49-562: Mutillinae Myrmillinae Pseudophotopsidinae Rhopalomutillinae Sphaeropthalminae Ticoplinae Velvet ants ( Mutillidae ) are a family of more than 7,000 species of wasps whose wingless females resemble large, hairy ants . Their common name velvet ant refers to their resemblance to an ant , and their dense pile of hair, which most often is bright scarlet or orange, but may also be black, white, silver, or gold. Their bright colors serve as aposematic signals. They are known for their extremely painful stings , (the sting of
98-405: A vascular strand in the nectary to assist in transport over a longer distance. Pollinators feed on the nectar and depending on the location of the nectary the pollinator assists in fertilization and outcrossing of the plant as they brush against the reproductive organs, the stamen and pistil , of the plant and pick up or deposit pollen . Nectar from floral nectaries is sometimes used as
147-457: A consensus over time. The naming of families is codified by various international bodies using the following suffixes: The taxonomic term familia was first used by French botanist Pierre Magnol in his Prodromus historiae generalis plantarum, in quo familiae plantarum per tabulas disponuntur (1689) where he called the seventy-six groups of plants he recognised in his tables families ( familiae ). The concept of rank at that time
196-446: A family, yet in the realm of plants, these classifications often rely on both the vegetative and reproductive characteristics of plant species. Taxonomists frequently hold varying perspectives on these descriptions, leading to a lack of widespread consensus within the scientific community for extended periods. The continual publication of new data and diverse opinions plays a crucial role in facilitating adjustments and ultimately reaching
245-477: A local predator of velvet ants in the black-headed Timulla and tropical mimicry rings. Male mutillids fly in search of females; after mating, the female enters a host insect nest, typically a ground-nesting bee or wasp burrow, and deposits one egg near each larva or pupa . Only a few species are known to parasitize other types of hosts; exceptions include the European velvet ant, Mutilla europaea , one of
294-427: A modified part or a novel structure. The different types of floral nectaries include: Most members of Lamiaceae have a nectariferous disc which surrounds the ovary base and derived from developing ovarian tissue. In most Brassicaceae , the nectary is at the base of the stamen filament. Many monocotyledons have septal nectaries, which are at the unfused margins of the carpels. These exude nectar from small pores on
343-412: A plethora of impressive defensive strategies in prey species to improve the likelihood of escape. Velvet ants avoid predation using the following defense mechanisms: a venomous sting (if female), aposematic coloration , a stridulatory organ in their abdomen, an alarm secretion from their mandibular gland, and a durable exoskeleton. This array of defenses has contributed to the velvet ants being attributed
392-617: A preference for plants with extrafloral nectaries, particularly some species of ants and wasps , which have been observed to defend the plants bearing them. Acacia is one example of a plant whose nectaries attract ants, which protect the plant from other insect herbivores . Among passion flowers , for example, extrafloral nectaries prevent herbivores by attracting ants and deterring two species of butterflies from laying eggs. In many carnivorous plants , extrafloral nectaries are also used to attract insect prey . Charles Darwin understood that extrafloral nectar "though small in quantity,
441-462: A protective function. The Nicotiana attenuata , a tobacco plant native to the US state of Utah , uses several volatile aromas to attract pollinating birds and moths. The strongest such aroma is benzylacetone , but the plant also adds bitter nicotine , which is less aromatic, so may not be detected by the bird until after taking a drink. Researchers speculate the purpose of this addition is to discourage
490-433: A reward to insects, such as ants , that protect the plant from predators. Many floral families have evolved a nectar spur . These spurs are projections of various lengths formed from different tissues, such as the petals or sepals. They allow for pollinators to land on the elongated tissue and more easily reach the nectaries and obtain the nectar reward. Different characteristics of the spur, such as its length or position in
539-437: A squeaking or chirping sound when alarmed. Both sexes of mutillids also bear hair-lined grooves on the side of the metasoma called felt lines. Only two other vespoid families ( Bradynobaenidae and Chyphotidae ) have felt lines, but the females of these families have a distinct pronotum , with a transverse suture separating it from the mesonotum ; in female mutillids, these two thoracic segments are completely fused. Members of
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#1732797687856588-416: A velvet ant it was exposed to. In both cases the velvet ants were exhibiting rapid lateral and vertical movements to ward off an attack. Once the attack occurred the velvet ants would immediately sting the lizards. This sting resulted in the dropping of the ants in both cases and avoidance for the remainder of the trial. The side-blotched lizard was found dead in its tank 24 hours later. The side-blotched lizard
637-423: Is a natural predator of velvet ants, while the whiptail is not. The aposematic coloration of velvet ants often corresponds to a specific Müllerian mimicry ring consisting of dozens of species. This offers protection because many local predators have learned to avoid prey with this same coloration. To test the aposematic coloration on birds, mealworms were painted to resemble a velvet ant. During these trials, none of
686-912: Is greedily sought by insects" but believed that "their visits do not in any way benefit the plant". Instead, he believed that extrafloral nectaries were excretory in nature ( hydathodes ). Their defensive functions were first recognized by the Italian botanist Federico Delpino in his important monograph Funzione mirmecofila nel regno vegetale (1886). Delpino's study was inspired by a disagreement with Darwin, with whom he corresponded regularly. Extrafloral nectaries have been reported in over 3941 species of vascular plants belonging to 745 genera and 108 families , 99.7% of which belong to flowering plants (angiosperms), comprising 1.0 to 1.8% of all known species. They are most common among eudicots , occurring in 3642 species (of 654 genera and 89 families), particularly among rosids which comprise more than half of
735-455: Is largely responsible for the divergent evolution of the African honey bee, A. m. scutellata and the western honey bee . Nectar is an economically important substance as it is the sugar source for honey . It is also useful in agriculture and horticulture because the adult stages of some predatory insects feed on nectar. For example, a number of predacious or parasitoid wasps (e.g.,
784-499: Is particularly rich in carbohydrates, proteins and lipids. While their function is not always clear, and may be related to regulation of sugars, in most cases they appear to facilitate plant insect relationships. In contrast to floral nectaries, nectar produced outside the flower generally has a defensive function. The nectar attracts predatory insects which will eat both the nectar and any plant-eating insects around, thus functioning as "bodyguards". Foraging predatory insects show
833-1216: Is variable due to many factors, including flower age, plant location, and habitat management. Extrafloral nectaries (also known as extranuptial nectaries) are specialised nectar-secreting plant glands that develop outside of flowers and are not involved in pollination , generally on the leaf or petiole (foliar nectaries) and often in relation to the leaf venation . They are highly diverse in form, location, size, and mechanism. They have been described in virtually all above-ground plant parts—including stipules , cotyledons , fruits , and stems , among others. They range from single-celled trichomes to complex cup-like structures that may or may not be vascularized . Like floral nectaries, they consist of groups of glandular trichomes (e.g., Hibiscus spp.) or elongated secretory epidermal cells. The latter are often associated with underlying vascular tissue. They may be associated with specialised pockets ( domatia ), pits or raised regions (e.g., Euphorbiaceae ). The leaves of some tropical eudicots (e.g., Fabaceae ) and magnoliids (e.g., Piperaceae ) possess pearl glands or bodies which are globular trichomes specialised to attract ants. They secrete matter that
882-535: The Genera Plantarum of George Bentham and Joseph Dalton Hooker this word ordo was used for what now is given the rank of family. Families serve as valuable units for evolutionary, paleontological, and genetic studies due to their relatively greater stability compared to lower taxonomic levels like genera and species. Nectar Nectar is a viscous , sugar -rich liquid produced by plants in glands called nectaries , either within
931-859: The Cenozoic , with weak support for a role played by arthropod herbivore diversifications. They are absent in bryophytes , gymnosperms , early angiosperms , magnoliids , and members of Apiales among the eudicots. Phylogenetic studies and the wide distribution of extrafloral nectaries among vascular plants point to multiple independent evolutionary origins of extrafloral nectaries in at least 457 independent lineages. The main ingredients in nectar are sugars in varying proportions of sucrose , glucose , and fructose . In addition, nectars have diverse other phytochemicals serving to both attract pollinators and discourage predators. Carbohydrates , amino acids , and volatiles function to attract some species, whereas alkaloids and polyphenols appear to provide
980-484: The flowers with which it attracts pollinating animals, or by extrafloral nectaries , which provide a nutrient source to animal mutualists , which in turn provide herbivore protection . Common nectar-consuming pollinators include mosquitoes , hoverflies , wasps , bees , butterflies and moths , hummingbirds , honeyeaters and bats . Nectar plays a crucial role in the foraging economics and evolution of nectar-eating species; for example, nectar foraging behavior
1029-584: The gynoecium from certain pathogens. Floral nectaries have evolved and diverged into the different types of nectaries due to the various pollinators that visit the flowers. In Melastomataceae , different types of floral nectaries have evolved and been lost many times. Flowers that ancestrally produced nectar and had nectaries may have lost their ability to produce nectar due to a lack of nectar consumption by pollinators, such as certain species of bees . Instead they focused on energy allocation to pollen production. Species of angiosperms that have nectaries use
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#17327976878561078-473: The Mutillidae are generally stenothermic and thermophilic ; they may not avoid light, but rather are active during temperatures that usually occur only after sunset. Predation is one of the strongest forces natural selection uses to drive the evolution of an organism's morphology, physiology, and behavior. During this coevolution, the prey either being consumed by the predator or escaping has resulted in
1127-576: The North American Southwest. North American Mutillidae have eight phenotypically distinct and geographically limited Müllerian mimicry rings (Desert, Eastern, Madrean, Texan, Red-headed Timulla , Black-headed Timulla , Tropical, and Western) making up one of the largest Müllerian mimicry complexes on the planet. These mimicry rings are the result of repeated convergent evolution of aposematic traits between co-occurring velvet ant species, rather than shared phylogenetic history. Through
1176-430: The abdomen is contracted. This mechanism is an auditory cue warning predators that are about to attack to stay away. In one experiment, every time a shrew got within 1 meter of a velvet ant, the velvet ant would begin stridulating. Stridulations became more frequent as the predator moved closer to the velvet ant, and the shrew never attempted to attack the velvet ant. However, different scenarios with shrews have shown that
1225-514: The book's morphological section, where he delved into discussions regarding the vegetative and generative aspects of plants. Subsequently, in French botanical publications, from Michel Adanson 's Familles naturelles des plantes (1763) and until the end of the 19th century, the word famille was used as a French equivalent of the Latin ordo (or ordo naturalis ). In zoology ,
1274-403: The evolution of aposematic traits in velvet ant species in the same ring, local predators have learned to avoid these well-defended wasps. The exoskeleton of all velvet ants is unusually tough (to the point that some entomologists have reported difficulty piercing them with steel pins when attempting to mount them for display in cabinets). This characteristic allows them to successfully invade
1323-508: The family Mutillidae contained one subfamily that was unrelated to the remainder, and this subfamily was removed to form a separate family Myrmosidae . Family (biology) Family ( Latin : familia , pl. : familiae ) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy . It is classified between order and genus . A family may be divided into subfamilies , which are intermediate ranks between
1372-399: The family Myrmosidae , formerly classified as a subfamily of mutillids, also have a distinct pronotum in females, but lack felt lines in both sexes. Adult mutillids feed on nectar . Although some species are strictly nocturnal, females are often active during the day. Females of Tricholabiodes thisbe are sometimes active up to two hours before sunset. Guido Nonveiller (1963) hypothesized
1421-540: The family as a rank intermediate between order and genus was introduced by Pierre André Latreille in his Précis des caractères génériques des insectes, disposés dans un ordre naturel (1796). He used families (some of them were not named) in some but not in all his orders of "insects" (which then included all arthropods ). In nineteenth-century works such as the Prodromus of Augustin Pyramus de Candolle and
1470-439: The flower, may determine the type of pollinator that visits the flower. Defense from herbivory is often one of the roles of extrafloral nectaries. Floral nectaries can also be involved in defense. In addition to the sugars found in nectar, certain proteins may also be found in nectar secreted by floral nectaries. In tobacco plants, these proteins have antimicrobial and antifungal properties and can be secreted to defend
1519-840: The forager after only a sip, motivating it to visit other plants, therefore maximizing the pollination efficiency gained by the plant for a minimum nectar output. Neurotoxins such as aesculin are present in some nectars such as that of the California buckeye . Nectar contains water, essential oils , carbohydrates , amino acids , ions , and numerous other compounds. Some insect pollinated plants lack nectaries, but attract pollinators through other secretory structures. Elaiophores are similar to nectaries but are oil secreting. Osmophores are modified structural structures that produce volatile scents. In orchids , these have pheromone qualities. Osmophores have thick domed or papillate epidermis and dense cytoplasm. Platanthera bifolia produces
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1568-429: The fracture of a velvet ant's exoskeleton, a total of 4 times resulted in the death of that velvet ant within 24 hours. Aside from protection from predators, the exoskeleton also helps control moisture. Due to these strong defense mechanisms, local predators generally avoid the velvet ants, so it has been difficult to determine their predators. One study found tropical and subtropical iguanian lizards ( Dactyloidae ) to be
1617-897: The known occurrences. The families showing the most recorded occurrences of extrafloral nectaries are Fabaceae , with 1069 species, Passifloraceae , with 438 species, and Malvaceae , with 301 species. The genera with the most recorded occurrences are Passiflora (322 species, Passifloraceae), Inga (294 species, Fabaceae), and Acacia (204 species, Fabaceae). Other genera with extrafloral nectaries include Salix ( Salicaceae ), Prunus ( Rosaceae ) and Gossypium ( Malvaceae ). Foliar nectaries have also been observed in 101 species of ferns belonging to eleven genera and six families, most of them belonging to Cyatheales (tree ferns) and Polypodiales . Fern nectaries appear to have evolved around 135 million years ago, nearly simultaneously with angiosperms. However, fern nectaries did not diversify remarkably until nearly 100 million years later, in
1666-410: The male carries the smaller female aloft while mating, which is also seen in the related family Thynnidae . As is the case for all aculeates , only female mutillids are capable of inflicting a sting. The stinger is a modified female organ called an ovipositor , which is unusually long and maneuverable in mutillids. In both sexes, a structure called a stridulitrum on the metasoma is used to produce
1715-454: The nectar to attract pollinators that consume the nectar, such as birds and butterflies . In Bromeliaceae , septal nectaries (a form of gynoecial nectary) are common in species that are insect or bird pollinated. In species that are wind pollinated, nectaries are often absent because there is no pollinator. In flowers that are generally pollinated by a long-tongued organism such as certain flies , moths , butterflies, and birds, nectaries in
1764-488: The nests of their prey and also helps them retain moisture. Mutillids exhibit extreme sexual dimorphism . As in some related families in the Vespoidea , males have wings, but females are wingless. The males and females are so distinct in their morphology that entomologists often find it very hard to determine whether a given male and female belong to the same species, unless they are captured while mating. In some species,
1813-471: The only species that attacks social bees (e.g., Bombus ), and the genus Pappognatha , whose hosts are tree-dwelling orchid bees . The mutillid larvae then develop as idiobiont ectoparasitoids , eventually killing their immobile larval/pupal hosts within a week or two. Velvet ants exhibit haplodiploid sex determination, as do other members of the superfamily Vespoidea . Recent classifications of Vespoidea sensu lato (beginning in 2008) concluded that
1862-428: The ovaries are common because they are able to reach the nectar reward when pollinating. Sepal and petal nectaries are often more common in species that are pollinated by short-tongued insects that cannot reach so far into the flower. Nectar secretion increases as the flower is visited by pollinators. After pollination, the nectar is frequently reabsorbed into the plant. The amount of nectar in flowers at any given time
1911-432: The painfulness of the sting of Dasymutilla klugii outscored 58 other species of stinging insects tested; the only species this researcher rated as having a more painful sting were Paraponera clavata (bullet ant), Synoeca septentrionalis (warrior wasp), and Pepsis and Hemipepsis spp. ( tarantula hawks ). In an experimental setting, only two lizard species (one whiptail and one side-blotched lizard ) attacked
1960-439: The painted mealworms were consumed, while all the control mealworms were consumed immediately. However, the painted mealworms were attacked by the birds, but the birds immediately ceased the attack. These experiments provide evidence that the aposematic coloration of velvet ants causes their predators to hesitate, acting as a visual defense mechanism. The stridulatory organ that velvet ants possess produces an audible squeaking when
2009-476: The pollinator that feeds on the plant's nectar. Nectar is secreted from epidermal cells of the nectaries, which have a dense cytoplasm , by means of trichomes or modified stomata . Adjacent vascular tissue conducts phloem bringing sugars to the secretory region, where it is secreted from the cells through vesicles packaged by the endoplasmic reticulum . The adjacent subepidermal cells may also be secretory. Flowers that have longer nectaries sometimes have
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2058-491: The ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to the family Juglandaceae , but that family is commonly referred to as the "walnut family". The delineation of what constitutes a family— or whether a described family should be acknowledged— is established and decided upon by active taxonomists . There are not strict regulations for outlining or acknowledging
2107-511: The social wasp species Apoica flavissima ) rely on nectar as a primary food source. In turn, these wasps then hunt agricultural pest insects as food for their young. Nectar is most often associated with flowering plants angiosperms , but it is also produced by other groups, including ferns . Nectar is derived from Greek νέκταρ , the fabled drink of eternal life. Some derive the word from νε- or νη- "not" plus κτα- or κτεν- "kill" , meaning "unkillable", thus "immortal". The common use of
2156-653: The species Dasymutilla klugii rated a 3 on the Schmidt pain index and lasts up to 30 minutes), and has resulted in the common name "cow killer" or "cow ant" being applied to the species Dasymutilla occidentalis . However, mutillids are not aggressive and sting only in defense. In addition, the actual toxicity of their venom is much lower than that of honey bees or harvester ants . Unlike true ants , they are solitary, and lack complex social systems. Mutillidae can be found worldwide with about 230 genera or subgenera and around 8,000 species worldwide. Over 400 species occur in
2205-436: The surface of the gynoecium. Nectaries may also vary in color, number, and symmetry. Nectaries can also be categorized as structural or non-structural. Structural nectaries refer to specific areas of tissue that exude nectar, such as the types of floral nectaries previously listed. Non-structural nectaries secrete nectar infrequently from non-differentiated tissues. The different types of floral nectaries coevolved depending on
2254-442: The title of "the indestructible insect." This title was bestowed on them after experimental interactions between velvet ants and their potential predators that resulted in the survival of the ant and the ultimate avoidance by the predator. The venom that velvet ants inject through their stinger has been investigated for five species of Dasymutilla , revealing that they are composed primarily of peptides. According to one researcher,
2303-419: The velvet ant would also stridulate after the shrew attacked it. Every time this occurred the shrew dropped the wasp. The exoskeleton of the velvet ant is remarkably strong. It required 11 times more force to crush than that of the honeybee. As well as being durable, the exoskeleton is also round, making it more difficult for predators to pierce it with attempted stings or bites. During all the trials that led to
2352-614: The word "nectar" to refer to the "sweet liquid in flowers", is first recorded in AD 1600. A nectary or honey gland is floral tissue found in different locations in the flower and is one of several secretory floral structures, including elaiophores and osmophores, producing nectar, oil and scent respectively. The function of these structures is to attract potential pollinators , which may include insects, including bees and moths , and vertebrates such as hummingbirds and bats . Nectaries can occur on any floral part, but they may also represent
2401-488: Was not yet settled, and in the preface to the Prodromus Magnol spoke of uniting his families into larger genera , which is far from how the term is used today. In his work Philosophia Botanica published in 1751, Carl Linnaeus employed the term familia to categorize significant plant groups such as trees , herbs , ferns , palms , and so on. Notably, he restricted the use of this term solely within
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