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.     

Colony interactions in <Emphasis Type="Italic">Reticulitermes grassei</Emphasis> population assessed by molecular genetic methods

The cryptic habits of subterranean termites impair studies of colony delimitation and the organization of foraging. Whereas feeding sites can be identified and the movements of foragers between them can be assessed using traditional mark-release techniques, the assignment of the termites found at feeding sites to their parent colonies remains problematical. Thus the extent and overlap of individual colony foraging territories are unknown. We used microsatellite markers to delineate colonies of R. grassei and compared the results with data from a mark-release-recapture study (Nobre et al., 2007) carried out in the same population and over the same period of time. The majority of colonies exhibited only a single-feeding site, but presented, even at a local scale, a high degree of variability in colony structure and no evidence of inbreeding. F-statistics for some colonies were consistent with pleometrosis. The population as a whole contained individual colonies that were separately identified as Mendelian, mixed family or pleometrotic families. Furthermore, microsatellite data suggest that what was represented by the MRR technique as a single foraging group could be drawn from more than one colony. Colony delineation is discussed as a tool for understanding overall population structure and termite feeding strategies. Received 5 July; revised 3 October 2007; accepted 19 October 2007.     

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。     

Factors affecting Grey‐headed Flying‐fox (Pteropus poliocephalus: Pteropodidae) foraging in the Melbourne metropolitan area,Australia

Abstract Factors affecting the foraging of mobile native fauna in highly fragmented urban landscapes have seldom been quantified at large spatial scales. We investigated factors affecting foraging by Grey‐headed Flying‐foxes (Pteropus poliocephalus; ‘flying‐foxes’) in the greater Melbourne metropolitan area. Flying‐foxes established a continuously occupied colony site in the Royal Botanic Gardens Melbourne in 1986, and the size of the colony has subsequently increased greatly. We used a stratified‐random sampling design to examine the importance of six variables on the detection of foraging flying‐foxes: (i) distance from the colony site (0–10, 10–20 and 20–30 km); (ii) distance from the Yarra River (0–5 and 5–20 km); (iii) the relative tree density of the municipality; (iv) whether the site was a park or street; (v) whether there was a relatively high or low density of trees at the site; and (vi) whether food was or was not detected at the site. We surveyed 240 sites within a 30‐km radius of the colony site for foraging flying‐foxes in both May and October 2002. The probability of detecting a foraging flying‐fox declined with increasing distance from the colony site, but increased with increasing tree cover, and was higher for parks compared with streets and when food was present. Flying‐foxes were observed foraging in a number of plant genera that have no species that naturally occur in the Melbourne area. Flying‐foxes in Melbourne thus forage on planted resources that are widely distributed within a fragmented landscape, and are an example of a positive response by a native species to the process of urbanization.     

黑翅土白蚁巢群觅食群体大小与工蚁体长呈正相关

【目的】明确黑翅土白蚁Odontotermes formosanus巢群的觅食群体大小与工蚁体长之间的关系,为进一步研究其生态学特点以及更好地开展防治提供依据。【方法】在杭州植物园内选择5处有黑翅土白蚁活动区域作试验点,诱捕并以中性红(neutral red)标记捕获的白蚁工蚁,通过“捕获-标记-释放-重捕”试验测定黑翅土白蚁的觅食群体大小。【结果】5个试验点中有4个成功实施了“捕获-标记-释放-重捕”试验,4个试验点黑翅土白蚁巢群的觅食群体个体数量分别为443 133±45 469, 495 360±67 429, 674 345±101 703和1 224 662±93 112头。黑翅土白蚁觅食群体大小(Y)与觅食工蚁的体长(X)呈正相关,拟合的指数函数方程式为：Y=1.8389e^0.7185X(R²=0.7834)。【结论】本研究明确了黑翅土白蚁的觅食群体大小以及工蚁体长之间的函数关系,为开展区域防治提供了依据。     

Response of Reticulitermes spp. (Isoptera: Rhinotermitidae) in northern California to baiting with hexaflumuron with sentricon termite colony elimination system

Colonies of Reticulitermes spp. were baited with prototype and commercial Sentricon stations (Dow AgroSciences LLC, Indianapolis, IN) to test the efficacy of hexaflumuron in different concentrations and bait matrices and to document reinvasion of the foraging territories vacated by eliminated colonies. Seven colonies of Reticuliternes spp. from two sites were characterized with cuticular hydrocarbon analyses and mark-release-recapture and agonistic behavioral studies. Three colonies were observed as controls and four colonies were baited. When a connection between the bait station and the monitoring station could not be confirmed by mark-release-recapture studies, the results of the baiting were equivocal. The monitoring stations of a colony at our wildland site were devoid of termites 406 d after baiting with one Sentricon station, but became reoccupied with the same species of termites approximately 6 mo after baiting. A colony at the residential site was baited with 0.5% hexaflumuron in the Recurit II bait matrix; 60 d later termites were absent from all monitoring stations. These monitoring stations remained unoccupied for > or = 18 mo. Foraging Reticulitermes spp. appeared in three of the seven monitoring stations 18, 24, and 36 mo after baiting, respectively. Using cuticular hydrocarbon analyses and agonistic behavior studies, we determined that the Reticulitermes spp. occupying these monitoring stations were from three different colonies; none were members of the original colony destroyed by baiting. Another colony at the residential site was baited using a noncommercial, experimental bait; 52 d later termites were absent from all monitoring stations. The monitoring stations remained unoccupied for > or = 9 mo. A different Reticulitermes sp. colony invaded one monitoring station 9 mo after baiting.     

Persistence of Allegheny woodrats Neotoma magister across the mid-Atlantic Appalachian Highlands landscape, USA

We examined a suite of macro-habitat and landscape variables around active and inactive Allegheny woodrat Neotoma magister colony sites in the Appalachian Mountains of the mid-Atlantic Highlands of Maryland, Virginia, and West Virginia using an information-theoretic modeling approach. Logistic regression analyses suggested that Allegheny woodrat presence was related positively to distance to the nearest occupied colony site and was influenced by location within physiographic subprovince. Colony sites were more likely to be active to the west (Allegheny Plateau) than the east (Blue Ridge/Piedmont), and colony sites were less likely to be active north of the Potomac River where land use and human disturbance patterns in the region were more intensive. Support also was generated for a presence-absence model that included forest cover within a 1-km radius of colony sites, although its importance was equivocal in this heavily forested region. Allegheny woodrats rely on emergent rock habitats for denning, and mast-bearing forest communities for foraging, and appear to display a metapopulation structure that is sensitive to a combination of natural and anthropogenically-induced isolation pressures that are recognizable but difficult to manage or mitigate.     

Hierarchical analysis of colony and population genetic structure of the eastern subterranean termite, Reticulitermes flavipes, using two classes of molecular markers

Termites (Isoptera) comprise a large and important group of eusocial insects, yet, in contrast to the eusocial Hymenoptera (ants, bees, wasps), the breeding systems of termites remain poorly understood. In this study, I inferred the breeding system of the subterranean termite Reticulitermes flavipes based on colony and population genetic structure as determined from microsatellite and mitochondrial DNA markers. Termites were sampled from natural wood debris from three undisturbed, forested sites in central North Carolina. In each site, two transects separated by 1 km were sampled at approximately 15-m intervals. A total of 1272 workers collected from 57 collection points were genotyped at six microsatellite loci, and mitochondrial DNA haplotype was determined for a subset of these individuals using either restriction fragment length polymorphism or sequence variation in the AT-rich region. Colonies appeared to be localized: workers from the 57 collection points represented 56 genetically distinct colonies with only a single colony occupying two collection points located 15 m apart. Genetic analysis of family structure and comparisons of estimates of F-statistics (F(IT), F(IC), F(CT)) and coefficients of relatedness (r) among nestmate workers with results of computer simulations of potential breeding systems suggested that 77% of all colonies were simple families headed by outbred monogamous pairs, whereas the remaining colonies were extended (inbred) families headed by low numbers of neotenics (about two females and one male) who were the direct offspring of the colony founders. There was no detectable isolation by distance among colonies along transects, suggesting that colony reproduction by budding is not common and that dispersal of reproductives during mating flights is not limited over this distance. Higher-level analysis of the microsatellite loci indicated weak but significant differentiation among sites (F(ST) = 0.06), a distance of 16-38 km, and between transects within sites (F(ST) = 0.06), a distance of 1 km. No significant differentiation at either the transect or site level was detected in the mitochondrial DNA sequence data. These results indicate that the study populations of R. flavipes have a breeding system characterized by monogamous pairs of outbred reproductives and relatively low levels of inbreeding because most colonies do not live long enough to produce neotenics, and those colonies that do generate neotenics contain an effectively small number of them.     

Evaluation of DNA Fingerprinting, Aggression Tests, and Morphometry as Tools for Colony Delineation of the Formosan Subterranean Termite

Multilocus DNA fingerprinting, aggression tests, and morphometry were compared to evaluate their potential for the delineation of colonies of Coptotermes formosanus Shiraki (Isoptera; Rhinotermitidae) in Hawaii. DNA fingerprinting segregates the termites from all collection sites and allows the assignment of all individuals to their original collection site. The genetic similarity of termites from different collection sites approaches the population's genetic background similarity, consequently collection sites represent independent colonies. Aggression between colonies is comparatively low and does not provide reliable colony delineation. Morphometry allows a 79% classification rate of termites to their colony of origin. No correlation among genetic similarities, aggression levels, and morphometric distances is found. Of the three investigated methods, we conclude that the genetic approach is the most useful tool for colony delineation in C. formosanus.     

Spatial patterns, temporal variability, and the role of multi-nest colonies in a monogynous Spanish desert ant

Abstract 1. The colonies of the Spanish desert ant Cataglyphis iberica are polydomous. This study describes the temporal and spatial patterns of the polydomy in this species at two different sites, and presents analyses of its role in reducing the attacks of the queen over sexual brood, and in allowing better habitat exploitation.
2. The spatial distribution of nests was clumped while colonies were distributed randomly. Mean nearest neighbour distance ranged from 3.4 to 7.0 m for nests and from 12.3 to 14.1 m for colonies. Distance of foragers searching for food varied among nests: mean values were between 6.1 and 12.6 m.
3. At both sites, the maximum number of nests per colony occurred in summer, during the maximum activity period of the species. Colonies regrouped at the end of this period but overwintered in several nests.
4. Nest renewal in C. iberica colonies was high and showed great temporal variability: nests changed (open, close, re-open) continuously through the activity season and/or among years. The lifetime of up to 55% of nests was only 1–3 months.
5. Polydomy in C. iberica might decrease the interactions between the queen and the sexual brood. In all colonies excavated just before the mating period, the nest containing the queen did not contain any virgin female. Females were in the queenless nests of the colony.
6. The results also suggest that polydomous C. iberica colonies may enhance habitat exploitation because foraging activity per colony increases with nest number. The relationship between total prey input and foraging efficiency and number of nests per colony attains a plateau or even decreases after a certain colony size (four to six nests). This value agrees with the observed mean number of nests per colony in C. iberica .     

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.     

Factors affecting the occupation of a colony site in Sydney,New South Wales by the Grey‐headed Flying‐fox Pteropus poliocephalus (Pteropodidae)

Previous authors have reported that Pteropus poliocephalus colony sites are occupied in response to blossom availability. However, in the present study it is reported that at the Gordon site in suburban Sydney, colony numbers were negatively correlated with the occurrence of pollen in the droppings. In addition, in contrast to reported occupational patterns at other colony sites, where flying‐foxes are not present at the site during winter and early spring, the Gordon site was occupied by substantial numbers of flying‐foxes throughout the entire period of 62 months from 1985 to 1990. As a result of the introduction of plants native to other parts of Australia and exotics from other continents, there is a variety of foods available throughout the year in the Sydney region, in comparison with less urbanized areas. This food supply permits the occupation of the Gordon colony site during winter and spring and reduces the migratory behaviour of flying‐foxes throughout the year. It is concluded that in the absence of a restrictive food supply, the occupational pattern of the Gordon colony of P. poliocephalus is the result of the reproductive requirements of the species modified by the vagaries of blossom production in the native forests outside the foraging range of the colony.     

Varying foraging patterns in response to competition? A multicolony approach in a generalist seabird

Reducing resource competition is a crucial requirement for colonial seabirds to ensure adequate self‐ and chick‐provisioning during breeding season. Spatial segregation is a common avoidance strategy among and within species from neighboring breeding colonies. We determined whether the foraging behaviors of incubating lesser black‐backed gulls (Larus fuscus) differed between six colonies varying in size and distance to mainland, and whether any differences could be related to the foraging habitats visited. Seventy‐nine incubating individuals from six study colonies along the German North Sea coast were equipped with GPS data loggers in multiple years. Dietary information was gained by sampling food pellets, and blood samples were taken for stable isotope analyses. Foraging patterns clearly differed among and within colonies. Foraging range increased with increasing colony size and decreased with increasing colony distance from the mainland, although the latter might be due to the inclusion of the only offshore colony. Gulls from larger colonies with consequently greater density‐dependent competition were more likely to forage at land instead of at sea. The diets of the gulls from the colonies furthest from each other differed, while the diets from the other colonies overlapped with each other. The spatial segregation and dietary similarities suggest that lesser black‐backed gulls foraged at different sites and utilized two main habitat types, although these were similar across foraging areas for all colonies except the single offshore island. The avoidance of intraspecific competition results in colony‐specific foraging patterns, potentially causing more intensive utilization of terrestrial foraging sites, which may offer more predictable and easily available foraging compared with the marine environment.     

Effects of concentration,distance, and application methods of Altriset (Chlorantraniliprole) on eastern subterranean termite (Isoptera: Rhinotermitidae)

The effects of various concentrations, distance, and application methods of Altriset (Chlorantraniliprole) were investigated against one of the most destructive termites, the eastern subterranean termite, Reticulitermes flavipes Kollar. Three laboratory experiments were conducted. First, we examined the concentration effect of treating the soil contiguously to established foraging tunnels at a fixed 1 m distance. The results demonstrated 100% termite control in 19 d posttreatment at 100 and 50 μg/g and 27% termite mortality at 25 μg/g. Second, we tested the distance effect of the soil treatment (2 and 4 m) on the efficacy of Altriset to the satellite termite populations at a fixed 50 μg/g concentration. This resulted in 100% termite control in 22 d posttreatment at both 2 and 4 m. Third, we examined the effect of differing application methods using 12.5 and 25 μg/g prior to the establishment of foraging tunnels at a fixed 1m distance. This illustrated 100% termite control in 9 d posttreatment at 25 μg/g and 12 d posttreatment at 12.5 μg/g. The third experiment demonstrated soil treatments that were applied prior to termite tunnel establishment had greater efficacy than applications made post tunnel construction. Our results provide a comprehensive understanding about the efficacy of Altriset treatments on eastern subterranean termites.     

Field evaluation of potential control strategies against the invasive eastern subterranean termite (Isoptera: Rhinotermitidae) in Chile

Four different termite control strategies consisting of two soil treatments with cypermethrin and fipronil and two bait treatments with hexaflumuron and sulfluramid were evaluated for their efficacy and potential for controlling the invasive eastern subterranean termite, Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae), in Quillota and Valparaiso, Chile. Monitoring stations were installed at all sites to measure the overall termite activity next to and at distances up to 30 m away from infested structures, most being 3-5 m. Foraging activity was extremely seasonal with the maximum consumption rates occurring from January to April. Termite foraging activity was low from July to September. After treatments in January 2003, there was a significant decrease in wood consumption, percentage of active stations, and numbers of termites in traps for the first 6 mo with hexaflumuron baits. No significant change in R. flavipes foraging activity was observed on wood consumption and percentage of active stations for sulfluramid bait or cypermethrin and fipronil soils treatments. The seasonal pattern of foraging activity remained unchanged for all treatments in 2004 and 2005, except structures baited with hexaflumuron. Feeding activity at the monitors in the area surrounding the hexaflumuron baits declined to 0 after January 2004. Alate flights occurred in approximately 38% of the structures treated with cypermethrin and sulfluramid bait in 2003 and in approximately 40% of the structures treated with cypermethrin, fipronil, and sulfluramid bait in 2004. Sulfluramid baits and cypermethrin soil treatments did not have any impact within a 15-m radius of the structures. Soils treatments of fipronil and cypermethrin prevented termites access in 75% of the homes. The number of active monitoring stations only declined within 2 m of the fipronil-treated zone. Hexaflumuron exhibited a reduction in activity for at least a 15-m radius.     

Spatial foraging patterns and colony energy status in the African honey bee,Apis mellifera scutellata

The relationship between changes in foraging patterns (inferred from waggle dance activity) and colony energy status (inferred from brood rearing activity, food storage, and colony weight) was examined for the African honey bee during a period of relative resource abundance and resource dearth. When resources were more abundant mean foraging distances (about 400 m) and foraging areas (4–5 km²) were small, and colonies recruited to 12–19 different sites per day. Colony foraging ranges and sites visited increased slightly during the dearth period, yet foraging continued to be concentrated within less than 10 km². The degree to which fluctuations in foraging patterns were correlated with colony energy status varied with the availability of floral resources. During periods of relative forage abundance, increases in foraging range and number of sites visited were significantly correlated with increases in brood rearing and colony weight. In contrast, colonies examined during periods of resource dearth exhibited no correlations between foraging areas, foraging distances, and fluctuations in brood rearing, food storage, or colony weight. Thus, during dearth periods colonies may not be able to coordinate foraging patterns with changes in colony energy status.     

Is food availability limiting African Penguins Spheniscus demersus at Boulders? A comparison of foraging effort at mainland and island colonies

The African Penguin Spheniscus demersus (Vulnerable) formed three new colonies during the 1980s, two on the South African mainland (Stony Point and Boulders) and one on Robben Island. One of the mainland colonies, at Boulders, Simon's Town, is in a suburban area, resulting in conflict with humans. Growth of the Boulders colony was initially rapid, largely through immigration, but has since slowed, possibly as a result of density‐dependent effects either on land (where there has been active management to limit the spread of the colony) or at sea. We test the latter hypothesis by comparing the foraging effort of Penguins feeding small chicks at island and mainland sites, and relate this to the foraging area available to birds. Three‐dimensional foraging paths of African Penguins were reconstructed using GPS and time–depth loggers. There were no intercolony differences in the rate at which birds dived during the day (33 dives/h), in diving depths (mean 17 m, max. 69 m) or in travelling speeds. The maximum speed recorded was 2.85 m/s, with birds travelling faster when commuting (average 1.18 m/s) than when foraging (0.93 m/s) or resting at sea (0.66 m/s during the day, 0.41 m/s at night). There were strong correlations between foraging trip duration, foraging range and total distance travelled. Foraging effort was correlated with chick age at Robben Island, but not at Boulders. Contrary to Ashmole's hypothesis, birds from Boulders (c. 1000 pairs) travelled further (46–53 km) and foraged for longer (13.2 h) than did birds from Robben Island (c. 7000 pairs) and Dassen Island (c. 21 000 pairs) (33 km, 10.3 h). The mean foraging range also differed significantly between mainland (18–20 km) and island colonies (9 km). The area available to central‐place‐foraging seabirds breeding on the mainland is typically less than that for seabirds breeding on islands, but the greater foraging range of Boulders birds results in an absolute foraging area roughly twice that of island colonies, and the area per pair is an order of magnitude greater for the relatively small Boulders colony. Ashmole's hypothesis assumes relatively uniform prey availability among colonies, but our results suggest this does not apply in this case. The greater foraging effort of Boulders birds probably reflects reduced prey availability in False Bay, and thus the recent slowing in growth at the colony may be the result of differential immigration rather than management actions to limit the spatial growth of the colony.     

Philopatry, Site Fidelity and Local Kin Associations within Grey Seal Breeding Colonies

Abstract Colonially breeding animals such as grey seals typically have a discrete number of breeding sites. The extent to which offspring are philopatric (return to breed at their natal site) and the prevalence of philopatry by sex of pup has fundamental consequences for the social and genetic structure within and between each colony.
Grey seals born and marked by tagging and cohort brands at colonies on North Rona, Outer Hebrides since 1960 and at the Isle of May, Firth of Forth since 1981 have shown philopatry. Overall re-sight rates of 1620 pups tagged at North Rona and 1667 pups at the Isle of May ranged from 0 to 17%. Although most evidence of philopatry relates to females, males at each colony displayed philopatry. In addition, females born at N. Rona pupped closer to their natal sites than would be expected by chance (p = 0.005), providing evidence of fine scale natal site fidelity. Females born at the Isle of May did not show the same degree of fine scale natal site fidelity, but there was considerable individual variation. The spatial scale of philopatry shown by seals differed according to the local area within each island. Fidelity of known adult females to pupping sites at both colonies was high. In addition, we found an instance of mother and daughter pupping together away from the daughter's natal site.
These data: (i) provide evidence of philopatry in both sexes at each site; (ii) describe occurrences of fine scale philopatry at both sites; (iii) show that mothers and offspring occupied the same areas on the colony, suggesting the possibility of highly related groups within colonies, which would provide conditions for kin-specific altruistic behaviours; (iv) suggest the possibility of individual or kin recognition, even though contact between mothers and offspring has been thought to cease after the 18 d lactation period.     

The spatial pattern of soil-dwelling termites in primary and logged forest in Sabah, Malaysia

Abstract.  1. Primary and logged lowland dipterocarp forest sites were sampled for subterranean termites using soil pits located on a grid system in order to detect any patchiness in their distribution.
2. A spatial pattern in termite distributions was observed in the primary and logged sites, but the response differed between soil-feeding and non-soil-feeding termites.
3. Spatial analysis showed that soil-feeding termites were homogeneously distributed in the primary forest but significantly aggregated in the logged forest. This pattern was reversed for non-soil-feeding termites and may result from differences in resource provisioning between the two sites.
4. Gaps in termite distribution comprised a greater area than patches for both feeding groups and sites, but gaps dominated the logged site.
5. A significant association between soil-feeding and non-soil-feeding termite distributions occurred at both sites. This arose from an association between patches in the primary forest and between gaps in the logged forest.
6. Termite spatial pattern was optimally observed at a minimum extent of 64 m and lag of 2 m.
7. The spatially explicit SADIE (Spatial Analysis by Distances IndicEs) analyses were more successful than (non-spatially explicit) multivariate analysis (Canonical Correspondence Analysis) at detecting associations between termite spatial distributions and that of other biotic and abiotic variables.     

Tunnel orientation and search pattern sequence of the formosan subterranean termite (Isoptera: Rhinotermitidae).

Foraging behavior of the Formosan subterranean termite, Coptotermes formosanus Shiraki, was studied in the laboratory by using two-dimensional foraging arenas containing multiple foraging sites. Within each arena, 16 foraging sites were arranged in a uniform grid pattern and foragers were introduced into the arena through a central initiation site. Chi-square analysis determined the frequency of tunnels was uniformly distributed around the perimeter of the initiation site but became significantly skewed toward the foraging sites at a distance where the foraging sites could be encountered. Tunnel distribution was similar whether wood was present or absent at the foraging sites, suggesting that foragers respond to structural anomalies in the substrate rather than simply to the presence of food. Also described is the generalized sequence of events as foragers tunnel throughout the arenas.