The Intricate World of Belonogaster petiolata
Delving into the fascinating social structures, behaviors, and life cycles of a primitively eusocial African wasp.
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Taxonomy and Phylogeny
Kingdom: Animalia
Belonogaster petiolata belongs to the animal kingdom, a vast group encompassing multicellular organisms that are heterotrophic and typically mobile.
Phylum: Arthropoda
As an arthropod, it possesses an exoskeleton, a segmented body, and jointed appendages, characteristic features shared with insects, spiders, and crustaceans.
Class: Insecta
Within the insect class, it exhibits the typical insect body plan: head, thorax, and abdomen, along with three pairs of legs and usually wings.
Order: Hymenoptera
This order includes ants, bees, wasps, and sawflies. Hymenoptera are known for their diverse social structures and often possess specialized stinging or egg-laying organs.
Family: Vespidae
Belonogaster petiolata is part of the Vespidae family, which comprises social wasps. This family displays a wide range of social organization, from solitary to highly eusocial species.
Subfamily: Polistinae
This subfamily includes paper wasps. They are known for constructing nests from chewed wood pulp and often exhibit complex social behaviors.
Tribe: Ropalidiini
A specific grouping within the Polistinae, characterized by certain morphological and behavioral traits.
Genus: Belonogaster
This genus contains primitively eusocial wasps. Species within Belonogaster share many characteristics, including social organization and nesting habits.
Species: B. petiolata
The specific species, Belonogaster petiolata, is closely related to species like B. juncea and B. grisea, exhibiting similar patterns in offspring production and queen development.
Description and Identification
Physical Characteristics
These wasps feature a distinct thorax and a elongated, striped gaster (abdomen) equipped with a sting. The gaster's first segment, the peduncle, connects it to the thorax. Legs are attached to the thorax.
Queen vs. Worker Morphology
A notable difference exists between queens and workers: queens possess a larger gaster and a smaller head, while workers have relatively larger heads and smaller gasters. Reproductive castes (queens, males) also have larger fat reserves for overwintering compared to workers.
Nest Construction
B. petiolata constructs paper nests from chewed pulp, often suspended in sheltered locations like caves. Wasps gather pulp from old cells or immature larvae. The nest structure includes combs, where the queen may reside or rest.
Distribution and Habitat
Geographic Range
This species is found in southern Africa, with a significant presence in South Africa. It has also been observed in areas like northern Johannesburg.
Climate Preferences
They thrive in temperate or moist subhumid climate zones. Preferred temperatures range from a January maximum of 25°C (77°F) and minimum of 14.9°C (58.8°F) to a June maximum of 15.7°C (60.3°F) and minimum of 5.3°C (41.5°F).
Preferred Environment
B. petiolata primarily resides in sheltered environments, most notably caves. The Sterkfontein Caves in South Africa host large populations. Nests are typically built hanging from cave walls.
Conservation Status
The species is abundant in southern Africa and is not considered threatened.
Colony Cycle
Nest Initiation
The annual nesting cycle begins around August. Overwintered foundress females initiate new nests in the spring. This overwintering period lasts approximately 4.5 months.
Foundress Associations
In about half of cases, additional females join the initial foundress, forming multiple-foundress colonies. Associations can include up to 16 foundresses. A dominance hierarchy forms, with one becoming the queen.
Colony Phases
The cycle includes a pre-emergence period (before first offspring), a post-emergence period (after offspring appear), a pre-matrifilial phase (subordinates present), a matrifilial phase (only queen and workers), and finally a reproductive phase (gynes and males born).
Colony Decline
Factors contributing to colony decline include a cessation of solid food supply during the reproductive phase, a decreasing worker-to-larva ratio, and adult priority for food over brood. This can lead to brood abortion and eventual abandonment of the nest.
Behavior
Dominance Hierarchy
A clear hierarchy exists. The dominant female becomes the queen, responsible for egg-laying. Workers perform most tasks: nest maintenance, foraging, and feeding larvae. Workers typically have undeveloped ovaries in the presence of a queen.
Interactions and Communication
Queens assert dominance through biting, hooking (grasping subordinates to receive regurgitated food), and soliciting (probing with antennae). Subordinates may ignore, submit, or solicit back. Dominance behaviors are usually directed downwards in the hierarchy.
Oophagy and Oviposition
Females may consume eggs, often those of subordinates (oophagy). Queens lay the majority of eggs in pre-emergence colonies. Queens also inspect eggs to identify those laid by subordinates.
Trophallaxis
Wasps engage in trophallaxis with larvae to obtain nutrient-rich salivary fluid. This involves soliciting larvae by antennating their heads and vibrating their bodies. Older larvae tend to provide more saliva.
Kin Selection and Castes
Caste System
Distinct worker and reproductive castes exist. Foundresses and gynes (reproductive females) are generally larger than workers. While workers start smaller, they grow, but reproductive castes maintain larger ovaries and fat reserves.
Winter Survival
A wasp's ability to survive winter is linked to its caste. Gynes and foundresses possess greater fat reserves, enabling them to endure colder periods, unlike most workers.
Queen Removal
If a queen dies, a dominant alpha worker or a beta-foundress may assume leadership. Queen loss can trigger ovary development in some workers, though this is less common than continued dominance by a replacement foundress.
Reproduction
Copulation
Mating involves mutual antennation, followed by the male mounting the female. Genital contact is established briefly, with copulation lasting only about five seconds.
Ovary Development
Ovarian width increases significantly in gynes during the overwintering period and leading up to spring. While workers have immature ovaries, they can develop if the queen is absent. Inseminated females are more likely to lay eggs.
Insemination Status
Workers are rarely inseminated. Foundresses and gynes, particularly those overwintering, show high rates of insemination. This is often a criterion for distinguishing reproductive females from workers.
Mate Recognition
Female wasps possess mate recognition signals, likely pheromones, within their venom, thorax, and head. These signals, spread through grooming, attract males.
Diet
Primary Food Sources
B. petiolata consumes both solid food, primarily small insects like caterpillars, and liquid nectar. They masticate solid food to feed themselves, others, and the young brood.
Water Requirement
Water is essential for survival, alongside nectar and solid food.
Food Allocation
Queens receive a disproportionately larger share of both liquid and solid food loads. They also retain a significant portion of the solid food for themselves rather than passing it all to larvae.
Seasonal Changes and Prey Availability
In late summer, reduced prey abundance, often due to poor rainfall, leads wasps to rely more heavily on nectar. A consistent supply of solid food is crucial for colony health.
Growth and Development
Early Development
Newly emerged wasps have black or brown eyes. As they mature, their eyes turn pink, and they develop wings. Young workers often huddle together for protection from aggression.
Body Size and Caste
Body size increases from January to March, particularly for workers born later in the season. The amount of food provided during larval stages influences caste determination and final body size.
Worker-to-Larva Ratio
A higher worker-to-larva ratio, observed in November and December, ensures sufficient feeding for developing gynes (the highest caste), indicating that adequate nutrition is key to caste differentiation.
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References
References
- Genus Belonogaster de Saussure. Catalog of Species in the Polistine Tribe Ropalidiini (Hymenoptera: Vespidae).
- Keeping, Malcolm G. "Social Behavior and Brood Decline in Reproductive-phase Colonies Of Belonogaster Petiolata (Degeer) (Hymenoptera: Vespidae)." Journal of Insect Behavior 10.2 (1997): 265-78. Web. 20 Sept. 2014.
- Keeping, Malcolm G. "Social Organization and Division of Labour in Colonies of the Polistine Wasp, Belonogaster Petiolata." Behavioral Ecology and Sociobiology 31.3 (1992): n. pag. Web. 20 Sept. 2014.
- "Animal Diversity Web." ADW: Belonogaster Petiolata: CLASSIFICATION. N.p., n.d. Web. 20 Sept. 2014.
- Keeping, Malcolm G. "Absence of Chemical Alarm in a Primitively Eusocial Wasp (Belonogaster Petiolata, Hymenoptera: Vespidae)." Insectes Sociaux 42.3 (1995): 317–20. Web. 20 Sept. 2014.
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