behavioral differences. These results lay the groundwork for functional studies of the locust for gene and its possible relations to locust phase polyphenism.

The importance of root-produced volatiles as foraging cues for entomopathogenic nematodes

Background Entomopathogenic nematodes (EPNs) are tiny parasitic worms that parasitize insects, in which they reproduce. Their foraging behavior has been subject to numerous studies, most of which have proposed that, at short distances, EPNs use chemicals that are emitted directly from the host as host location cues. Carbon dioxide (CO2) in particular has been implicated as an important cue. Recent evidence shows that at longer distances several EPNs take advantage of volatiles that are specifically emitted by roots in response to insect attack. Studies that have revealed these plant-mediated interactions among three trophic levels have been met with some disbelief. Scope This review aims to take away this skepticism by summarizing the evidence for a role of root volatiles as foraging cues for EPNs. To reinforce our argument, we conducted olfactometer assays in which we directly compared the attraction of an EPN species to CO2 and two typical inducible root volatiles. Conclusions The combination of the ubiquitous gas and a more specific root volatile was found to be considerably more attractive than one of the two alone. Hence, future studies on EPN foraging behavior should take into account that CO2 and plant volatiles may work in synergy as attractants for EPNs. Recent research efforts also reveal prospects of exploiting plant-produced signals to improve the biological control of insect pests in the rhizosphere.

Evolution of foraging behaviour in response to chronic malnutrition in Drosophila melanogaster.

Chronic exposure to food of low quality may exert conflicting selection pressures on foraging behaviour. On the one hand, more active search behaviour may allow the animal to find patches with slightly better, or more, food; on the other hand, such active foraging is energetically costly, and thus may be opposed by selection for energetic efficiency. Here, we test these alternative hypotheses in Drosophila larvae. We show that populations which experimentally evolved improved tolerance to larval chronic malnutrition have shorter foraging path length than unselected control populations. A behavioural polymorphism in foraging path length (the rover-sitter polymorphism) exists in nature and is attributed to the foraging locus (for). We show that a sitter strain (for(s2)) survives better on the poor food than the rover strain (for(R)), confirming that the sitter foraging strategy is advantageous under malnutrition. Larvae of the selected and control populations did not differ in global for expression. However, a quantitative complementation test suggests that the for locus may have contributed to the adaptation to poor food in one of the selected populations, either through a change in for allele frequencies, or by interacting epistatically with alleles at other loci. Irrespective of its genetic basis, our results provide two independent lines of evidence that sitter-like foraging behaviour is favoured under chronic larval malnutrition.

Ecology and evolution of soil nematode chemotaxis.

Plants influence the behavior of and modify community composition of soil-dwelling organisms through the exudation of organic molecules. Given the chemical complexity of the soil matrix, soil-dwelling organisms have evolved the ability to detect and respond to these cues for successful foraging. A key question is how specific these responses are and how they may evolve. Here, we review and discuss the ecology and evolution of chemotaxis of soil nematodes. Soil nematodes are a group of diverse functional and taxonomic types, which may reveal a variety of responses. We predicted that nematodes of different feeding guilds use host-specific cues for chemotaxis. However, the examination of a comprehensive nematode phylogeny revealed that distantly related nematodes, and nematodes from different feeding guilds, can exploit the same signals for positive orientation. Carbon dioxide (CO(2)), which is ubiquitous in soil and indicates biological activity, is widely used as such a cue. The use of the same signals by a variety of species and species groups suggests that parts of the chemo-sensory machinery have remained highly conserved during the radiation of nematodes. However, besides CO(2), many other chemical compounds, belonging to different chemical classes, have been shown to induce chemotaxis in nematodes. Plants surrounded by a complex nematode community, including beneficial entomopathogenic nematodes, plant-parasitic nematodes, as well as microbial feeders, are thus under diffuse selection for producing specific molecules in the rhizosphere that maximize their fitness. However, it is largely unknown how selection may operate and how belowground signaling may evolve. Given the paucity of data for certain groups of nematodes, future work is needed to better understand the evolutionary mechanisms of communication between plant roots and soil biota.

Expression of foraging and Gp-9 are associated with social organization in the fire ant Solenopsis invicta.

The aim of this study was to investigate levels of expression of two major genes, the odorant binding protein Gp-9 (general protein-9) and foraging, that have been shown to be associated with behavioural polymorphisms in ants. We analysed workers and young nonreproductive queens collected from nests of the monogyne (single reproductive queen per nest) and polygyne (multiple reproductive queens) social forms of Solenopsis invicta. In workers but not young queens, the level of foraging expression was significantly associated with social form and the task performed (ie localization in the nest or foraging area). The level of expression of Gp-9 was also associated with social form and worker localization. In addition there was a higher level of expression of the Gp-9(b) allele compared with the Gp-9(B) allele in the heterozygote workers and the young nonreproductive queens. Finally, in the polygyne colonies the level of expression of foraging was not significantly associated with the Gp-9 genotype for either workers or young nonreproductive queens, suggesting that both genes have independent non-epistatic effects on behaviour in S. invicta.

A review of evolution of oviposition guilds in the Bruchidae (Coleoptera)

Three guilds of bruchid beetles oviposit on seeds at different times and in different ways, i. e., in these guilds some species only oviposit on fruits while on the plant (Guild A), other species only oviposit on seeds exposed in fruits while still on the plant (Guild B) and some only oviposit on seeds once they are exposed on the substrate (Guild C). It has been established that one plant species may be oviposited upon by all three guilds, some only by two guilds and some by only one guild. Before and after the inception of this concept many papers have been published that seem to establish that early oviposition behavior of bruchids was probably onto fruits where they burrowed through the fruit wall and fed on seeds (Guild A). Then, as evolution of the fruits developed for dispersal of seeds and possible escape from bruchid predation, bruchids developed to feed in seeds in various other ways (Guilds B and C). Our data show that about 78% of extant bruchids oviposit on fruits, and the other 22% with behavior of Guilds B and C. A review of these papers and new data on oviposition guilds and bruchid evolution are presented and discussed here.

Foraging by Polybia (Trichothorax) ignobilis (Hymenoptera, Vespidae) on flies at animal carcasses

Proteins for brood nutrition of social wasps are obtained from many prey, including insects (even bees and other wasps), spiders and bits of decaying meat. After being captured and killed, prey are reduced to a shapeless mass and distributed to the brood. Little is known about the foraging activity, especially on this group. Herein we describe the sequence of foraging behaviours of the social wasp Polybia (Trichothorax) ignobilis for hunting flies (Diptera: Calliphoridae and Muscidae) over pig carcasses. To our knowledge, there are few scientific descriptions of prey foraging behaviour on this species.

Effect of Citrus floral extracts on the foraging behavior of the stingless bee Scaptotrigona pectoralis (Dalla Torre)

Effect of Citrus floral extracts on the foraging behavior of the stingless bee Scaptotrigona pectoralis (Dalla Torre). Stingless bees have an important role as pollinators of many wild and cultivated plant species in tropical regions. Little is known, however, about the interaction between floral fragrances and the foraging behavior of meliponine species. Thus we investigated the chemical composition of the extracts of citric (lemon and orange) flowers and their effects on the foraging behavior of the stingless bee Scaptotrigona pectoralis. We found that each type of flower has its own specific blend of major compounds: limonene (62.9%) for lemon flowers, and farnesol (26.5%), (E)-nerolidol (20.8%), and linalool (12.7%) for orange flowers. In the foraging experiments the S. pectoralis workers were able to use the flower extracts to orient to the food source, overlooking plates baited with hexane only. However, orange flower extracts were seemingly more attractive to these worker bees, maybe because of the particular blend present in it. Our results reveal that these fragrances are very attractive to S. pectoralis, so we can infer that within citric orchards they could be important visitors in the study area; however habitat destruction, overuse of pesticides and the competitive override by managed honeybees might have put at risk their populations and thus the ecological services they provide to us.

Foraging strategies of the ant Ectatomma vizottoi (Hymenoptera, Formicidae)

Foraging strategies of the ant Ectatomma vizottoi (Hymenoptera, Formicidae). Foraging activity may be limited by temperature, humidity, radiation, wind, and other abiotic factors, all of which can affect energy costs during foraging. Ectatomma vizottoi's biology has only recently been studied, and no detailed information is available on its foraging patterns or diet in the field. For this reason, and because foraging activity is an important part of the ecological success of social insects, the present study aimed to investigate E. vizottoi's foraging strategies and dietary habits. First, we determined how abiotic factors constrained E. vizottoi's foraging patterns in the field by monitoring the foraging activity of 16 colonies on eight different days across two seasons. Second, we characterized E. vizottoi's diet by monitoring another set of 26 colonies during peak foraging activity. Our results show that E. vizottoi has foraging strategies that are similar to those of congeneric species. In spite of having a low efficiency index, colonies adopted strategies that allowed them to successfully obtain food resources while avoiding adverse conditions. These strategies included preying on other ant species, a foraging tactic that could arise if a wide variety of food items are not available in the environment or if E. vizottoi simply prefers, regardless of resource availability, to prey on other invertebrates and especially on other ant species.