Beetles are a group of insects that form the order Coleoptera, in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, distinguishing them from most other insects; the Coleoptera, with about 400,000 species, is the largest of all orders, constituting 40% of described insects and 25% of all known animal life-forms. The largest of all families, the Curculionidae with some 70,000 member species, belongs to this order. Found in every habitat except the sea and the polar regions, they interact with their ecosystems in several ways: beetles feed on plants and fungi, break down animal and plant debris, eat other invertebrates; some species are serious agricultural pests, such as the Colorado potato beetle, while others such as Coccinellidae eat aphids, scale insects and other plant-sucking insects that damage crops. Beetles have a hard exoskeleton including the elytra, though some such as the rove beetles have short elytra while blister beetles have softer elytra; the general anatomy of a beetle is quite uniform and typical of insects, although there are several examples of novelty, such as adaptations in water beetles which trap air bubbles under the elytra for use while diving.
Beetles are endopterygotes, which means that they undergo complete metamorphosis, with a series of conspicuous and abrupt changes in body structure between hatching and becoming adult after a immobile pupal stage. Some, such as stag beetles, have a marked sexual dimorphism, the males possessing enormously enlarged mandibles which they use to fight other males. Many beetles are aposematic, with bright colours and patterns warning of their toxicity, while others are harmless Batesian mimics of such insects. Many beetles, including those that live in sandy places, have effective camouflage. Beetles are prominent in human culture, from the sacred scarabs of ancient Egypt to beetlewing art and use as pets or fighting insects for entertainment and gambling. Many beetle groups are brightly and attractively coloured making them objects of collection and decorative displays. Over 300 species are used as food as larvae. However, the major impact of beetles on human life is as agricultural and horticultural pests.
Serious pests include the boll weevil of cotton, the Colorado potato beetle, the coconut hispine beetle, the mountain pine beetle. Most beetles, however, do not cause economic damage and many, such as the lady beetles and dung beetles are beneficial by helping to control insect pests; the name of the taxonomic order, comes from the Greek koleopteros, given to the group by Aristotle for their elytra, hardened shield-like forewings, from koleos and pteron, wing. The English name beetle comes from the Old English word bitela, little biter, related to bītan, leading to Middle English betylle. Another Old English name for beetle is ċeafor, used in names such as cockchafer, from the Proto-Germanic *kebrô. Beetles are by far the largest order of insects: the 400,000 species make up about 40% of all insect species so far described, about 25% of all animals. A 2015 study provided four independent estimates of the total number of beetle species, giving a mean estimate of some 1.5 million with a "surprisingly narrow range" spanning all four estimates from a minimum of 0.9 to a maximum of 2.1 million beetle species.
The four estimates made use of host-specificity relationships, ratios with other taxa, plant:beetle ratios, extrapolations based on body size by year of description. Beetles are found in nearly all habitats, including freshwater and coastal habitats, wherever vegetative foliage is found, from trees and their bark to flowers and underground near roots - inside plants in galls, in every plant tissue, including dead or decaying ones; the heaviest beetle, indeed the heaviest insect stage, is the larva of the goliath beetle, Goliathus goliatus, which can attain a mass of at least 115 g and a length of 11.5 cm. Adult male goliath beetles are the heaviest beetle in its adult stage, weighing 70–100 g and measuring up to 11 cm. Adult elephant beetles, Megasoma elephas and Megasoma actaeon reach 50 g and 10 cm; the longest beetle is the Hercules beetle Dynastes hercules, with a maximum overall length of at least 16.7 cm including the long pronotal horn. The smallest recorded beetle and the smallest free-living insect, is the featherwing beetle Scydosella musawasensis which may measure as little as 325 µm in length.
The oldest known fossil insect that unequivocally resembles a Coleopteran is from the Lower Permian Period about 270 million years ago, though these members of the family Tshekardocoleidae have 13-segmented antennae, elytra with more developed venation and more irregular longitudinal ribbing, abdomen and ovipositor extending beyond the apex of the elytra. In the Permian–Triassic extinction event at the end of the Permian, some 30% of all insect species became extinct, so the fossil record of insects only includes beetles from the Lower Triassic 220 mya. Around this time, during the Late Triassic, fungus-feeding species such as Cupedidae appear in the fossil record. In the stages of the Upper Triassic, alga-feeding insects such as Triaplidae and Hydrophilidae begin to appear, alongside predatory water beetles; the first weevils, including the Obrienidae, appear alongside the first rove beetles, which resemb
In zoological nomenclature, a type species is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e. the species that contains the biological type specimen. A similar concept is used for suprageneric groups called a type genus. In botanical nomenclature, these terms have no formal standing under the code of nomenclature, but are sometimes borrowed from zoological nomenclature. In botany, the type of a genus name is a specimen, the type of a species name; the species name that has that type can be referred to as the type of the genus name. Names of genus and family ranks, the various subdivisions of those ranks, some higher-rank names based on genus names, have such types. In bacteriology, a type species is assigned for each genus; every named genus or subgenus in zoology, whether or not recognized as valid, is theoretically associated with a type species. In practice, there is a backlog of untypified names defined in older publications when it was not required to specify a type.
A type species is both a concept and a practical system, used in the classification and nomenclature of animals. The "type species" represents the reference species and thus "definition" for a particular genus name. Whenever a taxon containing multiple species must be divided into more than one genus, the type species automatically assigns the name of the original taxon to one of the resulting new taxa, the one that includes the type species; the term "type species" is regulated in zoological nomenclature by article 42.3 of the International Code of Zoological Nomenclature, which defines a type species as the name-bearing type of the name of a genus or subgenus. In the Glossary, type species is defined as The nominal species, the name-bearing type of a nominal genus or subgenus; the type species permanently attaches a formal name to a genus by providing just one species within that genus to which the genus name is permanently linked. The species name in turn is fixed, to a type specimen. For example, the type species for the land snail genus Monacha is Helix cartusiana, the name under which the species was first described, known as Monacha cartusiana when placed in the genus Monacha.
That genus is placed within the family Hygromiidae. The type genus for that family is the genus Hygromia; the concept of the type species in zoology was introduced by Pierre André Latreille. The International Code of Zoological Nomenclature states that the original name of the type species should always be cited, it gives an example in Article 67.1. Astacus marinus Fabricius, 1775 was designated as the type species of the genus Homarus, thus giving it the name Homarus marinus. However, the type species of Homarus should always be cited using its original name, i.e. Astacus marinus Fabricius, 1775. Although the International Code of Nomenclature for algae and plants does not contain the same explicit statement, examples make it clear that the original name is used, so that the "type species" of a genus name need not have a name within that genus, thus in Article 10, Ex. 3, the type of the genus name Elodes is quoted as the type of the species name Hypericum aegypticum, not as the type of the species name Elodes aegyptica.
Glossary of scientific naming Genetypes – genetic sequence data from type specimens. Holotype Paratype Principle of Typification Type Type genus
Nicrophorus vespillo is a burying beetle described by Linnaeus in 1758. Sikes, Derek S.. "A catalog of the Nicrophorinae of the world". Zootaxa. 65. Doi:10.11646/zootaxa.65.1.1. ISBN 0-9582395-1-7. Archived from the original on September 10, 2006
Eusociality, the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes called castes. Eusociality is distinguished from all other social systems because individuals of at least one caste lose the ability to perform at least one behavior characteristic of individuals in another caste. Eusociality exists in certain insects and mammals, it is observed and studied in the Hymenoptera and in termites. A colony has caste differences: Queens and reproductive males take the roles of the sole reproducers, while soldiers and workers work together to create a living situation favorable for the brood. In addition to Hymenoptera and Isoptera, there are two known eusocial vertebrates among rodents: the naked mole-rat and the Damaraland mole-rat.
Some shrimps, such as Synalpheus regalis, are eusocial. E. O. Wilson and others have claimed that humans have evolved a weak form of eusociality, but these arguments have been disputed; the term "eusocial" was introduced in 1966 by Suzanne Batra, who used it to describe nesting behavior in Halictine bees. Batra observed the cooperative behavior of the bees and females alike, as they took responsibility for at least one duty within the colony; the cooperativeness was essential as the activity of one labor division influenced the activity of another. For example, the size of pollen balls, a source of food, depended on when the egg-laying females oviposited. If the provisioning by pollen collectors was incomplete by the time the egg-laying female occupied a cell and oviposited, the size of the pollen balls would be small, leading to small offspring. Batra applied this term to species. Batra described other species, wherein the founder is accompanied by numerous helpers—as in a swarm of bees or ants—as "hypersocial."
In 1969, Charles D. Michener further expanded Batra’s classification with his comparative study of social behavior in bees, he observed multiple species of bees in order to investigate the different levels of animal sociality, all of which are different stages that a colony may pass through. Eusociality, the highest level of animal sociality a species can attain had three characteristics that distinguished it from the other levels: "Egg-layers and worker-like individuals among adult females" The overlap of generations Cooperative work on the cells of the bees' honeycomb E. O. Wilson extended the terminology to include other social insects, such as ants and termites, it was defined to include organisms that had the following three features: Reproductive division of labor Overlapping generations Cooperative care of youngAs eusociality became a recognized widespread phenomenon, however, it was discovered in a group of chordates, the mole-rats. Further research distinguished another important criterion for eusociality known as "the point of no return."
This is characterized by eusocial individuals that become fixed into one behavioral group, which occurs before reproductive maturity. This prevents them from transitioning between behavioral groups and creates an animal society, dependent on each other for survival and reproductive success. For many insects, this irreversibility has changed the anatomy of the worker caste, sterile and provides support for the reproductive caste. Most eusocial societies exist in arthropods; the order Hymenoptera contains the largest group of eusocial insects, including ants and wasps—those with reproductive "queens" and more or less sterile "workers" and/or "soldiers" that perform specialized tasks. For example, in the well-studied social wasp Polistes versicolor, dominant females perform tasks such as building new cells and ovipositing, while subordinate females tend to perform tasks like feeding the larvae and foraging; the task differentiation between castes can be seen in the fact that subordinates complete 81.4% of the total foraging activity, while dominants only complete 18.6% of the total foraging.
While only a moderate percentage of species in bees and wasps are eusocial, nearly all species of ants are eusocial. Some major lineages within these groups are or eusocial, as well, such as the bee tribes Apini, Bombini and Meliponini, the wasp subfamilies Polistinae and Vespinae. Eusociality in these families is sometimes managed by a set of pheromones that alter the behavior of specific castes in the colony; these pheromones may act across different species, as observed in Apis andreniformis, where worker bees responded to queen pheromone from the related Apis florea. Pheromones are sometimes used in these castes to assist with foraging. Workers of the Australian stingless bee Tetragonula carbonaria, for instance, mark food sources with a pheromone, helping their nest mates to find the food. Reproductive specialization involves the production of sterile members of the species, which carry out specialized tasks to care for the reproductive members, it can manifest in the appearance of individuals within a group whose behavior or morphology is modified for
Birds known as Aves, are a group of endothermic vertebrates, characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, a strong yet lightweight skeleton. Birds range in size from the 5 cm bee hummingbird to the 2.75 m ostrich. They rank as the world's most numerically-successful class of tetrapods, with ten thousand living species, more than half of these being passerines, sometimes known as perching birds. Birds have wings which are less developed depending on the species. Wings, which evolved from forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in flightless birds, including ratites and diverse endemic island species of birds; the digestive and respiratory systems of birds are uniquely adapted for flight. Some bird species of aquatic environments seabirds and some waterbirds, have further evolved for swimming; the fossil record demonstrates that birds are modern feathered dinosaurs, having evolved from earlier feathered dinosaurs within the theropod group, which are traditionally placed within the saurischian dinosaurs.
The closest living relatives of birds are the crocodilians. Primitive bird-like dinosaurs that lie outside class Aves proper, in the broader group Avialae, have been found dating back to the mid-Jurassic period, around 170 million years ago. Many of these early "stem-birds", such as Archaeopteryx, were not yet capable of powered flight, many retained primitive characteristics like toothy jaws in place of beaks, long bony tails. DNA-based evidence finds that birds diversified around the time of the Cretaceous–Palaeogene extinction event 66 million years ago, which killed off the pterosaurs and all the non-avian dinosaur lineages, but birds those in the southern continents, survived this event and migrated to other parts of the world while diversifying during periods of global cooling. This makes them the sole surviving dinosaurs according to cladistics; some birds corvids and parrots, are among the most intelligent animals. Many species annually migrate great distances. Birds are social, communicating with visual signals and bird songs, participating in such social behaviours as cooperative breeding and hunting and mobbing of predators.
The vast majority of bird species are monogamous for one breeding season at a time, sometimes for years, but for life. Other species have breeding systems that are polygynous or polyandrous. Birds produce offspring by laying eggs, they are laid in a nest and incubated by the parents. Most birds have an extended period of parental care after hatching; some birds, such as hens, lay eggs when not fertilised, though unfertilised eggs do not produce offspring. Many species of birds are economically important as food for human consumption and raw material in manufacturing, with domesticated and undomesticated birds being important sources of eggs and feathers. Songbirds and other species are popular as pets. Guano is harvested for use as a fertiliser. Birds prominently figure throughout human culture. About 120–130 species have become extinct due to human activity since the 17th century, hundreds more before then. Human activity threatens about 1,200 bird species with extinction, though efforts are underway to protect them.
Recreational birdwatching is an important part of the ecotourism industry. The first classification of birds was developed by Francis Willughby and John Ray in their 1676 volume Ornithologiae. Carl Linnaeus modified that work in 1758 to devise the taxonomic classification system in use. Birds are categorised as the biological class Aves in Linnaean taxonomy. Phylogenetic taxonomy places Aves in the dinosaur clade Theropoda. Aves and a sister group, the clade Crocodilia, contain the only living representatives of the reptile clade Archosauria. During the late 1990s, Aves was most defined phylogenetically as all descendants of the most recent common ancestor of modern birds and Archaeopteryx lithographica. However, an earlier definition proposed by Jacques Gauthier gained wide currency in the 21st century, is used by many scientists including adherents of the Phylocode system. Gauthier defined Aves to include only the crown group of the set of modern birds; this was done by excluding most groups known only from fossils, assigning them, instead, to the Avialae, in part to avoid the uncertainties about the placement of Archaeopteryx in relation to animals traditionally thought of as theropod dinosaurs.
Gauthier identified four different definitions for the same biological name "Aves", a problem. Gauthier proposed to reserve the term Aves only for the crown group consisting of the last common ancestor of all living birds and all of its descendants, which corresponds to meaning number 4 below, he assigned other names to the other groups. Aves can mean all archosaurs closer to birds than to crocodiles Aves can mean those advanced archosaurs with feathers Aves can mean those feathered dinosaurs that fly Aves can mean the last common ancestor of all the living birds and all of its descendants (a "c
Animals are multicellular eukaryotic organisms that form the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, grow from a hollow sphere of cells, the blastula, during embryonic development. Over 1.5 million living animal species have been described—of which around 1 million are insects—but it has been estimated there are over 7 million animal species in total. Animals range in length from 8.5 millionths of a metre to 33.6 metres and have complex interactions with each other and their environments, forming intricate food webs. The category includes humans, but in colloquial use the term animal refers only to non-human animals; the study of non-human animals is known as zoology. Most living animal species are in the Bilateria, a clade whose members have a bilaterally symmetric body plan; the Bilateria include the protostomes—in which many groups of invertebrates are found, such as nematodes and molluscs—and the deuterostomes, containing the echinoderms and chordates.
Life forms interpreted. Many modern animal phyla became established in the fossil record as marine species during the Cambrian explosion which began around 542 million years ago. 6,331 groups of genes common to all living animals have been identified. Aristotle divided animals into those with those without. Carl Linnaeus created the first hierarchical biological classification for animals in 1758 with his Systema Naturae, which Jean-Baptiste Lamarck expanded into 14 phyla by 1809. In 1874, Ernst Haeckel divided the animal kingdom into the multicellular Metazoa and the Protozoa, single-celled organisms no longer considered animals. In modern times, the biological classification of animals relies on advanced techniques, such as molecular phylogenetics, which are effective at demonstrating the evolutionary relationships between animal taxa. Humans make use of many other animal species for food, including meat and eggs. Dogs have been used in hunting, while many aquatic animals are hunted for sport.
Non-human animals have appeared in art from the earliest times and are featured in mythology and religion. The word "animal" comes from the Latin animalis, having soul or living being; the biological definition includes all members of the kingdom Animalia. In colloquial usage, as a consequence of anthropocentrism, the term animal is sometimes used nonscientifically to refer only to non-human animals. Animals have several characteristics. Animals are eukaryotic and multicellular, unlike bacteria, which are prokaryotic, unlike protists, which are eukaryotic but unicellular. Unlike plants and algae, which produce their own nutrients animals are heterotrophic, feeding on organic material and digesting it internally. With few exceptions, animals breathe oxygen and respire aerobically. All animals are motile during at least part of their life cycle, but some animals, such as sponges, corals and barnacles become sessile; the blastula is a stage in embryonic development, unique to most animals, allowing cells to be differentiated into specialised tissues and organs.
All animals are composed of cells, surrounded by a characteristic extracellular matrix composed of collagen and elastic glycoproteins. During development, the animal extracellular matrix forms a flexible framework upon which cells can move about and be reorganised, making the formation of complex structures possible; this may be calcified, forming structures such as shells and spicules. In contrast, the cells of other multicellular organisms are held in place by cell walls, so develop by progressive growth. Animal cells uniquely possess the cell junctions called tight junctions, gap junctions, desmosomes. With few exceptions—in particular, the sponges and placozoans—animal bodies are differentiated into tissues; these include muscles, which enable locomotion, nerve tissues, which transmit signals and coordinate the body. There is an internal digestive chamber with either one opening or two openings. Nearly all animals make use of some form of sexual reproduction, they produce haploid gametes by meiosis.
These fuse to form zygotes, which develop via mitosis into a hollow sphere, called a blastula. In sponges, blastula larvae swim to a new location, attach to the seabed, develop into a new sponge. In most other groups, the blastula undergoes more complicated rearrangement, it first invaginates to form a gastrula with a digestive chamber and two separate germ layers, an external ectoderm and an internal endoderm. In most cases, a third germ layer, the mesoderm develops between them; these germ layers differentiate to form tissues and organs. Repeated instances of mating with a close relative during sexual reproduction leads to inbreeding depression within a population due to the increased prevalence of harmful recessive traits. Animals have evolved numerous mechanisms for avoiding close inbreeding. In some species, such as the splendid fairywren, females benefit by mating with multiple males, thus producing more offspring of higher genetic quality; some animals are capable of asexual reproduction, which results
Nicrophorus vespilloides is a burying beetle described by Johann Friedrich Wilhelm Herbst in 1783. This is one of the most well studied of the burying beetles with over 1,000 citations found via Google Scholar. What had been considered Nicrophorus vespilloides in mid and eastern Canada and northeastern USA was determined by Sikes et al. in 2016 to be a separate, overlooked sister species of Nicrophorus vespilloides, named by Kirby in 1837. This sister species, Nicrophorus hebes Kirby, is restricted to Sphagnum marshes. Nicrophorus vespilloides occurs throughout the northern Palearctic and northwestern Canada where it is found in open forest habitats; the restriction of its sister species N. hebes to bogs in the North America has been attributed to competition with its related congener, N. defodiens which in this area is found in forest habitats. N. hebes reproduces in bogs in North America and is never found in adjacent forested habitat in the Mer Bleue bog area near Ottawa, Canada. There are a number of phoretic mites that are associated with N. vespilloides.
These include Pelzneria nr. crenulata, Macrocheles merderius, Uroobovella nr. novasimilis and the largest mite Poecilochirus carabi. P. carabi is not attached by any physical means to N. vespilloides. When the males or females of N. vespilloides have finished breeding on a carcass the deutonymphs of P. carabi roam about the body of the beetles as they search for new carcasses to reproduce. It had been proposed that P. carabi deutonymphs, on arrival at a new carcass dismounted from the beetles and consumed fly eggs and larvae which would have competed for the beetle larvae for food. This relationship which benefited the beetles has been described as mutualistic. However, it has been shown that adults of P. carabi consume the eggs of N. vespilloides and that this has direct and negative effects on the reproduction of this beetle species. N. Vespilloides is used as a model organism in the study of social immunity