Gymnadenia odoratissima is a species of orchid
Moths comprise a group of insects related to butterflies, belonging to the order Lepidoptera. Most lepidopterans are moths, there are thought to be 160,000 species of moth, many of which have yet to be described. Most species of moth are nocturnal, but there are crepuscular and diurnal species. While the butterflies form a monophyletic group, the moths, comprising the rest of the Lepidoptera, do not. Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic: Microlepidoptera and Macrolepidoptera and Rhopalocera, Jugatae and Frenatae and Ditrysia. Although the rules for distinguishing moths from butterflies are not well established, one good guiding principle is that butterflies have thin antennae and have small balls or clubs at the end of their antennae. Moth antennae are feathery with no ball on the end; the divisions are named by this principle: "club-antennae" or "varied-antennae". The modern English word "moth" comes from Old English "moððe" from Common Germanic.
Its origins are related to the Old English "maða" meaning "maggot" or from the root of "midge" which until the sixteenth century was used to indicate the larva in reference to devouring clothes. Moth larvae, or caterpillars, make cocoons from which they emerge as grown moths with wings; some moth caterpillars dig holes in the ground, where they live until they are ready to turn into adult moths. Moths evolved long before butterflies, with fossils having been found that may be 190 million years old. Both types of Lepidoptera are thought to have evolved along with flowering plants because most modern species feed on flowering plants, both as adults and larvae. One of the earliest species thought to be a moth-ancestor is Archaeolepis mane, whose fossil fragments show scaled wings similar to caddisflies in their veining; some moths their caterpillars, can be major agricultural pests in many parts of the world. Examples include corn bollworms; the caterpillar of the gypsy moth causes severe damage to forests in the northeastern United States, where it is an invasive species.
In temperate climates, the codling moth causes extensive damage to fruit farms. In tropical and subtropical climates, the diamondback moth is the most serious pest of brassicaceous crops. In sub-Saharan Africa, the African sugarcane borer is a major pest of sugarcane and sorghum. Several moths in the family Tineidae are regarded as pests because their larvae eat fabric such as clothes and blankets made from natural proteinaceous fibers such as wool or silk, they are less to eat mixed materials containing some artificial fibers. There are some reports that they may be repelled by the scent of wood from juniper and cedar, by lavender, or by other natural oils. Naphthalene is considered more effective. Moth larvae may be killed by freezing the items which they infest for several days at a temperature below −8 °C. Despite being notorious for eating clothing, most moth adults do not eat at all. Many, like the Luna, Atlas, Promethea and other large moths do not have mouth parts. While there are many species of adult moths that do eat, there are many.
Some moths are farmed for their economic value. The most notable of these is the larva of the domesticated moth Bombyx mori, it is farmed for the silk. As of 2002, the silk industry produces more than 130 million kilograms of raw silk, worth about 250 million U. S. dollars, each year. Not all silk is produced by Bombyx mori. There are several species of Saturniidae that are farmed for their silk, such as the ailanthus moth, the Chinese oak silkmoth, the Assam silkmoth, the Japanese silk moth; the larvae of many species are used as food in Africa, where they are an important source of nutrition. The mopane worm, the caterpillar of Gonimbrasia belina, from the family Saturniidae, is a significant food resource in southern Africa. Another saturniid used. In one country alone, more than 30 species of moth larvae are harvested; some are sold not only in the local village markets, but are shipped by the ton from one country to another. Nocturnal insectivores feed on moths. Moths are eaten by some species of lizards, dogs and some bears.
Moth larvae are vulnerable to being parasitized by Ichneumonidae. Baculoviruses are parasite double-stranded DNA insect viruses that are used as biological control agents, they are members of the Baculoviridae, a family, restricted to insects. Most baculovirus isolates have been obtained in particular from Lepidoptera. There is evidence that ultrasound in the range emitted by bats causes flying moths to make evasive maneuvers because bats eat moths. Ultrasonic frequencies trigger a reflex action in the noctuid moth that causes it to drop a few inches in its flight to evade attack. Tiger moths emit clicks which can foil bats' echolocation; the fungus Ophiocordyceps sinensis infects the larvae of many different species of moths. Some studies indicate that ce
Entomophily or insect pollination is a form of pollination whereby pollen of plants but not only of flowering plants, is distributed by insects. Flowers pollinated by insects advertise themselves with bright colours, sometimes with conspicuous patterns leading to rewards of pollen and nectar. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, in some species pollen baskets on their hind legs; this required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups. Many plants, including flowering plants such as grasses, are instead pollinated by other mechanisms, such as by wind; the early spermatophytes were dependent on the wind to carry their pollen from one plant to another, it was around 125 to 115 million years ago that a new pollination strategy developed and angiosperms first appeared. Before that, insect involvement in pollination was limited to "pollination assistants", insects which inadvertently carried the pollen between plants by their movements.
The real relationship between plants and insects began in the Early Cretaceous, with beetle-pollinated gymnosperms. The morphology of the first fossil basal angiosperms is similar to modern-day plants that are fertilised by beetles, it seems that beetles led the way in insect pollination, followed by flies. Among the twelve living families of basal angiosperms, six are predominantly pollinated by flies, five by beetles and only one by bees. Traits such as sapromyophily have evolved independently in several unrelated angiosperm families. Wind and water pollination require the production of vast quantities of pollen because of the chancy nature of its deposition. If they are not to be reliant on the wind or water, plants need pollinators to move their pollen grains from one plant to another, they need pollinators to choose flowers of the same species, so they have evolved different lures to encourage specific pollinators to maintain fidelity to the same species. The attractions offered are nectar, pollen and oils.
The ideal pollinating insect is hairy, spends time exploring the flower so that it comes into contact with the reproductive structures. Many insects are pollinators bees, wasps, flies and beetles. On the other hand, some plants are generalists, being pollinated by insects in several orders. Entomophilous plant species have evolved mechanisms to make themselves more appealing to insects, e.g. brightly coloured or scented flowers, nectar, or appealing shapes and patterns. Pollen grains of entomophilous plants are larger than the fine pollens of anemophilous plants, which has to be produced in much larger quantities because such a high proportion is wasted; this is energetically costly, but in contrast, entomophilous plants have to bear the energetic costs of producing nectar. Butterflies and moths have hairy bodies and long proboscides which can probe deep into tubular flowers. Butterflies fly by day and are attracted to pink and purple flowers; the flowers are large and scented, the stamens are so-positioned that pollen is deposited on the insects while they feed on the nectar.
Moths are nocturnal and are attracted by night-blooming plants. The flowers of these are tubular, pale in colour and fragrant only at night. Hawkmoths tend to visit larger flowers and hover. Other moths land on the smaller flowers, which may be aggregated into flowerheads, their energetic needs are not so great as those of hawkmoths and they are offered smaller quantities of nectar. Inflorescences pollinated by beetles tend to be flat with open corollas or small flowers clustered in a head with multiple, projecting anthers that shed pollen readily; the flowers are green or pale-coloured, scented with fruity or spicy aromas, but sometimes with odours of decaying organic matter. Some, like the giant water lily, include traps designed to retain the beetles in contact with the reproductive parts for longer periods. Unspecialised flies with short proboscides are found visiting primitive flowers with accessible nectar. More specialised flies like syrphids and tabanids can visit more advanced blooms, but their purpose is to nourish themselves, any transfer of pollen from one flower to another happens haphazardly.
The small size of many flies is made up for by their abundance, however they are unreliable pollinators as they may bear incompatible pollen, lack of suitable breeding habitats may limit their activities. Some Pterostylis orchids are pollinated by midges unique to each species. A decline, for whatever reason, to one side of this partnership can be catastrophic for the other. Flowers pollinated by bees and wasps vary in shape and size. Yellow or blue plants are visited, flowers may have ultra-violet nectar guides, that help the insect to find the nectary; some flowers, like sage or pea, have lower lips that will only open when sufficiently heavy insects, such as bees, land on them. With the lip depressed, the anthers may bow down to deposit pollen on the insect's back. Other flowers, like tomato, may only liberate their pollen by buzz pollination, a technique in which a bumblebee will cling on to a flower while vibrating its flight muscles
Large yellow underwing
The large yellow underwing is a moth, the type species for the family Noctuidae. It is an abundant species throughout the Palearctic ecozone, one of the most common and most familiar moths of the region. In some years the species is migratory with large numbers appearing in marginal parts of the range, it is present in Europe, North Africa, Canary Islands, Middle East, Iraq, Afghanistan, northwest India, Novosibirsk Oblast, Caucasus and Central Asia. It was introduced into North America at Nova Scotia. Since it has increased its range and has been recorded for Maine in 1985, spread throughout the northeast from Vermont and Massachusetts to New Hampshire, New York and Connecticut, it was first recorded in Pennsylvania in 1998, North Carolina and west to Colorado, California, British Columbia and Alaska and Ontario. This is a quite heavy moth with a wingspan of 50 -- 60 mm; the forewings are quite variable from light brown to black. The darker individuals have a pale streak along the costa; the hindwings are bright orange-yellow with a black sub-terminal band.
As with other Noctua species, this contrast of bland-on-land and bright-in-flight is used to confuse potential predators. This species flies at night from July to September and is attracted to light, sometimes in huge numbers, it will visit flowers such as Buddleia and red valerian. The larva is brown with two rows of black dashes along the back; this is one of the notorious "cutworms", causing fatal damage at the base of any herbaceous plant, sometimes severing it completely. This ubiquitous species is considered as a garden pest; the species feeds on mild days throughout the winter. ^ The flight season refers to the British Isles. This may vary in other parts of the range. See Robinson, G. S. et al. Chinery, Michael. Collins Guide to the Insects of Britain and Western Europe. Skinner, Bernard. Colour Identification Guide to Moths of the British Isles. Lepiforum Noctua pronuba at funet.fi Fauna Europaea
The Dinaric Alps commonly Dinarides, are a mountain range in Southern and Southeastern Europe, separating the continental Balkan Peninsula from the Adriatic Sea. They stretch from Italy in the northwest through Slovenia, Croatia and Herzegovina, Montenegro, Kosovo to Albania in the southeast; the Dinaric Alps extend for 645 kilometres along the Western Balkan Peninsula from the Julian Alps to the northwest in Italy, downwards to the Šar and Korab massif, where their direction changes. The Albanian Alps, or Prokletije, is the highest section of the entire Dinaric Alps. Maja Jezercë is the highest peak and is located in Albania, standing at 2,694 metres above the Adriatic; the Dinaric Alps are one of the most rugged and extensively mountainous areas of Europe, alongside the Caucasus Mountains, Alps and Scandinavian Mountains. They are formed of Mesozoic and Cenozoic sedimentary rocks of dolomite, limestone and conglomerates formed by seas and lakes that once covered the area. During the Alpine earth movements that occurred 50–100 million years ago, immense lateral pressures folded and overthrust the rocks in a great arc around the old rigid block of the northeast.
The Dinaric Alps were thrown up in more or less parallel ranges, stretching like necklaces from the Julian Alps as far as northern Albania and Kosovo, where the mountainous terrain subsides to make way for the waters of the Drin River and the plains of Kosovo. The Dinarides are named after Mount Dinara, a prominent peak in the center of the mountain range on the border with the Dalmatian part of Croatia and Bosnia and Herzegovina; the chain is called Alpet Dinaride or Alpet Dinarike in Albanian, Dinaridi/Динариди in Serbo-Croatian, Dinarsko gorstvo in Slovene and Alpi Dinariche in Italian. The Mesozoic limestone forms a distinctive region of the Balkans, notable for features such as the Karst, which has given its name to all such terrains of limestone eroded by groundwater; the Dinarides are known for being composed of karst — limestone rocks — as is Dinara, the mountain for which they were named. The Quaternary ice ages had little direct geologic influence on the Balkans. No permanent ice caps existed, there is little evidence of extensive glaciation.
Only the highest summits of Durmitor and Prenj have glacial valleys and moraines as low as 600 m. However, in the Prokletije, a range on the northern Albanian border that runs east to west, there is evidence of major glaciation. One geological feature of great importance to the present-day landscape of the Dinarides must be considered in more detail: that of the limestone mountains with their attendant faulting, they are hard and slow to erode, persist as steep jagged escarpments, through which steep-sided gorges and canyons are cleft by the rivers draining the higher slopes. The submerged western Dinaric Alps form the numerous islands and harbors along the Croatian coast; the most extensive example of limestone mountains in Europe are those of the Karst of the Dinaric Alps. Here, all the characteristic features are encountered again and again as one travels through this wild and underpopulated country. Limestone is a porous rock, yet hard and resistant to erosion. Water is the most important corrosive force, dissolving the limestone by chemical action of its natural acidity.
As it percolates down through cracks in the limestone it opens up fissures and channels of considerable depth, so that whole systems of underground drainage develop. During subsequent millennia these work deeper, leaving in their wake enormous waterless caverns and grottoes and forming underground labyrinths of channels and shafts; the roofs of some of these caverns may fall in, to produce great perpendicular-sided gorges, exposing the water to the surface once more. The Dinaric rivers carved many canyons characteristic for Dinaric Alps, in particular karst. Among largest and most well known are the Neretva, the Rakitnica, the Prača, the Drina, the Sutjeska, the Vrbas, the Piva, the Tara, the Komarnica, the Morača, the Cem/Ciijevna, the Lim, the Drin. Only along the Dinaric gorges is communication possible across the Karst, roads and railways tunnel through precipitous cliffs and traverse narrow ledges above roaring torrents. A number of springs and rivers rise in the Dinaric range, including Jadro Spring noted for having been the source of water for Diocletian's Palace at Split.
At the same time, the purity of these rocks is such that the rivers are crystal clear, there is little soil-making residue. Water quality testing of the Jadro River, for example, indicates the low pollutant levels present. Rock faces are bare of vegetation and glaring white, but what little soil there is may collect in the hollows and support lush lime-tolerant vegetation, or yield narrow strips of cultivation. Ruins of fortresses dot the mountainous landscape, evidence of centuries of war and the refuge the Dinaric Alps have provided to various armed forces. During the Roman period, the Dinarides provided shelter to the Illyrians resisting Roman conquest of the Balkans, which began with the conquest of the eastern Adriatic coast in the 3rd century BC. Rome conquered the whole of Illyria in 168 BC, but these mountains sheltered Illyrian resistance forces for many years until the area's complete subjugation by 14 AD. More the Ottoman Empire failed to subjugate the mountainous areas
The flowering plants known as angiosperms, Angiospermae or Magnoliophyta, are the most diverse group of land plants, with 64 orders, 416 families 13,164 known genera and c. 369,000 known species. Like gymnosperms, angiosperms are seed-producing plants. However, they are distinguished from gymnosperms by characteristics including flowers, endosperm within the seeds, the production of fruits that contain the seeds. Etymologically, angiosperm means a plant; the term comes from the Greek words sperma. The ancestors of flowering plants diverged from gymnosperms in the Triassic Period, 245 to 202 million years ago, the first flowering plants are known from 160 mya, they diversified extensively during the Early Cretaceous, became widespread by 120 mya, replaced conifers as the dominant trees from 100 to 60 mya. Angiosperms differ from other seed plants in several ways, described in the table below; these distinguishing characteristics taken together have made the angiosperms the most diverse and numerous land plants and the most commercially important group to humans.
Angiosperm stems are made up of seven layers. The amount and complexity of tissue-formation in flowering plants exceeds that of gymnosperms; the vascular bundles of the stem are arranged such that the phloem form concentric rings. In the dicotyledons, the bundles in the young stem are arranged in an open ring, separating a central pith from an outer cortex. In each bundle, separating the xylem and phloem, is a layer of meristem or active formative tissue known as cambium. By the formation of a layer of cambium between the bundles, a complete ring is formed, a regular periodical increase in thickness results from the development of xylem on the inside and phloem on the outside; the soft phloem becomes crushed, but the hard wood persists and forms the bulk of the stem and branches of the woody perennial. Owing to differences in the character of the elements produced at the beginning and end of the season, the wood is marked out in transverse section into concentric rings, one for each season of growth, called annual rings.
Among the monocotyledons, the bundles are more numerous in the young stem and are scattered through the ground tissue. They once formed the stem increases in diameter only in exceptional cases; the characteristic feature of angiosperms is the flower. Flowers show remarkable variation in form and elaboration, provide the most trustworthy external characteristics for establishing relationships among angiosperm species; the function of the flower is to ensure fertilization of the ovule and development of fruit containing seeds. The floral apparatus may arise terminally from the axil of a leaf; as in violets, a flower arises singly in the axil of an ordinary foliage-leaf. More the flower-bearing portion of the plant is distinguished from the foliage-bearing or vegetative portion, forms a more or less elaborate branch-system called an inflorescence. There are two kinds of reproductive cells produced by flowers. Microspores, which will divide to become pollen grains, are the "male" cells and are borne in the stamens.
The "female" cells called megaspores, which will divide to become the egg cell, are contained in the ovule and enclosed in the carpel. The flower may consist only of these parts, as in willow, where each flower comprises only a few stamens or two carpels. Other structures are present and serve to protect the sporophylls and to form an envelope attractive to pollinators; the individual members of these surrounding structures are known as petals. The outer series is green and leaf-like, functions to protect the rest of the flower the bud; the inner series is, in general, white or brightly colored, is more delicate in structure. It functions to attract bird pollinators. Attraction is effected by color and nectar, which may be secreted in some part of the flower; the characteristics that attract pollinators account for the popularity of flowers and flowering plants among humans. While the majority of flowers are perfect or hermaphrodite, flowering plants have developed numerous morphological and physiological mechanisms to reduce or prevent self-fertilization.
Heteromorphic flowers have short carpels and long stamens, or vice versa, so animal pollinators cannot transfer pollen to the pistil. Homomorphic flowers may employ a biochemical mechanism called self-incompatibility to discriminate between self and non-self pollen grains. In other species, the male and female parts are morphologically separated, developing on different flowers; the botanical term "Angiosperm", from the Ancient Greek αγγείον, angeíon and σπέρμα, was coined in the form Angiospermae by Paul Hermann in 1690, as the name of one of his primary divisions of the plant kingdom. This included flowering plants possessing seeds enclosed in capsules, distinguished from his Gymnospermae, or flowering plants with achenial or schizo-carpic fruits, the whole fruit or each of its pieces being here regarded as a seed and naked; the term and its antonym were maintained by Carl Linnaeus with the same sense, but with restricted application, in the names of the orders of his class Didynamia. Its use with any
A mycorrhiza is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in its root system. Mycorrhizae play important roles in soil biology and soil chemistry. In a mycorrhizal association, the fungus colonizes the host plant's root tissues, either intracellularly as in arbuscular mycorrhizal fungi, or extracellularly as in ectomycorrhizal fungi; the association is mutualistic, but in particular species or in particular circumstances, mycorrhizae may be variously parasitic in the host plants. A mycorrhiza is a symbiotic association between a fungus; the plant makes organic molecules such as sugars by photosynthesis and supplies them to the fungus, the fungus supplies to the plant water and mineral nutrients, such as phosphorus, taken from the soil. Mycorrhizas are located in the roots of vascular plants, but mycorrhiza-like associations occur in bryophytes and there is fossil evidence that early land plants that lacked roots formed arbuscular mycorrhizal associations.
Most plant species form mycorrhizal associations, though some families like Brassicaceae and Chenopodiaceae cannot. Different forms for the association are detailed in the next section; the most common is the arbuscular type, present in 70% of plant species, including many crop plants such as wheat and rice. Mycorrhizas are divided into ectomycorrhizas and endomycorrhizas; the two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual cells within the root, while the hyphae of endomycorrhizal fungi penetrate the cell wall and invaginate the cell membrane. Endomycorrhiza includes arbuscular and orchid mycorrhiza, while arbutoid mycorrhizas can be classified as ectoendomycorrhizas. Monotropoid mycorrhizas form a special category. Ectomycorrhizas, or EcM, are symbiotic associations between the roots of around 10% of plant families woody plants including the birch, eucalyptus, oak and rose families and fungi belonging to the Basidiomycota and Zygomycota.
Some EcM fungi, such as many Leccinum and Suillus, are symbiotic with only one particular genus of plant, while other fungi, such as the Amanita, are generalists that form mycorrhizas with many different plants. An individual tree may have 15 or more different fungal EcM partners at one time. Thousands of ectomycorrhizal fungal species hosted in over 200 genera. A recent study has conservatively estimated global ectomycorrhizal fungal species richness at 7750 species, although, on the basis of estimates of knowns and unknowns in macromycete diversity, a final estimate of ECM species richness would be between 20,000 and 25,000. Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and a Hartig net of hyphae surrounding the plant cells within the root cortex. In some cases the hyphae may penetrate the plant cells, in which case the mycorrhiza is called an ectendomycorrhiza. Outside the root, ectomycorrhizal extramatrical mycelium forms an extensive network within the soil and leaf litter.
Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from paper birch trees into Douglas-fir trees thereby promoting succession in ecosystems; the ectomycorrhizal fungus Laccaria bicolor has been found to lure and kill springtails to obtain nitrogen, some of which may be transferred to the mycorrhizal host plant. In a study by Klironomos and Hart, Eastern White Pine inoculated with L. bicolor was able to derive up to 25% of its nitrogen from springtails. When compared to non-mycorrhizal fine roots, ectomycorrhizae may contain high concentrations of trace elements, including toxic metals or chlorine; the first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete L. bicolor, has been published. An expansion of several multigene families occurred in this fungus, suggesting that adaptation to symbiosis proceeded by gene duplication. Within lineage-specific genes those coding for symbiosis-regulated secreted proteins showed an up-regulated expression in ectomycorrhizal root tips suggesting a role in the partner communication.
L. bicolor is lacking enzymes involved in the degradation of plant cell wall components, preventing the symbiont from degrading host cells during the root colonisation. By contrast, L. bicolor possesses expanded multigene families associated with hydrolysis of bacterial and microfauna polysaccharides and proteins. This genome analysis revealed the dual saprotrophic and biotrophic lifestyle of the mycorrhizal fungus that enables it to grow within both soil and living plant roots; this type of mycorrhiza involves plants of the Ericaceae subfamily Arbutoideae. It is however different from ericoid mycorrhiza and resembles ectomycorrhiza, both functionally and in terms of the fungi involved; the difference to ectomycorrhiza is that some hyphae penetrate into the root cells, making this type of mycorrhiza an ectendomycorrhiza. Endomycorrhizas are variable and have been further classified as arbuscular, arbutoid and orchid mycorrhizas. Arbuscular mycorrhizas, or AM, are mycorrhizas whose hyphae penetrate plant cells, producing structures that are either balloon-like or dichotomously branching invaginations as a means of nutrient exchange.
The fungal hyphae invaginate the cell membrane. The struct