Interspecific Agonistic Behavior of <Emphasis Type="Italic">Macrotermes gilvus</Emphasis> (Isoptera: Termitidae): Implication on Termite Baiting in the Philippines

The study investigated interspecific agonistic behavior of Macrotermes gilvus Hagen (Isoptera: Termitidae: Macrotermitinae) against three economically important subterranean termites in the Philippines, viz., Coptotermes gestroi Wasmann, Nasutitermes luzonicus Oshima and Microcerotermes losbanosensis Oshima. Termite-termite interactions after a 1:1 pairing experiment showed that M. gilvus workers and soldiers were highly aggressive against C. gestroi, N. luzonicus and M. losbanosensis leading to severe injury or death of the opponent termite species in a short period of time. The levels of agonism were caste and species specific. Worker termites of M. gilvus showed an equally aggressive behavior as soldiers contributing to the high mortality of opponent species used in this study. It is likely that the highly aggressive behavior of M. gilvus limits foraging activity of C. gestroi, N. luzonicus and M. losbanosensis around in-ground bait stations contributing to the low success of termite baits containing chitin synthesis inhibitors in the Philippines.     

Interspecific Interactions Between Coptotermes formosanus and Reticulitermes flavipes (Isoptera: Rhinotermitidae) in Laboratory Bioassays

Experiments were conducted to examine competitive interactions between the Formosan subterranean termite, Coptotermes formosanus Shiraki (FST), and the eastern subterranean termite, Reticulitermes flavipes (Kollar) (EST), using groups of termites with different worker:soldier proportions. Experiments were conducted using three connected test chambers: an FST chamber, an unoccupied center chamber, and an EST chamber. When groups of FST were comprised of 20% soldiers versus 2% EST soldiers, only 8% of center chambers were occupied exclusively by EST. When groups of FST were comprised of 10% soldiers versus 1% EST soldiers, 44% of center chambers were occupied exclusively by EST. When the only food source was located in the center chamber, 60% of center chambers were occupied by both species. FST did not completely displace EST in any of these experiments.     

Effectiveness of recruitment behavior in stingless bees (Apidae,Meliponini)

Summary We examined the ability of stingless bees to recruit nest mates to a food source (i) in group foraging species laying pheromone trails from the food to the nest (Trigona recursa Smith, T. hypogea Silvestri, Scaptotrigona depilis Moure), (ii) in solitary foraging species with possible but still doubtful communication of food location inside the nest (Melipona seminigra Friese, M. favosa orbignyi Guérin), and (iii) in species with a less precise (Nannotrigona testaceicornis Lep., Tetragona clavipes Fab.) or no communication (Frieseomelitta varia Lep.). The bees were allowed to collect food (sugar solution or liver in the necrophageous species) ad libitum and the forager number to accumulate, as it would do under normal unrestrained conditions. The median number of bees collecting differed considerably among the species (1.0–1436.5). It was highest in the species employing scent trails. The time course of recruitment was characteristic for most of the species and largely independent of the number of foragers involved. The two Melipona species recruited other bees significantly faster than T. recursa, S. depilis, and N. testaceicornis during the first 10 to 30 minutes of an experiment. In species laying a scent trail to guide nestmates to a food source the first recruits appeared with a delay of several minutes followed by a quick increase in forager number. The median time required to recruit all foragers available differed among the species between 95.0 and 240.0 min. These differences can at least partly be explained by differences in the recruitment mechanisms and do not simply follow from differences in colony biomass.     

Directional fidelity as a foraging constraint in the western harvester ant,Pogonomyrmex occidentalis

Summary Understanding the foraging behavior of an animal is critically dependent upon knowledge of the constraints on that animal. In this study, I tested whether fidelity to foraging direction acts as a behavioral constraint to foraging western harvester ants, Pogonomyrmex occidentalis. Individual P. occidentalis foragers showed strong fidelity to foraging route and direction. Directional fidelity in this population was not related to trunk trail use, food specialization, colony activity levels, or mortality risks. Directional fidelity constrained individual foraging decisions; when colonies were offered seeds of different quality in 2 directions, individuals did not switch directions to obtain the energetically more rewarding seeds. Colony-level recruitment was increased for energetically more profitable seeds, indicating that colonial responses may compensate for the constraints of directional fidelity on individual foragers.     

Localized treatments using commercial dust and liquid formulations of fipronil against Coptotermes formosanus （Isoptera： Rhinotermitidae） in the laboratory

Use of proper application methods and formulations of termiticides are im- portant to reduce their negative impact to the environment. In this study, we conducted laboratory experiments to determine the effect of localized treatments with commercial dust and liquid formulations of fipronil against Formosan subterranean termites, Coptoter- mes formosanus Shiraki. The test arena consisted of a specially designed 16-chambered structure with a center chamber connected to 5 foraging chambers that themselves were connected to 10 additional foraging chambers. One peripheral chamber received a liquid or dust treatment and termites were released in the center chamber. Results showed that 〉 91% of the termites were dead within the 9-d test period despite the localized treatment of only 1 foraging chamber. Termites that were still alive after 9 d were transferred to an untreated dish and held for 10 more days. The majority of those termites were dead and the rest were moribund on day 19. Regardless of the specific dish treated, both formulations of fipronil were found to be highly efficacious. Termites did not exhibit repellency to either formulation. Our results suggest that localized （or spot） treatment with either commer- cially available dust or liquid formulations of fipronil can be a viable option for control of a termite infestation where complete soil drenching is not desirable.     

Orientation and Communication in the Neotropical Ant Odontomachus bauri Emery (Hymenoptera,Formicidae, Ponerinae)

The Neotropical species Odontomachus bauri employs canopy orientation during foraging and homing. An artificial canopy pattern above the ants is much more effective as an orientation cue than horizontal landmarks or chemical marks. However, both horizontal visual cues and chemical marks on the ground can serve in localizing the nest entrance. Successful O. bauri foragers recruit nestmates to leave the nest and search for food. However, the recruitment signals do not contain directional information. Antennation bouts and pheromones from the pygidial gland most likely serve as stimulating recruitment signals. Secretions from the mandibular and poison gland elicit alarm and attack behavior.     

A comparison of two methods for sampling assemblages of subterranean,wood-eating termites (Isoptera)

Abstract Within a 50 × 50 m area of wandoo Eucalyptus capillosa woodland in the Western Australian wheatbelt, the diversity and frequency of occurrence of wood-eating termite species was assessed at two food types. Over a 12 month period, monthly termite activity was determined: (i) at sound/undecayed artificial baits (seasoned wooden stakes of Jarrah, Karri, Pine, Batu, Oregon; Jarrah sawdust; paper rolls); and (li) at naturally occurring timber, fallen logs and branches of wandoo, in varying stages of decay. Termite diversity was 11 species at baits, 18 species at wandoo out of an overall site richness of 21 species. Karri attracted the most species (9); sawdust attracted none. At wandoo, Nasutitermes exitiosus, Coptotermes acinaciformis and Occasitermes occasus accounted for 59% of samples where termites were recorded. At baits, Heterotermes occiduus accounted for a mean of 80% of samples across bait types, but was rarely sampled at wandoo (5% of samples). Only H. occiduus, C. acinaciformis and Amitermes neogermanus ate bait. Pine, Oregon and paper rolls were most effective in attracting foraging termites in terms of highest per cent of replicates showing bait consumption and highest consumption rates. Jarrah and Batu were least attractive to foraging termites. Samples from wandoo underestimated the relative frequency of occurrence of H. occiduus within the study site. Coptotermes acinaciformis, which attack large food items, and certain species of Amitermes, which forage on subterranean food, may have been underestimated by both sampling methods. These findings indicate that a proper understanding of the structure of wood-eating termite assemblages within a given area requires a composite sampling strategy which addresses termites that eat sound or decayed wood, as well as surface and subsurface foragers.     

Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars

Aggressive interactions, foraging behavior, habitat use and diet were studied in sympatric populations of white-sported char,Salvelinus leucomaenis, and Dolly Varden,Salvelinus malma, in a Japanese mountain stream. Underwater observations on individuals of both species revealed two distinct behavioral regimes: aggressive drift foragers and non-aggressive benthos foragers. Aggressive drift foragers defended partial territories around focal points from which they made forays to capture invertebrates drifting in the water column. Non-aggressive benthos foragers cruised around and beneath cobble in large foraging ranges that overlapped each other. Intra- and interspecific, size-dependent dominance hierarchies were recognized among aggressive drift foragers, whereas non-aggressive benthos foragers showed no such relationships. Terrestrial invertebrates were the most abundant prey in the diets of drift foragers, whereas a very small proportion of the diet of benthos foragers was made up of these taxa. Benthos foragers showed more complex diet composition than drift foragers. These results suggest that non-aggressive benthos foragers may avoid not only interference but also exploitative competition by using alternative foraging tactics. The proportion of drift foragers to benthos foragers among white-spotted char was more than 35 times that among Dolly Varden. The significant difference in the proportion of each species using the two types of foraging strategy results in interspecific food segregation in sympatric populations.     

Tarsomere and distal tibial glands: Structure and potential roles in termites (Isoptera: Rhinotermitidae,Termitidae)

Social insects have numerous exocrine glands, but these organs are understudied in termites compared to hymenopterans. The tarsomere and distal tibial glands of the termites Heterotermes tenuis, Coptotermes gestroi and Silvestritermes euamignathus were investigated by scanning and transmission electron microscopy. Pore plates are visible in scanning micrographs on the distal tibial surfaces and on the ventral surface of the first and second tarsomeres of workers of H. tenuis and C. gestroi. In contrast, workers of S. euamignathus have isolated pores spread throughout the ventral surfaces of the first, second, and third tarsomeres and the distal tibia. In all three species each pore corresponds to the opening of a class-3 secretory unit, composed of one secretory and one canal cell. Clusters of class-3 glandular cells are arranged side by side underneath the cuticle. The main characteristics of these exocrine glands include their presence on all the legs and the electron-lucent secretion in the secretory cells. Possible functions of these glands are discussed.     

Influence of environmental and colony factors on the initial commodity choice of foragers of the stingless bee Plebeia tobagoensis (Hymenoptera, Meliponini)

Novice foragers of social bees have to decide what food commodity to collect when they start foraging for the first time. In this decision making process two types of factors are involved: internal factors (the response threshold) and external factors (environmental and colony conditions). In this study we will focus on the importance of two external factors, pollen storage level and information from experienced foragers about food availability in the field, on the initial commodity choice of foragers of the stingless bee species Plebeia tobagoensis. We also studied the effect of the initial choice of individuals on their subsequent foraging career. This study was performed in a closed greenhouse compartment, where food availability and colony condition could be controlled. Information on food availability in the field from experienced foragers and pollen storage level both greatly influenced the initial commodity choice of individuals, with more choices for the commodity communicated by experienced foragers or lacking in storage. The initial choice of foragers is of importance for their future foraging career, although a substantial proportion of foragers did switch between food commodities. Because of the ability of novice foragers to become flexibly distributed over foraging tasks, social bees are able to react to changes in their environment without directly having to decrease foraging effort devoted to other foraging tasks. This, in combination with individual flexibility during foraging careers makes it possible for colonies of P. tobagoensis to forage efficiently in an ever-changing environment. Received 7 November 2005; revised 12 January 2006; accepted 16 February 2006.     

Foraging,recruitment and predation by Decamorium uelense (Sanstchi) (Formicidae: Myrmicinae) on termites in Southern Guinea Savanna,Nigeria

Summary The myrmicine ant Decamorium uelense is a predator of small termites which feed inside their food sources of roots and plant debris. The ant uses a specialized recruitment regime to secure its prey. A scout ant searches for foraging termites and returns to the nest where it recruits a column of 10–30 ants which attack and immobilize the termites. A mass recruitment phase is then instigated, with larger numbers of ants retrieving the prey. The major prey item is Microtermes (consumed at an annual rate of 632 termites m^-2), with other small termites accounting for 0.5% of the total predation. The annual predation of Microtermes in primary savanna woodland by D. uelense removes 74% of the standing population.     

Prey chemical discrimination and strike-induced chemosensory searching in lizards: Their absence in a crotaphytid lizard (Crotaphytus collaris) and a proposal for research in zoos

Lizards in seleroglossan families of actively foraging carnivores and the herbivorous iguanids use the tongue to gather chemical samples to detect, identify, and locate food prior to attack, and to relocate lost food. In contrast, previously studied iguanian families other than Iguanidae lack lingually mediated prey chemical discrimination (PCD) and do not exhibit lingually mediated chemosensory searching behavior for bitten prey that has escaped or been lost (SICS = strike-induced chemosensory searching). In the present study, experimental tests showed that PCD and SICS are absent in Crotaphytus collaris, a member of the previously unstudied family of iguanian ambush foragers, Crotaphytidae. Available data suggest that in active foragers natural selection favors use of chemical cues to locate hidden prey, whereas in ambush foragers natural selection favors immobility to avoid detection by predators and SICS precludes simultaneous ambush. In most families the states of PCD and SICS are retained from the ancestors, but when foraging mode shifts, a change in chemosensory behavior appears to be induced. A proposal is made for a research program involving herpetologists at zoos. © 1996 Wiley-Liss, Inc.     

Escape Behavior of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae) in Response to Disturbance

Disturbances to monitors and baits have often been noted as a confounding factor for the success of termite baiting systems. We studied escape behavior of Coptotermes formosanus Shiraki to a food source disturbance in the laboratory. Four peripheral chambers connected to the central food chamber through tubes served as escape routes from which escape patterns, escape and return times and differences in soldier and worker responses were recorded. We used two sand conditions: moist and dry to test whether favorable versus unfavorable conditions change escape behavior. The post disturbance responses were video recorded for 1 h. C. formosanus quickly escaped from the central chamber when disturbed. The time taken to reach the maximum number of termites that escaped ranged from 20 s to 210 s followed by a gradual return to the disturbed chamber. The majority of the escapees returned to the disturbed area within a few minutes after evacuation in both dry and wet sand conditions. A significantly higher percentage of termites escaped in wet sand arenas (43%) than in the dry sand arenas (31%). Our experiments demonstrate that C. formosanus is highly unlikely to abandon bait stations for extensive periods of time as a result of mechanical disturbances regardless of habitat quality.     

Tunnel geometry of the subterranean termite Reticulitermes grassei （Isoptera： Rhinotermitidae）in response to sand bulk density and the presence of food

The cryptic habits of subterranean termites restricts detailed analysis of their foraging patterns in situ, but the process is evidently dominated by tunnel constructions connecting the nest with woody resources discovered within the territory of each colony. In this study, tunnel formation and orientation were studied experimentally in the termite Reticulitermes grassei （Clement）, using 2-dimensional laboratory foraging arenas con- taining fine sand as the substratum. The building of exploratory tunnels over a 10-day period and the geometry of the resulting network are described. Fractal analysis showed that tunnel geometry had a fractal dimension, regardless of the total length tunnelled whether foragers encountered the food source or not. The bulk density of the sand in the arenas affected the distances tunnelled, with higher density reducing construction, but did not affect tunnel geometry. Tunnels were not discernibly orientated with respect to the positioning of the food source, even in a situation where termites had failed to find the food source at a distance of less than 50 mm, suggesting that volatiles from wood are not attractants.     

Foraging Behavior in Two Species of Ectatomma (Formicidae: Ponerinae): Individual Learning of Orientation and Timing

The foraging behavior of marked individuals of Ectatomma ruidum and E. tuberculatum (Formicidae: Ponerinae) was observed on Barro Colorado Island, Panama, in order to determine whether learning of orientation and timing is involved when foraging on patchy resources on the forest floor. Foraging experiments under seminatural conditions were designed to control for the spatiotemporal distribution of resources at one or two preying sites. Using both single- and multi–event past experience, individual foragers of E. ruidum and (although less significantly) E. tuberculatum made directional and temporal adjustments to their behavior in response to previous differential foraging success. In spite of considerable individual variation in foraging efficiency, it is hypothesized that the observed differences in cognitive abilities can be invoked to explain ecological differences among both species.     

Food acquisition by competing surfperch on a patchy environmental gradient

Synopsis Black surfperch, Embiotoca jacksoni, and striped surfperch, Embiotoca lateralis, coexisted along steep sloping rocky habitats at Santa Cruz Island, California. The range of depths occupied (to 15 m) was characterized by a strong gradient in abundance of prey and a changing mosaic of substrate types from which surfperch harvested food. Availability of prey and diversity of benthic substrates were greatest in shallowest areas and both declined with increasing depth. Individuals of both surfperch species were residential within a narrow range of depths, with the result that different segments of their populations were consistently exposed to different foraging environments. These two phenomena (residential behavior combined with a gradient in availability of resources) resulted in variation in foraging behaviors and diets among individuals that resided at different depths. The pattern of within-population variation differed between the surfperch species. Black surfperch individuals achieved similar taxonomic diets and expended similar foraging effort at all depths, but deep-water foragers captured much less prey biomass per unit effort. The taxonomic composition of striped surfperch diets differed among depths, and although similar amounts of prey biomass were captured everywhere, individuals in deep areas expended much greater effort to obtain that level of food return. For both species, habitat profitability (food return to foraging effort) declined with depth. The difference in habitat profitability appeared to influence fitness components of both surfperches. Individuals occupying deep habitats were about 5% shorter in standard length than conspecifics of the same chronological age living in shallow areas; the disparity in body size resulted in an estimated difference in clutch size of 10–18%.     

华南地区典型生境中红火蚁觅食行为及工蚁召集规律

观察研究了华南地区几种典型生境中红火蚁对不同类型食物的觅食行为和工蚁召集动态规律。研究结果表明红火蚁觅食行为存在搜寻、召集及搬运等主要过程。不同生境中红火蚁对食物的搜寻时间存在一定差异,荔枝园中搜寻时间明显长于其它生境;荔枝园、荒地、路边生境中红火蚁对蜂蜜的搜寻时间明显长于其它几种食物;同一生境中红火蚁对不同重量的同种食物搜寻时间无明显变化。发现食物后召集的工蚁数量随着时间延长呈现不断增加的趋势,对于较大的食物一般30min左右召集的工蚁数量达到最大,之后趋于稳定并逐渐减少,而对于可以直接搬动的食物发现后15min左右群体召集数量达到最大并很快将其搬走。召集工蚁数量与发现后时间的关系符合二次非线性方程,建立了火腿肠、花生油和蜂蜜等食物上红火蚁工蚁召集数量与发现时间的关系模型,分别为Na=4183.91e^-0.0327T-4231.48e^-0.0346T,Na=3253.78e^-0.0233T-3314.59e^-0.0271T,Na=117.97e^-0.0131T-163.93e^-0.0808T。对于不同食物红火蚁发现后召集的最大工蚁数量间有明显差异,其中花生油上最大,平均为176．3头,火腿肠上次之,为90．4头,蜂蜜上最少,为68．0头。对于不同重量的同种食物,工蚁的召集动态规律较为一致,发现食物后25-30min左右工蚁的召集数量均达到最大值,之后缓慢减少,发现食物后时间长度和食物上工蚁数量也符合房室模型函数Na=c1×exp（-c2T）-c3×exp（-c4T）。同种食物不同重量之间同一时间召集的工蚁数量存在较大差异,呈随食物重量增大而增大趋势。不同生境中对相同质量同种食物工蚁召集数量动态总体变化规律相近,但在发现食物后工蚁数量增长的速度和最大召集工蚁的数量存在明显差异,以路边工蚁数量增长最快、工蚁总数最多,苗圃次之,荔枝园、荒地中工蚁增长速度均较低。红火蚁对不同食物种类切割、搬运耗时存在很大差异,搬运完0．5g的火腿肠需要26．23h,而黄粉虫幼虫（重量约为0．1g）仅为15．6min。不同生境中红火蚁对相同重量同一种食物的搬运耗时也不同,荔枝园要长于其它生境。     

German Yellowjacket (Vespula germanica) Foragers Use Odors Inside the Nest to Find Carbohydrate Food Sources

Prior work has shown that yellowjacket waSPS remember food odors and use them as cues when foraging. There is also evidence they have mechanisms to recruit nest mates to highly rewarding food sources, as naïve individuals are more likely to go to food sources with scents similar to those visited by nest mates. We asked whether recruitment requires behavioral stimulation by returning foragers, as in honey bees, or if sampling the food source inside the nest is sufficient. We tested this by eliminating the behavior of returning foragers by inserting a scented sugar solution directly into a Vespula germanica nest. Exiting foragers were given a choice of the test scent and a control scent. WaSPS were more likely to choose the test scent. We conclude that behavioral interactions with returning foragers are not necessary to stimulate nest mates to associate an odor with a food source and search for a resource bearing that odor, and that experience with the scented reward inside the nest is sufficient to achieve this result.     

Experimental manipulation of termites (Isoptera,Macrotermitinae) foraging patterns in a Sahelo-Sudanese savanna: effect of litter quality

Summary The foraging pattern of four sympatric fungus-growing termite species was followed for five months of the dry season in a semi-arid savanna (Senegal). Experimental plots were established with and without termites (exclusion with insecticide) and amended with different litter quality: Acacia leaves, millet canes, ground millet (<500 µm), Combretum wood and cattle manure. Results showed different foraging patterns between the fungus-growing species. Odontotermes nilensis, for example, rapidly developed its foraging activity on the litter. In contrast, Macrotermes subhyalinus delayed its foraging activity but, extensively collected the food after its discovery. Differences in food preference were evident between the termite species: Odonto-termes nilensis foraged preferentially on millet, canes and Acacia leaves, Macrotermes subhyalinus preferred cane and ground millet. Ancistrotermes guineensis selected the millet, and Microtermes sp. was mostly recorded foraging on pieces of dead wood. Relationships between different foraging parameters were established to assess the relative intensity of this activity over the dry season. The comparison between untreated plots and plots treated with Fipronil® (Aventis) clearly demonstrated that the exclusion of termites from part of the savanna is possible and that this possibility can be used to study their role on the soil and organic matter cycle in situ.     

Recruitment Behavior During Foraging in the Neotropical Ant Gnamptogenys moelleri (Formicidae: Ponerinae): Does the Type of Food Matter?

Gnamptogenys moelleri nests in bromeliads and feeds on an array of food items, including dead and live animals, and nectar. Field data in Brazilian forests indicate that G. moelleri hunts solitarily, while retrieving is performed both by solitary workers for small items, or by a group of recruited workers for large items. This flexible foraging strategy was investigated in the laboratory through a series of experiments to assess the context in which recruitment is elicited. Three types of food were used: 50% honey solution, large insect prey, and cluster of small insects. For all food types the first encounter by a scout resulted in increased numbers of ants leaving the nest and finding the food in the arena. After finding liquid food or large prey, the forager returns to the nest and transmits information to nestmates about food location on the substrate. The successful scout repeatedly taps the sting on the ground, and recruited ants collectively retrieve the large insect to the nest. On the other hand, there is no transmission of information to nestmates about the location of small clumped prey, although the returning scout induces nestmates to leave the nest and hunt. Because foraging in G. moelleri is restricted mostly to the nest bromeliad, and small worker size (0.5 cm) precludes capturing large prey solitarily, recruitment behavior widens the spectrum of food items consumed by this ant species. Although recruitment behavior in ponerines has already been reported to vary with the type and size of a food source, this study also shows that the transmission of information about food location depends on the type of food found (large prey or liquid food versus cluster of small prey).