Apiculture: Honey Bee Diversity

 

A World of Diversity: Types and Characteristics of Honey Bees

Honey bees are distinguished by their construction of perennial, hexagonally celled nests made of secreted wax, their large colony sizes, and their unique method of storing digested carbohydrates as surplus food in the form of honey. A typical colony can range from several thousand to 60,000 workers, a single queen, and hundreds of drones, all clustered over wax combs used for food storage and rearing young. Their activities are highly seasonal, adapting to changes in temperature and food availability, from intensive brood rearing in spring to tight clustering for warmth in winter.  

While Apis mellifera, the Western Honey Bee, is the most common and commercially significant species globally, the genus Apis encompasses several other fascinating species, each with unique adaptations and geographical distributions.  

• The Western Honey Bee (Apis mellifera)

The Western Honey Bee (Apis mellifera) is the most widely distributed honey bee species, commonly kept by residential beekeepers due to its prolific honey production, typically ranging from 50 to 200 pounds annually, depending on environmental conditions and floral resources. These bees possess robust, hairy bodies with diverse color patterns and form large colonies, often comprising 20,000 to 80,000 individuals. Their role in agriculture is indispensable, as they are crucial for crop pollination, contributing to approximately one-third of the global food supply. Within the hive, they communicate vital information about food sources through complex dances, most notably the "waggle dance".  

Honey bees in the United States represent a heterogeneous blend of various races introduced from Europe, the Middle East, and Africa. Prominent among these are:  

• Italians: Introduced in 1859, Italian bees are the most popular race in the United States. They are characterized by their light yellowish or brown coloration with alternating stripes on the abdomen. They are known for starting brood rearing early in the spring and continuing late into the fall, which results in large populations throughout the active season. A significant advantage is their comparative resistance to European foulbrood (EFB), a major reason they replaced the original black or German bees.  
• Caucasians: Often regarded as the gentlest of all honey bees, Caucasians are dark-colored to black with grayish abdominal bands. They tend to construct burr comb and utilize large amounts of propolis to secure combs and reduce hive entrance size, though newer strains show less propolis use. While inclined to drifting and robbing, they are not prone to excessive swarming. Their colonies typically reach full strength later in the season, around midsummer, and they are noted for conserving honey stores more effectively than Italians.  
• Carniolans: These are dark bees, similar in appearance to Caucasians, but often exhibit brown spots or bands on their abdomen. Carniolan colonies overwinter as small clusters but experience rapid population increases in the spring once the first pollen becomes available. Their primary disadvantage is a strong inclination towards excessive swarming.  
• Hybrids: Commercial hybrids, such as Midnite and Starline, are developed by crossing inbred lines specifically for desirable characteristics like gentleness, productivity, or improved wintering capabilities. Buckfast bees, for instance, are a hybrid meticulously selected over a long period from various strains found in southwestern England. Other genetic stocks, including Russian, SMR (Suppressed Mite Reproduction), or Minnesota hybrids, have been selectively bred for enhanced mite resistance and/or improved hygienic behavior, which involves the efficient removal of dead or dying brood to prevent pathogen spread.  

The classification of Apis mellifera is notably complex, with 33 recognized subspecies distributed across Africa, Western Asia, and Europe, categorized into five evolutionary lineages. Identification relies on diverse methods, including morphometry (measuring body parts and analyzing wing shapes), allozyme-based techniques, and advanced mitochondrial and nuclear DNA analysis. The presence of transition zones between subspecies ranges results in gradual changes in characteristics, making precise discrimination challenging. The existence of numerous  

Apis mellifera subspecies, each with distinct traits like disease resistance or propolis use, highlights significant evolutionary pressures for local adaptation across diverse climates and ecological niches. This natural selection has been further shaped by human intervention, as evidenced by the development of hybrids specifically bred for traits like gentleness or mite resistance. The complex and sometimes contradictory taxonomy of  

A. mellifera, marked by transition zones where characteristics gradually blend, underscores a dynamic genetic landscape influenced by both natural gene flow and human-mediated introductions. This continuous evolution means that effective beekeeping practices must adapt to the specific traits of these diverse populations to ensure their health and productivity, especially in the face of emerging threats like mites.  

• The Asian Honey Bee (Apis cerana)

The Asian Honey Bee (Apis cerana) is smaller than A. mellifera and has remarkably adapted to warmer, tropical, and subtropical climates across Southeast Asia, India, and southern China. A key characteristic is their strong resistance to pests like Varroa mites, a significant challenge for Western honey bees. Their colonies are generally smaller, typically housing between 10,000 and 30,000 bees, and produce less honey compared to  

A. mellifera, usually yielding 20-50 pounds per year.  

Apis cerana colonies are known for constructing multi-comb nests within cavities that feature small entrances, presumably as a defense mechanism against invaders. They exhibit precise internal hive thermoregulation, maintaining optimal temperatures through collective fanning and water evaporation. A particularly fascinating defense is their unique "thermal defense" against invading hornets, where a cluster of bees vibrates their flight muscles to collectively raise the temperature around the hornet, heating it to death while staying within their own lethal limits. The remarkable resistance of  

Apis cerana to Varroa mites and its unique thermal defense against hornet invaders are not isolated phenomena. Instead, these represent sophisticated adaptations resulting from a long history of co-evolution with specific threats in their native Asian environments. Unlike its Western counterpart, which often struggles with Varroa,  

A. cerana has developed robust coping mechanisms. The collective thermal defense, where the colony acts as a unified entity to generate lethal heat for a predator, exemplifies a highly advanced form of social immunity. This demonstrates that the social organization of these bees extends beyond mere division of labor to encompass powerful collective defense strategies that significantly enhance species survival against formidable predators. Understanding these co-evolved defenses could offer invaluable pathways for developing more sustainable pest management for  

A. mellifera, perhaps through selective breeding for similar hygienic behaviors or by unraveling the genetic underpinnings of such resistance.

• The Giant Honey Bee (Apis dorsata)

The Giant Honey Bee (Apis dorsata) is the largest honey bee species, typically measuring 17-20 mm in length. They are renowned for building massive single, open combs that can reach up to 1.5 meters in width and 1 meter in depth, often in exposed locations high off the ground, such as on thick tree limbs, under cliff overhangs, or on buildings. Colonies can be exceptionally large, frequently exceeding 50,000 individuals, and are capable of producing 60-100 pounds of honey annually. These bees are known for their highly aggressive defensive strategies and vicious behavior when disturbed, with up to three-quarters of the colony forming a thick, protective curtain around the nest. Primarily tropical,  

A. dorsata colonies migrate seasonally, sometimes traveling up to 200 km to follow wet and dry seasons and ensure continuous access to floral resources. All species of giant honey bees are parasitized by  

Tropilaelaps mites, some strains of which can also parasitize Apis mellifera, posing a potential threat to Western honey bee populations. The impressive size, aggressive defensive posture, and preference for exposed nesting sites of  

Apis dorsata appear to be part of an intricate evolutionary trade-off. While open nests might seem vulnerable, the sheer scale of their colonies and the formation of a dense, aggressive 'curtain' of bees around the nest likely serve as a formidable deterrent to many predators. Their extensive seasonal migrations, sometimes covering hundreds of kilometers, are a direct response to the fluctuating availability of floral resources in tropical regions, ensuring a continuous supply of nectar and pollen. This migratory behavior illustrates that the collective functioning of these bees transcends the confines of a single hive, involving large-scale, coordinated movements across landscapes, all driven by environmental cues and the need for sustenance. The presence of  

Tropilaelaps mites, which can also affect A. mellifera, further highlights the interconnectedness of bee health across different species and geographical areas. The ability of  

A. dorsata to return to the same nesting sites year after year, even with a complete turnover of individual bees, suggests a remarkable collective memory and navigational capacity, offering a compelling area for further study into collective intelligence and ecological adaptation.

• The Dwarf Honey Bee (Apis florea)

Apis florea, also known as the dwarf honey bee, is the smallest honey bee species, with workers typically measuring 7-10 mm in body length and exhibiting a distinctive red-brown coloration. They construct a single, exposed comb, usually on small tree branches or shrubs. Colonies are relatively small, generally consisting of 5,000-10,000 bees, and produce limited honey, averaging 10-20 pounds annually. Unlike other  

Apis species, A. florea foragers perform their waggle dance on the horizontal upper surface of the comb, directly pointing to the food source. They display aggressive behaviors when competing for territory, compensating for their small size and limited flight range. Their social defense mechanisms include hissing and shimmering behaviors, and they often nest amidst dense foliage for camouflage. Uniquely, they salvage wax from old nests when building new ones nearby and apply sticky propolis barriers around their nests to deter ants. Drones of this species possess a distinctive thumb-like bifurcation called the basitarsus on their hind leg. Despite being considered one of the most primitive living  

Apis species, characterized by its smaller colony size and simpler nest construction, Apis florea exhibits a surprising array of complex social behaviors. These include sophisticated worker policing mechanisms, the unique practice of salvaging wax from old nests, and specialized defenses such as hissing, shimmering, and deploying sticky barriers against ants. The distinctive horizontal waggle dance, which directly points to food sources rather than relying on gravity as in  

A. mellifera, appears to be an adaptation to their exposed, single-comb nests, simplifying communication in their specific environment. Furthermore, their aggressive territorial behavior, compensating for a more limited flight range and smaller size, suggests a strategy focused on defending localized resources rather than engaging in extensive foraging over vast areas. This demonstrates that even in species considered 'primitive' within the eusocial spectrum, highly sophisticated adaptations for survival and resource management have evolved, precisely tailored to their unique environmental pressures.  

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Table: Comparative Characteristics of Key Honey Bee Species

Feature	Apis mellifera (Western Honey Bee)	Apis cerana (Asian Honey Bee)	Apis dorsata (Giant Honey Bee)	Apis florea (Dwarf Honey Bee)
Size	Robust body	Smaller, slender	Largest (17-20 mm)	Smallest (7-10 mm), slender
Coloration	Dark brown to golden yellow	Yellow or brown, black with 4 yellow stripes	Golden, black, pale bands	Red-brown
Colony Size	20,000 - 80,000 bees	10,000 - 30,000 bees	>50,000 bees	5,000 - 10,000 bees
Nest Type	Perennial, multi-comb in cavities	Multi-comb in cavities	Large, single open comb	Small, single exposed comb
Nest Location	Hives, tree cavities	Tree cavities, man-made structures	High tree branches, cliffs, buildings	Small tree branches, shrubs
Honey Production	High (50-200 lbs/yr)	Moderate (20-50 lbs/yr)	High (60-100 lbs/yr)	Limited (10-20 lbs/yr)
Aggression	Varies by subspecies, can be aggressive	Flexible defense, thermal defense	Highly aggressive	Aggressive for territory
Pest Resistance	Susceptible to Varroa mites	Strong resistance to Varroa mites	Parasitized by Tropilaelaps mites	-
Communication	Waggle dance (gravity-oriented)	Waggle dance	Waggle dance	Waggle dance (horizontal)
Geographical Distribution	Europe, N. America, Africa, Asia	Southeast Asia, India, S. China	South & Southeast Asia	South & Southeast Asia

This comparative table is valuable because it offers a concise, at-a-glance summary of the key features of different honey bee species. By presenting data points such as size, coloration, colony size, nest type, honey production, aggression, pest resistance, communication, and geographical distribution in a structured format, readers can quickly compare and contrast the unique adaptations and ecological roles of each species. This visual aid simplifies complex information, making the report more accessible and enhancing reader comprehension and retention.

References

Blythewood Bee Company. (n.d.). Honey bee types. Retrieved from https://blythewoodbeecompany.com/blogs/news/honey-bee-types

Delaware Cooperative Extension. (n.d.). Honey bee biology. MAAREC. Retrieved from https://canr.udel.edu/maarec/honey-bee-biology/

Idaho Fish and Game. (n.d.). Apis mellifera (Western Honey Bee). Retrieved from https://idfg.idaho.gov/species/taxa/23435

National Center for Biotechnology Information. (2020). Apis mellifera subspecies characteristics. PMC. Retrieved from https://pmc.ncbi.nlm.nih.gov/articles/PMC7714978/

National Institute of Food and Agriculture (NIFA). (n.d.). The social organization of honey bees. EDIS. Retrieved from https://edis.ifas.ufl.edu/publication/IN1102

Planet Bee Foundation. (n.d.). The three castes of honey bees and their vital roles in the hive. Retrieved from https://www.planetbee.org/post/the-three-castes-of-honey-bees-and-their-vital-roles-in-the-hive

Reddit. (n.d.). Does different type of bees make different honey?. Retrieved from https://www.reddit.com/r/askscience/comments/7qc47s/does_different_type_of_bees_make_different_honey/

The Holy Habibee. (n.d.). The superorganism model of social insects. Retrieved from https://theholyhabibee.com/superorganism-model/

University of California, Berkeley. (n.d.). Native bees vs. honey bees: There are big differences. Penasquitos.org. Retrieved from https://www.penasquitos.org/news/native-bees-vs-honey-bees-there-are-big-differences

Wikipedia. (n.d.). Apis cerana. Retrieved from https://en.wikipedia.org/wiki/Apis_cerana

Wikipedia. (n.d.). Apis dorsata. Retrieved from https://en.wikipedia.org/wiki/Apis_dorsata

Wikipedia. (n.d.). Apis florea. Retrieved from https://en.wikipedia.org/wiki/Apis_florea

Wikipedia. (n.d.). Honey bee. Retrieved from https://en.wikipedia.org/wiki/Honey_bee

YourDictionary. (n.d.). Eusociality. EBSCO. Retrieved from https://www.ebsco.com/research-starters/biology/eusociality

The Honeybee; a special Bee

Beyond the Hive: How Honey Bees Differ from Other Bees

 

The world of bees is incredibly diverse, with over 20,000 species worldwide. While all bees play vital roles as pollinators, honeybees (genus Apis) stand out due to several key distinctions, primarily their highly social organization and the scale of their honey production.

 Honeybees are different to other bees popularly as Polen bee

 Social Organization: Honeybees are highly social insects living in large, perennial colonies within hives, led by a single queen. This complex social structure is a hallmark of their species, characterized by distinct castes performing specialized roles. In contrast, most native bees, such as sweat bees, are solitary, meaning they do not have a queen or live in colonies. Some, like bumble bees, exhibit a less complex social structure, forming smaller, annual colonies. These solitary bees typically nest in the ground, dead wood, brush piles, or hollow flower stems.  

 Honey Production: Honey bees are unique in their ability to produce and store large amounts of honey in honeycombs, making it readily available for human consumption. This surplus storage is a defining characteristic of the Apis genus. While some other bees, like bumble bees and stingless bees found primarily in tropical regions, do produce small amounts of honey, it is generally not in volumes comparable to honey bees and often not easily accessible for human consumption. Carpenter bees and mason bees, for instance, do not produce honey at all. It is also important to note that the composition and taste of honey can vary significantly even among different Apis species or subspecies, largely influenced by their foraging preferences and the specific nectar sources available.  

 

Aggression and Stinging Behavior: Honey bees are known to be aggressive, particularly when defending their queen and hive. A worker honey bee's sting is barbed, meaning it embeds itself into the sting site, and the sting apparatus continues to deliver venom even after detachment, ultimately leading to the bee's death. This self-sacrificial defense is a testament to their highly social nature, where individual survival is secondary to colony protection. In contrast, most native bees are generally not aggressive. Male native bees typically lack stingers entirely, and females will only sting if their nest is directly threatened.  

 

Pollination Behavior: Honey bees primarily seek nectar, collecting pollen on their bodies and then brushing it into specialized pollen baskets (corbiculae) on their hind legs to transport back to the hive. Pollination occurs when these pollen baskets brush against the flower's stigma. They are widely used for crop pollination, with hives often transported for large-scale agricultural operations, such as for almond crops in Central California. Interestingly, native bees are often more efficient pollinators on an individual basis, capable of pollinating 60-100% more flowers. They tend to "wallow" around in flowers, getting completely covered in pollen, which then transfers to the next flower from any part of their body. It is estimated that native bees pollinate a significant 80% of flowering plants and 75% of the fruits, nuts, and vegetables grown in the U.S..  

 

Origin in North America: Honey bees (Apis mellifera) were introduced to North America from Europe in the 17th century. Consequently, all honey bees currently found in the U.S. are either imported or descendants of these imports. Conversely, many other bee species, such as bumble bees, are native to the U.S..  

The pronounced differences between honey bees and other bee species, particularly in their social complexity and capacity for large-scale honey production, underscore a significant evolutionary divergence. While many bee species thrive through solitary or primitively social lifestyles, optimizing individual reproduction, honey bees have specialized in a highly cooperative, colony-centric strategy. This strategic shift has driven the evolution of intricate communication systems, the ability to store vast quantities of resources, and advanced collective defense mechanisms. It is noteworthy that many native bees are individually more efficient pollinators than honey bees. This suggests that the immense ecological impact of honey bees, particularly in agriculture, stems not from individual efficiency but from their sheer numbers and the widespread management of their colonies. This indicates that honey bees occupy a distinct ecological niche, leveraging collective action to ensure survival and resource monopolization, a strategy that has proven exceptionally successful and valuable to human endeavors. The substantial reliance on introduced honey bees for large-scale agricultural pollination, while beneficial, also highlights a potential vulnerability if these populations face declines. This further emphasizes the critical importance of conserving diverse native bee populations, whose greater individual pollination efficiency contributes significantly to ecosystem resilience and food security, advocating for a balanced approach rather than an over-reliance on a single, introduced species.  

 The Hive's Heartbeat: Social Structure and Roles

At the core of a honey bee colony's extraordinary success lies its highly organized social structure, characterized by a strict division of labor among three distinct castes: the queen, the workers, and the drones. Each caste performs specific, vital roles that contribute to the collective survival and prosperity of the hive.  

 

The Queen: The Heart of the Hive

The queen is the sole reproductive female in the colony, and her primary function is to lay eggs, ensuring the continuity and growth of the colony. During peak season, she can lay an astonishing number of eggs, up to 2,000 per day. Beyond her reproductive duties, the queen plays a critical role in regulating the behavior and social order of the hive by emitting powerful pheromones. These chemical signals maintain harmony and cohesion among its members and also influence worker behavior, notably suppressing their reproductive capabilities. Worker bees are highly attentive to the queen's needs, providing her with a special diet of royal jelly and diligently protecting her. Her health and productivity are absolutely crucial for the colony's survival. Queens have a significantly longer lifespan than other bees, often living for several years. A remarkable aspect of her reproduction is her ability to choose whether to fertilize an egg; fertilized eggs develop into diploid females (either queens or workers), while unfertilized eggs develop into haploid males (drones). 

 

The Workers: The Backbone of the Colony

Worker bees are non-reproductive females and constitute the vast majority of the hive's population. They are responsible for nearly all tasks that keep the colony running smoothly, exhibiting a fascinating progression of duties based on their age, a phenomenon known as temporal polyethism.  

Ø  Early Life (Nurse Bees): In their early days, young worker bees serve as "nurse bees." Their primary responsibilities include cleaning the hive, tending to the queen, and feeding developing larvae with a nutrient-rich diet of royal jelly, honey, and pollen.  

Ø  Middle Life (Hive Maintenance): As worker bees age and their royal jelly-producing glands begin to atrophy, they transition to other vital duties within the hive. These tasks include building new comb cells from secreted wax, receiving nectar and pollen from returning foragers, storing food, and guarding the hive entrance against intruders.        

Ø  Late Life (Foragers): In the final stage of their lives, workers become foragers. They venture out of the hive to collect essential resources such as nectar, pollen, and water, which are crucial for the colony's sustenance and survival.  

Ø  Defense: Workers play a critical role in defending the hive from threats. They possess a barbed stinger, and while using it is an effective defense mechanism, it results in their death as the sting apparatus detaches from their body. This self-sacrificial act underscores the collective nature of the honey bee colony.  

 

 

  The Drones: The Males of the Hive

Drones are the male bees of the hive, and their sole purpose is reproduction: to mate with a virgin queen from another colony. They are haploid, meaning they develop from unfertilized eggs laid by the queen or, rarely, by an unfertilized laying worker. Drones are physically larger than workers, characterized by their large eyes that nearly meet at the top of their head and a blunt, thick abdomen. Unlike queens and workers, drones do not possess stingers, nor do they participate in any daily hive tasks such as foraging, cleaning, or brood care; their time is primarily spent eating and waiting for opportunities to fulfill their reproductive role. Mating occurs during "nuptial flights," where drones gather in specific congregation areas in the sky. After a successful mating, the drone dies. Drones that do not get the chance to mate are eventually expelled from the hive as winter approaches, a process known as "drone eviction." This is a necessary measure to conserve valuable resources, as drones do not contribute to the hive's winter survival and would otherwise consume precious honey stores.  
The strict division of labor and highly specialized roles within the honey bee colony illustrate a profoundly evolved system of resource allocation. The queen's exclusive role in reproduction ensures the genetic continuity of the colony. Workers, being non-reproductive, dedicate their entire lives to supporting the colony's growth, maintenance, and defense, progressing through their tasks in a predictable, age-based sequence. Drones, though seemingly unproductive within the hive, fulfill the crucial role of ensuring genetic diversity through mating with queens from other colonies. The seemingly harsh practice of expelling drones before winter is, in fact, an efficient resource management strategy, prioritizing the survival of the reproductive and working castes when food resources are scarce. This intricate, interdependent system demonstrates that the survival and prosperity of the colony as a whole take precedence, even at the cost of individual lives or reproductive potential. While this system confers incredible efficiency, it also introduces a vulnerability: if a key caste, particularly the queen, is compromised, the entire colony can be significantly set back. However, the colony possesses a crucial adaptive mechanism—the ability to 'supersede' or replace a failing queen—which mitigates this vulnerability and showcases the remarkable resilience of the colony as a unified entity.  

 

Table: Roles and Characteristics of Honey Bee Castes

Caste	Sex/Reproductive Status	Primary Role(s)	Key Characteristics
Queen	Female / Reproductive	Lay eggs (up to 2,000/day); emit pheromones to regulate colony	Largest bee in hive; long, plump abdomen; fed royal jelly; lives several years; mates with multiple drones from other colonies
Worker	Female / Non-reproductive	Brood care (nurse bee); hive cleaning & maintenance; wax production; food storage; foraging for nectar, pollen, water; hive defense	Smallest bee; specialized body for pollen/nectar collection (pollen press); barbed stinger (dies after stinging); progresses through tasks by age
Drone	Male / Reproductive	Mate with virgin queen from other colonies	Larger than workers; large eyes that meet at top of head; blunt abdomen; no stinger; does not forage or perform hive duties; dies after mating or expelled before winter

 
This table serves as an excellent reference, clearly outlining the distinct roles and physical characteristics of each honey bee caste. It allows for quick comprehension of the division of labor within a honey bee colony, highlighting how each specialized member contributes to the overall functioning and survival of the hive.

 

References
Blythewood Bee Company. (n.d.). Honey bee types. Retrieved from https://blythewoodbeecompany.com/blogs/news/honey-bee-types
Delaware Cooperative Extension. (n.d.). Honey bee biology. MAAREC. Retrieved from https://canr.udel.edu/maarec/honey-bee-biology/
Idaho Fish and Game. (n.d.). Apis mellifera (Western Honey Bee). Retrieved from https://idfg.idaho.gov/species/taxa/23435
National Center for Biotechnology Information. (2020). Apis mellifera subspecies characteristics. PMC. Retrieved from https://pmc.ncbi.nlm.nih.gov/articles/PMC7714978/
National Institute of Food and Agriculture (NIFA). (n.d.). The social organization of honey bees. EDIS. Retrieved from https://edis.ifas.ufl.edu/publication/IN1102
Planet Bee Foundation. (n.d.). The three castes of honey bees and their vital roles in the hive. Retrieved from https://www.planetbee.org/post/the-three-castes-of-honey-bees-and-their-vital-roles-in-the-hive
Reddit. (n.d.). Does different type of bees make different honey?. Retrieved from https://www.reddit.com/r/askscience/comments/7qc47s/does_different_type_of_bees_make_different_honey/
The Holy Habibee. (n.d.). The superorganism model of social insects. Retrieved from https://theholyhabibee.com/superorganism-model/
University of California, Berkeley. (n.d.). Native bees vs. honey bees: There are big differences. Penasquitos.org. Retrieved from https://www.penasquitos.org/news/native-bees-vs-honey-bees-there-are-big-differences
Wikipedia. (n.d.). Apis cerana. Retrieved from https://en.wikipedia.org/wiki/Apis_cerana
Wikipedia. (n.d.). Apis dorsata. Retrieved from https://en.wikipedia.org/wiki/Apis_dorsata
Wikipedia. (n.d.). Apis florea. Retrieved from https://en.wikipedia.org/wiki/Apis_florea
Wikipedia. (n.d.). Honey bee. Retrieved from https://en.wikipedia.org/wiki/Honey_bee

YourDictionary. (n.d.). Eusociality. EBSCO. Retrieved from https://www.ebsco.com/research-starters/biology/eusociality