Tunnel specificity and forager movement in subterranean termites (Isoptera: Rhinotermitidae and Termitidae)

The movement of foragers of two species of Australian, subterranean, mound-building termites, Coptotermes lacteus (Froggatt) (Rhinotermitidae) and Nasutitermes exitiosus (Hill) (Termitidae), was investigated in their natural habitat using artificial feeding sites along trenches dug to mimic natural forager tunnels that radiate out from the central mound-nests. Termites were dyed by self-feeding on cardboard soaked with histological fat-stains on one or two trenches and then termites were collected from other feeding sites at two and four weeks after the fat-stains were placed. At two and four weeks after marking commenced, 60-75% of marked termites were found in trenches containing the marked paper, and 2-16% were found in trenches on the opposite side of the nest. The proportion of marked termites in a sample was three to eight times greater in the trenches containing the marked paper relative to other trenches. Although difficulties with fat-stains used as markers might explain some of the observed patterns, it is evident that C. lacteus and N. exitiosus foragers do not move randomly between feeding sites in their natural habitat.     

Estimating the population size of specialised solitary bees

Abstract.  1. Reliable methods for quantifying population size are crucial for strategies to conserve endangered wild-bee species. Estimates of population size obtained through survey walks were compared with estimates obtained through mark–recapture studies in 10 populations of the red-listed solitary bee Andrena hattorfiana in southern Sweden.
2. The mean number of bees observed during survey walks was strongly correlated with estimates of population size obtained with mark–recapture. It was found that 5.5–23.4% (mean 13.4%) of the total population was observed during an average survey walk.
3. One component in mark–recapture analysis is the measure of survival of individuals. In the largest bee population, females of A. hattorfiana that emerged in early season were found to forage for pollen on average 18.4 days.
4. The findings suggest that during large-scale surveys, for example re-inventories for red-listed species, the population size of solitary bees can be quantified reliably and effectively by performing survey walks in a two-step process. The first step consists of survey walks to establish the relationship between number of bee observations per survey walk and mark–recapture population size for a small set of populations. In the second, simple observation survey walks can be performed for a large set of populations. In each population of A. hattorfiana , it is recommended that at least six survey walks are performed.     

Estimating relative decline in populations of subterranean termites (Isoptera: Rhinotermitidae) due to baiting.

Although mark-recapture protocols produce inaccurate population estimates of termite colonies, they might be employed to estimate a relative change in colony size. This possibility was tested using two Australian, mound-building, wood-eating, subterranean Coptotermes species. Three different toxicants delivered in baits were used to decrease (but not eliminate) colony size, and a single mark-recapture protocol was used to estimate pre- and postbaiting population sizes. For both species, the numbers of termites retrieved from bait stations varied widely, resulting in no significant differences in the numbers of termites sampled between treatments in either the pre- or postbaiting protocols. There were significantly fewer termites sampled in all treatments, controls included, in the postbaiting protocol compared with the pre-, suggesting a seasonal change in forager numbers. The comparison of population estimates shows a large decrease in toxicant treated colonies compared with little change in control colonies, which suggests that estimating the relative decline in population size using mark-recapture protocols might to be possible. However, the change in population estimate was due entirely to the significantly lower recapture rate in the control colonies relative to the toxicant treated colonies, as numbers of unmarked termites did not change between treatments. The population estimates should be treated with caution because low recapture rates produce dubious population estimates and, in some cases, postbaiting mark-recapture population estimates could be much greater than those at prebaiting, despite consumption of bait in sufficient quantities to cause population decline. A possible interaction between fat-stain markers and toxicants should be investigated if mark-recapture population estimates are used. Alternative methods of population change are advised, along with other indirect measures.     

Three ways of assessing metapopulation structure in the butterfly Plebejus argus

1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus . First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area.
2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m.
3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale.
4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations.
5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur.
6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.     

The allocation of foragers in red wood ants

Abstract. 1. We studied how colonies of the red wood ant, Formica polyctena , adjust the numbers of foragers allocated to different foraging trails. In a series of field experiments, foragers were marked and transferred from one nest to another, related nest, where they joined the foraging force. Transferred workers acted as a reserve of uncommitted, available foragers.
2. Previous work shows that each individual forager habitually uses one trail. We found that for an uncommitted forager, the influence of recruitment initially is stronger than that of directional fidelity. Transferred workers were likely to use trails leading to new food sources. When transferred to a new nest, foragers were not likely to use a trail in the same direction as their original trail in the donor nest.
3. After a week, transferred foragers tended to develop route fidelity. Even after bait was no longer present, they continued to use the trail that had formerly led to a bait source.
4. We examined how colonies adjust numbers on a trail by experimentally depleting some trails. Colonies usually did not compensate for depletion: foragers were not recruited to depleted trails.
5. In general, the dynamics of foraging in this species facilitate a consistent foraging effort rather than rapid adjustments of forager allocation.     

Survival, movement, and resource use of the butterfly Parnassius clodius

Abstract.  1. A mark–recapture study was conducted on the American Apollo butterfly Parnassius clodius Menetries during three field seasons (1998–2000) to examine its movement patterns over the course of a season within a sagebrush meadow in Grand Teton National Park, Wyoming, U.S.A. The study examined how resources affected butterfly distribution patterns and used mark–recapture data to gain insight into movement differences between sexes and over time.
2. The average straight-line movement of P. clodius was 202 m day⁻¹, adjusted for sampling effort at different distances. Movement estimates in all 3 years were highly correlated with the average distance between plots sampled.
3. Butterfly abundance was correlated positively with per cent cover of its host plant Dicentra uniflora , but this relationship decreased in importance during the peak of the flight period when individuals may be more interested in finding mates. There was a weak, positive correlation between butterfly abundance and the abundance of its primary nectar source, Eriogonum umbellatum in 1999, but no relationship in 2000.
4. Survival, recapture, and transition probabilities were estimated using open population, capture–recapture models. Survival and recapture probability decreased over the course of each season, while the probability of moving between plots increased. Recapture probability was significantly lower for females than for males among all 3 years, but there was no difference between the sexes in survival rate.     

The effects of colony characteristics on life span and foraging behavior of individual wasps (Polybia occidentalis,Hymenoptera: Vespidae)

Summary We studied the effects of intrinsic colony characteristics and an imposed contingency on the life span and behavior of foragers in the swarm-founding social waspPolybia occidentalis. Data were collected on marked, known-age workers introduced into four observation colonies.To test the hypothesis that colony demographic features affect worker life span, we examined the relationships of colony age and size with worker life span using survivorship analysis. Colony age and size had positive relationships with life span; marked workers from two larger, older colonies had longer life spans (¯X = 24.7 days) than those from two smaller, younger colonies (¯X = 20.1 days).We quantified the effects of experimentally imposed nest damage on forager behavior, to determine which of three predicted behavioral responses by foragers to this contingency (increased probability of foraging for building material, increased rate of foraging, or decrease in age of onset of foraging) would be employed. Increasing the colony level of need for materials used in nest construction (wood pulp and water) by damaging the nests of two colonies did not cause an increase in either the proportion of marked workers that gathered nest materials or in foraging rates of marked individuals, when compared with introduced workers in two simultaneously observed control colonies. Instead, nest damage caused a decrease in the age at which marked workers first foraged for pulp and water. The response to an increase in the need for building materials was an acceleration of behavioral development in some workers.     

Intercalibration of hydroacoustic and mark–recapture methods for assessing the spawning population size of a threatened fish species

Hydroacoustic counting and a three-year mark–recapture study with passive integrated transponders (PIT tags) were used to estimate the size of a spawning population of nase Chondrostoma nasus , a threatened potamodromous cyprinid that undertakes annual spawning migrations into a tributary of the Danube River. In 2005, the estimates of the size of the spawning population from the hydroacoustic counts ( N = 2234, 95% CL 1929–2538) and from the Jolly–Seber model ( N = 1198, 95% CL 461–5842) corresponded well. Estimates from the jackknife-estimator based on the hydroacoustic counts yielded slightly higher values ( N = 2783, 95% CL 2529–3037), but were still in the same order of magnitude as those from the hydroacoustic and mark–recapture approach. At low run-size, hydroacoustic counting was more time consuming and technically demanding than mark–recapture studies. At the same time, it was non-invasive, provided real-time data on a fine temporal scale, and estimates showed less variability than the Jolly–Seber model. Mark–recapture of fish in spawning streams involved substantial disturbance at a sensitive stage of the life cycle. Hence, hydroacoustics is highly suited for population estimates of threatened potamodromous fishes, where interference needs to be minimized.     

Sticking to their choice – honey bee subfamilies abandon declining food sources at a slow but uniform rate

Abstract. 1. The allocation of honey bee foragers among food patches is a result of decisions made by individual bees that are based on internal and external cues.
2. Decision-making processes are often based on internal thresholds. For example, if the quality of the food source is assessed by a forager as exceeding its internal threshold, the bee will continue foraging on that food source.
3. It is often assumed that all individuals have the same threshold and therefore use the same thresholds in decision-making, but because the honey bee queen mates with 12–30 males, the workers within a colony are genetically heterogeneous. Thus, the thresholds used by individual bees may be genetically variable within a colony.
4. Models of colony-level foraging behaviour of honey bees suggest that the rate of abandoning food sources is a critical parameter affecting foraging success. Moreover, these models show that variance among subfamilies in their abandonment rates may increase the colony's foraging efficiency.
5. Experimental data showing the relationship between the probability of abandoning a food source and its profitability are lacking, as is information on any variation in abandonment rates among subfamilies.
6. Abandonment rates were determined experimentally for four honey bee families for seven different sucrose concentrations. The results showed that abandonment rates appear to be invariant among (sub)families. The importance of forager fidelity to declining food sources is discussed with respect to foraging efficiency in a changing environment.     

Higher productivity at the cost of increased host-specificity when Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation

Abstract .1. Field studies were made of the benefits and costs of two feeding strategies in the genus Maculinea, whose final-instar larvae parasitise Myrmica ant colonies. Maculinea arion is an obligate predator of ant brood, whereas M. rebeli and M. alcon mimic ant larvae and are fed (like cuckoos) directly by the workers.
2. Samples of > 1500 Myrmica nests confirmed laboratory-based predictions that, by feeding at a lower trophic level, many (4.7-fold) more individuals of M. rebeli and M. alcon are supported per ant colony than M. arion.
3. Because of their efficient feeding, cuckoo species often occupied sites where their phytophagous early larval populations coincided to only a small extent (> 10%) with host Myrmica colonies, whereas all sites supporting M. arion had 50–100% of the phytophagous stages within foraging range of the host Myrmica species.
4. Greater host-specificity was identified as another consequence of cuckoo-feeding. The ecological cost of this is discussed .
5. The feeding of other Maculinea species had not been fully described: the data suggest that M. nausithous is a predator of ant brood and confirm that M. teleius is predacious .     

Intraspecific aggression, colony identity and foraging distances in Sudanese Microtermes spp. (Isoptera: Termitidae: Macrotermitinae)

Abstract. 1. Intraspecific aggression between termite major workers was used to obtain estimates of foraging distances for three Microtermes species in Sudan.
2. Maximum foraging distance recorded for M. sp. nr albopartitus (Sjöstedt) in Khartoum was 11.3 m, giving an estimated minimum colony area of 100 m². This is probably an underestimate. In the Tokar Delta, individuals from single colonies of M. najdensis Harris were encountered up to 42 m apart, giving a colony area of 1390 m².
3. Intraspecific aggression could not be used reliably to distinguish members of different colonies of M. lepidus Sjöstedt. Soil barrier formation between groups of workers in petri dishes may be of use as a supplementary technique, especially for species showing no clear aggression.
4. Experiments on isolated laboratory colonics of M. sp. Nr lepidus Sjöstedt, established from alates collected in Khartoum, further confirmed the value of inter-colony aggression for indicating colony identity.
5. The complications introduced into estimation of subterranean termite foraging areas by overlap and interdigitation of colonies are discussed.     

Diel changes in forager size, activity, and load selectivity in a tropical leaf-cutting ant, Atta cephalotes

Abstract. 1. The leaf-cutting ant Atta cephalotes (L.) in a Costa Rican tropical moist forest showed diel changes in foraging activity. In most colonies studied, foraging was primarily nocturnal, although in a few colonies it was primarily diurnal.
2. In all colonies studied, mean forager mass was larger at night than during the day.
3. At night, most foragers carried freshly cut leaf fragments, whereas during the day a large proportion carried dried fragments and other vegetable matter collected from along their trail.
4. Along one trail, where foraging was primarily nocturnal, the match between ant mass and load mass was compared for laden ants at night and during the day. Laden ants at night were larger, carried relatively heavier loads, and showed a higher degree of matching between their mass and load mass than those foraging during the day.
5. A comparison of load masses of ants coming down a local tree and of ants picking up marked fragments from along their trail suggested that the diel difference in load mass and in the match between ant mass and load mass were related to the greater proportion of ants carrying freshly cut leaf fragments at night. Fresh fragments weighed more due to higher water content, and the match between ant mass and load mass was greater for ants cutting fresh fragments than for ants picking up abandoned fragments from along their trail.
6. Possible explanations for the diel changes in forager size and activity are discussed.     

Worker populations of Formica lugubris Zett. nests in Irish plantation woods

Abstract. 1. Estimates of worker populations of Formica lugubris Zett. nests in Irish plantation woods were obtained using capture—mark—recapture methods. Forager numbers were estimated by paint-marking individuals on the foraging routes and 'recapturing' by traffic counts. Colony-size was estimated using Stradling's (1970) ³²P-radiolabelling technique. Worker brood was estimated by excavation and direct count.
2. Forager numbers ranged from 6906 to 64 686 (eleven nests), colony-size ranged from 9797 to 71 052 (five nests) and worker brood ranged from 9809 to 16 269 (four nests).
3. Forager number was highly correlated with the average traffic on individual routes and the nest forager populations declined rapidly in >30-year-old nests.     

Mark–recapture estimates of recruitment, survivorship and population growth rate for the screwworm fly, Cochliomyia hominivorax

Abstract.  Pradel model mark–release–recapture estimates of survivorship, φ , recruitment, f , and the rate of density-independent population growth, λ , are presented for eight mark–recapture studies of the screwworm Cochliomyia hominivorax (Coquerel) (Diptera: Calliphoridae) from Costa Rica, totalling 19 573 released and 4476 recaptured flies. Corroborative estimates of survivorship and the rate of population growth based on an extensive review of the literature are also reported. Weighted-mean ± standard error of the mean (SEM) mark–release–recapture estimates of survivorship, recruitment and the rate of population growth were φ   =  0.798 ± 0.008, f   = 0.193 ± 0.008 and λ  = 1.005 ± 0.002, respectively. Population doubling time was estimated from λ at 139 days. Estimates of φ and λ from the literature both exceeded those calculated by mark–recapture methods and estimates of population doubling times were consequently shorter.     

The influence of spatial scale on quantifying insect dispersal: an analysis of butterfly data

Abstract. 1. A linear relationship between mean movement distances recorded and the size of the study area was found in a comparison of five mark–release–recapture studies of the meadow brown butterfly.
2. The scale impact on mean butterfly movement distances was also apparent when comparing the results for different butterfly species reported in 21 mark–release–recapture studies.     

Population regulation and demography in a harvested freshwater crayfish from Madagascar

Despite advances in statistical techniques for investigating population dynamics based on mark–recapture data, the majority of our understanding about demography and regulation comes from relatively few taxa. Most proposed generalisations about the association between demography and variation in population size are based on data from vertebrates, there are few sufficiently detailed invertebrate studies to examine whether these generalisations are widely supported. The population biology of freshwater invertebrates is especially poorly known. We present a large-scale mark–recapture study of an endemic freshwater crayfish from Madagascar ( Astacoides granulimanus ). Variation in density, caused by difference in fishing pressure due to local taboos, allowed us to investigate density-dependent regulation. We found evidence of density dependence in fecundity operating through the proportion of reproductive females by size but no significant evidence of density dependence in growth. Using a prospective analysis based on the elasticities from a size-structured matrix model, we found that both recruitment rates and survival rates of large individuals were strongly associated with deterministic population growth – a result that differs from generalisations drawn from vertebrate studies. A central assumption in mark–recapture studies is that handling does not affect mortality. By treating the number of times an individual was captured as an individual covariate, easily done using the freeware program MARK, we were able to test for, and take account of, handling-induced mortality. Our results show interesting similarities, and important differences, to generalisations based on vertebrate studies and emphasise the importance of population studies on poorly known taxa.     

Measuring dispersal and detecting departures from a random walk model in a grasshopper hybrid zone

Abstract. 1. The grasshopper species Chorthippus brunneus and C. jacobsi form a complex mosaic hybrid zone in northern Spain. Two mark–release–recapture studies were carried out near the centre of the zone in order to make direct estimates of lifetime dispersal.
2. A model framework based on a simple random walk in homogeneous habitat was extended to include the estimation of philopatry and flying propensity. Each model was compared with the real data, correcting for spatial and temporal biases in the data sets.
3. All four data sets (males and females at each site) deviated significantly from a random walk. Three of the data sets showed strong philopatry and three had a long dispersal tail, indicating a low propensity to move further than predicted by the random walk model.
4. Neighbourhood size estimates were 76 and 227 for the two sites. These estimates may underestimate effective population size, which could be increased by the long tail to the dispersal function. The random walk model overestimates lifetime dispersal and hence the minimum spatial scale of adaptation.
5. Best estimates of lifetime dispersal distance of 7–33 m per generation were considerably lower than a previous indirect estimate of 1344 m per generation. This discrepancy could be influenced by prezygotic isolation, an inherent by-product of mosaic hybrid zone structure.     

Do bumblebees always forage as much as they could?

Summary Bumblebees must forage to provide food to the colony. However, foraging is costly as worker longevity is inversely related to foraging effort. Given this trade-off, workers from colonies with abundant food supplies could either maintain foraging to increase reserves for future use or forage less to avoid the associated costs. We tested these hypotheses over one summer, using 13 pairs of field colonies of Bombus impatiens. Half of the colonies were provided with a sucrose solution ad libitum and pollen at regular intervals throughout their entire development, while the other half served as controls. We measured the forager activity rates in colonies with infra-red motion detectors fit in nest box entrances. However, due to reasons beyond our control (loss of the queen, usurpation by Psithyrus, debris in the entrance tunnel, etc.), we could use data from only two pairs of colonies for the analysis. Food supplemented colonies had a forager activity rate per worker 25% lower than controls which supports the hypothesis that workers reduce risks when given the opportunity.Received 15 May 2003; revised 15 January 2004; accepted 19 February 2004.     

The Trail Less Traveled: Individual Decision-Making and Its Effect on Group Behavior

Social insect colonies are complex systems in which the interactions of many individuals lead to colony-level collective behaviors such as foraging. However, the emergent properties of collective behaviors may not necessarily be adaptive. Here, we examine symmetry breaking, an emergent pattern exhibited by some social insects that can lead colonies to focus their foraging effort on only one of several available food patches. Symmetry breaking has been reported to occur in several ant species. However, it is not clear whether it arises as an unavoidable epiphenomenon of pheromone recruitment, or whether it is an adaptive behavior that can be controlled through modification of the individual behavior of workers. In this paper, we used a simulation model to test how symmetry breaking is affected by the degree of non-linearity of recruitment, the specific mechanism used by individuals to choose between patches, patch size, and forager number. The model shows that foraging intensity on different trails becomes increasingly asymmetric as the recruitment response of individuals varies from linear to highly non-linear, supporting the predictions of previous work. Surprisingly, we also found that the direction of the relationship between forager number (i.e., colony size) and asymmetry varied depending on the specific details of the decision rule used by individuals. Limiting the size of the resource produced a damping effect on asymmetry, but only at high forager numbers. Variation in the rule used by individual ants to choose trails is a likely mechanism that could cause variation among the foraging behaviors of species, and is a behavior upon which selection could act.     

Comparison of social networks derived from ecological data: implications for inferring infectious disease dynamics

1.  Social network analyses tend to focus on human interactions. However, there is a burgeoning interest in applying graph theory to ecological data from animal populations. Here we show how radio-tracking and capture–mark–recapture data collated from wild rodent populations can be used to generate contact networks.
2.  Both radio-tracking and capture–mark–recapture were undertaken simultaneously. Contact networks were derived and the following statistics estimated: mean-contact rate, edge distribution, connectance and centrality.
3.  Capture–mark–recapture networks produced more informative and complete networks when the rodent density was high and radio-tracking produced more informative networks when the density was low. Different data collection methods provide more data when certain ecological characteristics of the population prevail.
4.  Both sets of data produced networks with comparable edge (contact) distributions that were best described by a negative binomial distribution. Connectance and closeness were statistically different between the two data sets. Only betweenness was comparable. The differences between the networks have important consequences for the transmission of infectious diseases. Care should be taken when extrapolating social networks to transmission networks for inferring disease dynamics.