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According to Jeanne and Downing , ants are the main predators of social wasps that build petioled nests without an envelope. Duration of the colony stage and density of broods and adults. The complete colony cycle of M. This duration is similar to the eight months reported by Litte for M. The mean duration of the pre-emergence stage in M. The egg, larval, and pupal sub-stages had mean durations of The duration of the pre-emergence stage was shorter than the In detail, we observed that the pre-emergence stage in pleometrotic foundations was longer than the haplometrotic.

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In pleometrosis the foundation phase was marked by antagonistic behaviors among the foundresses until the establishment dominance. Our results showed that in the pre-emergence stage, each colony contained 2. In this stage, the nests contained The post-emergence stage comprised most of the colonial cycle, lasting on average In this stage, the colonies had a mean of 6. In this stage, the nests contained a mean of The duration of the post-emergence stage in M.

The post-emergence stage in M. The first individuals produced were always females, precisely workers that were destined for the maintenance and defense of the colony. The few males seen in this study generally emerged at the end of the post-emergence stage, when the colonies contained a high density of adults. In this stage, the few cells that did not contain immatures were located on the periphery of the comb and were used mainly for nectar storage.

Colonial Switzerland

The declining stage had a mean duration of The duration of this stage was less than the In this stage, the colonies contained 5. In addition, the cells that did not contain immatures generally remained empty. As shown by West-Eberhard for Polistes , the declining stage was characterized by the addition of few or no cells to the comb, and an irreversible reduction in the numbers of adults and immatures in the colony, as well as progressive dysfunction of the social organization of the colony before the abandonment of the nest. Foundation patterns and colony success.

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The data this study showed that the foundation of colonies in M. Foundations with five females were unrecorded. The predominance of haplometrotic foundation shown for M. In the pleometrotic foundations, the association of females occurred in most cases after a female had begun the nest. In colonies in which the association occurred after the foundation, the queen was as a rule the founders that began the nest. The queen in the pleometrotic foundation was established mainly by differential oophagies. There was no relationship between the pattern of colony foundation and the success of the colony; contradictorily, haplometrotic foundation was the more common and the less successful.

It could be hypothesized that because of the need for physical combat for establishment of dominance in pleometrotic foundations, the haplometrotic foundation appears to be selected. The haplometrotic foundation could be preferred initially to preserve the physical integrity of the foundress, and to increase the chance of success of the colony.

However, the haplometrotic foundation is more susceptible to the attack of predators Gamboa ; Gamboa et al. Thus, the haplometrotic foundation would be selected only if the predation rate is low in a population Gamboa et al. Therefore, it is suggested that the decision for either haplo- or pleometrosis foundation is closely associated with the surrounding predation rate.

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Although it needs to be further explored, this hypothesis seems to be valid for M. All of the colonies observed were found on human-made structures, especially in locations protected from direct sunlight and rain. Although asbestos tiles were equally abundant in the study location, there was a certain preference for foundation on metal structures and concrete. The low number of foundations on asbestos tiles can be explained partly by the exposure of most of the tiles to the sun, which, depending on the degree of shade, provided unfavorable physical conditions for the maintenance of colonies; all the successful colonies established on tiles were partly protected from direct sunlight.

This suggests that the places selected for nesting are necessarily those that provide better climate conditions. The authors thank Janet W. Downing, H. The function and evolution of exocrine glands, p. Matthews eds. The social biology of wasps. Gadagkar, R. Belonogaster, Mischocyttarus, Parapolybia, and independent founding Ropalidia , p. Gamboa, G. Intraspecific defense: Advantage of social cooperation among paper wasp foundresses.

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Duffy; S. Defence against intraspecific usurpation by paper wasp cofoundresses Polistes fuscutus , Hymenoptera: Vespidae. Canadian Journal of Zoology 70 : Giannotti, E. Biology of the wasp Polistes epicnemius cinerascens Saussure Hymenoptera: Vespidae. The colony cycle of the social wasp, Mischocyttarus cerberus styx Richards, Hymenoptera, Vespidae. Revista Brasileira de Entomologia 41 : Antonialli-Junior I. This study describes some aspects of the colony cycle of the Neotropical social wasp Mischocyttarus consimilis , from data obtained under field conditions.

Our results showed that the colony cycle in M. The colonies remained active for approximately eight months. Most of the abandonments were associated with natural causes, and were most frequent in the pre-emergence stage. The nests were constructed preferentially in man-made structures, especially in sites protected from direct sunlight and rain. Colony foundation was either by haplometrosis or pleometrosis, being the first form predominant. Keywords: Colony development; natural history; neotropical wasp; phenology; Polistinae.

Os resultados desse estudo mostraram que o ciclo colonial em M. In polistine wasps, the periodicity of the foundation and abandonment of colonies differs between species of tropical and temperate climates Gadagkar ; Reeve In temperate areas, the colony cycle is synchronized with the seasons, beginning in the middle of spring and finishing at the end of autumn or beginning of winter West-Eberhard In these conditions there is a period of inactivity that extends from the abandonment of the colonies to the new foundations, during which the wasps hibernate Yoshikawa This period is characterized by the formation of aggregations, which are initially constituted by males and later by potentially reproductive females, which after they abandon the maternal nest, migrate to sheltered locations or old abandoned nests Spradbery In general, the mating of the potentially reproductive females occurs in their own aggregation, and during the winter Yoshikawa ; Spradbery In contrast, the colony cycle of social wasps in tropical areas is relatively longer, occasionally more than a year Giannotti Thus, colonies in different stages of development can occur in the same population in the same period, and this nesting cycle pattern is termed asynchronous Wilson However, under adverse climate conditions e.

In polistine wasps, the colonies are founded either independently or swarms form Jeanne In independent foundations, one or more foundresses unaccompanied by workers begin a new colony, which can occur either haplometrotic or pleometrotic forms Gadagkar Independent-foundation species generally contains few individuals in the colony, and the nests do not possess a covering envelope Gadagkar This foundation pattern can be found in four genera of Polistinae Polistes , Mischocyttarus, Belonogaster, and Parapolybia and in some species of Ropalidia Gadagkar Study site and climate characterization.

The study area is characterized by intense movement of people and sparse vegetation among the buildings. The climate of this region, according to Zavatini , is humid subtropical, with higher precipitation and temperatures in September through February Warm-Rainy Season , and less precipitation and mild temperatures in March through August Cold-Dry Season. The climate data were obtained from the Embrapa meteorological station in the Dourados district.

Colonial phenology. The colonial phenology was studied for one year, during which a total of 94 colonies were observed. The foundations and abandonments of colonies were determined through weekly censuses in the study area, from September through August Most of the colonies were observed from their foundation until they were abandoned. We collected data on established colonies three times a week on designated days.

We considered "foundation" to be the period from the construction of the petiole until the first egg was laid, "activity" the period from the first egg laid until the declining colony stage, and "abandonment" when the last wasp left the nest. The reasons for the abandonment of a colony were assessed, and were assigned to "natural causes" if the nest was complete in the nesting site, "human action" if the nest was totally removed from the nesting site and "predation" if the nest was partly destroyed or if ants were occupying it.

The term "natural causes" was only used for nests that were abandoned for no apparent reason and that were monitored rigorously. Duration of the colony stage. The duration of the colony stage and the densities of adults and immatures were determined by daily monitoring of each colony, from June through July The colony stage and sub-stage were defined according to Jeanne , as "pre-emergence stage" from the foundation of the nest to the emergence of the first imago; "egg sub-stage" from the first egg laid to the appearance of the first larva; "larval sub-stage" from the appearance of the first larva to the formation of the first pupa; "pupal sub-stage" from the formation of the first pupa to the emergence of the first imago; "post-emergence stage" from the emergence of the first imago to the beginning of the colony decline and "declining stage" from the beginning of irreversible reduction of the numbers of immatures to the total abandonment of the nest.

Foundation patterns. The foundation pattern and the colony success were observed for 59 colonies. We determined the pattern of colony foundation considering the "substrate" used to attach the nest and the "number of females" that participated in the foundation stage of the colony. We considered that the colony was a success when it reached at least the post-emergence stage, i. Statistical analysis. Correlation analysis was used to evaluate if climate variables air temperature and relative humidity influenced the numbers of foundations and abandonments.

The variable was considered significant when the p -values were less than 0. Richards mentioned that M. Our data showed that colonies of M. The nesting cycle was asynchronous in respect to the months of the year, and colonies in different stages occurred simultaneously in the population at any time of the year. This regular siting of new colonies by M.

Another factor that favors nesting cycle asynchrony in tropical wasps is the constant presence of males in the population, which allows the fertilization of reproductive females during the entire year Hunt Data from other studies with conspecific tropical wasps, for instance M. These data suggest that constancy of foundation and abandonment of colonies is associated with slight climate variation during the year, leading to the nesting cycle asynchrony. West-Eberhard , studying wasps of temperate regions, observed that the abandonments in those regions were motivated mainly by the negative variation of temperature.

Our data suggest that abandonments in warmer periods may be exceptional for tropical species. In any event, these data taken together strongly support the effect of temperature variation as the main environmental factor associated with colony abandonment in social wasps, in both temperate and tropical regions. Abandonments because of predation were uncommon, but were more frequent in pre-emerging or declining colonies, and occurred when these contained, soon before they were abandoned, a mean of In the abandonments from natural causes, the colonies contained a mean of Considering the colony stage, the abandoned colonies contained on average The abandonments motivated by predation occurred mainly because of attacks by ants.

According to Jeanne and Downing , ants are the main predators of social wasps that build petioled nests without an envelope. Duration of the colony stage and density of broods and adults. The complete colony cycle of M. Allocator - an allocator that is used to acquire memory to store the elements. The type must meet the requirements of Allocator. This type is used to form the skipfield which skips over erased T elements. The maximum size of element memory blocks is constrained by this type's bit-depth due to the nature of a jump-counting skipfield.

The default type, unsigned short , is bit on most platforms which constrains the size of individual memory blocks to a maximum of elements. In the case of small collections eg. In the case of very large collections millions where memory usage is not a concern and where erasure is less common, one might think that changing the skipfield bitdepth to a larger type may lead to slightly increased iteration performance due to the larger memory block sizes made possible by the largerbit depth and the fewer subsequent block transitions.

However in practice the performance benefits appear to be little-to-none, and introduce substantial performance issues if erasures are frequent, as with this approach it takes more erasures for a group to become empty - increasing the frequency of large skips between elements and subsequent cache misses, as well as skipfield update times. From this we can assume it is unlikely for a user to want to use types other than unsigned short and unsigned char. But since these scenarios are on a per-case basis, it has been considered best to leave control in the hands of the user.

If prior erasures have occurred, updating the skipfield may require a memmove operation, which creates a variable time complexity depending on the range of skipfield needing to be copied though in practice this will resolve to a singular raw memory block copy in most scenarios, and the performance impact is negligible. Average time complexity varies based on erasure pattern. With a random erasure pattern it will be closer to O 1 amortized.

Insert multiple : O N unless prior erasures have occurred. See Insertion single for rules in this case. Erase single : If erasures to elements consecutive with the element being erased have not occurred, or only consecutive erasures before the element being erased have occurred, O 1 amortised. If consecutive erasures after the element being erased have occurred, updating of the skipfield requires a memmove operation or vectorized update of O N complexity, where n is the number of consecutive erased elements after the element being erased.

Average time complexity varies based on erasure pattern, but with a random erasure pattern it's closer to O 1 amortized. Colony iterators cannot be random access due to the use of a skipfield. But member overloads for the standard library functions advance , next , prev and distance are available in the reference implementation, and are significantly faster than O N in the majority of scenarios. Thank you's to Glen Fernandes and Ion Gaztanaga for restructuring advice, Robert Ramey for documentation advice, various Boost and SG14 members for support, Baptiste Wicht for teaching me how to construct decent benchmarks, Jonathan Wakely for standards-compliance advice and critiques, Sean Middleditch, Patrice Roy and Guy Davidson for critiques, support and bug reports, Jason Turner and Phil Williams for cross-processor testing, that guy from Lionhead for annoying me enough to get me to get around to implementing the skipfield pattern, Jon Blow for some initial advice and Mike Acton for some influence.

Default constructor, but using a custom memory allocator eg. Setting the group sizes can be a performance advantage if you know in advance roughly how many objects are likely to be stored in your colony long-term - or at least the rough scale of storage. If that case, using this can stop many small initial groups being allocated.

Using an initialiser list to insert into the colony upon construction. Copy all contents from source colony, removes any empty erased element locations in the process. Size of groups created is either the total size of the source colony, or the maximum group size of the source colony, whichever is the smaller. Move all contents from source colony, does not remove any erased element locations or alter any of the source group sizes.

Source colony is now void of contents and can be safely destructed. All operators have O 1 amortised time-complexity. These have been transplanted to colony's advance , next , prev and distance member functions. Inserts the element supplied to the colony, using the object's copy-constructor. Will insert the element into a previously erased element slot if one exists, otherwise will insert to back of colony. Returns iterator to location of inserted element.


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Moves the element supplied to the colony, using the object's move-constructor. Will insert the element in a previously erased element slot if one exists, otherwise will insert to back of colony. Inserts n copies of val into the colony. Stops inserting once it reaches last. Copies elements from an initializer list into the colony. Removes the element pointed to by the supplied iterator, from the colony. Returns an iterator pointing to the next non-erased element in the colony or to end if no more elements are available. This must return an iterator because if a colony group becomes entirely empty, it may be removed from the colony, invalidating the existing iterator.

A group may either be removed when it becomes empty, or moved to the back of the colony for future insertions and made inactive. The decision to either remove or move should be largely implementation-defined, but testing has suggested that the best performance under high-modification occurs when groups are removed unless they meet the maximum group size, or are either of the last two active groups at the back of the colony. The reference implementation currently removes all groups when empty. Erases all contents of a given colony from first to the element before the last iterator.

Constructs new element directly within colony. Returns a boolean indicating whether the colony is currently empty of elements. Returns total number of elements currently stored in container. Returns the maximum number of elements that the allocator can store in the container. This is an approximation as it does attempt to measure the memory overhead of the container's internal memory structures. It is not possible to measure the latter because a copy operation may change the number of groups utilized for the same amount of elements, if the maximum or minimum group sizes are different in the source container.

Returns total number of elements currently able to be stored in container without expansion. Deallocates any unused groups retained during erase or unused since the last reserve.

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This function cannot invalidate iterators or pointers to elements, nor will it necessarily shrink the capacity to match size. In the implementation the maximum size for this number currently is limited to the maximum group size of the colony and will be truncated if necessary. This restriction will be lifted in a future version. This function is useful from a performance perspective when the user is inserting elements singly, but the overall number of insertions is known in advance.

By reserving, colony can forgo creating many smaller memory block allocations due to colony's growth factor and reserve a single memory block instead.

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Changes the minimum and maximum internal group sizes, in terms of number of elements stored per group. Changes the minimum internal group size only, in terms of minimum number of elements stored per group. Changes the maximum internal group size only, in terms of maximum number of elements stored per group. After the splice, the source colony is empty. Splicing is much faster than range-moving or copying all elements from one colony to another.

Note2: If the minimum group size of the source is smaller than the destination, the destination will change it's minimum group size to match the source. The same applies for maximum group sizes if source's is larger, the destination will adjust its. Sort the content of the colony. By default this compares the colony content using a less-than operator, unless the user supplies a comparison function ie. Append data from one colony container to the end of another.

This operation removes all elements from the source colony.