Caste developmental potentialities in the termite Nasutitermes novarumhebridarum

The normal development pathway and caste system of Nasutitermes novarumhebridarum (N. & K. Holmgren) (Isoptera; Termitidae) conform to the general Nasutitermes pattern. Queen-removal experiments in N. novarumhebridarum resulted in the production of replacement reproductives derived from workers (ergatoids), but never from the sexual brood (nymphs and alates). Three moults are necessary to complete the tranformation from the worker to the ergatoid. This pathway is longer than those previously konwn in other termites, in which two moults are sufficient. In a colony with functional ergatoids, some nymphs gave rise to worker-like individuals. The caste potentialities of N. novarumhebridarum differ from those of the sympatric species, N. princeps and N. polygynus, in which imaginal replacement reproductives are easily obtained. Colony reproduction in N. novarumhebridarum seems to be accomplished exclusively by independent alate founders, which colinize massive, but finite and widely spaced resoures such as dead or damaged tree trunks. N. princeps and N. polygynus societies abundantly use colony budding to expand their territory, and exploit various sources of dead wood scattered over their foraging range. We suggest that N. novarumhebridarum caste potentialities are adapted to channelling a maximum of resources into the production of dispersing alates that can colonize new nest sites.

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Résumé Les voies de développement et le système de castes chez Nasutitermes novarumhebridarum sont conformes au schéma général du genre Nasutitermes. L'enlèvement expérimental de la reine chez N. novarumhebridarum entraîne le développement de sexués de remplacement à partir d'ouvriers (ergatoïdes), mais jamais à partir de la lignée sexuée (nymphes et ailés). La transformation de l'ouvrier en ergatoîde fonctionnel nécessite trois mues. Ce développement est plus long que ceux décrits précédemment chez d'autres termites, où deux mues sont suffisantes. Dans une société où se sont développés des ergatoïdes fonctionnels, quelques nymphes se sont transformées en individus rappelant les ouvriers. Les potentialités des castes de N. novarumhebridarum diffèrent de celles des espèces sympatriques N. princeps et N. polygynus, chez lesquelles des sexués de remplacement imaginaux pauvent être obtenus aisément. La reproduction des sociétés de N. novarumhebridarum semble s'effectuer uniquement par des ailés essaimants, qui colonisent des sources de nourriture massives, mais limitées et largement espacées, tels les troncs d'arbres morts ou endomagés. N. princeps et N. polygynus étendent fréquemment leurs territoires par le bourgeonnement de nouvelles sociétés, et exploitent des sources de bois mort variées, dispersées dans leur champ d'exploitation. Nous suggérons que les potentialités des castes chez novarumhebridarum sont adaptées pour diriger un maximum de ressources vers la production d'ailés qui se dispersent et peuvent ainsi coloniser de nouveaux sites favorables.

     

An interspecific comparison of foraging range and nest density of four bumblebee (Bombus) species

Bumblebees are major pollinators of crops and wildflowers in northern temperate regions. Knowledge of their ecology is vital for the design of effective management and conservation strategies but key aspects remain poorly understood. Here we employed microsatellite markers to estimate and compare foraging range and nest density among four UK species: Bombus terrestris, Bombus pascuorum, Bombus lapidarius, and Bombus pratorum. Workers were sampled along a 1.5-km linear transect across arable farmland. Eight or nine polymorphic microsatellite markers were then used to identify putative sisters. In accordance with previous studies, minimum estimated maximum foraging range was greatest for B. terrestris (758 m) and least for B. pascuorum (449 m). The estimate for B. lapidarius was similar to B. pascuorum (450 m), while that of B. pratorum was intermediate (674 m). Since the area of forage available to bees increases as the square of foraging range, these differences correspond to a threefold variation in the area used by bumblebee nests of different species. Possible explanations for these differences are discussed. Estimates for nest density at the times of sampling were 29, 68, 117, and 26/km2 for B. terrestris, B. pascuorum, B. lapidarius and B. pratorum, respectively. These data suggest that even among the most common British bumblebee species, significant differences in fundamental aspects of their ecology exist, a finding that should be reflected in management and conservation strategies.     

Asexual queen succession in the higher termite Embiratermes neotenicus

Asexual queen succession (AQS), in which workers, soldiers and dispersing reproductives are produced sexually while numerous non-dispersing queens arise through thelytokous parthenogenesis, has recently been described in three species of lower termites of the genus Reticulitermes. Here, we show that AQS is not an oddity restricted to a single genus of lower termites, but a more widespread strategy occurring also in the most advanced termite group, the higher termites (Termitidae). We analysed the genetic structure in 10 colonies of the Neotropical higher termite Embiratermes neotenicus (Syntermitinae) using five newly developed polymorphic microsatellite loci. The colonies contained one primary king accompanied either by a single primary queen or by up to almost 200 neotenic queens. While the workers, the soldiers and most future dispersing reproductives were produced sexually, the non-dispersing neotenic queens originated through thelytokous parthenogenesis of the founding primary queen. Surprisingly, the mode of thelytoky observed in E. neotenicus is most probably automixis with central fusion, contrasting with the automixis with terminal fusion documented in Reticulitermes. The occurrence of AQS based on different mechanisms of ploidy restoration raises the hypothesis of an independent evolutionary origin of this unique reproductive strategy in individual lineages of lower and higher termites.     

Sampling subterranean termite species diversity and activity in tropical savannas: an assessment of different bait choices

Abstract.  1. Cellulose baits are commonly used for semi-quantitative sampling of subterranean wood-feeding termites, with a single food choice sampling programme implemented most often. In most situations, however, the composition and feeding preferences of the subterranean termite assemblage remain unknown.
2. The diversity, frequency, foraging activity, and intensity of attack of termites were assessed regularly at 144 baits representing 12 different bait choices over 8.5 months, in two northern Australian tropical savanna sites that differed in vegetation structure (closed vs open). Baits differed in type (paper rolls, cardboard, wooden stakes), position (surface, buried), and moisture status at installation (wet, dry).
3. Sixteen species were recorded, including 11 wood-feeders. Average species diversity, foraging activity, and bait consumption were greater at buried baits than at surface baits. Wooden baits were most attractive early in the experiment, and paper baits more attractive later. Mean species diversity was greatest at wooden stakes in the closed site. Species frequency of occurrence varied across bait choices.
4. A composite bait sampling protocol of stakes and paper rolls installed above and below ground gave an accurate assessment of the activity, diversity, and structure of the termite guild sampled across all baits over 8.5 months.
5. The choice of bait, its presentation, and time of examination are critical to the success of a termite baiting programme. If the aim is to characterise the structure and foraging activity of the subterranean termite assemblages that are attracted to baits, composite baiting protocols should be implemented.     

Population density of the greater mouse-eared bat (Myotis myotis), local diet composition and availability of foraging habitats

We studied the regional variation in population density of Myotis myotis (Borkhausen 1797) in south-eastern Bavaria, Germany, and its relations to diet composition and the availability of potential foraging habitats. We monitored colony size and juvenile mortality from 1991 to 2003, conducted faecal analyses in 1993 and determined land-use patterns around colonies. The numbers of individuals counted in the nursery colonies showed only small fluctuations over the years. However, data on colony size demonstrated a pronounced regional variation. Epigeic arthropods, mainly Carabidae, were the most important prey. The diet included prey taxa of forest as well as grassland habitats. The percentage of those prey taxa in the diet that originated in grassland managed with different intensity varied according to the availability of these potential foraging habitats around the nursery roosts. The calculated population density of the bats was positively correlated to forest area and especially to the area of mixed forest around the nursery roosts. Our results indicate that the availability of foraging habitats is a limiting factor for local population densities in the greater mouse-eared bat.     

Spatial models of foraging in clonal plant species

Ramets of some clonal plant species alter their internode lengths or their frequency of lateral branching in response to their immediate microenvironment. Such “plant foraging” responses are thought to allow clones to concentrate in favorable portions of their environment. Despite widespread interest among ecologists in plant foraging, few realistic models have been developed to examine conditions under which plant foraging responses are likely to provide clones with ecological benefit. In this paper, we develop spatially explicit, stochastic simulation models to examine consequences of both empirical and hypothetical plant foraging responses. We construct a hierarchical series of models in which we incorporate effects of resource heterogeneity on spacer lengths, angles of growth, and lateral branch production. We also vary the number, size, and arrangement of patches, and the presence or absence of ramet mortality. Simulations based on hypothetical data demonstrated the potential importance of shortening spacer lengths in favorable habitat. In these simulations, ramet crowding increased significantly, implying a potential cost to plant foraging responses whose magnitude is large enough to cause ramets to concentrate in favorable patches. Models calibrated with empirical data suggest that when clonal plants were able to concentrate in favorable habitat, this was usually caused by increased daughter ramet production in the favorable habitat. Variation in clonal growth angles had little impact on the ability of ramets or clones to locate favorable patches, but did increase the ability of clones to remain in favorable patches once found. Alterations in the number and size of patches strongly influenced the effectiveness of the foraging response. The spatial arrangement of patches also was important: clumped distributions of patches decreased the success with which plants located favorable patches, especially at the genet level and when the number of patches was low. Finally, when ramet mortality varied with patch quality, there was an increase in the percentage of ramets located in favorable patches; differential ramet mortality also lessened the impact of other effects, such as the decreased success of clones when patches are clumped. Overall, our models indicate that the effectiveness of plant foraging responses is variable and is likely to depend on a suite of environmental conditions.     

Colony genetic organization and colony fusion in the termite Reticulitermes flavipes as revealed by foraging patterns over time and space

Temporal and spatial analyses are seldom utilized in the study of colony genetic structure, but they are potentially powerful methods which can yield novel insights into the mechanisms underlying variation in breeding systems. Here we present the results of a study which incorporated both of these dimensions in an examination of genetic structure of subterranean termites in the genus Reticulitermes (primarily R. flavipes). Most colonies of this species (70%) were simple families apparently headed by outbred primary reproductives, while most of the remaining (27% of the total) colonies contained low effective numbers of moderately inbred reproductives. Mapping the spatial distribution of colony foraging sites over time revealed that despite the high colony density, the absolute foraging boundaries of most R. flavipes colonies were persistent and exclusive of other conspecific colonies, which suggests that this species is more territorial than has been implied by laboratory studies of intraspecific aggression. Nevertheless, we found a single colony (3% of all colonies) which contained the offspring of more than two unrelated reproductives. Although other studies have also described subterranean termite colonies with a similarly complex genetic composition, we demonstrate here that such colonies can form under natural conditions via the fusion of whole colonies. This study underscores how repeated sampling from individual colonies over time and space can yield information about colony spatial and genetic structure that cannot be obtained from conventional analyses or sampling methods.     

The effect of rearing history and aphid density on volatile‐mediated foraging behaviour of Diaeretiella rapae

1. Parasitoid females foraging for hosts rely on cues derived from the insect host, the host plant and/or their interaction, and all of these can be learned during the immature and adult stages. 2. The present study investigated the importance of rearing history on foraging behaviour of Diaeretiella rapae, an endoparasitoid often associated with aphids feeding on brassicaceous plant species. 3. Parasitoids were reared on one of the four possible combinations, comprising two brassicaceous host plant species, Brassica nigra or Raphanus sativus, and two aphid species Brevicoryne brassicae or Myzus persicae. These parasitoids were tested in a Y‐tube olfactometer and given the choice between volatiles emitted by an aphid‐infested plant (25 or 100 aphids per plant) and an uninfested control plant. The parasitoid's responses were compared when offered: (i) the same plant–aphid combination as the one on which it had been reared; (ii) the same host plant infested with the alternative aphid species; or (iii) an alternative plant with the alternative aphid species. 4. Aphid density did affect the behavioural responses to the various odour sources, but rearing history did not. Diaeretiella rapae only preferred aphid‐induced to non‐induced plant volatiles at low aphid infestation level, whereas they did not discriminate between volatiles at high aphid infestation level. 5. It is concluded that aphid‐induced volatiles of brassicaceous plants play an important role during host habitat location, but seem less important for parasitoids to locate the aphid host itself. The data are discussed in the light of manipulation of host plant defences by aphids.     

Queen pheromone regulates egg production in a termite

In social insects, resource allocation is a key factor that influences colony survival and growth. Optimal allocation to queens and brood is essential for maximum colony productivity, requiring colony members to have information on the total reproductive power in colonies. However, the mechanisms regulating egg production relative to the current labour force for brood care remain poorly known. Recently, a volatile chemical was identified as a termite queen pheromone that inhibits the differentiation of new neotenic reproductives (secondary reproductives developed from nymphs or workers) in Reticulitermes speratus. The same volatile chemical is also emitted by eggs. This queen pheromone would therefore be expected to act as an honest message of the reproductive power about queens. In this study, we examined how the queen pheromone influences the reproductive rate of queens in R. speratus. We compared the number of eggs produced by each queen between groups with and without exposure to artificial queen pheromone. Exposure to the pheromone resulted in a significant decrease in egg production in both single-queen and multiple-queen groups. This is the first report supporting the role of queen pheromones as a signal regulating colony-level egg production, using synthetically derived compounds in a termite.     

Threshold foraging by gray whales in response to fine scale variations in mysid density

The gray whale (Eschrichtius robustus) is a coastal species whose nearshore summer foraging grounds off the coast of British Columbia offer an opportunity to study the fine scale foraging response of baleen whales. We explore the relationship between prey density and gray whale foraging starting with regional scale (10 km) assessments of whale density (per square kilometer) and foraging effort as a response to regional mysid density (per cubic meter), between 2006 and 2007. In addition we measure prey density at a local scale (100 m), while following foraging whales during focal surveys. We found regional mysid density had a significant positive relationship with both gray whale density and foraging effort. We identify a threshold response to regional mysid density for both whale density and foraging effort. In 2008 the lowest average local prey density measured beside a foraging whale was 2,300 mysids/m³. This level was maintained even when regional prey density was found to be substantially lower. Similar to other baleen whales, the foraging behavior of gray whales suggests a threshold response to prey density and a complex appreciation of prey availability across fine scales.     

IDH knockdown alters foraging behavior in the termite Odontotermes formosanus in different social contexts

Foraging, as an energy-consuming behavior, is very important for colony survival in termites. How energy metabolism related to glucose decomposition and adenosine triphosphate (ATP) production influences foraging behavior in termites is still unclear. Here, we analyzed the change in energy metabolism in the whole organism and brain after silencing the key metabolic gene isocitrate dehydrogenase (IDH) and then investigated its impact on foraging behavior in the subterranean termite Odontotermes formosanus in different social contexts. The IDH gene exhibited higher expression in the abdomen and head of O. formosanus. The knockdown of IDH resulted in metabolic disorders in the whole organism. The dsIDH-injected workers showed significantly reduced walking activity but increased foraging success. Interestingly, IDH knockdown altered brain energy metabolism, resulting in a decline in ATP levels and an increase in IDH activity. Additionally, the social context affected brain energy metabolism and, thus, altered foraging behavior in O. formosanus. We found that the presence of predator ants increased the negative influence on the foraging behavior of dsIDH-injected workers, including a decrease in foraging success. However, an increase in the number of nestmate soldiers could provide social buffering to relieve the adverse effect of predator ants on worker foraging behavior. Our orthogonal experiments further verified that the role of the IDH gene as an inherent factor was dominant in manipulating termite foraging behavior compared with external social contexts, suggesting that energy metabolism, especially brain energy metabolism, plays a crucial role in regulating termite foraging behavior.     

Flock density, social foraging, and scanning: an experiment with starlings

Social foraging differs from individual foraging because italters both resource availability and the forager's behavior.We examined responses of starlings to the presence of conspecificsby manipulating foraging-group density experimentally, whileensuring that each subject's foraging opportunities were unchanged.To do this, we used individuals foraging simultaneously in fourbottomless enclosures placed at various separations in naturalforaging grounds. We measured foraging and scanning intensityand qualitative aspects of scanning of focal individuals. Additionally,we examined the temporal distribution of scanning between individuals.The focal individual analysis showed that (1) food-searchingactivity increased, while time spent scanning, time off theground and scanning bout length decreased with flock density;(2) food finding per unit of searching effort increased withdensity; (3) head orientation during scanning was sensitiveto companions' proximity: heads pointed away from the companionsat close distance, toward them at intermediate distance, andwas random farther away. The analysis of the (temporal overlappingin scanning) temporal distribution of scanning for the groupshowed that scanning was significantly synchronized when companionswere adjacent to each other but was not significantly differentfrom random at further separations. We conclude that behavioralresponses of individuals to the presence of others generateimportant changes in foraging performance even in the absenceof physical interference and, more generally, that assessingthe mechanisms that control the behavior of group members atdifferent flock densities offers a way to understand the functionaland ecological significance of foraging aggregations.     

Effects of pollen species composition on the foraging behaviour and offspring performance of the mason bee Osmia bicornis (L.)

The effects of floral species composition on offspring performance of solitary bees are rarely studied under conditions where foraging behaviour of mothers is allowed to play a role. In a semi-field experiment, we restricted foraging choices of the polylectic mason bee Osmia bicornis L. to flower species belonging to plant families presumably used to different extent: Borago officinalis L. (Boraginaceae), Centaurea cyanus L. (Asteraceae) and Brassica napus L. (Brassicaceae). We quantified the foraging behaviour and brood cell production by mother bees, and compared the quality of offspring in pure and mixed flower species stands. Offspring survival in pure stands was expected to reflect the mothers’ foraging preferences in the mixed stand. Pure stands of B. napus supported highest offspring survival, body mass and fraction of females produced. Offspring survival on C. cyanus and B. officinalis was very low. Larval mortality occurred earlier in brood cells provided with B. officinalis pollen than in brood cells provided with C. cyanus pollen suggesting different effects of pollen quality on early larval and later development. The time spent on different foraging activities correlated with lifetime reproductive output. However, in mixed stands, the proportion of time the bees were foraging on the different flower species did not differ significantly. Foraging behaviour may therefore not generally be a good proxy for the quality of floral resources for offspring production. Our results suggest that resources collected from one plant species may influence the usefulness of resources from another plant species. Bees may therefore overcome potentially deleterious effects of the suboptimal resources by mixing low- and high-quality resources. This may help generalist bees, such as O. bicornis, to cope with an unpredictable environment.     

Floral resource pulse decreases bumble bee foraging trip duration in central Wisconsin agroecosystem

1. Resource pulses, narrow periods of high resource availability, can elicit strong behavioural responses across diverse taxa. Mass‐flowering agricultural crops are an example of a resource pulse that insect pollinators exploit. However, the underlying mechanism behind changes in pollinator behaviour associated with mass‐flowering crops is still relatively unexplored. 2. The present study quantified the behavioural response of bumble bees, an important wild pollinator, to commercial cranberry bloom, an important mass‐flowering crop in Wisconsin, U.S.A. Over a 2‐year period, foraging trip duration was measured using radio frequency identification at 14 farms situated across landscape contexts, ranging from high to low natural area (woodland amount). Using transect surveys, floral resource abundance at a landscape scale was estimated. 3. It was found that bumble bees were highly sensitive to temporal changes in landscape‐level resource abundance associated with the onset of cranberry bloom, during which they decreased foraging trip duration by 22% and increased the number of foraging trips during bloom by 24% on average relative to the period before and after bloom. This phenomenon was consistent across colonies, individual bees, and landscape contexts, despite a higher abundance of flowers in low woodland landscapes. Bumble bee colonies growing in low‐ and high‐woodland landscapes exhibited a similar performance. 4. As mass‐flowering crops are probably a factor influencing bumble bee foraging behaviour in agricultural regions, investigations should continue into how variable resource landscapes, particularly those offering resource pulses, affect wild pollinators and the pollination services they provide.     

The effect of prey density on foraging mode selection in juvenile lumpfish: balancing food intake with the metabolic cost of foraging

1. In many species, individuals will alter their foraging strategy in response to changes in prey density. However, previous work has shown that prey density has differing effects on the foraging mode decisions of ectotherms as compared with endotherms. This is likely due to differences in metabolic demand; however, the relationship between metabolism and foraging mode choice in ectotherms has not been thoroughly studied. 2. Juvenile lumpfish Cyclopterus lumpus forage using one of two modes: they can actively search for prey while swimming, or they can 'sit-and-wait' for prey while clinging to the substrate using a ventral adhesive disk. The presence of these easily distinguishable foraging modes makes juvenile lumpfish ideal for the study of foraging mode choice in ectotherms. 3. Behavioural observations conducted during laboratory experiments showed that juvenile lumpfish predominantly use the 'cling' foraging mode when prey is abundant, but resort to the more costly 'swim' mode to seek out food when prey is scarce. The metabolic cost of active foraging was also quantified for juvenile lumpfish using swim-tunnel respirometry, and a model was devised to predict the prey density at which lumpfish should switch between the swim and cling foraging modes to maximize energy intake. 4. The results of this model do not agree with previous observations of lumpfish behaviour, and thus it appears that juvenile lumpfish do not try to maximize their net energetic gain. Instead, our data suggest that juvenile lumpfish forage in a manner that reduces activity and conserves space in their limited aerobic scope. This behavioural flexibility is of great benefit to this species, as it allows young individuals to divert energy towards growth as opposed to activity. In a broader context, our results support previous speculation that ectotherms often forage in a manner that maintains a minimum prey encounter rate, but does not necessarily maximize net energy gain.     

Structure and Dynamics of the Arboreal Termite Community in New Guinean Coconut Plantations21

Whereas ant mosaics have been widely recognized and described in tropical ecosystems, data on space partitioning among arboreal termite colonies are rudimentary. During a long term field study in New Guinea, the distribution of arboreal termite species in coconut plantations as well as the extent and dynamics of competition between them were investigated. The three dominant species, Mirrocerotermes biroi, Nasutitermes princeps and N. novarumhebridarum, feed on the same items but never exploit the same tree. The resulting distribution pattern is a mosaic with two peculiarities. First, some extended areas around N. princeps colonies appear unexploited, as this species practices interference competition on a wide scale, defending large territories inter- and intra-specifically. Second, interspecific relations are asymmetrical. In some plantations, large colonies of N. princeps expand their territory by destroying colonies of M. biroi, but when the pressure of N. princeps is relaxed, dense populations of colonies of M. biroi can recolonize the trees in a few years’ time. Territorial boundaries may thus change relatively fast. N. novarumhebridarum often colonizes dead trees and interferes less with the other species. These facts are consistent with each species’ reproductive investment strategy. Hypotheses are proposed to explain how the dominant species can coexist, even in long established plantations.