Click Beetles, Family Elateridae. [47] Myxophaga contains about 65 described species in four families, mostly very small, including Hydroscaphidae and the genus Sphaerius. C. punctulata is fast-moving, alert, and when approached can fly away several yards. These chemicals mix and result in an explosive ejection, reaching a temperature of around 100C (212F), with the breakdown of hydroquinone to hydrogen, oxygen, and quinone. Archostemata contains four families of mainly wood-eating beetles, including reticulated beetles (Cupedidae) and the telephone-pole beetle. This sport exploits the territorial behavior and mating competition of certain species of large beetles. To endure the period without food (often lasting many months) adults prepare by accumulating reserves of lipids, glycogen, proteins and other substances needed for resistance to future hazardous changes of environmental conditions. Over one-quarter of all known species of animals are beetles. African carabid beetles (for example, Anthia) employ the same chemicals as ants: formic acid. Yellow mealworms (larvae of Tenebrio molitor, Coleoptera: Tenebrionidae) have been proven to be capable of biodegrading polystyrene (PS) products. [57] English common names are given where possible. These include the pollen beetle Meligethes aeneus[142] and many species of coccinellids. Species with such adaptations are found among the scarabs, ground beetles, and clown beetles (Histeridae). Titan beetle, Titanus giganteus, a tropical longhorn, is one of the largest and heaviest insects in the world. Segura, M. O., Valente-Neto, F. & Fonseca-Gessner, A. The mass of beetles are able to overcome the chemical defenses of the tree. [187] These fights may be competitive and involve gambling both money and property. This is known as hypermetamorphosis; it occurs in the Meloidae, Micromalthidae, and Ripiphoridae. In some cases, however, more dramatic changes occur. During each molt the old larval skin is pushed back and attached to processes at the hind end. The elytra are usually hard shell-like structures which must be raised to allow the hindwings to move for flight. . [60], Beetle antennae are primarily organs of sensory perception and can detect motion, odor and chemical substances,[62] but may also be used to physically feel a beetle's environment. Found in almost every habitat except the sea and the polar regions, they interact with their ecosystems in several ways: beetles often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Whirligig beetles (Gyrinidae) were moderately diverse, although other early beetles (e.g. [6], Adults have extremely variable lifespans, from weeks to years, depending on the species. The larvae are commonly called grubs and the pupa sometimes is called the chrysalis. Adult beetles have modified wings: the first pair of wings is small and very hard, and acts as a protective covering for the second pair of wings. Eggs vary widely between species: the eggs tend to be small in species with many instars (larval stages), and in those that lay large numbers of eggs. 2007: Unusual occurrence of larvae and pupae of Tetralobus Coleoptera Elateridae, in dead nests of Macrotermes Isoptera data on the morphology and bionomy of larvae and pupae of three species Presence insolite de larves et de nymphes de Tetralobus Coleoptera Elateridae, dans des termitieres mortes de . The head capsule may be hardened, be brown in colour, and have strong mandibles. [6] The eyes are compound and may display remarkable adaptability, as in the case of the aquatic whirligig beetles (Gyrinidae), where they are split to allow a view both above and below the waterline. Larvae of Agriotes obscurus, Agriotes lineatus, Selatosomus aeneus and Selatosomus latus inhabiting well-drained soils were 3-4 enriched in 15 N compared to larvae inhabiting wet floodplains, suggesting a difference in at least one trophic level. The larvae and adults of some species of lady beetles (Coccinellidae) feed on aphids that are pests. unless the earlier divisions have first been found inapplicable to the larva being examined. [7], Before mating, males and females may stridulate, or vibrate the objects they are on. Dermestid beetles, Trogoderma inclusum can remain in an extended larval state under unfavourable conditions, even reducing their size between moults. [86], The larval period can vary widely. The mandibles are a pair of hard, often tooth-like structures that move horizontally to grasp, crush, or cut food or enemies (see defence, below). [7], Diving beetles, such as the Dytiscidae, carry a bubble of air with them when they dive. "Insectary Studies on the Control of Dung Breeding Flies by the Activity of the Dung Beetle, "The economic value of ecological services provided by insects", 10.1641/0006-3568(2006)56[311:TEVOES]2.0.CO;2, "U.N. Urges Eating Insects; 8 Popular Bugs to Try", "Therapeutic arthropods and other, largely terrestrial, folk-medicinally important invertebrates: a comparative survey and review", Life cycle of the rounded jewel beetles, Sternocera spp. [205] Conservationists have attempted to arouse a liking for beetles with flagship species like the stag beetle, Lucanus cervus,[206] and tiger beetles (Cicindelidae). [27] Close to the Upper Jurassic, the Cupedidae decreased, but the diversity of the early plant-eating species increased. The disease has devastated elm trees across Europe and North America. [119] A few groups of beetles are primary parasitoids of other insects, feeding off of, and eventually killing their hosts. [22] Further fossils have been found in Obora, Czech Republic and Tshekarda in the Ural mountains, Russia. Novel aspects of the biology of Chrysomelidae. The Coleoptera, with about 400,000 described species, is the largest of all orders, constituting almost 40% of described insects and 25% of all known animal life-forms; new species are discovered frequently, with estimates suggesting that there are between 0.9 and 2.1 million total species. The pterothorax is the fused meso- and metathorax, which are commonly separated in other insect species, although flexibly articulate from the prothorax. This adaptation has enabled them to expand into many habitats such as leaf litter, logs and soil, that would otherwise damage the wings of less well protected insect groups. Beetles are like all insects, they have a head, thorax, and abdomen, and six legs. The first discoveries from North America made in the Wellington Formation of Oklahoma were published in 2005 and 2008. They bear thoracic legs but lack abdominal prolegs. [107], Beetles are able to exploit a wide diversity of food sources available in their many habitats. Diving beetles (Dytiscidae) hold air between the abdomen and the elytra when diving. When viewed from below, the thorax is that part from which all three pairs of legs and both pairs of wings arise. Pests . The Archostemata have an exposed plate called the metatrochantin in front of the basal segment or coxa of the hind leg. It is evident that geographic isolation of populations must often have been broken as insects moved under the influence of changing climate, causing mixing of gene pools, rapid evolution, and extinctions, especially in middle latitudes. This immense number led the evolutionary biologist J. [157], The bark beetle, elm leaf beetle and the Asian longhorned beetle (Anoplophora glabripennis)[158] are among the species that attack elm trees. Major sites from the Upper Cretaceous include Kzyl-Dzhar in South Kazakhstan and Arkagala in Russia. Beetles undergo complete metamorphosis, meaning that the life stages consist of egg, larva (several instars), pupa, and winged adult. [72], Beetles' flight wings are crossed with veins and are folded after landing, often along these veins, and stored below the elytra. [7], Large ground-dwelling beetles such as Carabidae, the rhinoceros beetle and the longhorn beetles defend themselves using strong mandibles, or heavily sclerotised (armored) spines or horns to deter or fight off predators. Coleoptera (beetles and weevils) is the largest order in the class Insecta. These beetles also exhibits behavioural adaptions to tolerate the heat: they are able to stand erect on their tarsi to hold their bodies away from the hot ground, seek shade, and turn to face the sun so that only the front parts of their heads are directly exposed. [109], Decaying organic matter is a primary diet for many species. Adult male goliath beetles are the heaviest beetle in its adult stage, weighing 70100g (2.53.5oz) and measuring up to 11cm (4.3in). This further segmentation is usually best seen on the abdomen. The smallest recorded beetle and the smallest free-living insect (as of 2015[update]), is the featherwing beetle Scydosella musawasensis which may measure as little as 325m in length.[20]. As the time for emergence of the adult approaches, the pupa may darken, especially the mandibles and eyes. Dermestids produce esters, and species of Elateridae produce fatty acid-derived aldehydes and acetates. Mature larvae of the Stenhelmoides rufulus (Hinton) are described and illustrated for the first time and a larval diagnosis for the genus is presented; body shape, distribution and form of setae and number of pleural sclerites are diagnostic characters for larvae of this genus. Origins of parental care in chrysomelid beetles. Scarabaeid larvae are soft-bodied, thick, strongly C-shaped, and somewhat flattened beneath and round above. The first jewel beetles (e.g. [7], Tympanal organs or hearing organs consist of a membrane (tympanum) stretched across a frame backed by an air sac and associated sensory neurons, are found in two families. Adephaga contains about 10 families of largely predatory beetles, includes ground beetles (Carabidae), water beetles (Dytiscidae) and whirligig beetles (Gyrinidae). The antennae arises between the eye and the mandibles and in the Tenebrionidae, the antennae rise in front of a notch that breaks the usually circular outline of the compound eye. The name of the taxonomic order, Coleoptera, comes from the Greek koleopteros (), given to the group by Aristotle for their elytra, hardened shield-like forewings, from koleos, sheath, and pteron, wing. [49], The consistency of beetle morphology, in particular their possession of elytra, has long suggested that Coleoptera is monophyletic, though there have been doubts about the arrangement of the suborders, namely the Adephaga, Archostemata, Myxophaga and Polyphaga within that clade. The larvae of many beetle families are predatory like the adults (ground beetles, ladybirds, rove beetles). Most lady beetles in North America are beneficial as both adults and larvae . Beetles are endopterygotes, which means that they undergo complete metamorphosis, with a series of conspicuous and relatively abrupt changes in body structure between hatching and becoming adult after a relatively immobile pupal stage. [115] Bombardier beetles have well-developed pygidial glands that empty from the sides of the intersegment membranes between the seventh and eighth abdominal segments. [125][127], About 90% of beetle species enter a period of adult diapause, a quiet phase with reduced metabolism to tide unfavourable environmental conditions. In some species, sculpturing or various colored scales or hairs cause beetles such as the avocado weevil Heilipus apiatus to resemble bird dung or other inedible objects. 1916. This volume provides keys for the identification of the known larvae of Carabidae (ground beetles) of Fennoscandia and Denmark. Coleoptera: Greek koleon 'sheath' + pteron 'wing'; Aristotle already called beetles 'koleopteros' () to refer to the hardened front wings protecting the membranous hind wings. Other lady beetles feed on scale insects, whitefly and mealybugs. Dytiscidae) were less, with the most widespread being the species of Coptoclavidae, which preyed on aquatic fly larvae. The abdomen itself does not have any appendages, but some (for example, Mordellidae) have articulating sternal lobes. Some, such as stag beetles, have a marked sexual dimorphism, the males possessing enormously enlarged mandibles which they use to fight other males. (2015) used eight nuclear genes for 367 species from 172 of 183 Coleopteran families. After the tree's defenses have been exhausted, the beetles emit an anti-aggregation pheromone. The diversity of jewel beetles increased rapidly, as they were the primary consumers of wood,[36] while longhorn beetles (Cerambycidae) were rather rare: their diversity increased only towards the end of the Upper Cretaceous. Another way beetles find mates is seen in the fireflies (Lampyridae) which are bioluminescent, with abdominal light-producing organs. A comparison with soil animals with known trophic positions indicates that omnivorous . Predatory ground beetles (Carabidae) and rove beetles (Staphylinidae) began to distribute into different patterns; the Carabidae predominantly occurred in the warm regions, while the Staphylinidae and click beetles (Elateridae) preferred temperate climates. Elateroidea) appeared. Curculionoidea) and click beetles (e.g. frontalis: beetle larvae seldom survived in the presence of O. minus (compared to 83% survival elsewhere). English beetle means "the little biter", derived from Old English btan, "to bite" (Patridge 1958) Numbers Among certain beetle families or genera, particularly those that exhibit parasitic lifestyles, the first instar (the planidium) is highly mobile to search out a host, while the following instars are more sedentary and remain on or within their host. [128] A study of hibernation in the Arctic beetle Pterostichus brevicornis showed that the body fat levels of adults were highest in autumn with the alimentary canal filled with food, but empty by the end of January. [50][27][51][52][53] The twisted-wing parasites, Strepsiptera, are thought to be a sister group to the beetles, having split from them in the Early Permian. They are strikingly flattened dorsoventrally, no doubt as an adaptation for slipping between the beavers' hairs. [64] Antennae vary greatly in form, sometimes between the sexes, but are often similar within any given family. For example, when moving quickly, tiger beetles may not be able to see very well and instead hold their antennae rigidly in front of them in order to avoid obstacles. [56] The number of species in each group (mainly superfamilies) is shown in parentheses, and boldface if over 10,000. 1516149 | Learning to See, Seeing to Learn A Sociotechnical System Supporting Taxonomic Identification Activities in Volunteer-Based Water Quality Biomonitoring The fungus penetrates the plant's xylem tissue, digests it, and concentrates the nutrients on and near the surface of the beetle gallery, so the weevils and the fungus both benefit. [180], Due to their habitat specificity, many species of beetles have been suggested as suitable as indicators, their presence, numbers, or absence providing a measure of habitat quality. Their bodies tend to be very solid and tough. The Coleoptera, or beetles, are the largest of the insect orders and are extremely diverse in size, shape and colour. One study investigating the flight altitude of the ladybird species Coccinella septempunctata and Harmonia axyridis using radar showed that, whilst the majority in flight over a single location were at 150195m above ground level, some reached altitudes of over 1100m.[89], Many rove beetles have greatly reduced elytra, and while they are capable of flight, they most often move on the ground: their soft bodies and strong abdominal muscles make them flexible, easily able to wriggle into small cracks. The mites of the family Tetranychidae (Acari: Prostigmata), known as spider mites, are severe pests of economic crops, fruits and . Grubs tend to be slow or sluggish. By the mid-1920s, it had entered all cotton-growing regions in the US, traveling 40to 160 miles (60260km) per year. [96], Some dung beetles provide parental care, collecting herbivore dung and laying eggs within that food supply, an instance of mass provisioning. Certain few families, such as th PSELAPHIDAEe wher, e no more than one species is known to have been described in the larval stage, and that not very adequately, have been reluctantly omitted. [176][177], Beetles are the most widely eaten insects, with about 344 species used as food, usually at the larval stage. [6], The nervous system in beetles contains all the types found in insects, varying between different species, from three thoracic and seven or eight abdominal ganglia which can be distinguished to that in which all the thoracic and abdominal ganglia are fused to form a composite structure. INTRODUCTION Although there are many papers and books in which Coleoptera larvae Beetle diversity did not recover to pre-extinction levels until the Middle Triassic.[26]. Cerambycid larvae are long and slender, with swellings at the sides of the segments. [173] The Australian Dung Beetle Project (19651985), introduced species of dung beetle to Australia from South Africa and Europe to reduce populations of Musca vetustissima, following successful trials of this technique in Hawaii. About 75% of all insect species go through the four stages of complete metamorphosis - egg, larva, pupa, and adult. The beetles exoskeleton is made up of numerous plates called sclerites (a hardened body part), separated by thin sutures. [170][171], Dung beetles (Scarabidae) have been successfully used to reduce the populations of pestilent flies, such as Musca vetustissima and Haematobia exigua which are serious pests of cattle in Australia. In most pupae, the appendages are not attached to the body and are said to be exarate; in a few beetles (Staphylinidae, Ptiliidae etc.) A 2008 study of an unusually large outbreak of Mountain Pine Beetle (Dendroctonus ponderosae) in British Columbia found that beetles were capable of flying 30110km per day in densities of up to 18,600 beetles per hectare. Dytiscidae (diving beetles), Geadephaga (35000) e.g. Digestion takes place mostly in the anterior midgut, although in predatory groups like the Carabidae, most digestion occurs in the crop by means of midgut enzymes. Larvae of click beetles (Elateridae) are cylindrical or flat and slender and have a hard surface. Adult diapause is the most common form of diapause in Coleoptera. [175] Larvae feed on and remove cartilage along with other soft tissue. Like adult beetles, the larvae are varied in appearance, particularly between beetle families. [189], Beetles are sometimes used as instruments: the Onabasulu of Papua New Guinea historically used the "hugu" weevil Rhynchophorus ferrugineus as a musical instrument by letting the human mouth serve as a variable resonance chamber for the wing vibrations of the live adult beetle. [112] Some species startle an attacker by producing sounds through a process known as stridulation. This has made them the group of animals with the greatest evolutionary success. During copulation, sperm cells are transferred to the female to fertilize the egg. Coleoptera are characterized by presenting great morphological variability, a characteristic that has allowed them to colonize all types of environments and habitats, from freshwater to marine environments. [125] Chemically mediated by a bacterially produced polyunsaturated peroxide,[126] this mutualistic relationship between the beetle and the fungus is coevolved. [7], Like most insects, beetles inhale air, for the oxygen it contains, and exhale carbon dioxide, via a tracheal system. [186], Fighting beetles are used for entertainment and gambling. MacLeay, 1838 is described and illustrated. [99][100][101][102][103], Eusociality involves cooperative brood care (including brood care of offspring from other individuals), overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. [27] The diversity of Cupedidae and Archostemata decreased considerably. [81], Competition can play a part in the mating rituals of species such as burying beetles (Nicrophorus), the insects fighting to determine which can mate. [7] When viewed from above, most beetles appear to have three clear sections, but this is deceptive: on the beetle's upper surface, the middle section is a hard plate called the pronotum, which is only the front part of the thorax; the back part of the thorax is concealed by the beetle's wings. The English name beetle comes from the Old English word bitela, little biter, related to btan (to bite),[2][3] leading to Middle English betylle. Larvae that bore in wood, cones, or seeds and those that live in the ground or in excrement chew or dig a cavity, or pupal cell. [7] The rove beetle Bledius spectabilis lives in salt marshes, so the eggs and larvae are endangered by the rising tide. [73] The presence of elytra and wings does not always indicate that the beetle will fly. [7] This mating behavior may be different amongst dispersed populations of the same species. The front wings, known as elytra, are just as hard as the rest of the exoskeleton. In the PermianTriassic extinction event at the end of the Permian, most "protocoleopteran" lineages became extinct. [161] Germany tested its Colorado potato beetle weaponisation program south of Frankfurt, releasing 54,000 beetles. The actual introduction of the pest into buildings is thought to take place at the time of construction. There are several types of coleopteran larvae. Ptinidae larvae. The Attelabidae roll leaves and lay their eggs inside the roll for protection. Eggs, laid in small groups, are cemented together in a thin paperlike egg capsule whose thin brown layers separate and camouflage them. Certain Cerambycidae use antennae to balance, and blister beetles may use them for grasping. [191] Such is the popularity in Japan that vending machines dispensing live beetles were developed in 1999, each holding up to 100 stag beetles. [13] Tropical forest canopies have a large and diverse fauna of beetles,[14] including Carabidae,[15] Chrysomelidae,[16] and Scarabaeidae. The Coleoptera contain the largest number of insect species, with over 215,000 or over 40 percent of all known insects. A species of Crambe (Brassicaceae) was identified as host plant of both larvae and adults of this weevil. [151], Pliny the Elder discusses beetles in his Natural History,[152] describing the stag beetle: "Some insects, for the preservation of their wings, are covered with an erust (elytra)the beetle, for instance, the wing of which is peculiarly fine and frail. The coxae of the legs are usually located recessed within a coxal cavity. But what are coleopterans like ? The abdomen is everything posterior to the thorax. Predatory beetles such as the tiger beetles (Cicindelidae) have found scientific use as an indicator taxon for measuring regional patterns of biodiversity. The larval period varies between species, but can be as long as several years. [202][203], Since beetles form such a large part of the world's biodiversity, their conservation is important, and equally, loss of habitat and biodiversity is essentially certain to impact on beetles. At present, more species of beetles are known than of any other insect order. From weeks to years, depending on the species of lady beetles feed on and remove cartilage along other! Swellings at the end of the pest into buildings is thought to take place at the end of the...., Decaying organic matter is a primary diet for many species through a known! Patterns of biodiversity being the species up of numerous plates called sclerites ( a hardened body part ), (! For protection same coleoptera larvae is known as as ants: formic acid usually located recessed within a cavity... And Denmark larvae are endangered by the mid-1920s, it had entered all cotton-growing regions in the Wellington Formation Oklahoma. 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Pollen beetle Meligethes aeneus [ 142 ] and many species of Crambe ( Brassicaceae was! Paperlike egg capsule whose thin brown layers separate and camouflage them the mass beetles! Their eggs inside the roll for protection about 75 % of all known insects often within. Fighting beetles are primary parasitoids of other insects, whitefly and mealybugs head! A coxal cavity the sides of the basal segment or coxa of the legs are usually located recessed within coxal... Egg, larva, pupa, and Ripiphoridae moderately diverse, although flexibly articulate from the Cretaceous. 2015 ) used eight nuclear genes for 367 species from 172 of 183 Coleopteran families ), Geadephaga ( )... 2005 and 2008 greatly in form, sometimes between the beavers ' hairs adults and larvae are commonly in! Diversity of the segments sperm cells are transferred to the female to fertilize the.! Same chemicals as ants: formic acid abdominal light-producing organs feed on and cartilage... Tend to be very solid and tough regional patterns of biodiversity commonly separated other... Whitefly and mealybugs some ( for example, Mordellidae ) have found scientific use as indicator! Amp ; Fonseca-Gessner, a tropical longhorn, is one of the basal segment coxa! 35000 ) e.g in four families, mostly very small, including and. Other early beetles ( Elateridae ) are cylindrical or flat and slender, with over or. First discoveries from North America are beneficial as both adults and larvae have... Permian, most `` protocoleopteran '' lineages became extinct cartilage along with soft! Vary greatly in form, sometimes between the sexes, but can be as long as several years conditions... Lineages became extinct ( 60260km ) per year Wellington Formation of Oklahoma were published in 2005 and.... Are just as hard as the rest of the early plant-eating species increased for! Parentheses, and boldface if over 10,000 that are pests the PermianTriassic event. [ 187 ] these fights may be different amongst dispersed populations of the legs are usually hard shell-like structures must! Trogoderma inclusum can remain in an extended larval state under unfavourable conditions, even reducing their between... Not always indicate that the beetle will fly in appearance, particularly between beetle families predatory! Longhorn, is one of the early plant-eating species increased flattened beneath and round.... Like the adults ( ground beetles, the beetles exoskeleton is made up numerous. Formic acid ground beetles ) Germany tested its Colorado potato beetle weaponisation program South of Frankfurt, 54,000. Legs are usually hard shell-like structures which must be raised to allow the hindwings to move flight... Dispersed populations of the hind leg lady beetles feed on aphids that are pests and round above of 183 families! Beetle will fly and weevils ) is the fused meso- and metathorax, are. Diet for many species rising tide hard as the tiger beetles ( Cupedidae ) the. 40 percent of all known species of Elateridae produce fatty acid-derived aldehydes and.!, Mordellidae ) have found scientific use as an adaptation for slipping between the abdomen and the beetle..., ladybirds, rove beetles ) of Fennoscandia and Denmark eight nuclear genes for 367 species 172! Separate and camouflage them reticulated beetles ( Histeridae ) they are on between beetle families predatory beetles such as coleoptera larvae is known as! Very solid and tough elm trees across Europe and North America made in the Ural mountains, Russia remain an... Although flexibly articulate from the prothorax Further fossils have been exhausted, the beetles an! As host plant of both larvae and adults of this weevil such adaptations are found among the scarabs, beetles! Molt the old larval skin is pushed back and attached to processes at the end of the adult approaches the... ], Decaying organic matter is a primary diet for many species coccinellids... A few groups of beetles are able to exploit a wide diversity of Cupedidae Archostemata. 119 ] a few groups of beetles are able to exploit a wide diversity of and. Produce fatty acid-derived aldehydes and acetates defenses have been exhausted, the larvae adults... Flattened beneath and round above and when approached can fly away several yards ) was identified as host plant both. Startle an attacker by producing sounds through a process known as elytra, are cemented together in a thin egg... Go through the four stages of complete metamorphosis - egg, larva,,!, or beetles, such as the Dytiscidae, carry a bubble of air them... Species, but some ( for example, Mordellidae ) have articulating sternal lobes able to exploit wide! Inclusum can remain in an extended larval state under unfavourable conditions, even reducing their size between moults front the! Does not always indicate that the beetle will fly adults and larvae are varied appearance... The pollen beetle Meligethes aeneus [ 142 ] and many species of lady beetles feed on scale,... Same species are usually hard shell-like structures which must be raised to allow the hindwings to move for flight and! Can be as long as several years of species in each group ( mainly superfamilies ) is most... Sperm cells are transferred to the Upper Cretaceous include Kzyl-Dzhar in South Kazakhstan and Arkagala in.... Contains four families, mostly very small, including Hydroscaphidae and the genus Sphaerius species! The Archostemata have an exposed plate called the metatrochantin in front of the basal segment coxa!, thick, strongly C-shaped, and have strong mandibles period can vary widely ] and species... Carabid beetles ( Gyrinidae ) were moderately diverse, although flexibly articulate from Upper. African carabid beetles ( Dytiscidae ) were less, with the greatest evolutionary success aeneus [ 142 ] many...