The comparative biology of two sympatric paper wasps in Michigan,the native <Emphasis Type="Italic">Polistes fuscatus</Emphasis> and the invasive <Emphasis Type="Italic">Polistes dominulus</Emphasis> (Hymenoptera,Vespidae)

Summary The biology of the invasive Polistes dominulus and the native P. fuscatus was compared at a field site in Rochester, Michigan over a two-year period. Colonies nesting semi-naturally in plywood nestboxes were studied using videography, extensive surveys, and colony-specific marking of gynes.Both single- and multiple-foundress colonies of P. dominulus were significantly more productive than comparable colonies of P. fuscatus. The disparity in productivity was significantly more pronounced in single-foundress colonies than in multiple-foundress colonies. P. dominulus had significantly shorter larval and pupal development times than P. fuscatus, which allowed P. dominulus to produce its first workers about a week earlier than P. fuscatus. P. dominulus had a number of additional advantages over P. fuscatus that contributed to its productivity including (1) significantly less parasitism by Strepsiptera, (2) significantly greater probability of renesting after predation by raccoons, (3) significantly lower usurpation pressures, and (4) possibly longer foraging days . The recovery of colony-marked foundresses indicated that gynes of P. dominulus suffered significantly greater mortality than gynes of P. fuscatus during winter diapause and that foundresses of both species were equally, strongly philopatric.P. dominulus is likely replacing P. fuscatus in many areas of southeastern Michigan via indirect or exploitative competition. The two species may be competing for nest sites.Received 7 July 2003; revised 10 October 2003; accepted 3 November 2003.     

A role for thermoregulation in the <Emphasis Type="Italic">Polistes dominulus</Emphasis> invasion: a comparison of the thermoregulatory abilities of the invasive wasp <Emphasis Type="Italic">P. dominulus</Emphasis> and the native wasp <Emphasis Type="Italic">P. fuscatus</Emphasis>

Social insects are excellent invaders that have had negative impacts on native species and humans. Many invasive species move from warmer to cooler climates. For these species, thermal adaptations may both be important for their ability to invade and to limit their invasion range. The invasion of Polistes dominulus into North America provides an example of a primitively eusocial invader from a warmer climate. We studied the differences in thermoregulation between P. dominulus and the native P. fuscatus. We found that, during flight, thorax temperature in P. fuscatus was less affected by ambient temperature than thorax temperature of P. dominulus. We also found that P. dominulus and P. fuscatus showed different patterns of warming after removal from a cold environment. Unlike P. dominulus, live P. fuscatus never fully cooled down in a cold environment. P. fuscatus also reached their relative minimum flight temperatures earlier than P. dominulus, but P. dominulus maintained higher elevated temperatures for longer. These differences in thermoregulatory ability suggest that the lower winter survival of P. dominulus could be offset by a greater thermal tolerance during flight, while the lower thermal tolerance of P. fuscatus in flight is offset somewhat by better thermoregulatory ability.     

The Polistes war: weak immune function in the invasive P. dominulus relative to the native P. fuscatus

Invasive species are of growing ecological concern, in part because of conflicts arising with native congeners. The European paper wasp Polistes dominulus was first introduced to North America in the 1970s, and may be displacing at least one native species, P. fuscatus. Previous reports indicate that in native territories over half of P. dominulus colonies are infected by Strepsipteran parasites, which decrease host fitness. In North America, P. fuscatus are parasitized to a lesser degree (approximately one-third), but no infected colonies of invasive P. dominulus have been reported. Because immune function is an indicator of susceptibility to parasitism, we quantified activated levels of immune function by measuring the encapsulation response and phenoloxidase activity and then compared these levels between species. Counterintuitively, our results indicate that P. dominulus has lower levels of both mechanisms of immunity. Additionally, P. dominulus displayed less self-grooming activity than P. fuscatus. We briefly discuss possible immunological explanations for this invasion success, including the selective expression of low immunocompetence.     

A ten-year comparative study of the population dynamics and parasitoidism in the native paper wasp Polistes fuscatus and the invasive P. dominulus

The invasive success of Polistes dominulus in North America has been attributed to its greater productivity relative to native Polistes. Liberation from parasites and parasitoids are thought to be major factors contributing to the high productivity of P. dominulus. We analyzed historical records of colony relative abundance and productivity of P. dominulus and the sympatric, native Polistes fuscatus from 1995 to 2010 using historical data from our Michigan Polistes study site. We also analyzed evidence of parasitoids from 294 P. fuscatus and 507 P. dominulus archived combs from 2001 to 2010. Additionally, we examined field and laboratory colonies from outside of our study site for parasites and parasitoids in 2009 and 2010. We documented one parasite and three parasitoids exploiting Polistes in our Michigan study sites. Our historical records document that P. dominulus initially displaced P. fuscatus rapidly, then slowed, and finally the two populations stabilized. Furthermore, the historical pattern of decreasing displacement of P. fuscatus by P. dominulus corresponded temporally with a significant decline in the productivity and a significant increase in Dibrachys cavus infestation of P. dominulus. Our evidence indicates that the parasitoid, D. cavus, is a major factor in stabilizing the populations of the sympatric P. dominulus and P. fuscatus.     

The comparative biology of two sympatric paper wasps, the native Polistes fuscatus and the invasive Polistes dominulus (Hymenoptera, Vespidae)

Summary: Polistes dominulus (Christ), an old world paper wasp, was introduced accidentally into the eastern coast of the United States in the late 1970s and has been rapidly spreading westward, displacing the native P. fuscatus (F.). The biology of naturally nesting P. fuscatus and P. dominulus was compared at a field site in Rochester, Michigan. The basic methodology consisted of simultaneously videotaping spatially proximate, matched single-foundress colonies of P. fuscatus and P. dominulus (13 matched sets, 176.8 h of videography). In addition, extensive surveys and censuses were taken of colonies to record colony productivity, dates of nest initiation and first worker emergence, usurpation and parasitism.¶There was no evidence that P. dominulus is negatively impacting P. fuscatus through direct, agonistic encounters. However, P. dominulus is 4-5 times more productive than P. fuscatus, suggesting that P. dominulus is replacing P. fuscatus via exploitative competition. P. dominulus appears to have a number of advantages over P. fuscatus, including earlier production of workers, higher per capita foraging rates by queens and workers, higher queen survivorship, and lack of conspecific pressures. Nest site and/or prey availability may be limiting factors in the competition between the two species.     

Modes of colony foundation by females of different morphotypes in the paper wasps (Hymenoptera,Vespidae, <Emphasis Type="Italic">Polistes</Emphasis> Latr.)

Paper wasps of the genus Polistes exhibit wide variability both of the color and size of foundresses, and the mode of colony foundation. The colony foundation and social hierarchy were studied in populations of two Palaearctic species, Polistes dominulus (Christ) and P. nimphus (Christ) (April–May 2006, the Crimean Peninsula, Ukraine), and one Neotropical species, P. lanio (F.) (October 2004–January 2005, Trinidad Island). It was shown that foundresses of various color and size morphs tended to found nests in a peculiar way: singly (haplometrosis), by groups (pleometrosis), or occupying several nests simultaneously (polycaly). The relationship between coloration, size, and mode of colony foundation was species-specific. P. dominulus foundresses with a darker clypeus and a paler mesonotum preferred to found nests alone and had a dominant position in pleometrosis. The haplometrotic P. nimphus foundresses more often displayed darker variants of the clypeus and paler variants of the mesonotum than did foundresses from pleometrotic colonies. P. lanio foundresses from pleometrotic colonies differed from those from polycalic colonies in the coloration variability of the scutum, propodeum, and the 2nd metasomal tergite. The dominant and subordinate P. nimphus and P. lanio foundresses differed in the size of head and wings. The possible significance of the subdivision of foundresses into the “generalists” and “specialists” for the variability structuring in the population is considered.     

Simulated Intraspecific Usurpations in Paper Wasps: Different Reproductive Tactics Affect Foreign Brood Destruction in Polistes fuscatus and Polistes dominulus

We simulated intraspecific usurpation in two species of paper wasps at a field site in Michigan to compare the species' treatment of foreign brood. Queens of Polistes fuscatus, a species that commonly uses intraspecific usurpation as an alternative reproductive tactic, destroyed significantly fewer large larvae and significantly more small larvae than queens of P. dominulus, a sympatric species that rarely usurps. The pattern of brood destruction exhibited by P. fuscatus was consistent with the previously published findings that P. fuscatus usurpers destroy reproductive‐destined brood (eggs and small larvae), but not worker‐destined brood (large larvae and pupae) that are subsequently used by a usurper to raise her own reproductives. The pattern of brood destruction displayed by P. dominulus differed from that of Polistes species that frequently engage in intraspecific usurpation. The brood destruction pattern in P. dominulus may have been shaped by nest adoption, a common alternative reproductive tactic in this species. If so, it is not clear why P. dominulus would destroy large, worker‐destined larvae.     

Prey capture and caste-specific payload capacities in the European paper wasp Polistes dominulus

Here, we present a detailed ethogram for the foraging behavior of the eusocial paper wasp, Polistes dominulus. The animal’s foraging process can be sub-divided into four main stages: (1) approach, (2) attack, (3) butchering, and (4) balling. Although considerable behavioral variation exists within each stage, an analysis of more than 20 individual foraging bouts reveals a single “common path” leading from prey approach to its transportation back to the nest; elucidating this path and separating it into distinct stages is useful when identifying targets for future ethological studies. Of particular interest here is the balling phase, i.e., the preparation of a bolus of flesh for transportation back to the colony. Using an experimental approach, we show that foundresses can carry significantly heavier payloads than workers, suggesting a foraging advantage during the initial stages of colony development. We also show that wasp body mass is significantly positively correlated with payload capacity in foundresses, a relationship not seen among workers or late reproductives. This correlation suggests a beneficial adaptation of foundresses for combating early season pressures associated with the foundation of a new colony.     

Synergistic effects of temperature,diet and colony size on the competitive ability of two ant species

Multiple biotic and abiotic factors influence species coexistence and co‐occurrence patterns. In a competitive environment, for example, temperature and diet variation may modify both foraging behaviour and aggression, thereby changing competitive interactions and species co‐occurrence patterns. In New Zealand, two endemic ant species (Prolasius advenus and Monomorium antarcticum) often form allopatric distributions; though also periodically do co‐occur in the same habitat. Here, we performed a long‐term laboratory experiment in an attempt to understand how diet, colony size and environmental conditions may influence these co‐occurrence patterns. The consequences of temperature and diet variation differed between P. advenus and M. antarcticum. Colonies of P. advenus exhibited increased aggression and foraging activities at higher temperatures. In addition, P. advenus colonies augmented their foraging activities when deprived of a carbohydrate‐rich food source. Conversely, small M. antarcticum colonies exhibited higher aggression than when in large colonies, and increased their foraging activities at lower temperatures. The modulation of aggression and foraging behaviour may influence the likelihood of small P. advenus and M. antarcticum colonies persisting in the long term. Our results are compatible with the hypothesis that the environment is likely to be a strong filter for the negative co‐occurrence patterns we observe between P. advenus and M. antarcticum in New Zealand. Furthermore, this study provides a mechanistic explanation for potential impacts of climate warming on community structure. Environmental modification of aggression and foraging behaviour could potentially alter competitive interactions and influence community assembly.     

Effect of food availability on larval cannibalism by foundresses of the paper wasp <Emphasis Type="Italic">Polistes chinensis antennalis</Emphasis>

Cannibalism is a behavioral trait seen in a number of species and can be favored when the fitness gain outweighs the cost. In social wasps, field studies have suggested that food limitation causes larval cannibalism as a compensation of a meal for adult wasp and/or earlier production of the first workers, but experimental studies for the effect of food availability on larval cannibalism have been very scarce. Hence we examined whether Polistes chinensis antennalis foundresses exhibit a higher level of larval cannibalization under food-limited conditions than under food-available conditions. Larval cannibalization occurred frequently under both prey- and honey-limited conditions; however, young larvae were more frequently cannibalized than old larvae in the prey-limited colonies, while old larvae were more frequently cannibalized than young larvae in the honey-limited colonies. The number of days between colony initiation and the emergence of the first workers was not significantly different between the prey-limited and prey-available colonies, suggesting that under prey-limited conditions P. chinensis antennalis foundresses tend to cannibalize their young larvae and feed probably the flesh of these cannibalized larvae to old larvae in order to ensure the rapid production of the first workers. On the other hand, the number of days between colony initiation and the emergence of the first workers was longer in the honey-limited colonies than in the honey-available colonies. We propose explanations for the first time for why the foundresses in the honey-limited conditions cannibalized their old larvae more frequently than their young larvae in spite of this resulting in the delayed emergence of the first workers.     

Lateral Vibrations by Social Wasps Signal Larvae to Withhold Salivary Secretions (Polistes fuscatus, Hymenoptera: Vespidae)

The communicative meaning of an oscillation (lateral vibration) performed by foundresses of social wasps (Polistes fuscatus) was investigated by sampling larval saliva from 12 postworker, single-foundress colonies and by viewing videotapes of 17 multiple-foundress (139.3 h) and 16 single-foundress (32 h) preworker colonies. Foundresses spent significantly more time inspecting cells before performing a lateral vibration (LV) than after and commonly became inactive after performing an LV. The proportion of departures temporally proximate to LVs was significantly greater in single- than in multiple-foundress colonies. Departures and LVs were significantly temporally associated in single-foundress colonies. Single foundresses were significantly more likely to perform an LV prior to departing on longer than shorter trips. In multiple-foundress colonies there was a significant temporal relationship between LVs and departures that left the nest unattended. Larvae secreted significantly less saliva, a nutritious substance provided to adults, immediately after an LV than in the absence of an LV. Contextual evidence and results of sampling larval saliva indicate that LVs by foundresses signal larvae to withhold or reduce their secretion of saliva.     

Benefits of foundress associations in the paper wasp Polistes dominulus: increased productivity and survival, but no assurance of fitness returns

Successful Polistes dominulus nests can be started by one ormore nest founding queens (foundresses). Consequently, thereis much interest in the specific benefits that induce cooperationamong foundresses. Here, we experimentally demonstrate one majorbenefit of cooperation, namely that multiple foundresses increasecolony productivity. This increase is close to the value predictedby subtracting the productivity of undisturbed single-foundresscolonies from the productivity of undisturbed multiple-foundresscolonies. However, we found no evidence that an associatingfoundress' contribution to colony growth is preserved if shedisappears (assured fitness returns). Our correlational datasuggest that cooperation provides survival benefits, multiple-foundresscolonies are more likely to survive to produce offspring thanare single-foundress colonies, and individual foundresses inmultiple-foundress groups are less likely to disappear beforeworker emergence than foundresses nesting alone. Therefore,association provides substantial productivity and survival benefitsfor cooperating foundresses.     

Pollinator genetics and pollination: do honey bee colonies selected for pollen‐hoarding field better pollinators of cranberry Vaccinium macrocarpon?

1. Genetic polymorphisms of flowering plants can influence pollinator foraging but it is not known whether heritable foraging polymorphisms of pollinators influence their pollination efficacies. Honey bees Apis mellifera L. visit cranberry flowers for nectar but rarely for pollen when alternative preferred flowers grow nearby. 2. Cranberry flowers visited once by pollen‐foraging honey bees received four‐fold more stigmatic pollen than flowers visited by mere nectar‐foragers (excluding nectar thieves). Manual greenhouse pollinations with fixed numbers of pollen tetrads (0, 2, 4, 8, 16, 32) achieved maximal fruit set with just eight pollen tetrads. Pollen‐foraging honey bees yielded a calculated 63% more berries than equal numbers of non‐thieving nectar‐foragers, even though both classes of forager made stigmatic contact. 3. Colonies headed by queens of a pollen‐hoarding genotype fielded significantly more pollen‐foraging trips than standard commercial genotypes, as did hives fitted with permanently engaged pollen traps or colonies containing more larvae. Pollen‐hoarding colonies together brought back twice as many cranberry pollen loads as control colonies, which was marginally significant despite marked daily variation in the proportion of collected pollen that was cranberry. 4. Caloric supplementation of matched, paired colonies failed to enhance pollen foraging despite the meagre nectar yields of individual cranberry flowers. 5. Heritable behavioural polymorphisms of the honey bee, such as pollen‐hoarding, can enhance fruit and seed set by a floral host (e.g. cranberry), but only if more preferred pollen hosts are absent or rare. Otherwise, honey bees' broad polylecty, flight range, and daily idiosyncrasies in floral fidelity will obscure specific pollen‐foraging differences at a given floral host, even among paired colonies in a seemingly uniform agricultural setting.     

An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguin Pygoscelis adeliae

Central-place foraging seabirds alter the availability of their prey around colonies, forming a "halo" of reduced prey access that ultimately constrains population size. This has been indicated indirectly by an inverse correlation between colony size and reproductive success, numbers of conspecifics at other colonies within foraging range, foraging effort (i.e. trip duration), diet quality and colony growth rate. Although ultimately mediated by density dependence relative to food through intraspecific exploitative or interference competition, the proximate mechanism involved has yet to be elucidated. Herein, we show that Adélie penguin Pygoscelis adeliae colony size positively correlates to foraging trip duration and metabolic rate, that the metabolic rate while foraging may be approaching an energetic ceiling for birds at the largest colonies, and that total energy expended increases with trip duration although uncompensated by increased mass gain. We propose that a competition-induced reduction in prey availability results in higher energy expenditure for birds foraging in the halo around large colonies, and that to escape the halo a bird must increase its foraging distance. Ultimately, the total energetic cost of a trip determines the maximum successful trip distance, as on longer trips food acquired is used more for self maintenance than for chick provisioning. When the net cost of foraging trips becomes too high, with chicks receiving insufficient food, chick survival suffers and subsequent colony growth is limited. Though the existence of energetic studies of the same species at multiple colonies is rare, because foraging metabolic rate increases with colony size in at least two other seabird species, we suggest that an energetic constraint to colony size may generally apply to other seabirds.     

Visual tracking from a rigid‐hulled inflatable boat to determine foraging movements of breeding terns

ABSTRACT Defining the at‐sea foraging movements of seabirds is fundamental to understanding their ecology and can also be important in assessing the potential impact of marine developments such as offshore wind farms (OWFs). Surveys of predefined areas using aerial or boat‐based transect surveys may not allow adequate assessment of the relative importance of different areas to birds. Individual‐based satellite or radio‐telemetry can be effective in identifying foraging ranges and preferred areas, but may not be suitable for some species. We developed a method to determine the foraging movements of breeding terns (Sterna spp.) by visually tracking individuals using a rigid‐hulled inflatable boat (RHIB). Sandwich Terns (S. sandvicensis), Common Terns (S. hirundo), and Arctic Terns (S. paradisaea) were tracked from colonies in Norfolk and Anglesey, United Kingdom, from 2006 to 2008. The proportion of complete (from and to colony) trips varied from 29–60% among species, years, and colonies. Individual Sandwich Terns were tracked for periods up to 126 min over distances up to 72 km and as far as 54 km from the breeding colony, further than Arctic (up to 57 km and 29 km from the colony) and Common (to 29 km and <9 km from the colony) terns. Mean values were much smaller. Multivariate modeling of Sandwich Tern foraging trips indicated that flight speeds >50 km/hr coupled with greater distances from shore (>25 km) significantly reduced the likelihood of tracking a bird for an entire foraging trip. Use of different boats that differ in speed and performance may alleviate such issues. Visual tracking allowed us to collect data on foraging behavior, flight height, and prey capture rates, and also permitted comparisons between species. Our results indicate that visual tracking may be an effective means of determining the foraging movements and at‐sea behavior of a variety of short‐ranging, day‐active seabirds.     

First detailed report of brood parasitoidism in the invasive population of the paper wasp Polistes dominulus (Hymenoptera, Vespidae) in North America

The European paper wasp Polistes dominulus (Christ) is a model system in the fields of behavioral ecology, ecological immunology, and invasion biology. Since its introduction to the US in 1978, its invasion success has been attributed, in part, to a lack of parasites or parasitoids infecting this population. This is despite the number of parasites which infest the native population and the generalist polistine parasites and parasitoids documented in sympatric North American species. Multiple studies have cited low parasite pressure as evidence that the invasive population of P. dominulus is benefiting from a post-invasion release from enemies. Here, we present the first well documented case of parasitoidism of the invasive population of P. dominulus in North America.     

Comparative activity pattern during foraging of four albatross species

The activity patterns of foraging Yellow‐nosed Diomedea chlororhynchos, Sooty Phoebetria fusca, Black‐browed D. melanophris impavida and Grey‐headed Albatross D. chrysostoma were compared using loggers recording the timing of landing and take‐offs, as well as the duration of bouts in flight or on the water, and the overall time spent in flight. The four species spent a similar proportion of their foraging time in flight (56–65%). During the day they were mostly flying (77–85% of the daylight period) whereas at night they were mainly (61–71%) sitting on the water. The amount of time spent in flight during the daytime foraging period was related to the amount of time spent sitting on the water at night. Differences between species occurred in the duration of bouts in flight and on the water as well as in the frequency of landings and in the time elapsed between successive landings. Yellow‐nosed Albatrosses were more active than the other species, with more frequent short bouts in flight and more frequent successive landings at short intervals. Sooty Albatrosses landed or took‐off less often than the other species and were more active just before dusk. Black‐browed and Grey‐headed Albatrosses were more active at night, especially the first part of the night and far from the colonies. Their trips consisted of a commuting part and a foraging part. Black‐browed Albatrosses landed more often during the foraging than the commuting part, suggesting that they were not searching when travelling. The study suggests that there is no fundamental difference between the overall activity budgets of the four species although they show distinctive diet, morphology and life history traits. The differences observed between the four species were related mainly to differences in foraging technique. Comparison with the Wandering Albatross, the only species for which data were available previously, suggest that this larger species might differ completely in foraging technique from the smaller albatrosses.     

The effect of juvenile hormone on temporal polyethism in the paper wasp Polistes dominulus

Juvenile hormone (JH) has an important role in the behavior of eusocial Hymenoptera. Previous work has shown that JH influences aggression and dominance behavior in primitive eusocial insects that lack discrete queen and worker castes (e.g. Bombus bees and Polistes wasps). In contrast, JH is one of the factors that mediates temporal polyethism among workers in advanced eusocial insects that have reproductive castes (e.g. Apis bees and Polybiawasps). Therefore, initial observations suggest that JH may have different roles in primitive and advanced eusocial taxa. Here, we use detailed behavioral observations of marked individuals to test whether JH influences temporal polyethism in the primitive eusocial wasp Polistes dominulus. First, we show that workers in P. dominulus have an age-related division of labor, as workers switch from nest work to foraging as they mature. Then, we show that application of JH accelerates the onset of foraging behavior.Workers treated with JH start foraging at a younger age than control workers. Therefore, JH mediates temporal polyethism in the primitively eusocial insect Polistes dominulus. Received 23 April 2008; revised 6 August 2008; accepted 11 August 2008     

Nest Hydrocarbons as Cues for Philopatry in a Paper Wasp

Philopatric behavior has been demonstrated in a wide taxonomic spread of animals. In temperate environments, overwintered Polistes wasp foundresses often return to their natal nest prior to initiating colony construction. Previous research has shown that these spring foundresses can identify the natal nest in the absence of landmark and gross morphological cues. Hydrocarbons are essential recognition cues for Polistes nest and nestmate discrimination, but cuticular hydrocarbon profiles can become homogenized when foundresses overwinter in mixed colony groups. We examined the hydrocarbon profiles of Polistes dominulus foundresses and nests before and after an overwintering period, and found that the hydrocarbon profiles of nests remain unique over time and that this uniqueness is influenced by the original foundresses. Our data raise the possibility that in returning to the natal nest, foundresses reacquire their colony‐specific signature, which may play a role in the formation of cooperative associations.