The Honey Bee Apis mellifera: Comparison
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The honey bee Apis mellifera Linnaeus (1758) provides many benefits to humans and ecosystems. This species is an important pollinator in natural environments, which may help to preserve and restore the biodiversity of wild plants. On the other hand, pollination in agro-ecosystems by managed honey bee colonies may enhance crop yield and quality, meeting the increasing food demand.

Beekeeping is also a high-valued and income-generating activity, which provides humans with honey as high-quality food as well as substances used as raw materials and in pharmaceuticals.

In addition, the honey bee and its products are valuable bioindicators and bioaccumulators of environmental pollution: they provide valuable information on the impact of human activities, enabling the implementation of measures to mitigate risks to human and ecosystem health.

The honey bee is also linked to many cultural ecosystem services and has a longstanding tradition in human culture, mysticism, and religion. Its popularity may be therefore used for educational purposes and to raise public awareness of important issues, such as the conservation of pollinator habitats and biodiversity.

  • honey bee
  • ecosystem services
  • agro-ecosystems
  • bee products
  • provisioning services
  • regulating services
  • cultural services
  • biodiversity

1. Regulating Services Provided by the Honey bee: The Conservation of Plant Biodiversity and Enhancement of Crop Production

1.1. Plant biodiversity conservation

Wild and domesticated bees are the most important pollinator group and the role played by bees as pollinators within natural and agro-ecosystems is becoming increasingly evident and recognised [4,8,9].

Wild and domesticated bees are the most important pollinator group and the role played by bees as pollinators within natural and agro-ecosystems is becoming increasingly evident and recognised [1][2][3].
While the importance of

A. mellifera

for crop production is widely acknowledged, there is currently a great debate among researchers on the real benefits to natural ecosystems derived from the presence of managed bees. Managed bee colonies may endanger wild pollinators, including wild populations of

A. mellifera itself, due to floral resource limitation and potential pest and pathogen transmission [10,11]. This is especially true in the case of massive introduction of non-native honey bees in natural, protected areas [10,12]; therefore, according to some researchers, the best option should be to avoid high-density beekeeping and to increase spacing among neighbouring apiaries to guarantee abundant floral resources for all pollinators [10,13]. In view of this, laws and regulations to ban “intensive

itself, due to floral resource limitation and potential pest and pathogen transmission [4][5]. This is especially true in the case of massive introduction of non-native honey bees in natural, protected areas [4][6]; therefore, according to some researchers, the best option should be to avoid high-density beekeeping and to increase spacing among neighbouring apiaries to guarantee abundant floral resources for all pollinators [4][7]. In view of this, laws and regulations to ban “intensive beekeeping” in natural ecosystems, while, in any case, favouring more sustainable approaches also through financial incentives for beekeepers should be promoted.

beekeeping” in natural ecosystems, while, in any case, favouring more sustainable approaches also through financial incentives for beekeepers should be promoted.

On the other hand, studies point out the global importance of honey bees as pollinators in natural habitats and the need to ensure their conservation to maintain the genetic diversity of local subspecies and their ecological function [11,14,15]. In natural habitats, honey bees appear to be the most frequent pollinators, averaging 13% of floral visits, with 5% of plant species being exclusively visited by

On the other hand, it was pointed out that the global importance of honey bees as pollinators in natural habitats and the need to ensure their conservation to maintain the genetic diversity of local subspecies and their ecological function [5][8][9]. In natural habitats, honey bees appear to be the most frequent pollinators, averaging 13% of floral visits, with 5% of plant species being exclusively visited by

A. mellifera [15]. This confirms that honey bees may also aid in the maintenance of the biodiversity of native communities of flowering plants [14–16].

[9]. This confirms that honey bees may also aid in the maintenance of the biodiversity of native communities of flowering plants [8][9][10].
Another important aspect linking

A. mellifera and biodiversity is the possibility of using honey bees to understand the diversity status of flowering plants. Pollen richness is often used as a means to estimate the floristic richness of an ecosystem [17] and honey bees may provide useful information for monitoring purposes through, for example, analysis of the pollen grains packed into the pollen basket, as well as the analysis of pollen-contaminating

and biodiversity is the possibility of using honey bees to understand the diversity status of flowering plants. Pollen richness is often used as a means to estimate the floristic richness of an ecosystem [11] and honey bees may provide useful information for monitoring purposes through, for example, analysis of the pollen grains packed into the pollen basket, as well as the analysis of pollen-contaminating bee products, especially honey. The use of molecular tools may offer further advantages in terms of quality and quantity of information compared to pollen identification through microscopic analysis. For example, DNA metabarcoding applied to honey reveals the presence of DNA from both pollen- and nectar-providing plants [12][13].

bee products, especially honey. The use of molecular tools may offer further advantages in terms of quality and quantity of information compared to pollen identification through microscopic analysis. For example, DNA metabarcoding applied to honey reveals the presence of DNA from both pollen- and nectar-providing plants [18,19].

Of course, microscopic analysis and DNA metabarcoding applied to bee pollen and honey provide information on plant taxa on which honey bees forage, but, given that pollen composition in bee matrices is largely influenced by floristic local biodiversity and flowering phenology [12][13][14][15][16][17], these data may improve the understanding of the local biodiversity of flowering plants.

Of course, microscopic analysis and DNA metabarcoding applied to bee pollen and honey provide information on plant taxa on which honey bees forage, but, given that pollen composition in bee matrices is largely influenced by floristic local biodiversity and flowering phenology [18–23], these data may improve our understanding of the local biodiversity of flowering plants.

1.2. Crop pollination: quality and yield

 

1.2. Crop pollination: quality and yield

A. mellifera is easy to manage and transport, and the income the honey bee provides through the delivery of many products has made it the most valuable pollinator used to enhance agricultural production since ancient times. One of the best examples is almond pollination in California, where more than 70% of all honey bee colonies in the USA are moved to orchards for the promotion of pollination. To overcome the increasing demand for managed bees, growers are starting to breed almond varieties claimed as ‘pollinator-independent’ due to their presumed great capacity for self-pollination [33] [18]. However, researchers have

 found that even in this case if bees are not present in the orchards, growers obtain a lower crop yield, as pollinators guarantee a 20% increase in kernel yield [33] [18]. In kiwi and avocado, while anemophilous and/or self-pollination alone ensures 12% and up to 17% of fruit set, respectively, when flowers are also exposed to honey bees, the yield increases to 80% and 90% (Figure 1a) [34,35][19][20]. In apples, one of the most important fruit crops in the world, insect pollination is necessary to obtain marketable fruits, i.e., large and symmetric [36][21]. Symmetry is a classical aesthetic principle, and consumers usually prefer to opt for beautiful fruits because they evoke naturalness and seem more appetising and healthier as compared to asymmetric fruits [37][22]. Therefore, honey bee colonies are usually placed into large commercial orchards to ensure fruit quality and quantity, even if honey bees do not seem to be the most efficient pollinators of apple flowers [38,39][23][24]. Similarly, supplementing raspberry and blueberry crops with beehives is necessary to obtain marketable berries [40,41][25][26]. Indeed, raspberry and blueberry fruits are characterised by an aggregation of drupelets, and under-pollinated flowers develop into crumbly, misshapen, or small berries that are avoided by consumers.

Honey bee supplementation is also important for ensuring yield stability over space and time [42]. If growers place small apiaries throughout a farm, this may enhance the spatial stability of bee visits, ensuring a homogeneous rate of yield quality and quantity [43]. The presence of healthy colonies also guarantees fruit quality and quantity across seasons in both apple and pear crops [44]. The strength of the bee colonies is also decisive for promoting crop production in the northern highbush blueberry, which is self-fertile, but higher fruit set and yields occur following visitation by honey bees from healthy colonies [45]. For watermelon crops, as native solitary bees are effective pollinators but do not allow optimal yield, supplementary pollination services through A. mellifera are suggested, even if in this case native managed stingless bees are preferable because they compete less with native pollinators [46]. In wild blueberry, both honey bee and bumblebee abundance increases fruit set and reduces spatial heterogeneity in crop production [47].

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Figure 1. Benefits provided by A. mellifera in crop production. (a) Yield increase in almond, kiwi, avocado, and soybean crops; (b) increase in fruit weights in Cucurbita spp., fava bean, and sunflower.
Honey bee supplementation is also important for ensuring yield stability over space and time [27]. If growers place small apiaries throughout a farm, this may enhance the spatial stability of bee visits, ensuring a homogeneous rate of yield quality and quantity [28]. The presence of healthy colonies also guarantees fruit quality and quantity across seasons in both apple and pear crops [29]. The strength of the bee colonies is also decisive for promoting crop production in the northern highbush blueberry, which is self-fertile, but higher fruit set and yields occur following visitation by honey bees from healthy colonies [30]. For watermelon crops, as native solitary bees are effective pollinators but do not allow optimal yield, supplementary pollination services through A. mellifera are suggested, even if in this case native managed stingless bees are preferable because they compete less with native pollinators [31]. In wild blueberry, both honey bee and bumblebee abundance increases fruit set and reduces spatial heterogeneity in crop production [32].

is also known to improve yields in horticultural, legume, oilseed, and feed crops. In a recent study by Garibaldi et al. [48], the authors highlighted the fact that soybean productivity can significantly increase through insect pollination. Among soybean pollinators, it has been demonstrated that the honey bee effectively increases crop yield, pod set, and seed set [48]. In Brazil, crop yield has increased by up to 126% [48]. Honey bee supplementation enhances fruit weights of Cucurbita pepo, C. moschata, and C. maxima by approximately 26%, 70%, and 78%, respectively [49]; in fava bean (Vicia faba), the yield increased by 17% [50], and in sunflower (Helianthus annuus), about 50% [51].

Honey bee supplementation is also known to improve yields in horticultural, legume, oilseed, and feed crops. In a recent study by Garibaldi et al. [33], the authors highlighted the fact that soybean productivity can significantly increase through insect pollination. Among soybean pollinators, it has been demonstrated that the honey bee effectively increases crop yield, pod set, and seed set [33]. In Brazil, crop yield has increased by up to 126% [33]. Honey bee supplementation enhances fruit weights of Cucurbita pepo, C. moschata, and C. maxima by approximately 26%, 70%, and 78%, respectively [34]; in fava bean (Vicia faba), the yield increased by 17% [35], and in sunflower (Helianthus annuus), about 50% [36] (Figure 1b).

The effects of honey bees on fruit set and fruit/seed weight of biofuel crops, such as

The effects of honey bees on fruit set and fruit/seed weight of biofuel crops, such as 

Jatropha curcas and

 and 

Ricinus communis, have also been demonstrated [52,53]:

, have also been demonstrated [37][38]: 

J. curcas, for example, fruit set can increase up to 70% [53].

, for example, fruit set can increase up to 70% [38].

2. Use of Bee Products as Raw Materials and Medicinal Resources

Even if pollination is not unique to the honey bee, the delivery of a wide range of products to humans is exclusive to this insect. This has led to the development of beekeeping, a high-valued and income-generating activity, especially for honey production. In Europe the estimated number of hobby and professional beekeepers in 2010 was about 620,000, both with about 18.9 million hives and estimated honey production of more than 22,0000 tons [54][39]. Depending on the European country and distribution network, the price of honey can vary from a few euros to up to 40 euros/kg [54][39]. However, honey bees can provide not only food but also medicinal resources and raw materials.

2.1. Wax as Raw Material: New Perspectives

Beeswax is a secretion that adult bees aged between 12 and 18 days can produce from wax glands located in the abdomen. Once secreted, wax droplets solidify and are manipulated by the bee to build the nest, allowing food storage, brood rearing, and thermoregulation [59][40]. Beeswax is mainly composed of alkanes, fatty acids, long-chain esters, and trace compounds, including proteins and fragments of insects, plants, propolis, and pollen [60,61][41][42].
The use of beeswax by humans traces back to the Palaeolithic Age when early humans began to produce weapons for hunting by fixing stone tips to wooden shafts with a glue substance made of beeswax and resins [62][43]. Hunting was also enhanced by using poisonous substances obtained by mixing beeswax with Euphorbia toxic sap [63][44]. Since the Neolithic Age, beeswax has also been used for the waterproofing of furniture, rituals, and cosmetics, and its use in ancient medicine dates back to ancient Egypt [60,64,65][41][45][46]. Over time, the usage of beeswax has been documented in sculpture, ornaments, masks, and candles, and at present this substance is exploited for the production of comb foundations in beekeeping, but also in the food industry as a glazing agent in fruits. For example, in the European Union, beeswax is an authorised food additive (E901) (EU Commission Regulation No. 1147/2012, 4 December 2012).
Food packaging made from beeswax and other natural substances has been recently developed [66][47] and studies on the use of beeswax as a gelling agent in some food products are on-going [67][48].

2.2. Propolis

Propolis, commonly known as the ‘bee glue’, is a resinous substance that bees collect from plants and trees, buds, and exudates of plants, which are transformed in the presence of bee enzymes. Bees use propolis for the construction and adaptation of their nests, seal the holes in their honeycombs, smooth out internal walls, and cover carcasses of intruders who died inside the hive in order to avoid their decomposition [80][49]. Propolis also protects the colony from diseases because of its antiseptic and antimicrobial properties.
The use of propolis has a long history and goes back to ancient times, as a local medicine in many parts of the world. Egyptians, Greeks, and Romans reported the use of propolis for its general healing qualities and for the cure of skin problems [81][50]. Propolis has always been used as an anti-inflammatory agent and to heal sores, ulcers, wounds, and for tissue regeneration [82][51].
In general, propolis is composed of 30% wax, 50% resin and vegetable balsam, 10% essential and aromatic oils, 5% pollen, and other components [83,84][52][53]. Its chemical composition is very complex: more than 300 components have already been identified, and its composition is dependent on the vegetal source and the local flora (geographical origin), thus creating a problem for its medical use and standardisation [84,85,86][53][54][55]. The main components are phenolic compounds (flavonoids, aromatic acids, and benzopyranes), di- and triterpenes, and essential oils, among others [87,88,89][56][57][58].
The antimicrobial properties and activities of propolis have been widely investigated [90,91,92,93,94][59][60][61][62][63]. Propolis also shows antiviral [95[64][65],96], antifungal [97[66][67],98], and antiparasitic activities [99,100,101][68][69][70].
Owing to its properties, propolis is used in products for the protection of health and prevention of diseases, in bio-pharmaceuticals, and as a constituent of bio-cosmetics [102,103][71][72]. Propolis-based products are also marketed by the pharmaceutical industry and health-food supply chains [104][73]. However, further investigations are needed to better understand the effects of propolis on human health and to establish its potential dose levels and intake periods. The recent systematifoc review and meta-analysisus of randomised controlled clinical trials of Gheflati and colleagues [105][74] on the effects of propolis supplementation on metabolic parameters is worth mentioning. The current meta-analysis revealed that propolis supplementation can reduce aspartate aminotransferase.
Emerging directions are also given by the application of nanotechnologies to nutraceuticals and pharmaceuticals [106,107][75][76]. For instance, Botteon et al. [108][77] described the biosynthesis and characterisation of gold nanoparticles using Brazilian red propolis and evaluated their antimicrobial and anticancer activities.

2.3. Royal Jelly

Royal jelly is a secretion of the mandibular and hypopharyngeal glands of worker bees, A. mellifera. It is the food that regulates the distinction between reproductive and unreproductive females; only larvae exclusively fed on royal jelly develop into queens; otherwise, they develop into sterile workers [112][78].
Royal jelly is composed of 60–70% water, 9–18% protein, 7–18% simple sugars (monosaccharides), and 3–8% lipids [113][79]. It also contains trace minerals, pantothenic acid (vitamin B5), pyridoxine (vitamin B6), trace amounts of vitamin C, nucleotides, heterocyclic compounds, 10-hydroxy-2-decenoic acid (10-HDA), amino acids, and others [114][80]. Concerning the protein content, the major royal jelly proteins (MRJPs) [115][81] are a family of proteins secreted by the honeybees. Royal jelly has been used in traditional medicine since ancient times, and MRJPs are believed to be the main medicinal components. Other components include 10-acetoxydecanoic acid, trans-10-acetoxydec-2-enoic acid, 11- oxododecanoic acid, (11S)-hydroxydodecanoic acid, (10R,11R)-dihydroxydodecanoic acid, 3,11-dihydroxydodecanoic acid, and (11S),12-dihydroxydodecanoic acid [116][82].
Royal jelly has been widely used in commercial medical products, health foods, and cosmetics in many countries for more than 30 years [117][83]. A recent review bBy Guo et al. [118][84] summarised the biologically active role of royal jelly in the maintenance of biological functions, such as immunity, lifespan, memory, digestive system, blood glucose, obesity, antibacterial, and anti-cancer properties [116][82]. Ahmad et al. [114][80] provided new insights into the biological and pharmaceutical properties, such as antimicrobial, antioxidant, wound healing, immunomodulatory, anti-aging, anti-cancer, anti-inflammatory, anti-hypertension, anti-hyperlipidemic, oestrogenic, and neurotrophic effects of royal jelly.

2.4. Venom and Apitherapy

In the eusocial Aculeate Hymenoptera, the venom and stinging apparatus initially evolved as devices to immobilise prey, and then became weapons to defend the colony mainly from the attacks of invertebrate and vertebrate predators [119][85]. In particular, honey bee colonies are rewarding targets for predators and hunters because of the rich storage of honey and pollen, and the mass of immature broods and adults [119][85].
A. mellifera venom is a valuable product harvested from honeybees, with a price ranging between $30 and $300 per gram. However, bee venom is a marginal product of apiculture [120,121][86][87].
Bee venom is a natural toxin secreted from a specific venom gland located in the bee abdomen and is injected through the sting. Bee venom consists of simple organic molecules, peptides, proteins, and other bioactive elements [119,122][85][88]. In particular, bee venom contains polypeptides such as melittin, apamin, and mast cell degranulating peptides, amines, such as histamine, serotonin, dopamine, and norepinephrine, and enzymes, such as phospholipase, hyaluronidase, and histidine decarboxylase [120][86]. Melittin is a basic 26-amino-acid polypeptide that is the main component of A. mellifera venom and represents 40–60% of dry venom [123,124][89][90]. Melittin has several toxicological, pharmacological, and biological effects, such as haemolysin activity, antibacterial, and antifungal activities, anti-tumour properties, and intense surface activity on cell lipid membranes [123,125,126][89][91][92]. Nevertheless, ecological factors (temperature, flowering stage, and site) can influence the composition and diversity of the peptide and the weight of the bee venom [120][86].
Bee venom has been used in therapeutic applications in oriental traditional medicine since 1000–3000 BCE for treating inflammatory diseases and pain [127][93]. Recently, several situdies have has been proposed that bee venom as a promising neuroprotective therapy for Parkinson’s disease and as an effective treatment for patients with multiple sclerosis and other autoimmune diseases, such as rheumatoid arthritis [128,129,130][94][95][96].

3. Honey Bees and Bee Products to Safeguard Ecosystems from Pollution

A further role provided by honey bees is the possibility of delivering key information on the presence of pollutants in the environment. The first extensive study demonstrating that A. mellifera is an effective biological monitor of environmental contaminants over large geographic areas dates back to the 1980s [143][97].
This notable role is due to the morphological and behavioural characteristics of bees that, during their wide-ranging foraging activity, are highly exposed to organic and inorganic pollutants contaminating air, water, soil, and vegetation. Pollutants can also contaminate the bee products, such as pollen, honey, wax, propolis, and royal jelly.
The use of honey bees provides the following advantages over other pollution monitoring systems:
  • Very limited purchase costs and maintenance—beekeeping is an easy and low-cost activity, which provides a potentially unlimited supply of bioindicators in many environments;
  • Self-sustaining biosensors for the pollutant collection;
  • Reliable samplers of pollutants, as the bees can fly for more than 3 km around a barycentre (the hive), exploring flowers, vegetation, water, and air for a maximum of three weeks.
  • No environmental impact.
  • Simultaneous collection of a wide range of pollutants during the foraging behaviour;
  • Collection of evidence for pollutants to enter the food chain (e.g., through honey or other edible bee products) and to expose pollinators to pollutant ingestion.
In addition, as a living organism, the bee also offers the option to study lethal and sublethal effects of pollutant exposure on a biological system.
In the following paragraph, an overview of studies involving pollution in honey bees and bee products is provided.

Pollution in Bees and Bee Products

Biomonitors or bioindicators include organisms that provide information on the quality of the environment [144][98]. Importantly, a biomonitor is always a bioindicator, whereas a bioindicator does not necessarily meet the requirements of a biomonitor [144][98].
Bioindicators are used to assess both health and changes in the environment they inhabit [145][99]. Three types of bioindicators can be distinguished: plants (e.g., diatoms and lichens), animals (e.g., aquatic invertebrates), and microbes. Bioindicators can be further distinguished into four categories depending on the application, namely ecological, environmental, biodiversity, and pollution bioindicators [145][99]. A. mellifera represents, together with its products, the most complete biosensor (bioindicator and bioaccumulator), which can provide a considerable amount of data on the state of health of the environment [146,147,148,149,150,151,152,153,154,155][100][101][102][103][104][105][106][107][108][109]. Each forager bee manages to cover a foraging distance of more than 3 km from the hive and, in some cases, the area covered can be up to 100 km2 [156,157][110][111]. While passing from flower to flower, it comes into contact with a large number of pollutants.
Honeybees may accumulate pollutants in many ways. During flight and foraging activities, they collect airborne particulate matter and dust deposit on the surfaces on which the bee lands [158,159,160,161][112][113][114][115]. In addition, bees are exposed to pollutants through water used for both drinking and cooling the hive or to pollutants absorbed by the plants from the soil and accumulated in pollen and nectar [162][116].
Pollutants collected by bees can accumulate in honey, wax, pollen bullets, propolis, or other products (bees as collectors). Contaminants may also concentrate on the body of larvae or adults (bees as accumulators) [163,164][117][118].
The validity of the bee as a biological indicator has been demonstrated for the following pollutants:
-
Agrochemicals
-
Heavy metals
-
Polycyclic aromatic hydrocarbons
-
Radionuclides
-
Dioxin, polychlorinated biphenyls,
-
Particulate matter
In rural areas poor in wild vegetation, a bee is an excellent bioindicator of phytosanitary products; in this case, the insect is obliged to forage on or near cultivated species and will therefore come into contact with any sprayed active substance [165][119]. In these situations, a bee becomes a valid tool to identify times and ways of using substances at risk of toxicity and highlights the possible improper use of pesticides in real time [166][120].
Compared to rural areas, large urban agglomerations may be contaminated with pollutants from vehicular exhausts, which, in turn, may pollute the nectar and honeydew, the raw material for honey production [167][121]. Honey may thus contain pollutants that are characteristic of the environment [168][122], including minerals of natural or anthropogenic origin [169][123]. Exhaust fumes from vehicles may also interfere with the scents that ‘drive’ bees to the flowers they feed on [170][124]. Indeed, bees can perceive flower scents up to 1200 m away, although, due to pollution, this capacity reduces to 200–300 m.

4. Cultural Ecosystem Services and Bees

Cultural ecosystem services are the nonmaterial benefits people obtain from the ecosystem through spiritual enrichment, cognitive development, reflection, recreation, and aesthetics [1][125]. The ecosystem and its components, processes, and diversity provide the basis for education in many societies, influencing the types of social relations (e.g., agricultural societies differ in many respects from nomadic herding societies) and the diversity of cultures [1][125].
A brief analysis of Scopus of all the available literature from 2007 to 2021 (accessed 8/11/2021) can help understand the impact of cultural ecosystem services on ourthe community. From 2007 to 2021, we havthere are 963 documents mentioning ‘cultural ecosystem services’, either in the title, keyword, or abstract. One single document was found in 2007, whereas in the last five years, the annual number of documents exceeded 100, indicating a growing interest in this topic (Figure 2).
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Figure 2. Scopus trend of the documents on cultural ecosystem services from 2007 to 2021 (Data from Scopus online Database; accessed on 8 November 2021).
Besides provisioning and regulating ecosystem services, honey bees and beekeeping can also be linked to cultural services. Among them, the best-known examples are recreational/educational activities, such as api-tourism, a form of agro-tourism related to beekeeping, which includes a number of beekeeping-oriented activities (e.g., visits to apiaries, bee museums, honey tasting, and apitherapy), and art inspiration [135,186,187][126][127][128].
Here, we will focus is on the role of the honey bee in symbolic tradition and mysticism and the cultural values of the pollinator habitats.

3.1. The Role of the Honey Bee in the Ecosystem of the Symbolic Tradition

4.1. The Role of the Honey Bee in the Ecosystem of the Symbolic Tradition

Among the infinite hermeneutic streams of essays, studies, and interpretations of the Bible, there is no shortage of specific works on the role and meaning of honey bees [188][129]. Heirs of the medieval bestiaries, the modern dictionaries of symbols [189[130][131],190], at the entry ‘bee’, always carry a rich collection of passages, legends, and myths about this insect and its mystic role.
The highlights converge: in all cultural systems (from China to India, from Africa to Europe, from Babylonians to Christians), the honey bee has been respected, loved, and admired. This unanimity is noticeable: animals are usually ambiguous and have opposite meanings, even in the same cultural tradition. A lion at the portal of a Gothic cathedral represents the major power of redemption, but the same animal, holding a man in its paws, can signify the tremendous threat of sin. The positiveness of the bee was granted by generations of wisemen, who observed the presence of the bee in nature. They couldn’t keep from being seduced not only by the divine features of her products, but also by the dancing beauty of her trajectories. The symbolism always revolves around three main ambits: the sweetness of honey (and the pureness of wax), the organisation of the hive, and the nobleness of the queen bee.
It is easy to understand how honey was considered important and almost sacred in ancient times. Source of sweetness, medicines, and gods liquor (the legendary hydromel), unique to the golden transparency of its flow, even at the dawn of history, it was gathered in the wilderness, before the introduction of beekeeping. In the history of Israel, the Promised Land is presented by God as a «land of milk and honey» (Exodus 3:8) and John the Baptist ate «locusts and wild honey» (Matthew 3:4). However, for the Bible, bees—as all insects—are impure animals, honey is kasher: it is the miracle of a sacrality produced by profane beings, such as humans. The fascinating work of the bees, who draw the ingredients from the fragility of flowers without spoiling them, has been a symbol of chastity, a virtue which was considered in a wider sense than mere sexuality: the sweetest things, in life, are obtained without grab, through the lightness of a caress. Thanks to modern science,  we now know that life itself is known, through pollination, is granted by that caress.
The organisation of the hive, the laboriousness of the community, and the ability to work in the summer to ensure provisions for the winter, have often represented a perfect image of the ideal human society. The Church has observed in the storage of honey the treasure of good actions accumulated by the saints for the Heavenly Kingdom; the secular institutions have dreamed of the same ability to cooperate in such a harmonic way, even in large numbers. It is not surprising that bees and beehives have played an important role in the Western heraldry, carved on the shields of kings, nobles, and even monasteries, as in the coat of arms of the Cistercian abbey of Mellaray, which owes its name to honey (miel).
Finally, the queen bee’s majesty has provided a sense of unity and obedience, a perfect representation of the monarchic government (in which a single head guarantees peace for the whole community), but also an image of the Mother Church, who is responsible for the destiny of every single believer. In the Catholic Exsultet, a long chant delivered before the paschal candle during the Easter night, the ‘mother bee’ is thanked for having produced the wax that makes the celebration possible, enlightening the dark heart of the night. All these features are well known, and the comprehension of the symbolism of the honeybee through simple bibliographic research can be easily deduced.

3.2. The Values of Pollinator Habitats

4.2. The Values of Pollinator Habitats

Habitats for pollinators are wildflower meadows rich not only in native flowering plants, but also in host plants and nesting/resting sites for a range of animals, from invertebrates to reptiles, birds, and small mammals [194,195][132][133]. Some amphibians also thrive on the food and shelter provided by the meadow ecosystem [196][134]. From an ecological perspective, pollinator habitats act as ‘keystone habitats’, which are extremely valuable habitats that provide critical resources to native organisms and are therefore a sanctuary for biodiversity [196,197][134][135].
With their blooms throughout the growing season, their scents and natural sounds, pollinator habitats are generally highly appreciated. They boost the aesthetics of rural landscapes [194][132]. Greening interventions in cities promoting wilderness and spontaneous flowering vegetation have become widely applied [198][136]. Indeed, flower meadows are easy to manage and offer an extraordinary opportunity to reconnect citizens with the wilderness within the urban environment, boosting the awareness that nature is not just a decoration but provides a living infrastructure as important as power grids and public transport [199][137].
Pollinator habitats in cities also provide a great opportunity to emphasise their educational value. Environmental education is considered a cultural ecosystem service, as it may provide appreciation for nature and natural areas as well as educational enrichment, especially in school children [200,201][138][139]. Outdoor learning in school is known to provide cognitive, physical, social-emotional, and academic benefits [202][140] and pollinator habitats in schoolyards may facilitate such outcomes. It may make children aware that biodiversity is always around us and «if we look around with curious eyes, we can see biodiversity not as colourful, not as exuberant as the tropical one, but not less important. In the end, a flowery meadow with wild chicory and chamomile is sufficient to appreciate the biodiversity. These plants are visited by many insects, pollinators, predators, and even herbivores, both in the larval and adult stages. You will need to get a sweep-net, swinging it with strength on the plants of this lawn. The insects living on these plants will be collected at the bottom of the bag. You will have to be a quick observer because insects try to regain freedom quickly or you can transfer everything you have collected with the net into a glass jar to observe them more calmly, obviously making sure you release all the insects once you have satisfied your curiosity. » (M. Pellecchia, [203][141]).
Given that the honey bee is undoubtedly the most known pollinator species and holds the public interest, it may be used as a flagship species, i.e., an icon for pollinator habitats. Thus, actions to support pollinator habitats may include the creation of sites for bees. However, to avoid negative impacts on wild pollinators due to competition of floral resources with A. mellifera, apiaries should be small and food resources (flowers) abundant [10][4].
Many cases of cooperation among city councils and ecologists, zoologists, and entomologists to transform public green areas into open-air, living sanctuaries for pollinators, and other natural elements are available. One example is the ‘Ri-Natura’ project implemented in North Italy by the Sorbolo-Mezzani Municipality (Parma Province) to enhance citizens’ wellbeing and provide educational opportunities (Figure 3).
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Figure 3. The Ri-natura project of the Sorbolo-Mezzani Municipality. Areas of intervention (a,c) and projects to transform them into multiservice green areas (b,d) with wildflower meadows (*), recreation/educational areas (**), and a small apiary (***). Photo courtesy of Sorbolo-Mezzani Municipality.
Other examples of initiatives that improve well-being and nature education are urban food forests, i.e., small ecosystems planted with edible plants designed by humans to provide for their needs. Starting in 2010, food forests began to be included in municipal plans [204][142]. In many cases, food forests are also enriched by wildflower meadows to improve the presence of pollinators. One example is the Picasso Food Forest, in Parma, that, under the Erasmus Plus project From Seed to Spoon 2019-1-IT02-KA201-062392 involved high school students to plan, design, and seed a wildflower meadow to nourish pollinators [204][142] (Figure 4).
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Figure 4. High school students at the Picasso Food Forest of Parma. (a) Volunteers providing knowledge to students on the importance of seeds of nectariferous and polliniferous wild plants; (b) students seeding the wildflower meadow (Erasmus Plus project, From Seed to Spoon 2019-1-IT02-KA201-062392).
On the whole, the promotion of habitats for pollinators will, in turn, promote a range of cultural benefits. The educational, spiritual, recreational, and strongly inspiring value of nature is, in fact, a universally recognised value. Artists, who are well aware of this, often gain strength from the observation of natural beauty to produce their creations, and pollinator habitats are also frequently subject to naturalist illustrators (Figure 5).
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Figure 5.
Pollinator habitats in naturalist paintings by A. Ambrogio. Courtesy of A. Ambrogio.
Nature is the first inspiring muse of art. However, in reality, it is nature itself that is the great artwork.

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