98-409: The Nematocera (the name meaning "thread-horns") are a suborder of elongated flies with thin, segmented antennae and mostly aquatic larvae . This group is paraphyletic and contains all flies except for species from suborder Brachycera (the name meaning "short-horns"), which includes more commonly known species such as the housefly or the common fruit fly . The equivalent clade to Nematocera
196-465: A dramatic effect on community structure. Hunting of sea otters, for example, is thought to have led indirectly to the extinction of the Steller's sea cow ( Hydrodamalis gigas ). While the keystone species concept has been used extensively as a conservation tool, it has been criticized for being poorly defined from an operational stance. It is difficult to experimentally determine what species may hold
294-428: A dramatic impact on the ecology of individual species or whole ecosystems. For instance, the replacement of an ant species by another (invasive) ant species has been shown to affect how elephants reduce tree cover and thus the predation of lions on zebras . A trophic level (from Greek troph , τροφή, trophē, meaning "food" or "feeding") is "a group of organisms acquiring a considerable majority of its energy from
392-412: A flattened body relative to the main populations that live in open savanna. The population that lives in an isolated rock outcrop hides in crevasses where its flattened body offers a selective advantage. Habitat shifts also occur in the developmental life history of amphibians, and in insects that transition from aquatic to terrestrial habitats. Biotope and habitat are sometimes used interchangeably, but
490-612: A higher one." Small scale patterns do not necessarily explain large scale phenomena, otherwise captured in the expression (coined by Aristotle) 'the sum is greater than the parts'. "Complexity in ecology is of at least six distinct types: spatial, temporal, structural, process, behavioral, and geometric." From these principles, ecologists have identified emergent and self-organizing phenomena that operate at different environmental scales of influence, ranging from molecular to planetary, and these require different explanations at each integrative level . Ecological complexity relates to
588-444: A keystone role in each ecosystem. Furthermore, food web theory suggests that keystone species may not be common, so it is unclear how generally the keystone species model can be applied. Complexity is understood as a large computational effort needed to piece together numerous interacting parts exceeding the iterative memory capacity of the human mind. Global patterns of biological diversity are complex. This biocomplexity stems from
686-408: A keystone species can result in a community collapse just as the removal of the keystone in an arch can result in the arch's loss of stability. Sea otters ( Enhydra lutris ) are commonly cited as an example of a keystone species because they limit the density of sea urchins that feed on kelp . If sea otters are removed from the system, the urchins graze until the kelp beds disappear, and this has
784-741: A legacy niche that was constructed before their time. Biomes are larger units of organization that categorize regions of the Earth's ecosystems, mainly according to the structure and composition of vegetation. There are different methods to define the continental boundaries of biomes dominated by different functional types of vegetative communities that are limited in distribution by climate, precipitation, weather, and other environmental variables. Biomes include tropical rainforest , temperate broadleaf and mixed forest , temperate deciduous forest , taiga , tundra , hot desert , and polar desert . Other researchers have recently categorized other biomes, such as
882-456: A length of up to 7 cm (2.8 in), is generally considered to be the largest fly in the world, while the smallest is Euryplatea nanaknihali , which at 0.4 mm (0.016 in) is smaller than a grain of salt. Brachycera are ecologically very diverse, with many being predatory at the larval stage and some being parasitic. Animals parasitised include molluscs , woodlice , millipedes , insects, mammals , and amphibians . Flies are
980-439: A linear successional route, changes might occur quickly or slowly over thousands of years before specific forest successional stages are brought about by biological processes. An ecosystem's area can vary greatly, from tiny to vast. A single tree is of little consequence to the classification of a forest ecosystem, but is critically relevant to organisms living in and on it. Several generations of an aphid population can exist over
1078-439: A macroscopic view of the system. While the notion of trophic levels provides insight into energy flow and top-down control within food webs, it is troubled by the prevalence of omnivory in real ecosystems. This has led some ecologists to "reiterate that the notion that species clearly aggregate into discrete, homogeneous trophic levels is fiction." Nonetheless, recent studies have shown that real trophic levels do exist, but "above
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#17327663028881176-405: A meal. For visual course control, flies' optic flow field is analyzed by a set of motion-sensitive neurons. A subset of these neurons is thought to be involved in using the optic flow to estimate the parameters of self-motion, such as yaw, roll, and sideward translation. Other neurons are thought to be involved in analyzing the content of the visual scene itself, such as separating figures from
1274-516: A mix of herbivores and predators). Omnivores do not fit neatly into a functional category because they eat both plant and animal tissues. It has been suggested that omnivores have a greater functional influence as predators because compared to herbivores, they are relatively inefficient at grazing. Trophic levels are part of the holistic or complex systems view of ecosystems. Each trophic level contains unrelated species that are grouped together because they share common ecological functions, giving
1372-401: A mobile head, with a pair of large compound eyes , and mouthparts designed for piercing and sucking (mosquitoes, black flies and robber flies), or for lapping and sucking in the other groups. Their wing arrangement gives them great manoeuvrability in flight, and claws and pads on their feet enable them to cling to smooth surfaces. Flies undergo complete metamorphosis ; the eggs are often laid on
1470-420: A new appreciation for the influence that organisms have on the ecosystem and evolutionary process. The term "niche construction" is more often used in reference to the under-appreciated feedback mechanisms of natural selection imparting forces on the abiotic niche. An example of natural selection through ecosystem engineering occurs in the nests of social insects , including ants, bees, wasps, and termites. There
1568-435: A population-level phenomenon, as with the migration routes followed by plants as they occupied northern post-glacial environments. Plant ecologists use pollen records that accumulate and stratify in wetlands to reconstruct the timing of plant migration and dispersal relative to historic and contemporary climates. These migration routes involved an expansion of the range as plant populations expanded from one area to another. There
1666-401: A region), and sites are classed either as sources or sinks. A site is a generic term that refers to places where ecologists sample populations, such as ponds or defined sampling areas in a forest. Source patches are productive sites that generate a seasonal supply of juveniles that migrate to other patch locations. Sink patches are unproductive sites that only receive migrants; the population at
1764-517: A seasonal influx of new immigrants. A dynamic metapopulation structure evolves from year to year, where some patches are sinks in dry years and are sources when conditions are more favorable. Ecologists use a mixture of computer models and field studies to explain metapopulation structure. Community ecology examines how interactions among species and their environment affect the abundance, distribution and diversity of species within communities. Johnson & Stinchcomb (2007) Community ecology
1862-456: A species describes the environment over which a species is known to occur and the type of community that is formed as a result. More specifically, "habitats can be defined as regions in environmental space that are composed of multiple dimensions, each representing a biotic or abiotic environmental variable; that is, any component or characteristic of the environment related directly (e.g. forage biomass and quality) or indirectly (e.g. elevation) to
1960-402: A specific habitat, such as a cave or a pond, and principles gleaned from small-scale studies are extrapolated to larger systems. Feeding relations require extensive investigations, e.g. into the gut contents of organisms, which can be difficult to decipher, or stable isotopes can be used to trace the flow of nutrient diets and energy through a food web. Despite these limitations, food webs remain
2058-446: A valuable tool in understanding community ecosystems. Food webs illustrate important principles of ecology : some species have many weak feeding links (e.g., omnivores ) while some are more specialized with fewer stronger feeding links (e.g., primary predators ). Such linkages explain how ecological communities remain stable over time and eventually can illustrate a "complete" web of life. The disruption of food webs may have
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#17327663028882156-438: A variety of hosts. The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme environments including glaciers ( Diamesa sp., Chironomidae ), hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil ( Helaeomyia petrolei ). Adult hoverflies (Syrphidae) are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline scavengers inside
2254-399: A wide array of interacting levels of organization spanning micro-level (e.g., cells ) to a planetary scale (e.g., biosphere ) phenomena . Ecosystems, for example, contain abiotic resources and interacting life forms (i.e., individual organisms that aggregate into populations which aggregate into distinct ecological communities). Because ecosystems are dynamic and do not necessarily follow
2352-485: Is Spilomyia longicornis , which is a fly but mimics a vespid wasp. Flies have a mobile head with a pair of large compound eyes on the sides of the head, and in most species, three small ocelli on the top. The compound eyes may be close together or widely separated, and in some instances are divided into a dorsal region and a ventral region, perhaps to assist in swarming behaviour. The antennae are well-developed but variable, being thread-like, feathery or comb-like in
2450-485: Is a central concept in the ecology of organisms and is sub-divided into the fundamental and the realized niche. The fundamental niche is the set of environmental conditions under which a species is able to persist. The realized niche is the set of environmental plus ecological conditions under which a species persists. The Hutchinsonian niche is defined more technically as a " Euclidean hyperspace whose dimensions are defined as environmental variables and whose size
2548-504: Is a function of the number of values that the environmental values may assume for which an organism has positive fitness ." Biogeographical patterns and range distributions are explained or predicted through knowledge of a species' traits and niche requirements. Species have functional traits that are uniquely adapted to the ecological niche. A trait is a measurable property, phenotype , or characteristic of an organism that may influence its survival. Genes play an important role in
2646-414: Is a larger taxonomy of movement, such as commuting, foraging, territorial behavior, stasis, and ranging. Dispersal is usually distinguished from migration because it involves the one-way permanent movement of individuals from their birth population into another population. In metapopulation terminology, migrating individuals are classed as emigrants (when they leave a region) or immigrants (when they enter
2744-461: Is an emergent homeostasis or homeorhesis in the structure of the nest that regulates, maintains and defends the physiology of the entire colony. Termite mounds, for example, maintain a constant internal temperature through the design of air-conditioning chimneys. The structure of the nests themselves is subject to the forces of natural selection. Moreover, a nest can survive over successive generations, so that progeny inherit both genetic material and
2842-404: Is another statistical approach that is often used in conservation research . Metapopulation models simplify the landscape into patches of varying levels of quality, and metapopulations are linked by the migratory behaviours of organisms. Animal migration is set apart from other kinds of movement because it involves the seasonal departure and return of individuals from a habitat. Migration is also
2940-438: Is greater than respiration) by photosynthesis or chemosynthesis . Heterotrophs are organisms that must feed on others for nourishment and energy (respiration exceeds production). Heterotrophs can be further sub-divided into different functional groups, including primary consumers (strict herbivores), secondary consumers ( carnivorous predators that feed exclusively on herbivores), and tertiary consumers (predators that feed on
3038-547: Is one of the major insect orders and of considerable ecological and human importance. Flies are major pollinators, second only to the bees and their Hymenopteran relatives. Flies may have been among the evolutionarily earliest pollinators responsible for early plant pollination . Fruit flies are used as model organisms in research, but less benignly, mosquitoes are vectors for malaria , dengue , West Nile fever , yellow fever , encephalitis , and other infectious diseases ; and houseflies , commensal with humans all over
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3136-590: Is the Park Grass Experiment , which was initiated in 1856. Another example is the Hubbard Brook study , which has been in operation since 1960. Holism remains a critical part of the theoretical foundation in contemporary ecological studies. Holism addresses the biological organization of life that self-organizes into layers of emergent whole systems that function according to non-reducible properties. This means that higher-order patterns of
3234-460: Is the natural science of the relationships among living organisms and their environment . Ecology considers organisms at the individual, population , community , ecosystem , and biosphere levels. Ecology overlaps with the closely related sciences of biogeography , evolutionary biology , genetics , ethology , and natural history . Ecology is a branch of biology , and is the study of abundance , biomass , and distribution of organisms in
3332-413: Is the biosphere: the total sum of ecosystems on the planet. Ecological relationships regulate the flux of energy, nutrients, and climate all the way up to the planetary scale. For example, the dynamic history of the planetary atmosphere's CO 2 and O 2 composition has been affected by the biogenic flux of gases coming from respiration and photosynthesis, with levels fluctuating over time in relation to
3430-522: Is the study of the interactions among a collection of species that inhabit the same geographic area. Community ecologists study the determinants of patterns and processes for two or more interacting species. Research in community ecology might measure species diversity in grasslands in relation to soil fertility. It might also include the analysis of predator-prey dynamics, competition among similar plant species, or mutualistic interactions between crabs and corals. These ecosystems, as we may call them, are of
3528-643: Is the whole Diptera, with Brachycera as a subclade. Families in Nematocera include mosquitoes , crane flies , gnats , black flies , and multiple families commonly known as midges . The Nematocera typically have fairly long, fine, finely-jointed antennae. In many species, such as most mosquitoes, the female antennae are more or less threadlike, but the males have spectacularly plumose antennae. The larvae of most families of Nematocera are aquatic, either free-swimming, rock-dwelling, plant-dwelling, or luticolous . Some families however, are not aquatic; for instance
3626-556: Is used for managing wildlife stocks and setting harvest quotas. In cases where basic models are insufficient, ecologists may adopt different kinds of statistical methods, such as the Akaike information criterion , or use models that can become mathematically complex as "several competing hypotheses are simultaneously confronted with the data." The concept of metapopulations was defined in 1969 as "a population of populations which go extinct locally and recolonize". Metapopulation ecology
3724-524: Is widely thought to be a member of Mecopterida , along with Lepidoptera (butterflies and moths), Trichoptera (caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly Strepsiptera (twisted-wing flies). Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been confirmed by molecular studies. Diptera were traditionally broken down into two suborders, Nematocera and Brachycera , distinguished by
3822-631: The Jurassic , some 180 million years ago. A third radiation took place among the Schizophora at the start of the Paleogene , 66 million years ago. The phylogenetic position of Diptera has been controversial. The monophyly of holometabolous insects has long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and Diptera, and it is the relationships between these groups which has caused difficulties. Diptera
3920-1071: The Mecopterida , alongside the Mecoptera , Siphonaptera , Lepidoptera and Trichoptera . The possession of a single pair of wings distinguishes most true flies from other insects with "fly" in their names. However, some true flies such as Hippoboscidae (louse flies) have become secondarily wingless. The cladogram represents the current consensus view. Hymenoptera (sawflies, wasps, ants, bees) [REDACTED] Raphidioptera (snakeflies) [REDACTED] Megaloptera (alderflies and allies) [REDACTED] Neuroptera (Lacewings and allies) [REDACTED] Coleoptera (beetles) [REDACTED] Strepsiptera (twisted-wing parasites) [REDACTED] Trichoptera (caddisflies) [REDACTED] Lepidoptera (butterflies, moths) [REDACTED] Diptera [REDACTED] Mecoptera (scorpionflies) [REDACTED] Siphonaptera (fleas) [REDACTED] The first true dipterans known are from
4018-666: The Middle Triassic (around 240 million years ago), and they became widespread during the Middle and Late Triassic . Modern flowering plants did not appear until the Cretaceous (around 140 million years ago), so the original dipterans must have had a different source of nutrition other than nectar . Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed on honeydew produced by sap-sucking bugs which were abundant at
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4116-742: The Strepsiptera . In contrast to the flies, the Strepsiptera bear their halteres on the mesothorax and their flight wings on the metathorax. Each of the fly's six legs has a typical insect structure of coxa, trochanter, femur, tibia and tarsus, with the tarsus in most instances being subdivided into five tarsomeres . At the tip of the limb is a pair of claws, and between these are cushion-like structures known as pulvilli which provide adhesion. Ecological Ecology (from Ancient Greek οἶκος ( oîkos ) 'house' and -λογία ( -logía ) 'study of')
4214-594: The Tipulidae tend to be soil-dwelling and the Mycetophilidae feed on fungi such as mushrooms. Unlike most of the Brachycera, the larvae of Nematocera have distinct heads with mouthparts that may be modified for filter feeding or chewing, depending on their lifestyles. The pupae are orthorrhaphous which means that adults emerge from the pupa through a straight, longitudinal seam in the dorsal surface of
4312-405: The abundance or biomass at each level. When the relative abundance or biomass of each species is sorted into its respective trophic level, they naturally sort into a 'pyramid of numbers'. Species are broadly categorized as autotrophs (or primary producers ), heterotrophs (or consumers ), and Detritivores (or decomposers ). Autotrophs are organisms that produce their own food (production
4410-436: The logistic equation by Pierre Verhulst : where N(t) is the number of individuals measured as biomass density as a function of time, t , r is the maximum per-capita rate of change commonly known as the intrinsic rate of growth, and α {\displaystyle \alpha } is the crowding coefficient, which represents the reduction in population growth rate per individual added. The formula states that
4508-570: The order Diptera , the name being derived from the Greek δι- di- "two", and πτερόν pteron "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced mechanosensory organs known as halteres , which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing more than 150,000 species including horse-flies , crane flies , hoverflies , mosquitoes and others. Flies have
4606-548: The thorax , bears the wings and contains the flight muscles on the second segment, which is greatly enlarged; the first and third segments have been reduced to collar-like structures, and the third segment bears the halteres , which help to balance the insect during flight. The third tagma is the abdomen consisting of 11 segments, some of which may be fused, and with the 3 hindmost segments modified for reproduction. Some Dipterans are mimics and can only be distinguished from their models by very careful inspection. An example of this
4704-768: The Brachycera includes broader, more robust flies with short antennae. Many nematoceran larvae are aquatic. There are estimated to be a total of about 19,000 species of Diptera in Europe, 22,000 in the Nearctic region, 20,000 in the Afrotropical region, 23,000 in the Oriental region and 19,000 in the Australasian region. While most species have restricted distributions, a few like the housefly ( Musca domestica ) are cosmopolitan. Gauromydas heros ( Asiloidea ), with
4802-551: The Earth and atmospheric conditions within a narrow self-regulating range of tolerance. Population ecology studies the dynamics of species populations and how these populations interact with the wider environment. A population consists of individuals of the same species that live, interact, and migrate through the same niche and habitat. A primary law of population ecology is the Malthusian growth model which states, "a population will grow (or decline) exponentially as long as
4900-1344: The Nematocera. The construction of a phylogenetic tree has been the subject of ongoing research. The following cladogram is based on the FLYTREE project. Ptychopteromorpha (phantom and primitive crane-flies) [REDACTED] Culicomorpha (mosquitoes, blackflies and midges) [REDACTED] Blephariceromorpha (net-winged midges, etc) [REDACTED] Bibionomorpha (gnats) [REDACTED] Psychodomorpha (drain flies, sand flies, etc) [REDACTED] Tipulomorpha (crane flies) [REDACTED] Stratiomyomorpha (soldier flies, etc) [REDACTED] Xylophagomorpha (stink flies, etc) [REDACTED] Tabanomorpha (horse flies, snipe flies, etc) [REDACTED] Nemestrinoidea [REDACTED] Asiloidea (robber flies, bee flies, etc) [REDACTED] Empidoidea (dance flies, etc) [REDACTED] Aschiza (in part) Phoroidea (flat-footed flies, etc) [REDACTED] Syrphoidea (hoverflies) [REDACTED] Hippoboscoidea (louse flies, etc) [REDACTED] Muscoidea (house flies, dung flies, etc) [REDACTED] Oestroidea (blow flies, flesh flies, etc) [REDACTED] Acalyptratae (marsh flies, etc) [REDACTED] Flies are often abundant and are found in almost all terrestrial habitats in
4998-519: The antennae and the maxillary palps bear the main olfactory receptors, while the gustatory receptors are in the labium, pharynx, feet, wing margins and female genitalia, enabling flies to taste their food by walking on it. The taste receptors in females at the tip of the abdomen receive information on the suitability of a site for ovipositing. Flies that feed on blood have special sensory structures that can detect infrared emissions, and use them to home in on their hosts, Many blood-sucking flies can detect
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#17327663028885096-744: The context of the environment. It encompasses life processes, interactions, and adaptations ; movement of materials and energy through living communities; successional development of ecosystems; cooperation, competition, and predation within and between species ; and patterns of biodiversity and its effect on ecosystem processes. Ecology has practical applications in conservation biology , wetland management, natural resource management ( agroecology , agriculture , forestry , agroforestry , fisheries , mining , tourism ), urban planning ( urban ecology ), community health , economics , basic and applied science , and human social interaction ( human ecology ). The word ecology ( German : Ökologie )
5194-404: The differences in antennae. The Nematocera are identified by their elongated bodies and many-segmented, often feathery antennae as represented by mosquitoes and crane flies. The Brachycera have rounder bodies and much shorter antennae. Subsequent studies have identified the Nematocera as being non-monophyletic with modern phylogenies placing the Brachycera within grades of groups formerly placed in
5292-431: The different families. The mouthparts are adapted for piercing and sucking, as in the black flies, mosquitoes and robber flies, and for lapping and sucking as in many other groups. Female horse-flies use knife-like mandibles and maxillae to make a cross-shaped incision in the host's skin and then lap up the blood that flows. The gut includes large diverticulae , allowing the insect to store small quantities of liquid after
5390-714: The diversity of life from genes to ecosystems and spans every level of biological organization. The term has several interpretations, and there are many ways to index, measure, characterize, and represent its complex organization. Biodiversity includes species diversity , ecosystem diversity , and genetic diversity and scientists are interested in the way that this diversity affects the complex ecological processes operating at and among these respective levels. Biodiversity plays an important role in ecosystem services which by definition maintain and improve human quality of life. Conservation priorities and management techniques require different approaches and considerations to address
5488-530: The dynamic resilience of ecosystems that transition to multiple shifting steady-states directed by random fluctuations of history. Long-term ecological studies provide important track records to better understand the complexity and resilience of ecosystems over longer temporal and broader spatial scales. These studies are managed by the International Long Term Ecological Network (LTER). The longest experiment in existence
5586-456: The ecology and evolution of plants and animals. Ecological theory has also been used to explain self-emergent regulatory phenomena at the planetary scale: for example, the Gaia hypothesis is an example of holism applied in ecological theory. The Gaia hypothesis states that there is an emergent feedback loop generated by the metabolism of living organisms that maintains the core temperature of
5684-434: The energy is transferred through a chain of organisms by consumption. The simplified linear feeding pathways that move from a basal trophic species to a top consumer is called the food chain . Food chains in an ecological community create a complex food web. Food webs are a type of concept map that is used to illustrate and study pathways of energy and material flows. Empirical measurements are generally restricted to
5782-460: The environment experienced by all individuals in the population remains constant." Simplified population models usually starts with four variables: death, birth, immigration , and emigration . An example of an introductory population model describes a closed population, such as on an island, where immigration and emigration does not take place. Hypotheses are evaluated with reference to a null hypothesis which states that random processes create
5880-451: The equilibrium, r / α {\displaystyle r/\alpha } as K , which is known as the "carrying capacity." Population ecology builds upon these introductory models to further understand demographic processes in real study populations. Commonly used types of data include life history , fecundity , and survivorship, and these are analyzed using mathematical techniques such as matrix algebra . The information
5978-749: The flies, principally in the family Cecidomyiidae (gall midges). Many flies (most importantly in the family Agromyzidae) lay their eggs in the mesophyll tissue of leaves with larvae feeding between the surfaces forming blisters and mines. Some families are mycophagous or fungus feeding. These include the cave dwelling Mycetophilidae (fungus gnats) whose larvae are the only diptera with bioluminescence. The Sciaridae are also fungus feeders. Some plants are pollinated by fungus feeding flies that visit fungus infected male flowers. The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint and shoe polish. The Exorista mella (Walker) fly are considered generalists and parasitoids of
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#17327663028886076-589: The flux of energy and matter through an environment. Ecosystems have biophysical feedback mechanisms that moderate processes acting on living ( biotic ) and abiotic components of the planet. Ecosystems sustain life-supporting functions and provide ecosystem services like biomass production (food, fuel, fiber, and medicine), the regulation of climate , global biogeochemical cycles , water filtration , soil formation , erosion control, flood protection, and many other natural features of scientific, historical, economic, or intrinsic value. The scope of ecology contains
6174-400: The former applies to a community's environment, whereas the latter applies to a species' environment. Definitions of the niche date back to 1917, but G. Evelyn Hutchinson made conceptual advances in 1957 by introducing a widely adopted definition: "the set of biotic and abiotic conditions in which a species is able to persist and maintain stable population sizes." The ecological niche
6272-544: The former relates only to the physical modifications of the habitat whereas the latter also considers the evolutionary implications of physical changes to the environment and the feedback this causes on the process of natural selection. Ecosystem engineers are defined as: "organisms that directly or indirectly modulate the availability of resources to other species, by causing physical state changes in biotic or abiotic materials. In so doing they modify, maintain and create habitats." The ecosystem engineering concept has stimulated
6370-518: The full ecological scope of biodiversity. Natural capital that supports populations is critical for maintaining ecosystem services and species migration (e.g., riverine fish runs and avian insect control) has been implicated as one mechanism by which those service losses are experienced. An understanding of biodiversity has practical applications for species and ecosystem-level conservation planners as they make management recommendations to consulting firms, governments, and industry. The habitat of
6468-527: The ground using motion parallax. The H1 neuron is responsible for detecting horizontal motion across the entire visual field of the fly, allowing the fly to generate and guide stabilizing motor corrections midflight with respect to yaw. The ocelli are concerned in the detection of changes in light intensity, enabling the fly to react swiftly to the approach of an object. Like other insects, flies have chemoreceptors that detect smell and taste, and mechanoreceptors that respond to touch. The third segments of
6566-408: The herbivore trophic level, food webs are better characterized as a tangled web of omnivores." A keystone species is a species that is connected to a disproportionately large number of other species in the food-web . Keystone species have lower levels of biomass in the trophic pyramid relative to the importance of their role. The many connections that a keystone species holds means that it maintains
6664-406: The human and oceanic microbiomes . To a microbe , the human body is a habitat and a landscape. Microbiomes were discovered largely through advances in molecular genetics , which have revealed a hidden richness of microbial diversity on the planet. The oceanic microbiome plays a significant role in the ecological biogeochemistry of the planet's oceans. The largest scale of ecological organization
6762-404: The interplay among ecological processes that operate and influence patterns at different scales that grade into each other, such as transitional areas or ecotones spanning landscapes. Complexity stems from the interplay among levels of biological organization as energy, and matter is integrated into larger units that superimpose onto the smaller parts. "What were wholes on one level become parts on
6860-412: The interplay of development and environmental expression of traits. Resident species evolve traits that are fitted to the selection pressures of their local environment. This tends to afford them a competitive advantage and discourages similarly adapted species from having an overlapping geographic range. The competitive exclusion principle states that two species cannot coexist indefinitely by living off
6958-431: The larval food-source and the larvae, which lack true limbs, develop in a protected environment, often inside their food source. Other species are ovoviviparous , opportunistically depositing hatched or hatching larvae instead of eggs on carrion , dung, decaying material, or open wounds of mammals. The pupa is a tough capsule from which the adult emerges when ready to do so; flies mostly have short lives as adults. Diptera
7056-402: The latter focuses on materials and energy fluxes. System behaviors must first be arrayed into different levels of the organization. Behaviors corresponding to higher levels occur at slow rates. Conversely, lower organizational levels exhibit rapid rates. For example, individual tree leaves respond rapidly to momentary changes in light intensity, CO 2 concentration, and the like. The growth of
7154-424: The lifespan of a single leaf. Each of those aphids, in turn, supports diverse bacterial communities. The nature of connections in ecological communities cannot be explained by knowing the details of each species in isolation, because the emergent pattern is neither revealed nor predicted until the ecosystem is studied as an integrated whole. Some ecological principles, however, do exhibit collective properties where
7252-432: The lower adjacent level (according to ecological pyramids ) nearer the abiotic source." Links in food webs primarily connect feeding relations or trophism among species. Biodiversity within ecosystems can be organized into trophic pyramids, in which the vertical dimension represents feeding relations that become further removed from the base of the food chain up toward top predators, and the horizontal dimension represents
7350-558: The most various kinds and sizes. They form one category of the multitudinous physical systems of the universe, which range from the universe as a whole down to the atom. Tansley (1935) Ecosystems may be habitats within biomes that form an integrated whole and a dynamically responsive system having both physical and biological complexes. Ecosystem ecology is the science of determining the fluxes of materials (e.g. carbon, phosphorus) between different pools (e.g., tree biomass, soil organic material). Ecosystem ecologists attempt to determine
7448-456: The nature of the unique physical environments that shapes the biodiversity within each. A more recent addition to ecosystem ecology are technoecosystems , which are affected by or primarily the result of human activity. A food web is the archetypal ecological network . Plants capture solar energy and use it to synthesize simple sugars during photosynthesis . As plants grow, they accumulate nutrients and are eaten by grazing herbivores , and
7546-419: The nests of social insects. Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases. Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of the fly, the head, bears the eyes, the antennae , and the mouthparts (the labrum, labium, mandible, and maxilla make up the mouthparts). The second tagma,
7644-406: The neural ganglia , and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder. Some flies such as the ectoparasitic Nycteribiidae and Streblidae are exceptional in having lost their wings and become flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are
7742-573: The niche is called the ecotope , which is defined as the full range of environmental and biological variables affecting an entire species. Organisms are subject to environmental pressures, but they also modify their habitats. The regulatory feedback between organisms and their environment can affect conditions from local (e.g., a beaver pond ) to global scales, over time and even after death, such as decaying logs or silica skeleton deposits from marine organisms. The process and concept of ecosystem engineering are related to niche construction , but
7840-531: The number of nitrogen fixers , can lead to disproportionate, perhaps irreversible, changes in the system properties." Biodiversity refers to the variety of life and its processes. It includes the variety of living organisms, the genetic differences among them, the communities and ecosystems in which they occur, and the ecological and evolutionary processes that keep them functioning, yet ever-changing and adapting. Noss & Carpenter (1994) Biodiversity (an abbreviation of "biological diversity") describes
7938-399: The observed data. In these island models, the rate of population change is described by: where N is the total number of individuals in the population, b and d are the per capita rates of birth and death respectively, and r is the per capita rate of population change. Using these modeling techniques, Malthus' population principle of growth was later transformed into a model known as
8036-406: The organization and structure of entire communities. The loss of a keystone species results in a range of dramatic cascading effects (termed trophic cascades ) that alters trophic dynamics, other food web connections, and can cause the extinction of other species. The term keystone species was coined by Robert Paine in 1969 and is a reference to the keystone architectural feature as the removal of
8134-468: The published work of George Perkins Marsh ("Man and Nature"). Within an ecosystem, organisms are linked to the physical and biological components of their environment to which they are adapted. Ecosystems are complex adaptive systems where the interaction of life processes form self-organizing patterns across different scales of time and space. Ecosystems are broadly categorized as terrestrial , freshwater , atmospheric, or marine . Differences stem from
8232-729: The pupal cuticle. The bodies and legs of most adult Nematocera are elongated, and many species have relatively long abdomens. Males of many species form mating swarms like faint pillars of smoke, competing for females that visit the cloud of males to find a mate. These families belong to the suborder Nematocera: Fly Housefly (Muscidae) (top left) Haematopota pluvialis (Tabanidae) (top right) Ctenophora pectinicornis (Tipulidae) (mid left) Ochlerotatus notoscriptus (Culicidae) (mid right) Milesia crabroniformis (Syrphidae) (bottom left) Holcocephala fusca (Asilidae) (bottom right) Nematocera ( paraphyletic ) (inc Eudiptera ) Brachycera Flies are insects of
8330-405: The raised concentration of carbon dioxide that occurs near large animals. Some tachinid flies (Ormiinae) which are parasitoids of bush crickets , have sound receptors to help them locate their singing hosts. Diptera have one pair of fore wings on the mesothorax and a pair of halteres , or reduced hind wings, on the metathorax . A further adaptation for flight is the reduction in number of
8428-502: The rate of change in population size ( d N ( t ) / d t {\displaystyle \mathrm {d} N(t)/\mathrm {d} t} ) will grow to approach equilibrium, where ( d N ( t ) / d t = 0 {\displaystyle \mathrm {d} N(t)/\mathrm {d} t=0} ), when the rates of increase and crowding are balanced, r / α {\displaystyle r/\alpha } . A common, analogous model fixes
8526-424: The same limiting resource ; one will always out-compete the other. When similarly adapted species overlap geographically, closer inspection reveals subtle ecological differences in their habitat or dietary requirements. Some models and empirical studies, however, suggest that disturbances can stabilize the co-evolution and shared niche occupancy of similar species inhabiting species-rich communities. The habitat plus
8624-568: The second largest group of pollinators after the Hymenoptera (bees, wasps and relatives). In wet and colder environments flies are significantly more important as pollinators. Compared to bees, they need less food as they do not need to provision their young. Many flowers that bear low nectar and those that have evolved trap pollination depend on flies. It is thought that some of the earliest pollinators of plants may have been flies. The greatest diversity of gall forming insects are found among
8722-399: The site will disappear unless rescued by an adjacent source patch or environmental conditions become more favorable. Metapopulation models examine patch dynamics over time to answer potential questions about spatial and demographic ecology. The ecology of metapopulations is a dynamic process of extinction and colonization. Small patches of lower quality (i.e., sinks) are maintained or rescued by
8820-524: The study of ecology into a conceptually manageable framework, the biological world is organized into a nested hierarchy , ranging in scale from genes , to cells , to tissues , to organs , to organisms , to species , to populations , to guilds , to communities , to ecosystems , to biomes , and up to the level of the biosphere . This framework forms a panarchy and exhibits non-linear behaviors; this means that "effect and cause are disproportionate, so that small changes to critical variables, such as
8918-432: The sum of the components explain the properties of the whole, such as birth rates of a population being equal to the sum of individual births over a designated time frame. The main subdisciplines of ecology, population (or community ) ecology and ecosystem ecology , exhibit a difference not only in scale but also in two contrasting paradigms in the field. The former focuses on organisms' distribution and abundance, while
9016-606: The time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues. The basal clades in the Diptera include the Deuterophlebiidae and the enigmatic Nymphomyiidae . Three episodes of evolutionary radiation are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in the Triassic , about 220 million years ago. Many lower Brachycera appeared in
9114-670: The tree responds more slowly and integrates these short-term changes. O'Neill et al. (1986) The scale of ecological dynamics can operate like a closed system, such as aphids migrating on a single tree, while at the same time remaining open about broader scale influences, such as atmosphere or climate. Hence, ecologists classify ecosystems hierarchically by analyzing data collected from finer scale units, such as vegetation associations , climate, and soil types , and integrate this information to identify emergent patterns of uniform organization and processes that operate on local to regional, landscape , and chronological scales. To structure
9212-415: The underlying causes of these fluxes. Research in ecosystem ecology might measure primary production (g C/m^2) in a wetland in relation to decomposition and consumption rates (g C/m^2/y). This requires an understanding of the community connections between plants (i.e., primary producers) and the decomposers (e.g., fungi and bacteria). The underlying concept of an ecosystem can be traced back to 1864 in
9310-439: The use of a location by the animal." For example, a habitat might be an aquatic or terrestrial environment that can be further categorized as a montane or alpine ecosystem. Habitat shifts provide important evidence of competition in nature where one population changes relative to the habitats that most other individuals of the species occupy. For example, one population of a species of tropical lizard ( Tropidurus hispidus ) has
9408-473: The world apart from Antarctica. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. More than 150,000 have been formally described and the actual species diversity is much greater, with the flies from many parts of the world yet to be studied intensively. The suborder Nematocera include generally small, slender insects with long antennae such as mosquitoes, gnats, midges and crane-flies, while
9506-607: The world, spread foodborne illnesses . Flies can be annoyances especially in some parts of the world where they can occur in large numbers, buzzing and settling on the skin or eyes to bite or seek fluids. Larger flies such as tsetse flies and screwworms cause significant economic harm to cattle. Blowfly larvae, known as gentles , and other dipteran larvae, known more generally as maggots , are used as fishing bait , as food for carnivorous animals, and in medicine in debridement , to clean wounds . Dipterans are holometabolans , insects that undergo radical metamorphosis. They belong to
9604-593: Was coined in 1866 by the German scientist Ernst Haeckel . The science of ecology as we know it today began with a group of American botanists in the 1890s. Evolutionary concepts relating to adaptation and natural selection are cornerstones of modern ecological theory . Ecosystems are dynamically interacting systems of organisms, the communities they make up, and the non-living ( abiotic ) components of their environment. Ecosystem processes, such as primary production , nutrient cycling , and niche construction , regulate
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