**4. The bumble bees and the stingless bees: the other bee domestications**

About 90% of world's plant species are pollinated by animals [130–132], and the main animal pollinators in most ecosystems are bees [88]. Although other taxa like butterflies, flies, beetles, wasps, or vertebrates can be important pollinators in certain habitats or for particular plants [133, 134], none achieves the numerical dominance as flower visitors worldwide as bees [130, 131]. The pollination efficiency of bees has been used by humans to improve their crop yields. The western honey bees is the most commonly used species in managed pollination service [76, 135]. This species pollinates nearly half of the top 115 global food commodities and is capable of increasing the yields of 96% of animal-pollinated crops [117, 136]. However, the lack of sufficient stocks of honey bees to ensure pollination service [115, 137], the aggressiveness of Africanized honey bees (i.e., obtained by man-made hybridization between African and European subspecies of *A. mellifera* to breed a strain of bees that would produce more honey and be better adapted to tropical conditions) in Neotropics [138], and the poor pollination efficiency of *A. mellifera* for some plants, as well as the requirement of maintaining the honey bee colonies outside the flowering period of valuable crops [139] have triggered or restarted the domestication of other bee species: the bumble bees and the stingless bees.

#### **4.1. The bumble bees**

sedentary [106]. This context presumably triggered the beekeeping development by providing hives to honey bees that make it easier to harvest their honey and wax by humans [105]. At the beginnings of beekeeping, honey bees were not "bred" so much as "kept": humans provided rudimentary containers (often destroyed during honey harvesting) and hoped that wild bee colonies would take up residence without later swarming [105]. Over time, humans increased their control on bees by developing swarming control device (i.e., queen excluder [96]), reproduction control (e.g., artificial insemination [97]), mass breeding (e.g., [107]), selective breeding programs (e.g., [108–110]), and new strains (e.g., Buckfast strain [111] or Africanized honey bees [112]).

The honey bees' domestication concerns only *A. mellifera* and *A. cerana* (see details about the later species in [89]) most likely because they display intrinsic features that facilitated the domestication process: (i) cavity-nesting habit making hives suitable for these species, (ii) hygienic behavior (i.e., detection and removal of diseased brood and wastes) limiting diseases, and (iii) adaptations to tropical and temperate climate facilitating the apiculture development across the world [89, 110], for example, *A. mellifera*. Moreover, differentiations in traits facilitating beekeeping are observed at the subspecies level. Subsequently, some particular subspecies were preferably domesticated by humans. For instance, non-African subspecies have been more widely used by most beekeepers since they can survive in temperate

Domestication history of honey bees has been investigated through molecular datasets that highlight several domestication events followed by introgression between subspecies [90, 113, 114]. Although the honey bee domestication history has been regarded as a directed pathway [10], the evolution from early beekeeping practices to modern apiculture practices can been seen as similar to the prey pathway in which game-keeping strategies turns into control over movements, feeding, and reproduction. However, it is likely than directed and prey pathways occurred during honey bee domestication history since several domestication events hap-

Many authors acknowledge (often without justification) the domesticated status of *A. mellifera* (e.g., [10, 16, 47, 58, 89, 102, 115–117]). In contrast, *A. mellifera* has been considered as never properly domesticated but only as managed species by other authors (e.g., [110, 114]; however, some of these scientists acknowledge an ongoing domestication process) because (i) their biology, physiology, and behavior are seen as largely unchanged from their wild counterparts [114], (ii) honey bees are able to survive without human's help [118], (iii) there is extensive gene flow between wild/feral and managed bees in native range due to the difficulties to achieve controlled mating [119]. However, these points should be reconsidered. First, the comparison of phenotypes between "wild" and "nonwild" populations is difficult in a large a part of the species range. Indeed, colonies that are found in the wild may have escaped from a managed colony, and therefore, they may not be wild [120]. In Europe, it is unlikely that there are any truly wild subpopulations left due to this gene flow [120]. This means that the differentiation fostered by the domestication process can be blurred by the large amount of feral populations in the wild. Nevertheless, there are significant behavioral changes observed in man-controlled honey bees stocks such as multiple queen colonies (i.e., colonies conserved several queens without

regions, have a low tendency to swarm, and low aggressiveness [101].

pened [90, 113, 114].

42 Animal Domestication

**3.3. Is** *Apis mellifera* **domesticated?**

Bumble bees (Hymenoptera, Apidae, *Bombus* spp.) are social insects with a nearly worldwide distribution with their largest species diversity in temperate and cold areas [75, 140]. Except in tropical regions, bumble bees produce annual colonies (i.e., colony dies after the production of new queens and males). They have several adaptations such as their ability to "buzz pollinate" (i.e., sonication) and their insulated bodies that make them ideal pollinators for many valuable crops (e.g., raspberry and tomatoes) for which honey bees are quite inefficient pollinators [139–141]. Moreover, bumble bee colonies can be easily shipped and can be maintained without specialists' help in crops fields [139]. However, they do not produce honey or hive material suitable for mass market sales [117]. Therefore, their commercial importance is only based on their pollination efficiency on particular crops, mainly in greenhouses [117]. First trials of bombiculture were attempted by researchers at the beginning of the twentieth century [142, 143], but bumble bee mass production started during the 1980s [139, 144]. At least five species have underwent domestication process for pollinator production (*B. ignites*, *B. impatiens*, *B. lucorum*, *B. occidentalis*, and *B. terrestris*) and many more for research purpose [139, 142–145]. Their domestication history follows a directed pathway with several domestication events since several bumble bee breeders started independently the production of the same taxon [139, 144]. *Bombus terrestris* is the most traded species: more than 2 million colonies are yearly produced and shipped throughout the world (review in [146]). The species is mass produced in indoor facilities in which the whole life cycle is controlled by humans [139]. The choice of the most effective bumble bee species for mass production has been made through the test of several species [139]. Most of them have been proved to be poorer choices (e.g., more aggressive, hard to feed, low production success), and their domestication programs have been abandoned [139, 144]. This means that the domestication process of such species (e.g., *B. lapidarius* [139]) has regressed after a period of human interest.

species, and *Ericerus* species. These species are economically valuable for the substances that they produce under proper farming [152]. *Dactylopius coccus* and *Kerria lacca* are among the

Insects: The Disregarded Domestication Histories http://dx.doi.org/10.5772/intechopen.81834 45

Cochineal is an important source of red for dyes, lake pigments, cosmetics, and food/pharmaceutical colorants [151, 153]. Indeed, the red dye is mainly composed of carmine, which is a pigment obtained from the scale insects belonging to *Dactylopius* genus (Hemiptera, Dactyloidae) or some *Porphyrophora* species (Hemiptera, Margarodidae) [151, 153–155]. Nowadays, most of the carmine production is based on the farming of *D. coccus* inhabiting

The species is used as a source of carmine in Mesoamerica and South America since the pre-Columbian times [156]. The earliest known cochineal-dyed textiles dates back to the twelfth century, but first evidence of cochineal farming is estimated to the tenth century [155–157]. The center of domestication is thought to be in Mexico [157]. Carmine became an important export good during the Spanish colonial period [156]. Later, the species was introduced in other areas such as Australia, Canary Islands, South Africa, and South Asia [156]. In the middle of the nineteenth century, the production of cochineal fell sharply due to the development of artificial red dyes. Consequently, the cochineal trade almost totally disappeared in the twentieth century. Since the 1970s, cochineal production was restarted due to the discov-

*Dactylopius coccus* females are wingless sessile parasites of cacti in the genus *Opuntia* [151, 153, 155]. After mating with winged males, females give birth to nymph that are airborne transported by the wind to new host plants. Since the *D. coccus* females depend on *Opuntia* plants, their production takes place in cactus farms producing domesticated plant species [156, 158]. Cochineals are produced by infesting cacti plants and harvesting the insects by hand 90 days later. During this period, humans actively control potential predators [159]. At the end of the process, some cochineals are left to reproduce, while others are collected for

*Dactylopius coccus* has been considered as a domesticated species [47, 58, 151], since it is reliant on human propagation and protection for survival at least in some regions [159]. Moreover, most of *D. coccus* is produced in man-controlled environments (i.e., cactus farms). Nevertheless, humans poorly control the species reproduction and gene flow with wild populations is frequent; yet more recent developments are improving this control (e.g., environment-control microtunnels in Mexico [58]). Overall, the domestication process of *D. coccus* (level 3, **Figure 1**) is far behind the ones of silkworms and honey bees. Since comparison between farmed and

Lac is an important commercial resin of several utilities (e.g., material construction, cosmetics, medicine). It is a resinous secretion of lac insect species from Asia and Central America

ery of carcinogenic and hazardous properties of synthesized dyes [155].

wild cochineals is lacking, potential domestication syndrome is unknown.

most produced scale insects.

(sub)tropical South and Central America [151, 153].

**5.1.** *Dactylopius coccus*

carmine extraction.

**5.2.** *Kerria lacca*

Although domestication of bumble bees has been acknowledged by various authors (e.g., [139, 147]), comparison between breeders' stocks and wild populations is still lacking to highlight potential domestication syndrome in *Bombus* species. Nevertheless, the domestication programs of the five most produced species are already quite advanced (Level 4, **Figure 1**) since current bumble bee breeders' stocks experience low gene flow with their wild counterparts. However, no selective breeding program has been reported to date.

#### **4.2. The stingless bees**

Stingless bees (Hymenoptera, Apidae, Meliponini) are social bees with perennial colonies (i.e., nest can remain active for more than 50 years) occurring in most tropical or subtropical areas [75, 148]. They are known for their pollen/honey production and their pollination efficiency for several valuable crops (e.g., coffee, Avocado, Strawberry, Rambutan) [138, 148]. Meliponiculture dates back to the Maya civilization and is nowadays practiced in Australia and Central/South America [148–150]. Nevertheless, their domestication process has not progressed so far (Level 2, **Figure 1**) since most of the meliponiculture is mainly a capture production that consists in attracting stingless bee swarms and maintaining the colonies in artificial wooden hives [148, 150].

### **5. Cochineal insects**

Scale insects (Hemiptera, Coccoidea) are the third large insect groups including species that are, sometimes, considered as domesticated [47, 58, 151]: cochineals, lac scales, *Cerplastes*

species, and *Ericerus* species. These species are economically valuable for the substances that they produce under proper farming [152]. *Dactylopius coccus* and *Kerria lacca* are among the most produced scale insects.

#### **5.1.** *Dactylopius coccus*

tropical regions, bumble bees produce annual colonies (i.e., colony dies after the production of new queens and males). They have several adaptations such as their ability to "buzz pollinate" (i.e., sonication) and their insulated bodies that make them ideal pollinators for many valuable crops (e.g., raspberry and tomatoes) for which honey bees are quite inefficient pollinators [139–141]. Moreover, bumble bee colonies can be easily shipped and can be maintained without specialists' help in crops fields [139]. However, they do not produce honey or hive material suitable for mass market sales [117]. Therefore, their commercial importance is only based on their pollination efficiency on particular crops, mainly in greenhouses [117]. First trials of bombiculture were attempted by researchers at the beginning of the twentieth century [142, 143], but bumble bee mass production started during the 1980s [139, 144]. At least five species have underwent domestication process for pollinator production (*B. ignites*, *B. impatiens*, *B. lucorum*, *B. occidentalis*, and *B. terrestris*) and many more for research purpose [139, 142–145]. Their domestication history follows a directed pathway with several domestication events since several bumble bee breeders started independently the production of the same taxon [139, 144]. *Bombus terrestris* is the most traded species: more than 2 million colonies are yearly produced and shipped throughout the world (review in [146]). The species is mass produced in indoor facilities in which the whole life cycle is controlled by humans [139]. The choice of the most effective bumble bee species for mass production has been made through the test of several species [139]. Most of them have been proved to be poorer choices (e.g., more aggressive, hard to feed, low production success), and their domestication programs have been abandoned [139, 144]. This means that the domestication process of such species (e.g., *B. lapidarius* [139]) has regressed after a period of human interest. Although domestication of bumble bees has been acknowledged by various authors (e.g., [139, 147]), comparison between breeders' stocks and wild populations is still lacking to highlight potential domestication syndrome in *Bombus* species. Nevertheless, the domestication programs of the five most produced species are already quite advanced (Level 4, **Figure 1**) since current bumble bee breeders' stocks experience low gene flow with their wild counter-

parts. However, no selective breeding program has been reported to date.

Stingless bees (Hymenoptera, Apidae, Meliponini) are social bees with perennial colonies (i.e., nest can remain active for more than 50 years) occurring in most tropical or subtropical areas [75, 148]. They are known for their pollen/honey production and their pollination efficiency for several valuable crops (e.g., coffee, Avocado, Strawberry, Rambutan) [138, 148]. Meliponiculture dates back to the Maya civilization and is nowadays practiced in Australia and Central/South America [148–150]. Nevertheless, their domestication process has not progressed so far (Level 2, **Figure 1**) since most of the meliponiculture is mainly a capture production that consists in attracting stingless bee swarms and maintaining the colonies in artificial wooden hives [148, 150].

Scale insects (Hemiptera, Coccoidea) are the third large insect groups including species that are, sometimes, considered as domesticated [47, 58, 151]: cochineals, lac scales, *Cerplastes*

**4.2. The stingless bees**

44 Animal Domestication

**5. Cochineal insects**

Cochineal is an important source of red for dyes, lake pigments, cosmetics, and food/pharmaceutical colorants [151, 153]. Indeed, the red dye is mainly composed of carmine, which is a pigment obtained from the scale insects belonging to *Dactylopius* genus (Hemiptera, Dactyloidae) or some *Porphyrophora* species (Hemiptera, Margarodidae) [151, 153–155]. Nowadays, most of the carmine production is based on the farming of *D. coccus* inhabiting (sub)tropical South and Central America [151, 153].

The species is used as a source of carmine in Mesoamerica and South America since the pre-Columbian times [156]. The earliest known cochineal-dyed textiles dates back to the twelfth century, but first evidence of cochineal farming is estimated to the tenth century [155–157]. The center of domestication is thought to be in Mexico [157]. Carmine became an important export good during the Spanish colonial period [156]. Later, the species was introduced in other areas such as Australia, Canary Islands, South Africa, and South Asia [156]. In the middle of the nineteenth century, the production of cochineal fell sharply due to the development of artificial red dyes. Consequently, the cochineal trade almost totally disappeared in the twentieth century. Since the 1970s, cochineal production was restarted due to the discovery of carcinogenic and hazardous properties of synthesized dyes [155].

*Dactylopius coccus* females are wingless sessile parasites of cacti in the genus *Opuntia* [151, 153, 155]. After mating with winged males, females give birth to nymph that are airborne transported by the wind to new host plants. Since the *D. coccus* females depend on *Opuntia* plants, their production takes place in cactus farms producing domesticated plant species [156, 158]. Cochineals are produced by infesting cacti plants and harvesting the insects by hand 90 days later. During this period, humans actively control potential predators [159]. At the end of the process, some cochineals are left to reproduce, while others are collected for carmine extraction.

*Dactylopius coccus* has been considered as a domesticated species [47, 58, 151], since it is reliant on human propagation and protection for survival at least in some regions [159]. Moreover, most of *D. coccus* is produced in man-controlled environments (i.e., cactus farms). Nevertheless, humans poorly control the species reproduction and gene flow with wild populations is frequent; yet more recent developments are improving this control (e.g., environment-control microtunnels in Mexico [58]). Overall, the domestication process of *D. coccus* (level 3, **Figure 1**) is far behind the ones of silkworms and honey bees. Since comparison between farmed and wild cochineals is lacking, potential domestication syndrome is unknown.

#### **5.2.** *Kerria lacca*

Lac is an important commercial resin of several utilities (e.g., material construction, cosmetics, medicine). It is a resinous secretion of lac insect species from Asia and Central America [160, 161]. *Kerria lacca* (Hemiptera, Kerriidae) is one of the main species used for lac production [160, 161]. Its life cycle is similar to *D. coccus* with winged males and wingless sessile females that parasite several hundred host plants [161, 162]. For several centuries, lac yields were collected from the wild on infested host plants by local human populations [161]. During the nineteenth century, the increase of exportation from Asia triggered the development of artificial inoculation and mass production [161] through a domestication history that can be interpreted as a prey pathway (i.e., human control on the species was triggered by the need of increasing lac supply). Similarly to *D. coccus*, the domestication process of *K. lacca* is at an early stage (level 3, **Figure 1**) since the current production involved only host plant, lac crop, and lac pest management.

**7. Biological control agents and sterile insect technique**

yearlong basis to response to demand across the world [178, 182].

control of pests [175].

**8. Insects as pets**

Addressing the needs of the increasing human population will require a 60% increase in global food production by 2050 [174]. Insects could aid in achieving this objective by providing food production [19, 164] as well as pollination service (see Section 4) and biological

Insects: The Disregarded Domestication Histories http://dx.doi.org/10.5772/intechopen.81834 47

Biological control is a method of controlling pests such as arthropods, weeds, and plant diseases using predator (e.g., ladybugs to control aphids [176], herbivorous, or parasite species [175]). Parasitoids are among the most widely used biological control agents (e.g., [177, 178]). In these species, female deposits its egg inside or outside a host where emerged parasitoid larva continues to feed resulting in the host death [178–180]. This parasitic way of life is used by humans to target hosts that are pests. Whiteflies parasitoids (Hymenoptera, Aphelinidae, Encyrtidae, Eulophidae, Platygastridae, Pteromalidae, and Signiphoridae) are an example of insects used in greenhouses to control major crop pests (i.e., the whiteflies; Hemiptera: Aleyrodidae) [177, 180]. As many other parasitoids (e.g., fly *Eucelatoria*, the beetle *Chrysolina*, and the wasp *Aphytis*), they are massively produced in captive conditions by humans before being shipped across the world [180]. The full control of their life cycle by humans is needed in order to ensure that the production (i) matches with the appropriate release dates when susceptible host species is at a suitable phase of development [181] and (ii) is available on a

The sterile insect technique (SIT) is an alternative approach to control main pests (e.g., [183–185]) or disease vectors (e.g., [186–188]). This method implies to massively release sterile males (sterilized through the effects of irradiation on the reproductive cells) of an insect species into a target environment to compete with wild males for reproduction [183–185]. Ultimately, mass releases allow limiting offspring production of a particular pest and promoting its eradication (e.g., [184]). Mass-rearing production with a life cycle fully controlled by

The required full control of life cycle of pest insects for SIT or biological control agents means that an advanced domestication process is reached (up to 5 since some patented strains are available [189]). In the context of SIT, several studies have investigated the differences between wild and mass-produced males in order to ensure that released sterile males are able to compete with wild males (e.g., [183, 190]). These studies show that the domestication process has triggered several ecological and behavioral divergences between produced and wild populations as well as a decreased fitness of produced populations in the wild (e.g., [183, 190]).

Archeological pieces of evidence show that insects have been used as pets for centuries [191]. Nowadays, crickets, grasshoppers, beetles, cockroaches, silkworms, ants, honey bees, bumble bees, mantises, and stick/leaf insects are bred by humans as a pleasing activity or for teaching purpose [192–194]. Conversely to vertebrates [8, 195–197], there is no, to my knowledge, scientific

humans is needed to produce the large quantity of insect required by SIT [183].
