Colony Members Research Articles

ХУДОЖНИК А. Н. МОКРИЦКИЙ В ИТАЛИИ В 1840-Е ГГ.: ОПЫТ РАБОТЫ НАД ПЕЙЗАЖЕМ

The paper explores the art works of A. N. Mokritsky, the painter who lived in Italy in the 1840s and had a strong passion for landscape painting. Being taught by A. G. Venecianov first and then graduating from the Imperial Academy of Arts under K. P. Bryullov, he himself followed the path of teaching and became an outstanding person in the Russian Art History of the second third of the 19th century. Mokritsky came to be known as a painter of an average talent who didn’t leave a distinctive mark on the national art. However, it was him who as a presumable representative of the artistic milieu became an indicator of the changes taking place in this art environment. The article provides a picture of years Mokritsky spent in Italy which is the most important period of his professional development and a prominent time of the Roman colony of the Russian artists as well. The author considers the artist’s close interaction not only with members of the Russian colony in Rome, but also with representatives of European art schools. Involving of archival materials and literary sources allowed to substantially supplement information about the life and work of Mokritsky during his trip abroad. Upon analysis of a significantly expanded list of landscape works created by the artist in this period, the author identified a number of characteristic features of the Italian landscape of the 40s of the 19th century taking into account the works of other painters.

Distribution pattern of swarm-founding eusocial wasps in the Indonesian Archipelago in comparison of that of parrots, one of sedentary bird groups

In the colony founding modes, eusocial wasps are divided into “independent-founders”, which initiate a colony by one to several inseminated females, and “swarm-founders”, which found their new colony by a number of “workers” accompanying one to many “queens”. As the swarm-founders put their trail pheromone on a guide to lead their colony members to a new nesting site, they would not disperse across large water bodies, such as sea and wide rivers. The present-day distribution pattern of swarm-founding wasps on the islands that are currently separated from each other and from continents should reflect the historical geology of these islands. The distribution patterns of the swarm-founding eusocial wasps in the Indonesian Archipelago are characterized by (1) occurrence of Asian continental elements in Borneo and Sumatra and their adjacent small islands, with a few species extending eastwards to Flores of Lesser Sunda Islands; (2) restriction of Australasian elements to New Guinea and its adjacent small islands including Aru; and (3) absence on most islands in Wallacea such as Sulawesi, Moluccas and eastern parts of Lesser Sunda Islands including Timor. These islands where no swarm-founders occur have never merged with any continental (Asian or Australasian) land mass. In the current distribution patterns in the Indonesian Archipelago, the swarm-founding eusocial wasps and the parrots are superficially similar, but their historical biogeography would be quite different.

ANALYSIS OF FISHING ACTIVITY AND MANAGEMENT OF RESOURCES IN THE ARTISAN FISHERMAN COLONY IN BAHIA SEMIARID

In this article we seek to analyze the fishing activity of the Colony of Fishermen and Aquaculture Z-86, in the municipality of Gloria, Bahia. Currently with 246 associates, where randomly, 20 families with low educational level and mostly represented by women were interviewed. The study focuses on the analysis of fishing activity, and the management of its resources against large-scale production. The methodology used was bibliographic research, websites and books, as well as technical visits, where two types of semi-structured questionnaires were applied, one for the colony members, the other for the president. The questionnaires contained questions regarding social, technical and economic aspects. The results indicate that artisanal fishing is going through difficult times due to the changes that man, in general, make in the environment, causing environmental impacts and consequently affecting the community economy.

Two aggressive neighbours living peacefully: the nesting association between a stingless bee and the bullet ant

Stingless bees often associate with termites, while association with ants is uncommon due to the high costs related to the aggressiveness of ants. Few combative genera of stingless bees can associate with a larger number of hosts, including ants. Here, we describe for the first time the association between a stingless bee (Partamona testacea) and the aggressive predator bullet ant (Paraponera clavata). In the study area, the colonies of P. testacea we spotted were all associated with P. clavata or Atta sp. ants. COI mitochondrial gene sequences of bees associated with bullet ants and leafcutter ants did not show any divergence, thus revealing no evidence that the two nesting strategies represent a case of cryptic speciation and specialisation on specific hosts. Bees are not unarmed respect to the bullet ants; when ants attempted to penetrate in the colony entrance, they were dragged inside the nest and covered by a resin-like substance. Behavioural experiments focused on ants in arenas and focused on bees at their nest entrance proved that the ants are significantly less aggressive toward associated bees and that guard bees are less alarmed when associated ants are presented. We verified by Gas Chromatography and Mass Spectrometry that P. testacea maintains its species-specific cuticular signature in the association with different ant species and that ants and bees possess typical colony signatures. The differential behavioural responses expressed toward associated colony members by both species are likely based on learning these heterospecific cuticular signatures as it occurs in ant parabiotic associations.

Sight in a Clique, Scent in Society: Plasticity in the Use of Nestmate Recognition Cues Along Colony Development in the Social Wasp Polistes dominula

Nestmate recognition, i.e., the ability to discriminate nestmates from foreign individuals, is a crucial feature of insect societies, and it has been traditionally considered to be predominantly based on chemical cues. Recent empirical evidence, however, suggests a relevant plasticity in the use of different communication channels according to cue availability and reliability in different contexts. In particular, visual cues have been shown to influence various types of social recognition in several social insects, but their role in nestmate recognition is still under-investigated. We tested the hypothesis of plasticity in the use of visual and chemical recognition cues in the primitively eusocial wasp Polistes dominula, in which the availability and reliability of recognition cues vary across the colony cycle. Indeed, before the emergence of workers, P. dominula colonies are rather small (one to few individuals), and the variability in the facial pattern might allow resident wasps to use visual cues for nestmate recognition. After workers' emergence, the increase in the number of colony members reduces the reliability of visual cues, thus leaving chemical cues as the most reliable nestmate recognition cues. We thus predict a differential use of chemical and visual cues along colony life. We experimentally separated visual and chemical cues of nestmates and non-nestmates and presented them alone or in combination (with coherent or mismatched cues) to resident wasps to test which communication channel was used in the two stages and, in case, how visual and chemical cues interacted. Our results show, for the first time in a social insect, the differential use of visual and chemical cues for nestmate recognition in two different phases of colony, which supports the hypothesis of a plastic, reliability-based use of recognition cues in this species according to the different colonial contexts.

Facing death: How does the subterranean termite Coptotermes gestroi (Isoptera: Rhinotermitidae) deal with corpses?

Social insects display a range of sophisticated behaviors to deal with cadavers, which together act to guarantee the health and social homeostasis of their colonies. In termites, cadaver management involves an entire repertoire of activities, such as antennation, grooming, retreat, alarm and agonistic behaviors. We performed different bioassays to evaluate whether postmortem-age, origin and caste of a cadaver influence the behavior of Coptotermes gestroi. Quantitative analyses of corpse management behaviors indicated that C. gestroi is able to discriminate the origin and caste of cadavers. Cadavers of worker nestmates were preferentially cannibalized while corpses of alien workers were buried. In addition, soldiers that had been dead for 24hours were buried while freshly dead soldiers were either buried, consumed or ignored, indicating a plastic behavioral response that depends on postmortem age. Corpse consumption is an important component of corpse management repertoire because this process eliminates contaminant and enables nutrient recycling. The physical isolation of the termite corpse through burial after grooming also helps to prevent other colony members from contacting the cadaver, further mitigating against the potential spread of disease.

Worker demography and behavior in a supercolonial ant colony: The case of the desert antCataglyphis niger

AbstractInsect societies headed by multiple queens (polygyny) raise evolutionary questions, such as how does genetic heterogeneity among colony members affect in‐nest interactions; or, are all queens equally reproductive or equally treated by workers? Answering such questions requires intensive and continuous observations of in‐nest behavior. Here, we addressed these questions in the polygyne supercolonial ant,Cataglyphis niger,using a barcoding system that enables the tracking of individual interactions, together with polymorphic DNA microsatellite markers that indicate the matriline and patriline of all individuals. Our findings that both queens and workers have low interrelatedness corroborate earlier reports regarding the supercolony structure ofC. niger. Ovary inspection and worker genotyping revealed that all the queens contribute similarly to nest demography. Tracking positions of individual workers through time revealed that only a small proportion of them are constantly engaged in tending the queens and can be considered as retinue workers. However, genotyping these workers and the attended queens revealed no relationship to genetic relatedness, again typical of a true polygyne and supercolonial species. Unlike invasive supercolonial species,C. nigeris native to Israel, enabling us to address questions regarding the driving forces, other than kin selection, that stabilize this society.

Wild zebra finches that nest synchronously have long-term stable social ties.

Many animals live and breed in colonies, and yet, with just a few exceptions, the value of the social bonds between colony members has rarely been examined. Social ties are important for group coordination at collective tasks, and social coordination can facilitate synchronized reproduction among colony members. Synchronized reproduction in turn can amplify the benefits of coloniality, such as social foraging and predator avoidance. We conducted a field study to investigate whether synchronized reproduction among individuals in replicated colonies is linked to the strength of their social bond, and whether these strong bonds are maintained beyond the reproductive period. We PIT-tagged wild zebra finches (Taeniopygia guttata), monitoring their reproduction and social foraging over two consecutive years. We then used social network analysis to characterize the strength of social bonds among birds in the population. We show that birds that are more synchronized in their reproductive timing (and breed in the same colony) had significantly stronger social ties both during and after reproduction than expected by chance. Our long-term sampling also revealed that the strong social ties between synchronized breeders were carried over across years. Our study reveals a strong correspondence between synchronized breeding and the social structure of the breeding colony. This suggests that the synchrony between pairs is not a simple process based on opportunity, but a mechanism underpinned by more complex sociality, which could be carried over to other behavioural contexts. The maintenance of cross-contextual social ties across years suggests that social structure could have extensive consequences on the overall life history of individuals in addition to playing a key role for the reproductive dynamics of colonial breeders.

Transcriptome changes reveal the genetic mechanisms of the reproductive plasticity of workers in lower termites

BackgroundThe reproductive plasticity of termite workers provides colonies with tremendous flexibility to respond to environmental changes, which is the basis for evolutionary and ecological success. Although it is known that all colony members share the same genetic background and that differences in castes are caused by differences in gene expression, the pattern of the specific expression of genes involved in the differentiation of workers into reproductives remains unclear. In this study, the isolated workers of Reticulitermes labralis developed into reproductives, and then comparative transcriptomes were used for the first time to reveal the molecular mechanisms underlying the reproductive plasticity of workers.ResultsWe identified 38,070 differentially expressed genes and found a pattern of gene expression involved in the differentiation of the workers into reproductives. 12, 543 genes were specifically upregulated in the isolated workers. Twenty-five signal transduction pathways classified into environmental information processing were related to the differentiation of workers into reproductives. Ras functions as a signalling switch regulates the reproductive plasticity of workers. The catalase gene which is related to longevity was up-regulated in reproductives.ConclusionWe demonstrate that workers leaving the natal colony can induce the expression of stage-specific genes in the workers, which leads to the differentiation of workers into reproductives and suggests that the signal transduction along the Ras-MAPK pathway crucially controls the reproductive plasticity of the workers. This study also provides an important model for revealing the molecular mechanism of longevity changes.

Phenotypic Plasticity of Nest-Mate Recognition Cues in Formica exsecta Ants

It is well established that many ant species have evolved qualitatively distinct species-specific chemical profile that are stable over large geographical distances. Within these species profiles quantitative variations in the chemical profile allows distinct colony-specific odours to arise (chemotypes) that are shared by all colony members. This help maintains social cohesion, including defence of their colonies against all intruders, including con-specifics. How these colony -level chemotypes are maintained among nest-mates has long been debated. The two main theories are; each ant is able to biochemically adjust its chemical profile to ‘match’ that of its nest-mates and or the queen, or all nest-mates share their individually generated chemical profile via trophollaxis resulting in an average nest-mate profile. This ‘mixing’ idea is better known as the Gestalt model. Unfortunately, it has been very difficult to experimentally test these two ideas in a single experimental design. However, it is now possible using the ant Formica exsecta because the compounds used in nest-mate recognition compounds are known. We demonstrate that workers adjust their profile to ‘match’ the dominant chemical profile within that colony, hence maintaining the colony-specific chemotype and indicates that a ‘gestalt’ mechanism, i.e. profile mixing, plays no or only a minor role.

The survival of the shyest: a computational model shows the effect of web structure on the origins of social spiders

For social life to emerge, the benefits of group living must overcome its costs, a balance that depends on the life history of the organisms and may occur more easily in some taxa than in others. Among spiders, sociality is uncommon. Social species evolved several times independently through the subsocial route, a process connected to a mean reduction in the aggressiveness and boldness of the individuals, with a simultaneous increase in the variability of personality. Personality variability is at the root of spider's division of labour, but the mechanism for its increase is yet poorly studied. One possible mechanism for the evolution of the variability in personality would be the creation of a new selective regime by the newly evolved, larger colonial webs, which could offer a mix of more protected and predator-exposed areas. To evaluate this hypothesis, we present an ecological-evolutionary model in which heterogeneous, spatially explicit variation in predatory selective pressures appear through changes in niche construction. Our model shows that social webs, by providing areas of greater protection against predators, increase the survival of the shyest variants, leading to increased intracolonial variability in personality. We speculate that this greater protection allowed social species the possibility of colonizing otherwise inhospitable environments, and that larger intracolonial personality variance, associated with social web spatial heterogeneity provided the basis for social organization through division of tasks among colony members.

Colony entropy-Allocation of goods in ant colonies.

Allocation of goods is a key feature in defining the connection between the individual and the collective scale in any society. Both the process by which goods are to be distributed, and the resulting allocation to the members of the society may affect the success of the population as a whole. One of the most striking natural examples of a highly successful cooperative society is the ant colony which often acts as a single superorganism. In particular, each individual within the ant colony has a “communal stomach” which is used to store and share food with the other colony members by mouth to mouth feeding. Sharing food between communal stomachs allows the colony as a whole to get its food requirements and, more so, allows each individual within the colony to reach its nutritional intake target. The vast majority of colony members do not forage independently but obtain their food through secondary interactions in which food is exchanged between individuals. The global effect of this exchange is not well understood. To gain better understanding into this process we used fluorescence imaging to measure how food from a single external source is distributed and mixed within a Camponotus sanctus ant colony. Using entropic measures to quantify food-blending, we show that while collected food flows into all parts of the colony it mixes only partly. We show that mixing is controlled by the ants’ interaction rule which implies that only a fraction of the maximal potential is actually transferred. This rule leads to a robust blending process: i.e., neither the exact food volume that is transferred, nor the interaction schedule are essential to generate the global outcome. Finally, we show how the ants’ interaction rules may optimize a trade-off between fast dissemination and efficient mixing. Our results regarding the distribution of a single food source provide a baseline for future studies on distributed regulation of multiple food sources in social insect colonies.

Behavioural response of workers to repeated intergroup encounters in the harvester ant Messor barbarus

The evolution of cooperation in animal societies is often associated with the evolution of hostility towards members of other groups. It is usually predicted that groups under attack from outsiders should respond by becoming more cohesive or cooperative. However, the responses of individuals to real or simulated intergroup encounters vary widely, for reasons that are poorly understood. We tested how groups of workers of the harvester ant, Messor barbarus, responded to exposure to members of a different colony versus members of their own colony, and how previous exposure to an intruder affected the intensity of the within-group response. We found that workers increased in activity and had more contact with one another immediately following exposure to an ant from a different colony, but also showed a similar behavioural response to presentations involving an ant from their own colony. However, exposure to an intruder from a different colony resulted in much stronger behavioural responses to a second intruder, encountered shortly afterwards. Our results are consistent with studies of social vertebrates which suggest that exposure to intruders results in increased social cohesion. Our results also show that exposure to an intruder primes group members to respond more strongly to future intrusions. Our findings highlight a disconnect between the assumptions of theoretical models which study the effect of intergroup conflict on social evolution over many generations, and the short-term behavioural responses that are the usual focus of studies of intergroup conflict in insects and vertebrates.

Food for Sex in Bats Revealed as Producer Males Reproduce with Scrounging Females

Food sharing is often evolutionarily puzzling, because the provider's benefits are not always clear. Sharing among kin may increase indirect fitness [1], but when non-kin are involved, different mechanisms were suggested to act. Occasionally, "tolerated theft" [2, 3] is observed, merely because defending a resource is not cost effective. Sharing may also be explained as "costly signaling" [4, 5], where individuals signal their high qualities by distributing acquired resources, as has been suggested to occur in certain human cultures [6]. Alternatively, a transferred food item might be compensated for in later interactions [7]. In vampire bats, blood sharing reflects reciprocity between non-kin colony members [8-10], and long-term social bonds affect food sharing in chimpanzees [11]. Food may also be exchanged for other goods or social benefits [12-14]. One reciprocity-based explanation for intersexual food sharing is the food-for-sex hypothesis [15-17]. This hypothesis proposes that males share food with females in exchange for mating opportunities. Studies on human hunter-gatherer societies suggest that males with increased foraging success have higher reproductive success [18, 19]. Male chimpanzees, which in contrast to humans do not maintain pair bonds, were suggested to share food with females to increase their mating opportunities [16] (but see [20]). Bats, which are long-lived social mammals [21, 22], provide an opportunity to study long-term social reciprocity mechanisms. We monitored producer-scrounger interactions of a captive Egyptian fruit bat (Rousettus aegyptiacus) colony for more than a year and genetically determined the paternity of the pups that were born in the colony. We found that females carry the young of males from which they used to scrounge food, supporting the food-for-sex hypothesis in this species.

Bumblebees adjust protein and lipid collection rules to the presence of brood.

Animals have evolved foraging strategies to acquire blends of nutrients that maximize fitness traits. In social insects, nutrient regulation is complicated by the fact that few individuals, the foragers, must address the divergent nutritional needs of all colony members simultaneously, including other workers, the reproductives, and the brood. Here we used 3D nutritional geometry design to examine how bumblebee workers regulate their collection of 3 major macronutrients in the presence and absence of brood. We provided small colonies artificial nectars (liquid diets) and pollens (solid diets) varying in their compositions of proteins, lipids, and carbohydrates during 2 weeks. Colonies given a choice between nutritionally complementary diets self-selected foods to reach a target ratio of 71% proteins, 6% carbohydrates, and 23% lipids, irrespective of the presence of brood. When confined to a single nutritionally imbalanced solid diet, colonies without brood regulated lipid collection and over-collected protein relative to this target ratio, whereas colonies with brood regulated both lipid and protein collection. This brood effect on the regulation of nutrient collection by workers suggests that protein levels are critical for larval development. Our results highlight the importance of considering bee nutrition as a multidimensional phenomenon to better assess the effects of environmental impoverishment and malnutrition on population declines.

Liquid exchange via stomodeal trophallaxis in the ponerine ant Diacamma sp. from Japan

Trophallaxis plays a major role in the sharing of food in colonies of many social insects, and two modes of this are known: stomodeal (oral) and proctodeal (abdominal) trophallaxis. In social Hymenoptera, only a small proportion of colony members perform the task of food collection, and oral trophallaxis is predominant in their social sharing of food. Typically, foragers distribute liquid food stored in their crop to nestmates via oral trophallaxis. Similar to bees, some ants (Formicidae) forage for liquid food from plant secretions (nectars) and insect exudates (honeydew). While regurgitation is common in ants, it has been documented in only two species of the Ponerinae. Here, we report the ability of Diacamma sp. from Japan to perform trophallaxis. After thirsty ants had been paired with ants provided with colored water, the abdomens of both groups of ants were dissected. The digestive organ was colored red in half of the receivers. In addition, we observed mouth-to-mouth interactions in the laboratory, not “social bucket” behavior (i.e., exchange of liquid held between mandibles). Our results suggest that Diacamma sp. can exchange liquid by true oral trophallaxis and shed new light on social organization via liquid exchange.

Time‐frequency decomposition of seismic signals via quantum swarm evolutionary matching pursuit

ABSTRACTMatching pursuit belongs to the category of spectral decomposition approaches that use a pre‐defined discrete wavelet dictionary in order to decompose a signal adaptively. Although disengaged from windowing issues, matching point demands high computational costs as extraction of all local structure of signal requires a large size dictionary. Thus in order to find the best match wavelet, it is required to search the whole space. To reduce the computational cost of greedy matching pursuit, two artificial intelligence methods, (1) quantum inspired evolutionary algorithm and (2) particle swarm optimization, are introduced for two successive steps: (a) initial estimation and (b) optimization of wavelet parameters. We call this algorithm quantum swarm evolutionary matching pursuit. Quantum swarm evolutionary matching pursuit starts with a small colony of population at which each individual, is potentially a transformed form of a time‐frequency atom. To attain maximum pursuit of the potential candidate wavelets with the residual, the colony members are adjusted in an evolutionary way. In addition, the quantum computing concepts such as quantum bit, quantum gate, and superposition of states are introduced into the method. The algorithm parameters such as social and cognitive learning factors, population size and global migration period are optimized using seismic signals. In applying matching pursuit to geophysical data, typically complex trace attributes are used for initial estimation of wavelet parameters, however, in this study it was shown that using complex trace attributes are sensitive to noisy data and would have lower rate of convergence.The algorithm performance over noisy signals, using non‐orthogonal dictionaries are investigated and compared with other methods such as orthogonal matching pursuit. The results illustrate that quantum swarm evolutionary matching pursuit has the least sensitivity to noise and higher rate of convergence. Finally, the algorithm is applied to both modelled seismograms and real data for detection of low frequency anomalies to validate the findings.

Social immunity behaviour among ants infected by specialist and generalist fungi.

Social insects are distinguished by their lifestyle of living in groups with division of labour, cooperative brood care, and reproduction limited to a few colony members. Social insects often build large colonies with remarkable densities of highly related individuals and this can lead to an increased pathogen pressure. Our review focuses on interactions of ants with two important taxonomic groups of fungi infecting ants: Hypocreales (Ascomycota) and Entomophthorales (Entomophthoromycotina), and their different infection strategies, including host manipulation for optimal spore dispersal in the specialised ant pathogens. In social insects such as ants, resistance to pathogens is present at the colony level, with social immunity in addition to the individual resistance. We describe how ants use both organizational and behavioural defence strategies to combat fungal pathogens, with emphasis on highly specialised fungi from the genera Ophiocordyceps and Pandora.

Red Wood Ants Display Natural Aversive Learning Differently Depending on Their Task Specialization.

The adaptive benefits of individual specialization and how learning abilities correlate with task performance are still far from being well-understood. Red wood ants are characterized by their huge colonies and deep professional specialization. We hypothesized that red wood ants Formica aquilonia form aversive learning after having negative encounters with hoverfly larvae differently, depending on their task specialization. We tested this hypothesis, first, by examining whether hunters and aphid milkers learn differently to avoid the nuisance of contacts with syrphid larvae, and, second, by analyzing the difference between learning in “field” and laboratory-reared (naïve) foragers. During the first interaction with the syrphid larva in their lives the naïve foragers showed a significantly higher level of aggressiveness than the members of a natural colony. Naïve foragers applied the “mortal grip,” “prolonged bites,” and “nibbling” toward the enemy with a significantly higher frequency, whereas members of both “field” groups behaved more carefully and tried to avoid encounters with the larva. The aphid milkers, who had a negative experience of interaction with the larva, being “glued” with its viscous secretion, behaved much less aggressively in the follow-up experiments after 10 min and even 3 days, thus exhibiting the shaping of both short- and long-term memories. However, both “field” hunters and naïve foragers demonstrated no signs of aversive learning. These data provide some new insights into the relationship between task specialization and learning performance in ants. Given our previous results, we speculate that scouts and aphid milkers are the most cognitively gifted specialists in red wood ants, whereas hunters and guards are rather brave than smart.

Caste- and pesticide-specific effects of neonicotinoid pesticide exposure on gene expression in bumblebees.

Social bees are important insect pollinators of wildflowers and agricultural crops, making their reported declines a global concern. A major factor implicated in these declines is the widespread use of neonicotinoid pesticides. Indeed, recent research has demonstrated that exposure to low doses of these neurotoxic pesticides impairs bee behaviours important for colony function and survival. However, our understanding of the molecular‐genetic pathways that lead to such effects is limited, as is our knowledge of how effects may differ between colony members. To understand what genes and pathways are affected by exposure of bumblebee workers and queens to neonicotinoid pesticides, we implemented a transcriptome‐wide gene expression study. We chronically exposed Bombus terrestriscolonies to either clothianidin or imidacloprid at field‐realistic concentrations while controlling for factors including colony social environment and worker age. We reveal that genes involved in important biological processes including mitochondrial function are differentially expressed in response to neonicotinoid exposure. Additionally, clothianidin exposure had stronger effects on gene expression amplitude and alternative splicing than imidacloprid. Finally, exposure affected workers more strongly than queens. Our work demonstrates how RNA‐Seq transcriptome profiling can provide detailed novel insight on the mechanisms mediating pesticide toxicity to a key insect pollinator.