Estimating population density of the Formosan subterranean termite, Coptotermes formosanus(Isoptera: Rhinotermitidae) using the effective sampling area of in-ground monitoring stations

The effective sampling area of a monitoring station, alpha, was calculated for several Coptotermes formosanus Shiraki colonies in Broward County, Florida, USA. A simple mark-recapture protocol provided data on termite station catch within a foraging range of a colony. Average recapture probability was 0.005 close to the release point (< 5 m) and declined to 0.0008 at a distance of 51 to 60 m. The relation between the log % termites recaptured was fitted with log distance, to determine P(x), the average proportion of captured termites that started at distance x from the release point. The effective sampling area was estimated by using P(x) and the equation, alpha 2 pi integral [ x P(x)] dx. Integrating this equation, an average estimate alpha that ranged from 0.607 to 14.5 m2 was obtained. Factors influencing the variation of alpha among the colonies are discussed. The effective sampling area estimated should be taken as a reliable estimator that translates subterranean termite catches into termite population density.     

Interaction between the subterranean termite Reticulitermes flavipes (Isoptera: Rhinotermitidae) and the entomopathogenic fungus Metarhizium anisopliae in foraging arenas

The entomopathogenic fungus Metarhizium anisopliae (Metsch.) Sorokin was tested in the laboratory against field-collected groups of eastern subterranean termite, Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae), in foraging arenas to determine the potential effect of a "trap and treat" protocol (trapping a part of the population, treating it with a biological control agent and releasing it back into the original population). Individual termites were treated with a suspension of M. anisopliae conidia and released back into the arenas containing untreated termites. After 5 d, 90% of the treated termites died in the arena, but untreated termites did not exhibit a significant increase in mortality within 90 d after release, indicating no transfer of viable M. anisopliae and no epizootic. Although M. anisopliae was isolated from the arenas after 90 d, the average number of fungal colony-forming units recovered was <0.1% of the conidia introduced.     

Directional change in tunneling of subterranean termites (Isoptera: Rhinotermitidae) in response to decayed wood attractants

Wood discs decayed with brown rot fungi and polymer discs impregnated with acetone extract of decayed wood were embedded in sand of a two-dimensional foraging arena to evaluate their attractant potential in directing termite tunnels toward them. Termites were released near one arena corner and were guided to follow the physical guideline of the arena edges. In the absence of the attractants, termites generally formed a relatively linear tunnel along the edges. When decayed wood discs or treated polymer discs were placed in wet sand near one side of the arena, termite tunnels departed from the arena edges and were oriented toward them. The attraction distance ranged from 12 to 18 cm. The attractant properties were most likely water soluble and permeated through wet sand to cause termites to change their orientation. The results demonstrated that when such attractants are placed near a bait station in the field, they may be used to direct termite foraging toward the station.     

Tunneling activity of subterranean termites (Isoptera: Rhinotermitidae) in sand with moisture gradients

Tunnel formation by Coptotermes formosanus and Reticulitermes flavipes was studied in a two-dimensional foraging arena with a moisture gradient. Moisture did not appear to affect tunneling when termite first emerged from the central release chamber. But as termites of both species moved further away from the chamber and into a moisture gradient, they tunneled significantly (P < 0.05) more in sand with a higher moisture content than in sand with a lower moisture content. Over the 10 d test period, both termite species tunneled more in sand with a higher moisture content. Fractal analysis indicated that regardless of the sand moisture content, termite tunnel geometry had a fractal dimension and termites generally tunneled more in higher moisture sand.     

黑胸散白蚁取食群体大小与活动范围研究

为明确黑胸散白蚁Reticulitermes chinensis在绿地取食群体大小与活动范围,选取白蚁活动频繁、数量较多的5个诱集点进行研究。试验采用三重标记重捕法,染色标记物为0.2%耐尔兰滤纸(wt/wt)。经统计和测量发现,诱集点A、B、C和D处的取食个体数量分别为78.8、11.3、21.4和8.3万头,距诱集箱的最大取食直线距离分别为11.8 m(A)、5.5 m(B)和2.3 m(D)。由于C点的3次重捕均在诱集箱处完成,因此未能获得该白蚁群体的活动范围数据。而在诱集点E处,2次释放(诱集箱和7号饵站)均未能在释放点和周围饵站内重捕到标记白蚁。黑胸散白蚁4个群体的取食群体数量为8-79万头,连续监测30 d,其距诱集箱的取食直线距离为0-12 m,而实际群体数量和活动范围均大于试验中所得的数据。     

Foraging populations and distances of the desert subterranean termite, Heterotermes aureus (Isoptera: Rhinotermitidae), associated with structures in southern Arizona

Mark-release-recapture studies were conducted on foraging populations of Heterotermes aureus (Snyder) (Isoptera: Rhinotermitidae) associated with three structures in Tucson, AZ. Foraging population estimates ranged from 64,913 to 307,284 termites by using the Lincoln Index and from 75,501 to 313,251 termites using the weighted mean model. The maximum distance between monitors ranged from 26 to 65 m, with minimum total foraging distance ranging between 297 and 2,427 m. Characterizations of the cuticular hydrocarbons of foraging groups were qualitatively identical. Quantitative similarities within sites and differences among sites suggested that each site was occupied by a single colony during the sampling period. The colony at each site had a proportion of soldiers (0.135, 0.069, and 0.040) that was significantly different from the colonies at each of the other sites. From this study, we question the assumption of equal mixing of marked H. aureus foragers throughout the occupied collars around structures.     

Density estimation in live-trapping studies

Unbiased estimation of population density is a major and unsolved problem in animal trapping studies. This paper describes a new and general method for estimating density from closed-population capture–recapture data. Many estimators exist for the size (N) and mean capture probability ( p ) of a closed population. These statistics suffer from an unknown bias due to edge effect that varies with trap layout and home range size. The mean distance between successive captures of an individual (     ) provides information on the scale of individual movements, but is itself a function of trap spacing and grid size. Our aim is to define and estimate parameters that do not depend on the trap layout. In the new method, simulation and inverse prediction are used to estimate jointly the population density (D) and two parameters of individual capture probability, magnitude (g₀) and spatial scale (σ), from the information in     , p and     . The method uses any configuration of traps (e.g. grid, web or line) and any choice of closed-population estimator. It is assumed that home ranges have a stationary distribution in two dimensions, and that capture events may be simulated as the outcome of competing Poisson processes in time. The method is applied to simulated and field data. The estimator appears unusually robust and free from bias.     

The role of predation in the decline and extirpation of woodland caribou

To select appropriate recovery strategies for endangered populations, we must understand the dynamics of small populations and distinguish between the possible causes that drive such populations to low numbers. It has been suggested that the pattern of population decline may be inversely density-dependent with population growth rates decreasing as populations become very small; however, empirical evidence of such accelerated declines at low densities is rare. Here we analyzed the pattern of decline of a threatened population of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Using information on the instantaneous rate of increase relative to caribou density in suitable winter foraging habitat, as well as on pregnancy rates and on causes and temporal distribution of mortalities from a sample of 349 radiocollared animals from 15 subpopulations, we tested 3 hypothesized causes of decline: (a) food regulation caused by loss of suitable winter foraging habitat, (b) predation-sensitive foraging caused by loss of suitable winter foraging habitat and (c) predation with caribou being secondary prey. Population sizes of caribou subpopulations ranged from <5 to >500 individuals. Our results showed that the rates of increase of these subpopulations varied from −0.1871 to 0.0496 with smaller subpopulations declining faster than larger subpopulations. Rates of increase were positively related to the density of caribou in suitable winter foraging habitat. Pregnancy rates averaged 92.4% ±2.24 and did not differ among subpopulations. In addition, we found predation to be the primary cause of mortality in 11 of 13 subpopulations with known causes of mortality and predation predominantly occurred during summer. These results are consistent with predictions that caribou subpopulations are declining as a consequence of increased predation. Recovery of these woodland caribou will thus require a multispecies perspective and an appreciation for the influence of inverse density dependence on population trajectories.     

Foraging arena size and structural complexity affect the dynamics of food distribution in ant colonies

Food acquisition by ant colonies is a complex process that starts with acquiring food at the source (i.e., foraging) and culminates with food exchange in or around the nest (i.e., feeding). While ant foraging behavior is relatively well understood, the process of food distribution has received little attention, largely because of the lack of methodology that allows for accurate monitoring of food flow. In this study, we used the odorous house ant, Tapinoma sessile (Say) to investigate the effect of foraging arena size and structural complexity on the rate and the extent of spread of liquid carbohydrate food (sucrose solution) throughout a colony. To track the movement of food, we used protein marking and double-antibody sandwich enzyme-linked immunosorbent assay, DAS-ELISA. Variation in arena size, in conjunction with different colony sizes, allowed us to test the effect of different worker densities on food distribution. Results demonstrate that both arena size and colony size have a significant effect on the spread of the food and the number of workers receiving food decreased as arena size and colony size increased. When colony size was kept constant and arena size increased, the percentage of workers testing positive for the marker decreased, most likely because of fewer trophallactic interactions resulting from lower worker density. When arena size was kept constant and colony size increased, the percentage of workers testing positive decreased. Nonrandom (clustered) worker dispersion and a limited supply of food may have contributed to this result. Overall, results suggest that food distribution is more complete is smaller colonies regardless of the size of the foraging arena and that colony size, rather than worker density, is the primary factor affecting food distribution. The structural complexity of foraging arenas ranged from simple, two-dimensional space (empty arenas) to complex, three-dimensional space (arenas filled with mulch). The structural complexity of foraging arenas had a significant effect on food distribution and the presence of substrate significantly inhibited the spread of food. Structural complexity of foraging arenas and the resulting worker activity patterns might exert considerable influence on socioecological processes in ants and should be considered in laboratory assays.     

Estimating abundance of mountain lions from unstructured spatial sampling

Mountain lions (Puma concolor) are often difficult to monitor because of their low capture probabilities, extensive movements, and large territories. Methods for estimating the abundance of this species are needed to assess population status, determine harvest levels, evaluate the impacts of management actions on populations, and derive conservation and management strategies. Traditional mark–recapture methods do not explicitly account for differences in individual capture probabilities due to the spatial distribution of individuals in relation to survey effort (or trap locations). However, recent advances in the analysis of capture–recapture data have produced methods estimating abundance and density of animals from spatially explicit capture–recapture data that account for heterogeneity in capture probabilities due to the spatial organization of individuals and traps. We adapt recently developed spatial capture–recapture models to estimate density and abundance of mountain lions in western Montana. Volunteers and state agency personnel collected mountain lion DNA samples in portions of the Blackfoot drainage (7,908 km²) in west-central Montana using 2 methods: snow back-tracking mountain lion tracks to collect hair samples and biopsy darting treed mountain lions to obtain tissue samples. Overall, we recorded 72 individual capture events, including captures both with and without tissue sample collection and hair samples resulting in the identification of 50 individual mountain lions (30 females, 19 males, and 1 unknown sex individual). We estimated lion densities from 8 models containing effects of distance, sex, and survey effort on detection probability. Our population density estimates ranged from a minimum of 3.7 mountain lions/100 km² (95% CI 2.3–5.7) under the distance only model (including only an effect of distance on detection probability) to 6.7 (95% CI 3.1–11.0) under the full model (including effects of distance, sex, survey effort, and distance × sex on detection probability). These numbers translate to a total estimate of 293 mountain lions (95% CI 182–451) to 529 (95% CI 245–870) within the Blackfoot drainage. Results from the distance model are similar to previous estimates of 3.6 mountain lions/100 km² for the study area; however, results from all other models indicated greater numbers of mountain lions. Our results indicate that unstructured spatial sampling combined with spatial capture–recapture analysis can be an effective method for estimating large carnivore densities. Published 2012. This article is a U.S. Government work and is in the public domain in the USA.     

Food distance and its effect on nutrient balancing in a mobile insect herbivore

We investigated how the distance between foods of differing nutrient content affects macronutrient (protein and digestible carbohydrate) regulation and the patterns of food acquisition, movement and feeding activity in fifth-instar nymphs of Locusta migratoria (L.) (Orthoptera: Acrididae). We placed individual insects into one of three differently sized circular arenas (20, 40 or 80 cm diameter) that contained four dishes of chemically defined synthetic food. One of these dishes contained high-protein, low-carbohydrate food (P), and the other three dishes contained low-protein, high-carbohydrate food (C). Alone, these foods are nutritionally unbalanced, but together they are complementary. Regardless of arena size, locusts regulated their protein-carbohydrate intake to similar points, and in all three arenas they ate preferentially from the dish containing the P-food. We also recorded the patterns of foraging behaviour for 12 h on days 1 and 4 of the experiment, which allowed us to determine the effect of distance between foods and whether behaviour was modified with experience in the different arenas. Locusts' foraging behaviour in small arenas was similar on days 1 and 4, and the time they spent in different parts of the arena did not differ from that predicted by the simplest random probability model. In contrast, the foraging behaviour of locusts in the medium and large arenas changed between days 1 and 4, including spending more time in the area of the arena containing the P-food and visiting fewer of the C-food dishes. We discuss how distance between foods influences foraging strategies, and the possible role of learning in its development. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

The predatory behavior of Pheidole megacephala

We studied the foraging and predatory behaviors of the invasive African myrmicine ant, Pheidole megacephala (F.) in its native range. Workers can singly capture a wide range of insects, including relatively large prey items. For still larger prey, they recruit at short range those nestmates situated within reach of an alarm pheromone and together spread-eagle the insect. These behaviors are complimented by a long-range recruitment (of nestmates remaining in the nest) based on prey size. P. megacephala scouts also use long-range recruitment when they detect the landmarks of termites and competing ant species, thus permitting them to avoid confronting these termites and ants solitarily.     

Development of empirical feeding models for a benthic predator

Feeding on Chironomus riparius, Glyptotendipes paripes (Chironomidae). and Daphnia magna by the predatory freshwater leech, Nephelopsis obscura , was investigated in the laboratory and predictive models developed to estimate prey capture rate in the field. Experimental arena (container) size significantly affected predation rates when arena diameter was less than twice the total body length of the predator. Both chironomid and Daphnia magna capture rates were influenced by predator size, temperature and prey density.     

Modeling species dispersal with occupancy urn models

Models for species dispersal make various simplifications to facilitate analysis, such as ignoring spatial correlations or assuming equal probability of colonization among all sites within a dispersal neighborhood. Here we introduce a variation of the basic contact process (BCP) which allows us to separate the number of offspring produced from the neighborhood size, which are confounded in the original BCP. We then use classical results arising from probability models involving placing balls in urns to study our modified BCP, obtaining bounds for the critical value of the survival probability needed for the population to persist. We also use the probability urn calculations with a local-dispersal mean-field approximation to estimate equilibrium population density. These methods are able to include features such as unequal dispersal probabilities to different sites in the neighborhood, e.g., as would arise when dispersers have a fixed rate of mortality per distance traveled from the parent site. We also show how urn models allow one to generalize these results to two species competing for space.     

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.     

Interactions of Metarhizium anisoplae and tree-based mulches in repellence and mycoses against Coptotermes formosanus (Isoptera: Rhinotermitidae)

The use of mulch in urban landscapes has increased in the United States for the past decade. Tree-based organic mulches can supply Coptotermes formosanus Shiraki with food, moisture, and shelter. The current research contributes to mulch management technology in termite control. A choice test arena was designed to determine the repellence and mortality caused by commercial mulches treated with different concentrations of the fungus Metarhizium anisopliae (Metschnikoff) against C. formosanus. Each of six tree-based mulches (pine bark, pine straw, bald cypress, eucalyptus, water oak, and melaleuca) was coated with six conidial concentrations ranging from 1 x 10(3) to 1 x 10(8) conidia/ml. The foragers of C. formosanus were repelled significantly by the fungal-treated mulch substrates; the proportion of termites on fungal-treated mulch was usually <20% during the 28-d test. By day 28, >99% of the termites were killed in test arenas containing a chamber with mulch treated with 10(7) or 10(8) conidia/ml. M. anisopliae significantly reduced mulch consumption by 34-71%. Mulch consumption by the termites was negatively correlated with fungal concentration, and the type of mulch also affected consumption. The differences in termite foraging activities, mortality, and food consumption among mulches were usually confounded by differences in fungal concentrations of M. anisopliae. The results indicate that repellence and virulence of M. anisopliae conidia should significantly reduce the suitability of these six mulches as a habitat for C. formosanus.     

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.     

Long-term field trial to control the invasive Argentine ant (Hymenoptera: Formicidae) with synthetic trail pheromone

Previous short-term experiments showed that trail following behavior of the Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae), can be disrupted by a high concentration of synthetic trail pheromone component (Z)-9-hexadecenal. In this study, a long-term field trial was conducted in 100-m2 plots of house gardens in an urban area of Japan to see whether the control effect on Argentine ants can be obtained by permeating synthetic trail pheromone from dispensers. The dispensers were placed in the experimental plots during the ant's active season (April-November) for 2 yr with monthly renewal. To estimate Argentine ant population density, foraging activity of Argentine ants in the study plots was monitored by monthly bait surveys. Throughout the study period, Argentine ant foraging activity was suppressed in the presence of the dispensers, presumably via trail forming inhibition. In contrast, the level of foraging activity was not different between treatment and no-treatment plots when the dispensers were temporarily removed, suggesting that treatment with pheromone dispensers did not suppress Argentine ant density in the treatment plots. Population decline may be expected with larger-scale treatment that covers a significant portion of the ant colony or with improvement in the potency of the disruptant.     

Directional selection by the subterranean termite Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) at tunnel intersections

Subterranean termites construct complicated tunnel networks for foraging below ground. Thus, they often encounter tunnel intersections during foraging activity. Directional selection by termites at intersections is likely to affect foraging efficiency because depending on their selection, the path length from food resources to the nest can vary significantly. In order to understand how termites are guided to choose the most economical path without the use of pheromones, we artificially excavated two tunnels of varying widths (W₁ and W₂; 2, 3, or 4 mm) that intersected at a 90° angle in each of nine soil‐filled arenas. We observed the directional selection of termites at intersections in arenas with combinations of W₁ (mm) and W₂ (mm) (W₁, W₂). For (W₁, W₂) = (2, 2) and (4, 4), termites statistically equally chose the three directions left, right, and straight, while for the (3, 3) arena combination, termites preferentially decided to go straight. On the other hand, for (W₁, W₂) = (2, 3) (2, 4), and (3, 4), termites advancing from narrower tunnels into intersections tended to turn right or left, while termites coming from the wider tunnel were more inclined to go straight. On the assumption that a wider tunnel is more frequently used by termites in the field, we can deduce that the wider tunnel may represent the most efficient/shortest path. Thus, this simple selection mechanism can prevent termites from deviating from the most economical path.     

Unpacking chimpanzee (Pan troglodytes) patch use: Do individuals respond to food patches as predicted by the marginal value theorem?

The marginal value theorem is an optimal foraging model that predicts how efficient foragers should respond to both their ecological and social environments when foraging in food patches, and it has strongly influenced hypotheses for primate behavior. Nevertheless, experimental tests of the marginal value theorem have been rare in primates and observational studies have provided conflicting support. As a step towards filling this gap, we test whether the foraging decisions of captive chimpanzees (Pan troglodytes) adhere to the assumptions and qualitative predictions of the marginal value theorem. We presented 12 adult chimpanzees with a two-patch foraging environment consisting of both low-quality (i.e., low-food density) and high-quality (i.e., high-food density) patches and examined the effect of patch quality on their search behavior, foraging duration, marginal capture rate, and its proxy measures: giving-up density and giving-up time. Chimpanzees foraged longer in high-quality patches, as predicted. In contrast to predictions, they did not depress high-quality patches as thoroughly as low-quality patches. Furthermore, since chimpanzees searched in a manner that fell between systematic and random, their intake rates did not decline at a steady rate over time, especially in high-quality patches, violating an assumption of the marginal value theorem. Our study provides evidence that chimpanzees are sensitive to their rate of energy intake and that their foraging durations correlate with patch quality, supporting many assumptions underlying primate foraging and social behavior. However, our results question whether the marginal value theorem is a constructive model of chimpanzee foraging behavior, and we suggest a Bayesian foraging framework (i.e., combining past foraging experiences with current patch sampling information) as a potential alternative. More work is needed to build an understanding of the proximate mechanisms underlying primate foraging decisions, especially in more complex socioecological environments.