Taste-rejection by predators and the evolution of unpalatability in prey

Aposematic species advertise their unpalatability to potential predators using conspicuous warning colouration. The initial evolution of aposematism is thought to occur by warningly coloured mutants emerging in an already unpalatable cryptic species. However, possessing defence chemicals is often costly, and it is difficult to understand what the selective benefits might be for a mutation causing its bearer to be defended in a population of otherwise palatable cryptic prey. One solution to this problem is that chemically defended individuals are tasted and rejected by predators, and are, therefore, more likely to survive predatory attacks than undefended individuals. Using naïve domestic chicks Gallus gallus domesticus as predators and cryptic green chick crumbs as prey, we asked whether the accuracy with which birds discriminated between palatable and unpalatable prey was affected by the palatability of the unpalatable prey (moderately or highly defended), or their frequency in the population (10 or 25%). Birds could discriminate between green prey on the basis of their defences, and showed better discrimination between palatable and unpalatable prey when defended crumbs were highly unpalatable, compared to when they were moderately unpalatable. Although there was no detectable effect of the frequency of unpalatable prey in the population on predator taste-rejection behaviour in our main analysis, frequency did appear to affect the strategies that birds used in their foraging decisions when prey were only moderately unpalatable. How birds used taste to reject prey also suggests that birds may be able to monitor and regulate their chemical intake according to the frequency and defence levels of the unpalatable prey. Taken together, these results show that avian predators can generate selection for unpalatability in cryptic prey by sampling and taste-rejecting prey, but that a relatively large chemical difference between palatable and unpalatable prey may be necessary before unpalatable prey can enjoy a selective advantage. The exact nature of this evolutionary dynamic will depend on other environmental factors, such as defence costs and prey availability, but it provides a mechanism by which defences can evolve in a cryptic population.     

Warning signals confer advantage to prey in competition with predators: bumblebees steal nests from insectivorous birds

Aposematic (warning) signals of prey help predators to recognize the defended distasteful or poisonous prey that should be avoided. The evolution of aposematism in the context of predation has been in the center of modern ecology for a long time. But, the possible roles of aposematic signals in other ecological contexts have been largely ignored. Here we address the role of aposematic signals in competition between prey and predators. Bumblebees use visual and auditory aposematic signals to warn predators about their defenses. For 2 years, we observed competition for nestboxes between chemically defended insects, Bombus ardens (and possibly also Bombus ignitus), and cavity nesting birds (Parus minor and Poecile varius). Bumblebees settled in 16 and 9 % of nestboxes (in 2010 and 2011 breeding seasons, respectively) that contained bird nests at the advanced stage of nest building or at the stage of egg laying. Presence of bumblebees prevented the birds from continuing the breeding activities in the nestboxes, while insects took over the birds’ nests (a form of kleptoparasitism). Playback experiments showed that the warning buzz by bumblebees contributed to the success in ousting the birds from their nests. This demonstrates that aposematic signals may be beneficial also in the context of resource competition.     

Growth-selective survival of young jack mackerel Trachurus japonicus during transition from pelagic to demersal habitats in the East China Sea

To examine the growth-selective survival in early life stages of jack mackerel from the East China Sea, we compared growth rates of young fish collected from the surface layer in April with those of survivors collected from the near-bottom layer in May?CJune. We performed this analysis from 3?year classes with different levels of relative abundance: low in 2005, intermediate in 2008 and high in 2009. Growth rates during the larval stage of the original population were slower than the survivors in all survey years, indicating that faster-growing individuals in the surface layer had a higher probability of survival to the near-bottom habitat than slower-growing individuals. Selection intensity, defined as differences in growth rates between the two groups, was higher in year classes with a lower abundance. Hence, growth rates during the early life stages of jack mackerel are considered to be a critical determinant of successful survival to the demersal habitat.     

Variation in pelagic larval growth of Atlantic billfishes: the role of prey composition and selective mortality

Atlantic blue marlin (Makaira nigricans) and sailfish (Istiophorus platypterus) larvae were collected from 10 monthly cruises (June–October 2003 and 2004) across the Straits of Florida to test (1) whether growth differed between the more productive western region near the Florida shelf, and the less productive eastern region toward the Bahamas, and (2) whether growth was related to prey consumption. Examination of larval sagittal otoliths revealed that instantaneous growth and daily growth during the first 2–3 weeks of life did not vary significantly between the two regions for either species. However, recent growth during the last two full days prior to collection was greater in the west for blue marlin larvae. Recent growth of blue marlin larvae <9 mm SL (primarily zooplanktivorous) was significantly related to prey composition (faster growth when higher proportions of Farranula copepods were consumed). Western larvae grew faster and had higher proportions of Farranula in their guts. Trends for sailfish larvae were not significant. In both species, comparison of early growth between <9 and ≥9 mm SL size groups indicated that growth trajectories diverged around 5–8 mm SL, the time when billfish larvae become capable of piscivory. Significantly faster growth of larger (older) larvae suggests that mortality was selective for fast growers and that the transition to piscivory may be a critical point in the early life of billfish.     

Horizontal distribution patterns of presettlement reef fish: are they influenced by the proximity of reefs?

The distribution patterns of presettlement reef fish and how they were influenced by the proximity of reefs were investigated off the coast of Northland, New Zealand, from 1981–1986. We used ichthyoplankton tows and visual counts of fish. Distributions of presettlement fish of some species were influenced by the proximity of reefs, regardless of whether reefs were on the coast of the mainland or islands across the shelf. Presettlement fish of families that lay demersal eggs were most abundant near reefs: Gobiescocidae, Acanthoclinidae, Tripterygiidae, Eleotridae, and Gobiidae. The distribution of presettlement sparids, mullids (pelagic eggs), and blenniids and monacanthids (demersal eggs) was not determined in a predictable way by the proximity of reefs. High-frequency sampling over three days suggested that patches of presettlement sparids of 1 to 2 km in dimension may move quickly through a study area. High abundance of presettlement gobiescocids and tripterygiids were found in 0 to 2 m of water over rocky reefs at high and low tides. Presettlement eleotrids were associated with reefs in deeper water (3 to 20 m) and in some habitats with aggregations of mysids. The lack of general patterns of distribution for presettlement reef fish suggests that modelling patterns of drift of these fish as a single group is inappropriate; this concurs with evidence from tropical waters.     

Use of chemical cues in detection of conspecific predators and prey by newts, <Emphasis Type="Italic">Notophthalmus viridescens</Emphasis>

Summary. Detection of heterospecific predators and prey via chemical cues is well known, but only a few studies have examined the potential for such discrimination in cannibalistic systems. In newts, Notophthalmus viridescens, adults are opportunistic predators of conspecific larvae. I used a laboratory bioassay to determine whether larvae and adults distinguish between chemical stimuli from members of the different age classes. Larvae distinguished between chemical stimuli from larvae and adults by decreasing their activity only following exposure to stimuli from adults. Decreased activity is consistent with an antipredator response in many prey species, including larval newts. In contrast, adults increased their activity and increased time spent in open areas in response to stimuli from larvae, but not to stimuli from adults. Increased activity is consistent with a feeding response; adults also showed increased activity and increased time in open areas in response to chemical stimuli from familiar heterospecific prey (brine shrimp). The proximate cue that allowed the newts to distinguish between the different age classes is not associated with short-term dietary differences because all stimulus animals were fed the same prey. Stimulus strength was controlled by diluting the stimulus solutions according to the volume of the stimulus animal. Therefore, there appear to be intrinsic differences in the chemical signatures of larval and adult newts.     

Differences in feeding ecology among three co-occurring species of wrasse (Teleostei: Labridae) on rocky reefs of temperate Australia

The foraging behaviours and dietary compositions of three co-occurring labrids (Ophthalmolepis lineolatus, Notolabrus gymnogenis and Pictilabrus laticlavius), which are conspicuous on rocky reefs in temperate south-eastern Australia, were investigated between 2003 and 2005. SCUBA observations at two locations showed that the feeding intensity, and hence the associated effects of these fishes on rocky reef invertebrate prey, was temporally consistent. Relative differences in the contributions of ingested prey and use of different feeding microhabitats demonstrated that the feeding ecology differed significantly among the three species. Thus, O. lineolatus fed on proportionately higher volumes of polychaetes, polyplacophorans, marginellid gastropods (especially Austroginella sp.), bivalves and echinoids, which were sighted opportunistically in a wide selection of microhabitats, but particularly in sand/rubble. Ambush hunting was used regularly by smaller N. gymnogenis and all sizes of P. laticlavius to forage on amphipods, small decapods and small gastropods at algal bases or fronds and Diopatra dentata tubes. Amphipods were similarly important in the diet of smaller O. lineolatus. Larger N. gymnogenis foraged opportunistically over an increased reef area and made greater use of microhabitats that offered minimal prey refuge (e.g. sand/rubble, bare rock/steel) from which common prey, in particular decapods, were obtained. The significant intra- and inter-specific differences in dietary compositions, allied with differences in the use of feeding microhabitats, would facilitate co-occurrence of these three conspicuous species and contribute to maintaining high richness of labrid species in reef systems. Echinoids were regularly consumed by each species but they made a moderate contribution to the diet of only O. lineolatus, which suggests that only one of the three labrids is likely to play a significant role in regulation of echinoid densities in these rocky reef habitats. However, the broad diets and diverse forging strategies employed by these labrid species imply that they have a system-wide influence on invertebrate prey on rocky reefs.     

Distribution patterns of sea urchins and barrens in shallow Mediterranean rocky reefs impacted by the illegal fishery of the rock-boring mollusc<Emphasis Type="Italic"> Lithophaga lithophaga</Emphasis>

Shallow rocky habitats in SW Apulia (SE Italy, Mediterranean Sea) were surveyed in late spring 2002 to assess distribution patterns of sea urchins (Paracentrotus lividus and Arbacia lixula) and barren habitats (coralline barrens and bare substrates) in rocky reefs impacted by the destructive fishery of the rock-boring date-mussel Lithophaga lithophaga. Sea urchin density, test size-structure and biomass, and the percent cover of barrens were evaluated at four locations (5–6 km apart from each other), two heavily impacted by the date-mussel fishery and two controls. Sea urchin density and barren habitat cover were assessed at two and three sites (100–300 m apart), respectively, within each location. Sea urchin biomass was evaluated only at the scale of locations. Average density of P. lividus did not significantly change between impacted locations and controls, whereas A. lixula showed a greater density at the impacted locations. Distribution patterns of A. lixula, in addition, differed at the spatial scale of a few metres between impacted locations and controls, being generally more aggregated at the controls. The size-frequency distribution (test diameter) of P. lividus showed a mode at 3–4 cm at the impacted locations compared to a mode at 2–3 cm in the controls. The size-frequency of A. lixula was bimodal at the damaged locations (with modes at 1–2 and 4–5 cm, respectively) and unimodal (with the mode at 4–5 cm) at the controls. Average biomass of both sea urchins (P. lividus and A. lixula) was two- to fourfold greater at the impacted locations (~600 g wet wt m^–2) than at the controls (150–250 g wet wt m^–2). Barren habitats had a far greater average cover (mainly of macroalgae) at the impacted locations (from 79% to 96%) than at control locations (from 7% to 21%). These results show that the date-mussel fishery may have the potential to affect distribution patterns of sea urchins and to greatly enhance the percent cover of barren grounds in shallow Mediterranean rocky reefs.Communicated by R. Cattaneo-Vietti, Genova     

Isotopic evidence of distinct feeding ecologies and movement patterns in two migratory predators (yellowfin tuna and swordfish) of the western Indian Ocean

Ecologists primarily use δ¹⁵N values to estimate the trophic level of organisms, while δ¹³C, and even recently δ¹⁵N, are utilized to delineate feeding habitats. However, many factors can influence the stable isotopic composition of consumers, e.g. age, starvation or isotopic signature of primary producers. Such sources of variability make the interpretation of stable isotope data rather complex. To examine these potential sources of variability, muscle tissues of yellowfin tuna (Thunnus albacares) and swordfish (Xiphias gladius) of various body lengths were sampled between 2001 and 2004 in the western Indian Ocean during different seasons and along a latitudinal gradient (23°S to 5°N). Body length and latitude effects on δ¹⁵N and δ¹³C were investigated using linear models. Both latitude and body length significantly affect the stable isotope values of the studied species but variations were much more pronounced for δ¹⁵N. We explain the latitudinal effect by differences in nitrogen dynamics existing at the base of the food web and propagating along the food chain up to top predators. This spatial pattern suggests that yellowfin and swordfish populations exhibit a relatively unexpected resident behaviour at the temporal scale of their muscle tissue turnover. The body length effect is significant for both species but this effect is more pronounced in swordfish as a consequence of their different feeding strategies, reflecting specific physiological abilities. Swordfish adults are able to reach very deep water and have access to a larger size range of prey than yellowfin tuna. In contrast, yellowfin juveniles and adults spend most of their time in the surface waters and large yellowfin tuna continue to prey on small organisms. Consequently, nitrogen isotopic signatures of swordfish tissues are higher than those of yellowfin tuna and provide evidence for different trophic levels between these species. Thus, in contrast to δ¹³C, δ¹⁵N analyses of tropical Indian Ocean marine predators allow the investigation of complex vertical and spatial segregation, both within and between species, even in the case of highly opportunistic feeding behaviours. The linear models developed in this study allow us to make predictions of δ¹⁵N values and to correct for any body length or latitude differences in future food web studies.     

Chemical defense in pelagic octopus paralarvae: Tetrodotoxin alone does not protect individual paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) from common reef predators

Some pelagic marine larvae possess anti-predator chemical defenses. Occasionally, toxic adults imbue their young with their own defensive cocktails. We examined paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) for the deadly neurotoxin tetrodotoxin (TTX), and if present, whether TTX conferred protection to individual paralarvae. Paralarvae of H. lunulata possessed 150 ± 17 ng TTX each. These paralarvae appeared distasteful to a variety of fish and stomatopod predators, yet food items spiked with 200 ng TTX were readily consumed by predators. We conclude that TTX alone does not confer individual protection to paralarvae of H. lunulata, and that they possess an alternative defense. In larger doses, tetrodotoxin is a deterrent to the predatory stomatopod Haptosquilla trispinosa (mean dose = 3.97 μg/g). This corresponds to 12–13 paralarvae per predator based on the TTX levels of the clutch we examined. Thus, the basic assumption that individual paralarvae of H. lunulata are defended by TTX alone was disproved. Instead, functionality of TTX levels in paralarvae may arise through alternative selective pathways, such as deterrence to parasites, through kin selection, or against predator species not tested here.     

Larval durations and recruitment patterns of two Caribbean gobies (Gobiidae): contrasting early life histories in demersal spawners

The population dynamics of small tiger prawns (Penaeus esculentus and P. semisulcatus) were studied at three sites around north-western Groote Eylandt, Gulf of Carpentaria, Australia, between August 1983 and August 1984. Seagrasses typical of open-coastline, reef-flat and river-mouth communities were found in the shallow depths (2.5 m) at these sites. The temperature and salinity of the bottom waters did not differ among the shallowest depths of the three sites and mean values at night ranged from 21.9 to 32.0 °C, and from 30.1 to 37.5% S. Data from fortnightly sampling with beam trawls showed that virtually all post-larvae (90%) were caught in the intertidal and shallow subtidal waters (2.0 m deep). At one site, where the relationship between seagrass biomass, catches and depth could be studied in detail, high catches were confined to seagrass in shallow water, within 200 m of the high-water mark. This was despite the fact that seagrass beds of high biomass (>100 g m^-2 between August and February) were found nearby, in only slightly deeper water (2.5 m). It is likely, therefore, that only the seagrass beds in shallow waters of the Gulf of Carpentaria act as important settlement and nursery areas for tiger prawns. In general, catches of tiger prawn postlarvae (both P. esculentus and P. semisulcatus) and juvenile P. esculentus on the seagrass in the shallowest waters at each site were higher in the tropical prewet (October–December) and wet (January–March) seasons than at other times of the year. Juvenile P. semisulcatus catches were highest in the pre-wet season. While seasonal differences accounted for the highest proportion of variation in catches of tiger prawn postlarvae and juvenile P. semisulcatus, site was the most important factor for juvenile P. esculentus. In each season, catches of juvenile P. esculentus were highest in the shallow, open-coastline seagrass, where the biomass of seagrass was highest. The fact that the type of seagrass community appears to be more important to juvenile P. esculentus than to postlarvae, suggests that characteristics of the seagrass community may affect the survival or emigration of postlarval tiger prawns. Few prawns (<10%) from the seagrass communities in shallow waters exceeded 10.5 mm in carapace length. Despite the intensive sampling, growth was difficult to estimate because postlarvae recruited to the seagrass beds over a long period, and the residence times of juveniles in the sampling area were relatively short (8 wk).     

Fine-scale distribution of larval fishes: patterns and processes adjacent to coral reefs in Kaneohe Bay,Hawaii

Plankton samples were taken from January to June 1987 in Kaneohe Bay, Oahu, Hawaiian Islands, with a free-fall plankton net, to investigate the fine-scale distribution of larval fishes around coral reefs. Daytime samples indicated that the postflexion larvae of two gobiids (Psilogobius mainlandi and an unidentified species) were significantly more abundant at stations immediately adjacent to reefs (near-reef) than at stations in open water off the reef (off-reef). These postflexion gobiid larvae appeared to be capable of resisting advection and dispersal while remaining in the water column near suitable adult habitats. The larvae of Foa brachygramma (Apogonidae) and Encrasicholina purpurea (Engraulidae) were significantly more abundant at off-reef stations than at near-reef stations. Nighttime samples indicated that the gobiid larvae depend on visual cues to remain near the reef. The horizontal distributions of F. brachygramma and E. purpurea larvae appeared to be related to their vertical positioning. These data suggest that typical ichthyoplankton surveys which do not sample close to adult fish habitats would greatly underestimate the abundances of larvae such as the gobiids.     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.     

Movement patterns of the seastar Heliaster helianthus in central Chile: relationship with environmental conditions and prey availability

The rates and patterns of feeding and displacement of predators constitute two of the most important plastic behavioral responses that allow individuals to respond quickly to changes in abundance of their prey, predation risks and to rapid alterations in environmental conditions. In this study, we quantified seasonal and spatial variation in displacement (net changes in location in 12 or 24 h periods) and prey consumed of marked individuals of the keystone seastar Heliaster helianthus at six sites spanning 600 km along the coast of north-central Chile. We evaluated the hypotheses that: (1) at sites with low availability (cover) of the main prey, the mussel Perumytilus purpuratus, Heliaster displays larger displacements and consumes a greater proportion of other prey (e.g. mobile species) than at sites with high mussel cover, (2) daily displacements will be correlated with sea surface temperature (SST) and (3) increased wave action will reduce seastar daily displacement. Our results show that Heliaster displacement is higher at sites with lower availability of P. purpuratus; and at these sites, a larger proportion of Heliaster individuals are observed feeding, mostly on other prey (e.g. limpets), which could offset the higher costs associated with increased movement. In addition, wave forces affected the activity of Heliaster negatively. Contrary to our expectations, the daily displacements did not show any relationship with SST measured on the day or the previous days of the surveys, despite the fact that average displacement was generally higher in summer than in winter months. Future studies should examine Heliaster movement during single foraging excursions and determine whether these responses affect the growth and reproductive output of individuals. Such information is vital to understand how changes in prey abundance and environmental conditions alter the behavior and energy budget of this predator and its ability to control prey populations.     

Feeding by larvae of intertidal invertebrates: assessing their position in pelagic food webs

One of the leading determinants of the structure and dynamics of marine populations is the rate of arrival of new individuals to local sites. While physical transport processes play major roles in delivering larvae to the shore, these processes become most important after larvae have survived the perils of life in the plankton, where they usually suffer great mortality. The lack of information regarding larval feeding makes it difficult to assess the effects of food supply on larval survival, or the role larvae may play in nearshore food webs. Here, we examine the spectrum of food sizes and food types consumed by the larvae of two intertidal barnacle species and of the predatory gastropod Concholepas concholepas. We conducted replicated experiments in which larvae were exposed to the food size spectrum (phytoplankton, microprotozoan and autotrophic picoplankton) found in nearshore waters in central Chile. Results show that barnacle nauplii and gastropod veligers are omnivorous grazers, incorporating significant fractions of heterotrophs in their diets. In accordance with their feeding mechanisms and body size, barnacle nauplii were able to feed on autotrophic picoplankton (<5 microm) and did not consume the largest phytoplankton cells, which made the bulk of phytoplankton biomass in spring-summer blooms. Balanoid nauplii exhibited higher ingestion rates than the smaller-bodied chthamaloid larvae. Newly hatched C. concholepas larvae also consumed picoplankton cells, while competent larvae of this species ingested mostly the largest phytoplankton cells and heterotrophic protozoans. Results suggest that persistent changes in the structure of pelagic food webs can have important effects on the species-specific food availability for invertebrate larvae, which can result in large-scale differences in recruitment rates of a given species, and in the relative recruitment success of the different species that make up benthic communities.     

Trophic importance of subtidal metazoan meiofauna: evidence from in situ exclusion experiments on soft and rocky substrates

In coastal marine ecosystems, predation might affect spatial distribution and population dynamics of benthic assemblages. Here, by means of experimental exclusion of potential predators, we compared the effects of epibenthic predation on metazoan meiofaunal assemblages on soft and rocky substrates. Different patterns of abundance were observed in uncaged versus caged plots, across habitats. In caged soft substrates, the abundance of Nematodes, Copepods and Polychaetes increased by 56, 45, 57%, respectively, in the first 3 months. An increase in the number of meiofaunal taxa was also observed. The exclusion of predators from rocky substrates showed less clear patterns. It did not affect the number of taxa while a decrease in meiofaunal abundance was observed. Our results suggest that the exclusion of epibenthic predators had clear effect on total metazoan meiofaunal abundance and on the number of taxa, only in soft bottoms. The different impact of predation across habitats can be potentially explained by differences in terms of spatial variability and substrate complexity. We estimated that, coarsely, more than 75% of total metazoan meiofaunal production can be channeled to higher trophic levels through predation on soft-bottoms. Among meiofaunal taxa, Polychaetes and Nematodes provided the major contribution to benthic energy transfers. These results suggest the trophic relevance of metazoan meiofauna in coastal food webs and claim for the refinement of further experiments for the quantification of its role in different ecological systems.     

Teams in social insects: group retrieval of prey by army ants (Eciton burchelli,Hymenoptera: Formicidae)

Summary Eciton burchelli workers frequently form groups to retrieve large prey items. Such groups have a definite structure. There is a constant relationship between total ant dry weight and prey item dry weight for both individual porters and groups, and this relation is such that a larger weight of ant or ants can carry disproportionately heavy items. Furthermore, all prey items are carried at a standard retrieval speed. This means that groups are superefficient; they can carry items that are so large that if they were fragmented the original members of the group would be unable to carry all the fragments. Groups also have a distinct caste distribution. There is a significant tendency for each group to have a single submajor, the specialist porter caste in Eciton burchelli. These findings which were verified by experiments suggest that groups fulfill the criteria of efficient teams. The biomechanics are proposed to explain the superefficiency of groups. The organization of co-operation is considered as is the role of teams in the economy of these societies. Colony caste profiles can only be understood by examining the role of teams which form a plastic supercaste.