document_title: Earthworm

question_text: what is the last segment of earthworm called and what organ does it contain

root

text: Earthworm

text: Anatomy

text: Form and function

text: Depending on the species, an adult earthworm can be from 10 mm (0.39 in) long and 1 mm (0.039 in) wide to 3 m (9.8 ft) long and over 25 mm (0.98 in) wide, but the typical Lumbricus terrestris grows to about 360 mm (14 in) long.[4] Probably the longest worm on confirmed records is Amynthas mekongianus that extends up to 3 m (10 ft) [5] in the mud along the banks of the 4,350 km (2,703 mi) Mekong River in Southeast Asia.

tag: p

text: From front to back, the basic shape of the earthworm is a cylindrical tube, divided into a series of segments (called metamerisms) that compartmentalize the body. Furrows are generally[6] externally visible on the body demarking the segments; dorsal pores and nephridiopores exude a fluid that moistens and protects the worm's surface, allowing it to breathe. Except for the mouth and anal segments, each segment carries bristle-like hairs called lateral setae[7] used to anchor parts of the body during movement;[8] species may have four pairs of setae on each segment or more than eight sometimes forming a complete circle of setae per segment.[7] Special ventral setae are used to anchor mating earthworms by their penetration into the bodies of their mates.[citation needed]

text: Generally, within a species, the number of segments found is consistent across specimens, and individuals are born with the number of segments they will have throughout their lives. The first body segment (segment number 1) features both the earthworm's mouth and, overhanging the mouth, a fleshy lobe called the prostomium, which seals the entrance when the worm is at rest, but is also used to feel and chemically sense the worm's surroundings. Some species of earthworm can even use the prehensile prostomium to grab and drag items such as grasses and leaves into their burrow.

text: An adult earthworm develops a belt-like glandular swelling, called the clitellum, which covers several segments toward the front part of the animal. This is part of the reproductive system and produces egg capsules. The posterior is most commonly cylindrical like the rest of the body, but depending on the species, may also be quadrangular, octagonal, trapezoidal, or flattened. The last segment is called the periproct; the earthworm's anus, a short vertical slit, is found on this segment.[7]

text: The exterior of an individual segment is a thin cuticle over skin, commonly pigmented red to brown, which has specialized cells that secrete mucus over the cuticle to keep the body moist and ease movement through soil. Under the skin is a layer of nerve tissue, and two layers of muscles—a thin outer layer of circular muscle, and a much thicker inner layer of longitudinal muscle.[9] Interior to the muscle layer is a fluid-filled chamber called a coelom[10] that by its pressurization provides structure to the worm's boneless body. The segments are separated from each other by septa (the plural of "septum")[11] which are perforated transverse walls, allowing the coelomic fluid to pass between segments.[12] A pair of structures called nephrostomes are located at the back of each septum; a nephric tubule leads from each nephrostome through the septum and into the following segment. This tubule then leads to the main body fluid filtering organ, the nephridium or metanephridium, which removes metabolic waste from the coelomic fluid and expels it through pores called nephridiopores on the worm's sides; usually two nephridia (sometimes more) are found in most segments.[13] At the center of a worm is the digestive tract, which runs straight through from mouth to anus without coiling, and is flanked above and below by blood vessels (the dorsal blood vessel and the ventral blood vessel as well as a subneural blood vessel) and the ventral nerve cord, and is surrounded in each segment by a pair of pallial blood vessels that connect the dorsal to the subneural blood vessels.

text: Many earthworms can eject coelomic fluid through pores in the back in response to stress; Australian Didymogaster sylvaticus (known as the "blue squirter earthworm") can squirt fluid as high as 30 cm (12 in).[12]

text: Nervous system

text: The earthworm's nervous system has three parts: the central nervous system (CNS), peripheral nervous system and the sympathetic nervous system.[14][15]

text: Central nervous system

text: The CNS consists of a bilobed brain (cerebral ganglia, or supra-pharyngeal ganglion), sub-pharyngeal ganglia, circum-pharyngeal connectives and a ventral nerve cord.

text: Earthworms' brains consist of a pair of pear-shaped cerebral ganglia. These are located in the dorsal side of the alimentary canal in the third segment, in a groove between the buccal cavity and pharynx.

text: A pair of circum-pharyngeal connectives from the brain encircle the pharynx and then connect with a pair of sub-pharyngeal ganglia located below the pharynx in the fourth segment. This arrangement means the brain, sub-pharyngeal ganglia and the circum-pharyngeal connectives form a nerve ring around the pharynx.

text: The ventral nerve cord (formed by nerve cells and nerve fibres) begins at the sub-pharyngeal ganglia and extends below the alimentary canal to the most posterior body segment. The ventral nerve cord has a swelling, or ganglion, in each segment, i.e. a segmental ganglion, which occurs from the fifth to the last segment of the body. There are also three giant axons, one medial giant axon (MGA) and two lateral giant axons (LGAs) on the mid-dorsal side of the ventral nerve cord. The MGA is 0.07 mm in diameter and transmits in an anterior-posterior direction at a rate of 32.2 m/s. The LGAs are slightly wider at 0.05 mm in diameter and transmit in a posterior-anterior direction at 12.6 m/s. The two LGAs are connected at regular intervals along the body and are therefore considered one giant axon.[16][17]

text: Peripheral nervous system

text:
Eight to ten nerves arise from the cerebral ganglia to supply the prostomium, buccal chamber and pharynx.
Three pairs of nerves arise from the subpharyangeal ganglia to supply the 2nd, 3rd and 4th segment.
Three pairs of nerves extend from each segmental ganglia to supply various structures of the segment.

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text: Sympathetic nervous system

text: The sympathetic nervous system consists of nerve plexuses in the epidermis and alimentary canal. (A plexus is a web of nerve cells connected together in a two dimensional grid.) The nerves that run along the body wall pass between the outer circular and inner longitudinal muscle layers of the wall. They give off branches that form the intermuscular plexus and the subepidermal plexus. These nerves connect with the circumpharyngeal connective.

text: Movement

text: On the surface, crawling speed varies both within and among individuals. Earthworms crawl faster primarily by taking longer "strides" and a greater frequency of strides. Larger Lumbricus terrestris worms crawl at a greater absolute speed than smaller worms. They achieve this by taking slightly longer strides but with slightly lower stride frequencies.[18]

text: Touching an earthworm, which causes a "pressure" response as well as (often) a response to the dehydrating quality of the salt on human skin (toxic to earthworms), stimulates the subepidermal nerve plexus which connects to the intermuscular plexus and causes the longitudinal muscles to contact, thereby the writhing movements when we pick up an earthworm. This behaviour is a reflex and does not require the CNS; it occurs even if the nerve cord is removed. Each segment of the earthworm has its own nerve plexus. The plexus of one segment is not connected directly to that of adjacent segments. The nerve cord is required to connect the nervous systems of the segments.[19]

text: The giant axons carry the fastest signals along the nerve cord. These are emergency signals that initiate reflex escape behaviours. The larger dorsal giant axon conducts signals the fastest, from the rear to the front of the animal. If the rear of the worm is touched, a signal is rapidly sent forwards causing the longitudinal muscles in each segment to contract. This causes the worm shorten very quickly as an attempt to escape from a predator or other potential threat. The two medial giant axons connect with each other and send signals from the front to the rear. Stimulation of these causes the earthworm to very quickly retreat (perhaps contracting into its burrow to escape a bird).

text: The presence of a nervous system is essential for an animal to be able to experience nociception or pain. However, other physiological capacities are also required such as opioid sensitivity and central modulation of responses by analgesics.[20] Enkephalin and α-endorphin-like substances have been found in earthworms. Injections of naloxone (an opioid antagonist) inhibit the escape responses of earthworms. This indicates that opioid substances play a role in sensory modulation, similar to that found in many vertebrates.[21]

text: Senses

text: Photosensitivity

text: Earthworms do not have eyes (although some worms do), however, they do have specialised photosensitive cells called "light cells of Hess". These photoreceptor cells have a central intracellular cavity (phaosome) filled with microvilli. As well as the microvilli, there are several sensory cilia in the phaosome which are structurally independent of the microvilli.[22] The photoreceptors are distributed in most parts of the epidermis but are more concentrated on the back and sides of the worm. A relatively small number occur on the ventral surface of the 1st segment. They are most numerous in the prostomium and reduce in density in the first three segments; they are very few in number past the third segment.[19]

text: Digestive system

text: The gut of the earthworm is a straight tube which extends from the worm's mouth to its anus. It is differentiated into a buccal cavity (generally running through the first one or two segments of the earthworm), pharynx (running generally about four segments in length), esophagus, crop, gizzard (usually) and intestine.[23]

text: Food enters in the mouth. The pharynx acts as a suction pump; its muscular walls draw in food. In the pharynx, the pharyngeal glands secrete mucus. Food moves into the esophagus, where calcium (from the blood and ingested from previous meals) is pumped in to maintain proper blood calcium levels in the blood and food pH. From there the food passes into the crop and gizzard. In the gizzard, strong muscular contractions grind the food with the help of mineral particles ingested along with the food. Once through the gizzard, food continues through the intestine for digestion. The intestine secretes Pepsin to digest proteins, Amylase to digest polysaccharides, Cellulase to digest cellulose, and lipase to digest fats.[2] Earthworms use, in addition to the digestive proteins, a class of surface active compounds called drilodefensins, which help digest plant material.[24] Instead of being coiled like a mammalian intestine, an earthworm's intestine increases surface area to increase nutrient absorption by having many folds running along its length. The intestine has its own pair of muscle layers like the body, but in reverse order—an inner circular layer within an outer longitudinal layer.[25]

text: Circulatory system

text: The earthworm has a dual circulatory system in which both the coelomic fluid and a closed circulatory system carry the food, waste, and respiratory gases. The closed circulatory system has five main blood vessels: the dorsal (top) vessel, which runs above the digestive tract; the ventral (bottom) vessel, which runs below the digestive tract; the subneural vessel, which runs below the ventral nerve cord; and two lateroneural vessels on either side of the nerve cord.[26] The dorsal vessel moves the blood forward, while the other four longitudinal vessels carry the blood rearward. In segments seven through eleven, a pair of aortic arches rings the coelom and acts as hearts, pumping the blood to the ventral vessel that acts as the aorta. The blood consists of ameboid cells and hemoglobin dissolved in the plasma. The second circulatory system derives from the cells of the digestive system that line the coelom. As the digestive cells become full, they release non-living cells of fat into the fluid-filled coelom, where they float freely but can pass through the walls separating each segment, moving food to other parts and assist in wound healing.[27]

text: Excretory system

text: The excretory system contains a pair of nephridia in every segment, except for the first three and the last ones.[28] The three types of nephridia are: integumentary, septal, and pharyngeal. The integumentary nephridia lie attached to the inner side of the body wall in all segments except the first two. The septal nephridia are attached to both sides of the septa behind the 15th segment. The pharyngeal nephridia are attached to fourth, fifth and sixth segments.[28] The waste in the coelom fluid from a forward segment is drawn in by the beating of cilia of the nephrostome. From there it is carried through the septum (wall) via a tube which forms a series of loops entwined by blood capillaries that also transfer waste into the tubule of the nephrostome. The excretory wastes are then finally discharged through a pore on the worm's side.[29]

text: Respiration

text: Earthworms have no special respiratory organs. Gases are exchanged through the moist skin and capillaries, where the oxygen is picked up by the hemoglobin dissolved in the blood plasma and carbon dioxide is released. Water, as well as salts, can also be moved through the skin by active transport.

text: Reproduction

text: Mating occurs on the surface, most often at night. Earthworms are hermaphrodites; that is, they have both male and female sexual organs. The sexual organs are located in segments 9 to 15. Earthworms have one or two pairs of testes contained within sacs. The two or four pairs of seminal vesicles produce, store and release the sperm via the male pores. Ovaries and oviducts in segment 13 release eggs via female pores on segment 14, while sperm is expelled from segment 15. One or more pairs of spermathecae are present in segments 9 and 10 (depending on the species) which are internal sacs that receive and store sperm from the other worm during copulation. As a result, segment 15 of one worm exudes sperm into segments 9 and 10 with its storage vesicles of its mate. Some species use external spermatophores for sperm transfer.

text: In Hormogaster samnitica and Hormogaster elisae transcriptome DNA libraries were sequenced and two sex pheromones, Attractin and Temptin, were detected in all tissue samples of both species.[30] Sex pheromones are probably important in earthworms because they live in an environment where chemical signaling may play a crucial role in attracting a partner and in facilitating outcrossing. Outcrossing would provide the benefit of masking the expression of deleterious recessive mutations in progeny.[31] (Also see complemenation.)

text: Copulation and reproduction are separate processes in earthworms. The mating pair overlap front ends ventrally and each exchanges sperm with the other. The clitellum becomes very reddish to pinkish in color. Some time after copulation, long after the worms have separated, the clitellum (behind the spermathecae) secretes material which forms a ring around the worm. The worm then backs out of the ring, and as it does so, it injects its own eggs and the other worm's sperm into it. As the worm slips out of the ring, the ends of the cocoon seal to form a vaguely lemon-shaped incubator (cocoon) in which the embryonic worms develop. They emerge as small, but fully formed earthworms, but lack their sex structures, which develop in about 60 to 90 days. They attain full size in about one year. Scientists predict that the average lifespan under field conditions is four to eight years, while most garden varieties live only one to two years. Several common earthworm species are mostly parthenogenetic.

text: Among lumbricid earthworms, parthenogenesis arose from sexual relatives many times.[32] Parthenogenesis in some Aporrectodea trapezoides lineages arose 6.4 to 1.1 million years ago from sexual ancestors.[33]

text: Regeneration

text: Earthworms have the ability to regenerate lost segments, but this ability varies between species and depends on the extent of the damage. Stephenson (1930) devoted a chapter of his monograph to this topic, while G.E. Gates spent 20 years studying regeneration in a variety of species, but “because little interest was shown”, Gates (1972) only published a few of his findings that, nevertheless, show it is theoretically possible to grow two whole worms from a bisected specimen in certain species. Gates’s reports included:

text:
Eisenia fetida (Savigny, 1826) with head regeneration, in an anterior direction, possible at each intersegmental level back to and including 23/24, while tails were regenerated at any levels behind 20/21, i.e., two worms may grow from one.[34]
Lumbricus terrestris Linnaeus, 1758 replacing anterior segments from as far back as 13/14 and 16/17 but tail regeneration was never found.
Perionyx excavatus Perrier, 1872 readily regenerated lost parts of the body, in an anterior direction from as far back as 17/18, and in a posterior direction as far forward as 20/21.
Lampito mauritii Kinberg, 1867 with regeneration in anterior direction at all levels back to 25/26 and tail regeneration from 30/31; head regeneration was sometimes believed to be caused by internal amputation resulting from Sarcophaga sp. larval infestation.
Criodrilus lacuum Hoffmeister, 1845 also has prodigious regenerative capacity with ‘head’ regeneration from as far back as 40/41.[35]

text: An unidentified Tasmanian earthworm shown growing a replacement head has been reported.[36]

text: Locomotion and importance to soil

text: Earthworms travel underground by the means of waves of muscular contractions which alternately shorten and lengthen the body (peristalsis). The shortened part is anchored to the surrounding soil by tiny claw-like bristles (setae) set along its segmented length. In all the body segments except the first, last and clitellum, there is a ring of S-shaped setae embedded in the epidermal pit of each segment (perichaetine). The whole burrowing process is aided by the secretion of lubricating mucus. Worms can make gurgling noises underground when disturbed as a result of their movement through their lubricated tunnels. Earthworms move through soil by expanding crevices with force; when forces are measured according to body weight, hatchlings can push 500 times their own body weight whereas large adults can push only 10 times their own body weight.[37]

text: Earthworms work as biological "pistons" forcing air through the tunnels as they move. Thus earthworm activity aerates and mixes the soil, and is conducive to mineralization of nutrients and their uptake by vegetation. Certain species of earthworm come to the surface and graze on the higher concentrations of organic matter present there, mixing it with the mineral soil. Because a high level of organic matter mixing is associated with soil fertility, an abundance of earthworms is generally considered beneficial by farmers and gardeners.[38][39] In fact, as long ago as 1881 Charles Darwin wrote: "It may be doubted whether there are many other animals which have played so important a part in the history of the world, as have these lowly organized creatures."[40]

text: Benefits

text: The major benefits of earthworm activities to soil fertility can be summarized as:

text: Biological: In many soils, earthworms play a major role in the conversion of large pieces of organic matter into rich humus, thus improving soil fertility. This is achieved by the worm's actions of pulling below the surface deposited organic matter such as leaf fall or manure, either for food or to plug its burrow. Once in the burrow, the worm will shred the leaf and partially digest it and mingle it with the earth. Worm casts (see bottom right) can contain 40 percent more humus than the top 9" (23 cm) of soil in which the worm is living.[3]

text:
Chemical: In addition to dead organic matter, the earthworm also ingests any other soil particles that are small enough—including sand grains up to 1/20 of an inch (1.25 mm)—into its gizzard, wherein those minute fragments of grit grind everything into a fine paste which is then digested in the intestine. When the worm excretes this in the form of casts, deposited on the surface or deeper in the soil, minerals and plant nutrients are changed to an accessible form for plants to use. Investigations in the United States show that fresh earthworm casts are five times richer in available nitrogen, seven times richer in available phosphates, and 11 times richer in available potassium than the surrounding upper 6 inches (150 mm) of soil. In conditions where humus is plentiful, the weight of casts produced may be greater than 4.5 kg (10 lb) per worm per year.[3]
Physical: The earthworm's burrowing creates a multitude of channels through the soil and is of great value in maintaining the soil structure, enabling processes of aeration and drainage. Permaculture co-founder Bill Mollison points out that by sliding in their tunnels, earthworms "act as an innumerable army of pistons pumping air in and out of the soils on a 24-hour cycle (more rapidly at night)".[41] Thus, the earthworm not only creates passages for air and water to traverse the soil, but also modifies the vital organic component that makes a soil healthy (see Bioturbation). Earthworms promote the formation of nutrient-rich casts (globules of soil, stable in soil [mucus]) that have high soil aggregation and soil fertility and quality.[3]

text: Earthworms accelerate nutrient cycling in the soil-plant system through fragmentation & mixing of plant debris – physical grinding & chemical digestion.[3] The earthworm's existence cannot be taken for granted. Dr. W. E. Shewell-Cooper observed "tremendous numerical differences between adjacent gardens", and worm populations are affected by a host of environmental factors, many of which can be influenced by good management practices on the part of the gardener or farmer.[42]

text: Darwin estimated that arable land contains up to 53,000 worms per acre (13/m2), but more recent research from Rothamsted Experimental Station has produced figures suggesting that even poor soil may support 250,000/acre (62/m2), whilst rich fertile farmland may have up to 1,750,000/acre (432/m2), meaning that the weight of earthworms beneath a farmer's soil could be greater than that of the livestock upon its surface.

text: The ability to break down organic materials and excrete concentrated nutrients makes the earthworm a functional contributor in restoration projects. In response to ecosystem disturbances, some opencast mining sites have utilized earthworms to prepare soil for the return of native flora. Sites using this method have observed advances in the return of ecosystem services that previously took much longer to re-establish.[43] Research from the Station d'écologie Tropicale de Lamto asserts that the earthworms positively influence the rate of macroaggregate formation, an important feature for soil structure.[44] The stability of aggregates in response to water was also found to be improved when constructed by earthworms.[44]

text: As an invasive species

text: From a total of around 7,000 species, only about 150 species are widely distributed around the world. These are the peregrine or cosmopolitan earthworms.[45]

text: Special habitats

text: While, as the name earthworm suggests, the main habitat of earthworms is in soil, the situation is more complicated than that. The brandling worm Eisenia fetida lives in decaying plant matter and manure. Arctiostrotus vancouverensis from Vancouver Island and the Olympic Peninsula is generally found in decaying conifer logs. Aporrectodea limicola, Sparganophilus spp., and several others are found in mud in streams. Some species are arboreal, some aquatic and some euryhaline (salt-water tolerant) and littoral (living on the sea-shore, e.g. Pontodrilus litoralis). Even in the soil species, special habitats, such as soils derived from serpentine, have an earthworm fauna of their own.

text: Ecology

text: Earthworms are classified into three main ecophysiological categories: (1) leaf litter- or compost-dwelling worms that are nonburrowing, live at soil-litter interface and eat decomposing OM (called Epigeic) e.g. Eisenia fetida; (2) topsoil- or subsoil-dwelling worms that feed (on soil), burrow and cast within soil, creating horizontal burrows in upper 10–30 cm of soil (called Endogeics); and (3) worms that construct permanent deep vertical burrows which they use to visit the surface to obtain plant material for food, such as leaves (called Anecic (meaning "reaching up")), e.g. Lumbricus terrestris.[46]

text: Earthworm populations depend on both physical and chemical properties of the soil, such as temperature, moisture, pH, salts, aeration, and texture, as well as available food, and the ability of the species to reproduce and disperse. One of the most important environmental factors is pH, but earthworms vary in their preferences. Most favor neutral to slightly acidic soils. Lumbricus terrestris is still present in a pH of 5.4 and Dendrobaena octaedra at a pH of 4.3 and some Megascolecidae are present in extremely acidic humic soils. Soil pH may also influence the numbers of worms that go into diapause. The more acidic the soil, the sooner worms go into diapause and the longer they remain in diapause at a pH of 6.4.

text: Earthworms form the base of many food chains. They are preyed upon by many species of birds (e.g. starlings, thrushes, gulls, crows, European robins and American robins), snakes, mammals (e.g. bears, foxes, hedgehogs, pigs, moles) and invertebrates (e.g. ground beetles and other beetles, snails, slugs). Earthworms have many internal parasites, including protozoa, platyhelminthes, and nematodes; they can be found in the worms' blood, seminal vesicles, coelom, or intestine, or in their cocoons.

text: Nitrogenous fertilizers tend to create acidic conditions, which are fatal to the worms, and dead specimens are often found on the surface following the application of substances such as DDT, lime sulphur, and lead arsenate. In Australia, changes in farming practices such as the application of superphosphates on pastures and a switch from pastoral farming to arable farming had a devastating effect on populations of the giant Gippsland earthworm, leading to their classification as a protected species.

text: The addition of organic matter, preferably as a surface mulch, on a regular basis will provide earthworms with their food and nutrient requirements, and will create the optimum conditions of temperature and moisture that will stimulate their activity.

text: Economic impact

text: Various species of worms are used in vermiculture, the practice of feeding organic waste to earthworms to decompose food waste. These are usually Eisenia fetida (or its close relative Eisenia andrei) or the Brandling worm, commonly known as the tiger worm or red wiggler. They are distinct from soil-dwelling earthworms. In the tropics, the African nightcrawler Eudrilus eugeniae and the Indian blue Perionyx excavatus are used.

text: Earthworms are sold all over the world; the market is sizable. According to Doug Collicut, "In 1980, 370 million worms were exported from Canada, with a Canadian export value of $13 million and an American retail value of $54 million."

text: Earthworms are also sold as food for human consumption. Noke is a culinary term used by the Māori of New Zealand, and refers to earthworms which are considered delicacies for their chiefs.

text: Taxonomy and distribution

text: Within the world of taxonomy, the stable 'Classical System' of Michaelsen (1900) and Stephenson (1930) was gradually eroded by the controversy over how to classify earthworms, such that Fender and McKey-Fender (1990) went so far as to say, "The family-level classification of the megascolecid earthworms is in chaos."[47] Over the years, many scientists developed their own classification systems for earthworms, which led to confusion, and these systems have been and still continue to be revised and updated. The classification system used here, developed by Blakemore (2000), is a modern reversion to the Classical System that is historically proven and widely accepted.[48]

text: Categorization of a megadrile earthworm into one of its taxonomic families under suborders Lumbricina and Moniligastrida is based on such features as the makeup of the clitellum, the location and disposition of the sex features (pores, prostatic glands, etc.), number of gizzards, and body shape.[48] Currently, over 6,000 species of terrestrial earthworms are named, as provided in a species name database,[49] but the number of synonyms is unknown.

text: The families, with their known distributions or origins:[48]

text:
Acanthodrilidae – (Gondwanan or Pangaean?)
Ailoscolecidae – Pyrenees and southeast USA
Almidae – tropical equatorial (South America, Africa, Indo-Asia)
Benhamiinae – Ethiopian, Neotropical (a possible subfamily of Octochaetidae)
Criodrilidae – southwestern Palaearctic: Europe, Middle East, Russia and Siberia to Pacific coast; Japan (Biwadrilus); mainly aquatic
Diplocardiinae/-idae – Gondwanan or Laurasian? (a subfamily of Acanthodrilidae)
Enchytraeidae – cosmopolitan but uncommon in tropics (usually classed with Microdriles)
Eudrilidae – Tropical Africa south of the Sahara
Exxidae – Neotropical: Central America and Caribbean
Glossoscolecidae – Neotropical: Central and South America, Caribbean
Haplotaxidae – cosmopolitan distribution (usually classed with Microdriles)
Hormogastridae – Mediterranean
Kynotidae – Malagasian: Madagascar
Lumbricidae – Holarctic: North America, Europe, Middle East, Central Asia to Japan
Lutodrilidae – Louisiana southeast USA
Megascolecidae – (Pangaean?)
Microchaetidae – Terrestrial in Africa especially South African grasslands
Moniligastridae – Oriental and Indian subregion
Ocnerodrilidae – Neotropics, Africa; India
Octochaetidae – Australasian, Indian, Oriental, Ethiopian, Neotropical
Octochaetinae – Australasian, Indian, Oriental (subfamily if Benhamiinae is accepted)
Sparganophilidae – Nearctic, Neotropical: North and Central America
Tumakidae – Colombia, South America


document_title: JetBlue Mint

question_text: first jetblue mint flight between jfk and lax

text: JetBlue Mint

text: Mint is a premium airline product offered by JetBlue on some flights between the West Coast and the East Coast of the contiguous United States, and seasonally on some flights between the United States and Caribbean destinations. The service is operated with a fleet of Airbus A321 aircraft in a premium configuration, with lie-flat seats in premium class. As of May 2016, JetBlue is the only low cost airline in the Americas offering a dedicated premium product.[1]

text: History[edit]

text: JetBlue first announced its intention to introduce premium seats on some of its transcontinental flights in March 2013.[2] The company officially announced its Mint premium class product on September 30, 2013; the aforementioned service was to be available on flights between New York and San Francisco and New York and Los Angeles, with fares for Mint-class seats starting at US$599 each way.[3] Dave Barger, JetBlue CEO at the time, said that the move was necessary for the airline to better compete with its competitors,[4] which were operating lie-flat seats in transcontinental markets, leaving JetBlue with an all-economy class cabin.[1] The routes were to be operated with new Airbus A321 aircraft, and feature Wi-Fi connectivity. JetBlue's all seven daily round-trip flights between New York and Los Angeles were to feature the service by the fourth quarter of 2014, and all five daily round-trip flights between New York and San Francisco by early 2015. The first Mint flight, from New York–JFK to Los Angeles, was to begin on June 15, 2014.[4]

text: On March 15, 2015, JetBlue announced seasonal expansion of its Mint service to Caribbean routes, with service from JFK Airport to Aruba and Barbados. The service was to operate once daily per destination during the winter holiday season, and once weekly per destination for the rest of the winter season.[5] JetBlue announced expansion of its Mint service to Boston in June 2015, with year-round service from Boston to Los Angeles and San Francisco,[6] and seasonal Saturday-only service from Boston to Barbados.[7] Flights from Boston to San Francisco began on March 24, 2016,[8] and flights from Boston to Los Angeles were to begin on October 20, 2016.[9]

text: On April 12, 2016, JetBlue announced an expansion of its Mint service. New routes utilizing the Mint configuration were to be launched to Fort Lauderdale, Las Vegas, San Diego and Seattle, with service beginning during 2017 and 2018.[10] The announcement was made almost immediately after Alaska Air Group announced its intention to acquire Virgin America and merge it with Alaska Airlines. Either Alaska or Virgin operated in 4 out of 7 of JetBlue's newly announced markets.[1]

text: Service[edit]

text: The Mint subfleet consists of A321 aircraft with 159 seats,[11] including 16 flat-bed Mint seats. The Mint seat is a version of the Thompson Aero Vantage business class seat, customized to include extra storage and dual 110V+USB power outlets at every seat.[12][13] 4 of the 16 Mint seats are mini-suites which include a sliding door for privacy.[12] The seats have a width of up to 22.3 inches and can convert to a flat bed of up to 6'8" in length. Pairs of non-suite Mint seats alternate with single mini-suite seats.[14] The seats use air cushioning, allowing passengers to adjust the firmness of the seats.[13][14] Economy class on Mint A321 aircraft uses B/E Aerospace Pinnacle slimline[15] seats, with a seat pitch of 33 inches in the main cabin and up to 37 inches in the "Even More Space" extra legroom product.[12] There are two power outlets per row of economy class seats.[15]

text: A 15.6 inch video screen provides Live TV 4.0, SiriusXM Radio, and a selection of films as in-flight entertainment.[14] The TV system provides about 100 DirecTV channels.[13] However, on flights outside the coverage range of satellite TV, such as flights to Caribbean destinations, TV channels and radio are not provided.[16] In-flight Wi-Fi is provided via a Ka-band satellite system.[12]

text: Passengers are provided with a choice of five offerings for meal service, including hot and cold options, and may choose three.[12][13] Amenity kits are provided by Hayward & Hopper, a cosmetics company based in New York.[17]


document_title: List of most lethal American battles

question_text: which battle lost the most soldiers in ww2

text: List of most lethal American battles

text: The United States military has existed for over two centuries. It was birthed out of necessity during fighting with its ruler, Great Britain, for independence (1775–83). It continued through the bloody American Civil War (1861–65) and, after collaborating in triumph during both World War I (1917-1918) and World War II (1941–45). Since WWII the United States military has been constantly at war in such notable conflicts as the Korean War (1950-1953), Vietnam War (1955-1975) and the present day ongoing conflicts in the Middleast and War in Afghanistan (2001–present). For most of its existence, America has been involved in one or another military conflict.[1]

text: The term Casualty in warfare can often be confusing. It often does not refer to those that are killed on the battlefield rather it refers to those who can no longer fight. This can include disabled by injuries, disabled by psychological trauma, captured, deserted, or missing. A casualty is just a soldier who is no longer available for the immediate battle or campaign, the major consideration in combat; the number of casualties is simply the number of members of a unit who are not available for duty. For example during the Seven Days Battles in the American Civil War (June 25 to July 1, 1862) there were 5,228 killed, 23,824 wounded and 7,007 missing or taken prisoner for a total of 36,059 casualties.[2] [A 1] The word casualty has been used in a military context since at least 1513.[3] In this article the numbers killed refer to those killed in action, killed by disease or someone who died from their wounds.

text: The following is a list of the most lethal battles fought by the military of the United States of America. This list shows battles where the number of Americans killed was higher than 1000. The battles that reached that number of deaths in the field are so far limited to the Civil War, WWI, WWII, Korean War and one battle during the Vietnam War the Tet Offensive (January 30 to September 23, 1968). The bloodiest battle with the most American soldiers killed was and still is the Battle of Normandy (June 6 to August 30, 1944) where 29,198 soldiers were killed trying to gain a foothold in Nazi-occupied France.[4] The bloodiest single day, of the Battle of Normandy, with the most Americans killed was the first day, D-day (June 6, 1944) with 2,499 killed.[5] The bloodiest single day in American military history was during the Battle of Antietam when 3,654 soldiers were killed on September 17, 1862.[6] [A 2] [7]

text:
Battle, Siege, or Offensive
Conflict
Date
Estimated Number killed
References

Battle of Normandy / Northern France
World War II
1944–06-06 !June 6 to September 14, 1944
34,137 killed [A 3]
[8]

Meuse-Argonne Offensive
World War I
1918–09-26 !September 26 to November 11, 1918
26377 !26,277 killed
[9]

Battle of Okinawa
World War II
1945–04-01 !April 1 to June 21, 1945
20195 !20,195 killed
[10]

Battle of the Bulge
World War II
1944–12-16 !December 16, 1944, to January 25, 1945
19276 !19,276 killed
[11]

Battle of Hürtgen Forest
World War II
1944–09-19 !September 19, 1944, to February 10, 1945
Over 12,000 killed
[A 4]

Battle of Luzon
World War II
1945–01-09 !January 9 to August 15, 1945
10640 !10,640 killed
[12]

Battle of Guadalcanal
World War II
1942–08-07 !August 7, 1942, to February 9, 1943
7100 !7,100 killed
[13]

Battle of Gettysburg
American Civil War
1863–07-06 !July 1 to July 3, 1863
7,058 killed [A 5]
[14]

Operation Nordwind
World War II
1944–12-31 !December 31, 1944, to January 25, 1945
7,000 killed [15]
[15]

Battle of Iwo Jima
World War II
1945–02-16 !February 16 to March 26, 1945
6,821 killed
[16]

Lorraine Campaign
World War II
1944–09-01 !September 1 to December 18, 1944
6,657 killed
[17]

Battle of Anzio
World War II
1944–01-22 !January 22 to June 5, 1944
5,538 killed
[8]

Seven Days Battles
American Civil War
1862–06-25 !June 25 to July 1, 1862
5,228 killed [A 6]
[2]

Battle of Pusan Perimeter
Korean War
1950–08-04 !August 4 to September 18, 1950
4,599 killed
[18]

Chinese Second Phase Offensive in North Korea
Korean War
1950–11-25 !November 25 to December 15, 1950
4,538 killed [A 7]
[19]

Battle of Spotsylvania
American Civil War
1864–05-08 !May 8 to May 21, 1864
4,240 killed [A 8]
[20]

Battle of Chickamauga
American Civil War
1863–09-19 !September 19 to September 20, 1863
3,968 killed [A 9]
[20]

Battle of the Wilderness
American Civil War
1864–05-05 !May 5 to May 7, 1864
3,741 killed [A 10]
[20]

Battle of Antietam
American Civil War
1862–09-17 !September 17, 1862
3,654 killed [A 11]
[20]

Battle of Leyte
World War II
1944–10-17 !October 17-December 26, 1944
3,593 killed
[A 12]

Operation Diadem
World War II
1944–05-11 !May 11–18, 1944
3,145 killed
[22]

Battle of Leyte Gulf
World War II
1944–10-23 !October 23–25, 1944
2,800 killed
[23]

Battle of Shiloh
American Civil War
1862–04-06 !April 6 to April 7, 1862
3,482 killed [A 13]
[20]

Battle of Saipan
World War II
1944–06-15 !June 15 to July 9, 1944
3,426 killed
[24]

Battle of Chancellorsville
American Civil War
1863–04-30 !April 30 to May 6, 1863
3,330 killed [A 14]
[20]

Tet Offensive
Vietnam War
1968–01-30 !January 30 to September 23, 1968
3,178 Killed
[A 15]

Battle of Stones River
American Civil War
1862–12-31 !December 31, 1862 to January 2, 1863
3,024 killed [A 16]
[20]

Battle of Second Manassas
American Civil War
1862–08-26 !August 26 to August 30, 1862
3,052 killed [A 17]
[20]

Battle of Chosin Reservoir
Korean War
1950–11-27 !November 27 to December 13, 1950
2,840~ killed [A 18]
[26]

Battle of Sicily
World War II
1943–07-09 !July 9 to August 17, 1943
2,811 killed
[27]

Battle of Cold Harbor
American Civil War
1864–05-21 !May 21 to June 12, 1864
2,632 killed [A 19]
[20]

D-day (first day of Operation Overlord)
World War II
1944–06-06 !June 6, 1944
2,499 killed
[5]

Pearl Harbor Attack
World War II
1941–12-07 !December 7, 1941
2,403 killed [A 20]
[28]

Battle of Peleliu
World War II
1944–09-15 !September 15 to November 25, 1944
2,336 killed
[29]

Battle of Franklin
American Civil War
1864–11-30 !November 30, 1864
1,939 killed [A 21]
[30][31]

Second Battle of the Marne
World War I
1918–07-15 !July 15 to August 6, 1918
1,926 killed [A 22]
[33]

Battle of Fredericksburg
American Civil War
1862–12-11 !December 11 to December 15, 1862
1,892 killed [A 23]
[34]

Battle of Guam
World War II
1944–07-21 !July 21 to August 10, 1944
1,777 killed
[35]

Battle of Salerno
World War II
1943–09-03 !September 3–16, 1943
1,635~ killed [A 24]
[36]

Battle of Seven Pines
American Civil War
1862–05-31 !May 31 to June 1, 1862
1,770 killed [A 25]
[37][38]

Battle of Tarawa
World War II
1943–11-20 !November 20 to November 23, 1943
1,759 killed [A 26]
[39]

Battle of the Ch'ongch'on River
Korean War
1950–11-25 !November 25 to December 2, 1950
1,489 killed [A 27]
[40]

Task Force Faith
Korean War
1950–11-27 !November 27 to December 2, 1950
1,450~ killed [A 28]
[41]

Battle of Perryville
American Civil War
1862–10-08 !October 8, 1862
1,422 killed [A 29]
[43][42]

Battle of Chattanooga
American Civil War
1863–09-21 !September 21 to November 25, 1863
1,114 killed [A 30]
[44]

Battle of Belleau Wood
World War I
1918–06-01 !June 1 to June 26, 1918
1,062 killed
[45]

Battle for Manila
World War II
1945–02-03 !February 3 to March 3, 1945
1,010 killed
[46]

tag: table

text: Annotations[edit]

text:
Jump up ^ Union and Confederate numbers added together
Jump up ^ Union and Confederate numbers added together
Jump up ^ Source adds Battle of Normandy (June 6 - July 24, 1944) and "Northern France (July 25 - September 14, 1944)
Normandy KIA 13,959 + Northern France 15,239 = 29,198
Total deaths Normandy June 6 - July 24, 1944 16,293 + Northern France July 25 - September 14, 1944 17,844 = 34,137 [8]
Jump up ^ 50,410 Americans died in the Rhineland from September 1944 - March 21, 1945 [4]
Jump up ^ 3,155 Union and 3,903 Confederate
Jump up ^ 1,734 Union and 3,494 Confederacy[2]
Jump up ^ U.S. Army KIA+Army POW died+Army MIA died+Marine KIA+Navy KIA
1183+1167+1410+763+15=4,538
Jump up ^ 2,725 Union[20] and 1,515 Confederacy
Jump up ^ 1,656 Union[20] and 2,312 Confederacy
Jump up ^ 2,246 Union[20] and 1,495 Confederacy
Jump up ^ 2,108 Union[20] and 1,546 Confederacy
Jump up ^ 16,233 died in Leyete, Luson, and Southern Philippines during October 17, 1944 - July 4, 1945 [21]
Jump up ^ 1,754 Union[20] and 1,728 Confederacy
Jump up ^ 1,606 Union[20] and 1,724 Confederacy
Jump up ^ 16,592 American died in 1968 [25]
Jump up ^ 1,730 Union[20] and 1,294 Confederacy
Jump up ^ 1,747 Union[20] and 1,305 Confederacy
Jump up ^ 836 Marines killed + 2,000 US Army killed[26]
Jump up ^ 1,844 Union[20] and 788 Confederacy
Jump up ^ 2,008 Navy KIA + 109 Marines + 218 Army + 68 civilians
2008+109+218+68 [28]
Jump up ^ 189 Union[30] and 1,750 Confederacy [31]
Jump up ^ 12,000 casualities including KIA, WIA, POW [32]
Jump up ^ 1,284 Union and 608 Confederacy[34]
Jump up ^ 5th Army 788 killed + US Navy 296 killed + 551 missing presumed drowned [36]
Jump up ^ 790 Union [37] and 980 Confederacy[38]
Jump up ^ 984 Marines + 88 MIA + 687 Navy
984+ 88+ 687 [39]
Jump up ^ 676 killed + 813 Missing presumed dead
Jump up ^ Of the 2,500 soldiers in Task Force Faith only 1,050 made it back. Of those only 385 were able-bodied[41]
Jump up ^ 890[42] Union and 532 Confederacy[43]
Jump up ^ 753 Union and 361 Confederacy[44]

tag: ol

text: Bibliography[edit]

text: Notes

text:
Jump up ^ Kelly 2017
^ Jump up to: a b c Tucker 2013, p. 892
Jump up ^ Oxford English Dictionary, 2nd ed gives a 1513 reference for military casualty, and an 1844 reference for civilian use
^ Jump up to: a b Statistical and accounting branch office of the adjutant general 1953, p. 93
^ Jump up to: a b D-Day Museum and Overlord Embroidery 2017
Jump up ^ Tucker 2013, p. 903
Jump up ^ History.com 2017
^ Jump up to: a b c Statistical and accounting branch office of the adjutant general 1953, p. 92
Jump up ^ Brown 2013, p. 191
Jump up ^ Traynor 2017
Jump up ^ Graham 2007, p. 159
Jump up ^ Willmott 2005, p. 22
Jump up ^ Tucker 2014, p. 213
Jump up ^ Burke & Roth 2014, p. 7
^ Jump up to: a b Corson 1995
Jump up ^ Smith 2009, p. II
Jump up ^ Zabecki 1999, p. 1560
Jump up ^ Varhola 2000, p. 6
Jump up ^ Ecker 2005, p. 62
^ Jump up to: a b c d e f g h i j k l m n o p q r Allen 2017
Jump up ^ Statistical and accounting branch office of the adjutant general 1953, p. 94
Jump up ^ Fisher 2014, p. 222
Jump up ^ Tucker 2013, p. 1668
Jump up ^ Hearn 2007, p. 88
Jump up ^ United States 2010
^ Jump up to: a b Hickman 2017
Jump up ^ Hart 2015, p. 627
^ Jump up to: a b Alison 2016
Jump up ^ Stamford Historical Society 2009
^ Jump up to: a b Eicher 2002, p. 774
^ Jump up to: a b United States War Department 1886, p. 344
Jump up ^ Sondhaus 2011, p. 413
Jump up ^ Surgeon General 1920, pp. 43–44
^ Jump up to: a b Tucker 2013, p. 919
Jump up ^ Ramirez 2017
^ Jump up to: a b Konstam 2007, p. 157
^ Jump up to: a b Sears 2014, pp. 146–147
^ Jump up to: a b Tucker 2013, p. 888
^ Jump up to: a b Tucker 2013, p. 1605
Jump up ^ Ecker 2005, p. 62.
^ Jump up to: a b Daily 1999, p. 68
^ Jump up to: a b United States War Department 1886, pp. 1033–1036
^ Jump up to: a b United States War Department 1886, p. 1112
^ Jump up to: a b Tucker 2013, p. 966
Jump up ^ Tucker 2013, p. 1323
Jump up ^ Tucker 2009, p. 370

text: References

text:
Alison (Jun 14, 2016). "The Count of Pearl Harbor Deaths". VisitPearlHarbor.org. Retrieved November 7, 2017.
Allen, Jonathan R. (2017). "How Many Died in the Civil War?". nellaware.com. Retrieved November 6, 2017.
Brown, Meredith Mason (2013). Touching America's History: From the Pequot War Through World War II. Indiana University Press. ISBN 9780253008336.  - Total pages: 271
Burke, Patrick; Roth, Jack (2014). Ghost Soldiers of Gettysburg: Searching for Spirits on America's Most Famous Battlefield. Llewellyn Worldwide. ISBN 9780738741352.  - Total pages: 264
Corson, Captain William quoting Danny S. Parker's article 'War's Last Eruption' is Military History Magazine: Great Battles, September 1992, pp. 42–9. 3 (June 1995). Captured in Hatten: Captain William Corson's speech (Speech). Retrieved November 13, 2017.
Daily, Edward L. (1999). MacArthur's X Corps in Korea: Inchon to the Yalu, 1950. Turner Publishing Company. ISBN 9781563114397.  - Total pages: 152
D-Day Museum and Overlord Embroidery (2017). "D-Day and the Battle of Normandy: Your Questions Answered". D-Day Museum and Overlord Embroidery. Retrieved November 8, 2017.
Ecker, Richard E. (2005). Korean Battle Chronology: Unit-by-unit United States Casualty Figures and Medal of Honor Citations. McFarland & Company. ISBN 9780786419807.  - Total pages: 207
Eicher, David J (2002). The Longest Night: A Military History of the Civil War. Simon and Schuster. ISBN 9780743218467.  - Total pages: 992
Fisher, Ernest F. Jr. (2014). United States Army in WWII - the Mediterranean - Cassino to the Alps: [Illustrated Edition]. Pickle Partners Publishing. ISBN 9781782894117.  - Total pages: 602
Graham, Roger D; Colonel, USAF-Retired (2007). The Nimrods. AuthorHouse. ISBN 9781467830966.  - Total pages: 248
Hart, B. H. Liddell (2015). A History of the Second World War. Pan Macmillan. ISBN 9781447209676.  - Total pages: 848
Hearn, Chester G. (2007). Marines: An Illustrated History : the U.S. Marine Corps from 1775 to the 21st Century. Zenith Press. ISBN 9780760332115.  - Total pages: 192
Hickman, Kennedy (April 28, 2017). "Korean War: Battle of Chosin Reservoir". Thoughtco. Retrieved November 7, 2017.
History.com (2017). "The Battle of Antietam". History.com. Retrieved November 8, 2017.
Kelly, Martin (April 4, 2017). "American Involvement in Wars from Colonial Times to the Present". Thoughtco. Retrieved November 8, 2017.
Konstam, Angus (2007). Salerno 1943: The Allied Invasion of Italy. Pen and Sword. ISBN 9781844155170.  - Total pages: 192
Ramirez, Fernando (August 9, 2017). "Historic WWII photos show gritty U.S. Marine struggle over Guam". Houston Chronicle. Retrieved November 8, 2017.
Sears, Stephen W. (2014). To the Gates of Richmond: The Peninsula Campaign. Houghton Mifflin Harcourt. ISBN 9780547527550.  - Total pages: 512
Smith, Larry (2009). Iwo Jima: World War II Veterans Remember the Greatest Battle of the Pacific. W. W. Norton & Company. ISBN 9780393285635.  - Total pages: 384
Sondhaus, Lawrence (2011). World War One: The Global Revolution. Cambridge University Press. ISBN 9781107782501.
Stamford Historical Society (2009). "The Stamford Historical Society Presents". Stamford Historical Society. Retrieved November 8, 2017.
Statistical and accounting branch office of the adjutant general (1953). Army battle casualties and nonbattle deaths in World War II. Federal government of the United States. OCLC 220594130.  - Total pages: 121
Surgeon General (1920). "Report of the Office of the Surgeon General". War Department Annual Reports. 1920 (1): 1106. Retrieved November 7, 2017.
Traynor, John & Lisa Early, Clare Horrie (2017). "Background: Find out more about the events leading up to Truman's decision to drop the bomb". The National Archives (United Kingdom). Retrieved November 6, 2017.
Tucker, Spencer C. (2009). The Encyclopedia of the Spanish-American and Philippine-American Wars: A Political, Social, and Military History, Volume 1. ABC-CLIO. ISBN 9781851099511.  - Total pages: 370
Tucker, Spencer C. (2013). Almanac of American Military History, Volume 1. ABC-CLIO. ISBN 9781598845303.  - Total pages: 2524
Tucker, Spencer C. (2014). Battles That Changed American History: 100 of the Greatest Victories and Defeats. ABC-CLIO. ISBN 9781440828621.  - Total pages: 355
United States, Government (2010). "Statistical information about casualties of the Vietnam War". National Archives and Records Administration. Archived from the original on 26 January 2010. Retrieved March 6, 2010.
United States War Department, John Sheldon Moody, Calvin Duvall Cowles, Frederick Caryton Ainsworth, Robert N. Scott, Henry Martyn Lazelle, George Breckenridge Davis, Leslie J. Perry, Joseph William Kirkley (1886). The war of the rebellion: a compilation of the official records of the Union and Confederate armies. ; Series 1 - Volume 16 (Part I). United States Government Publishing Office.
Varhola, Michael O. Varhola (2000). Fire And Ice: The Korean War, 1950- 53. Da Capo Press. ISBN 9781882810437.  - Total pages: 317
Willmott, H. P. (2005). The Battle of Leyte Gulf: The Last Fleet Action. Indiana University Press. ISBN 9780253003515.  - Total pages: 416
Zabecki, David T. Zabecki (1999). World War Two in Europe. Taylor & Francis. ISBN 9780824070298.  - Total pages: 1920

text: Lists of battlesUnited States military-related listsBattles involving KoreaWar casualtiesAmerican Civil War-related listsBattles of the American Civil War


document_title: Mahanati

question_text: what is the role of samantha in mahanati movie

text: Mahanati

text: Plot[edit]

text: In Bangalore, India, popular yesteryear actress Savitri is found unconscious by her young son and hospitalised, spending a year in coma. The incident receives heavy media attention, but Savitri's initial life and whereabouts remain unknown to the public. A newspaper editor appoints Madhuravani, a stuttering journalist, and Vijay Anthony, a photographer, to write a memoir on Savitri. Madhuravani recovers a letter written by Savitri, in which she wishes to take her son to someone called Shankaraiya. Madhuravani continues her research even after she is relieved from writing the memoir by her editor, and sneaks into Savitri's house, overhearing her aunt talk about her childhood.

text: In 1942 Vijayawada, a widowed Subhadramma and her daughter Savitri are taken in by Subhadramma's sister, Durgamamba, and husband, K. V. Chowdary, much to the latter's chagrin. Magnanimous in speech, thought and action, Savitri befriends Susheela, and they become best friends. She becomes an accomplished dancer and later earns a profit for Chowdary by acting in plays with Susheela. On one such occasion, she is praised by the Bollywood actor Prithviraj Kapoor for her performance.

text: The increasing popularity of cinema leads to less attendance for plays, and Chowdary incurs huge losses. He tries to have Savitri join the film industry centred in Madras, but her young age and inability to speak Tamil hinder her prospects. She, however, succeeds in meeting Ramasamy "Gemini" Ganesan who works in the casting department and has her photograph taken for a casting directory. A year later, a director comes across the directory and wants to launch Savitri, but she gets the dialogue wrong and is thrown out. However, she successfully earns a role in a film song, and her distinguished performance helps her in getting major roles.

text: Savitri is offered a role opposite Ganesan, now an actor, and he teaches her the Tamil language. The two actors together rise as superstars and Savitri falls in love with the married Ganesan. They secretly marry, which leads to a clash with Savitri's family. Reluctant to act in films after marriage, Savitri gives in to the persuasion of filmmakers and Ganesan, and the films turn out to be successful. Savitri gives birth two children– a girl and a boy. Initially supportive of her career, Ganesan starts to feel jealous, with Savitri's fame overshadowing his, and starts consuming alcohol. Savitri tries to stop him, but Ganesan instead introduces her to drinking.

text: A few ladies urge Savitri to both direct and star in an all-woman cast and crew film. On the day of her film's release, Savitri finds Ganesan in their guesthouse with another woman. Feeling cheated, she angrily refuses to talk to him and shuts him out of their house. Savitri becomes an alcoholic, and also gains weight. Meanwhile, her mother dies, and poor financial decisions lead to a raid by the Income Tax Department, resulting in the loss of her wealth. One day, Savitri fights with her daughter about her excessive alcohol consumption and faints, accidentally setting their house on fire. They learn that Savitri has diabetes.

text: Chowdary visits Savitri, and she asks both her daughter and uncle for forgiveness and gets her daughter married. Savitri stops drinking and loses weight. She begins living in a smaller house with her son and starts acting in low-budget films. She sets aside money for opening a rehabilitation centre but discovers that she still owes income tax, strongly refusing to part with any more money. She visits Bangalore with her son the next day; feeling lost, she calls Ganesan but cuts the line before speaking. Angry with him and with herself, she drinks alcohol again, before collapsing into a coma.

text: Madhuravani realizes her love for Anthony via Savitri's story. She interrupts what is supposedly Anthony's engagement and speaks strongly without a stutter, but discovers it is his sister's engagement and that Anthony loves her. Madhuravani writes the memoir about Savitri, which reveals that Shankaraiya is a statue in her hometown that she talked to as though he was her father and that she always had an unsatisfied wish to see her father. Madhuravani visits the comatose Savitri, and places a picture of her father in her hands, retrieved from a newspaper archive.

text: Cast[edit]

text:
Keerthy Suresh as Savitri
Dulquer Salmaan as Gemini Ganesan
Samantha Akkineni as Madhuravani
Vijay Devarakonda as Vijay Anthony
Rajendra Prasad as K.V. Chowdary
Divya Vani as Subhadramma
Bhanupriya as Durgamamba
Prakash Raj as Aluri Chakrapani
Tanikella Bharani as the editor of Prajavani newspaper
Malavika Nair as Alamelu Ganesan
Shalini Pandey as Susheela
Tulasi as Madhuravani’s mother
Mahesh Achanta as Satyam

text:
Akkineni Naga Chaitanya as Akkineni Nageswara Rao
Mohan Babu as S. V. Ranga Rao
Naresh as Cameraman Keshava
Krish as K. V. Reddy
Srinivas Avasarala as L. V. Prasad
Manobala as P. Pullaiah
Sandeep Vanga as Vedantam Raghavaiah
Tharun Bhascker Dhaassyam as Singeetham Srinivasa Rao
Sai Madhav Burra as Pingali Nagendra Rao

text: Production[edit]

text: Development[edit]

text: In May 2016, director Nag Ashwin, who had previously made Yevade Subramanyam (2015), announced that his next film would be a biopic of South Indian actress Savitri and that it would be shot simultaneously in Telugu, Malayalam and Tamil. Titled Mahanati, Ashwin revealed that he had finished work on the script and was in the middle of casting actors to be a part of the project.[4] During the pre-production phase, Ashwin spent six months in research by reading articles and books, while meeting old journalists and actors who had been associated with the actress. Explaining why he chose to turn Savitri's career into a film script, he stated "from a penniless origin to the first female superstar, her life is quite a story" and that it was a "moment in time where it has been long enough that film legends have left us" and "that we will have to revisit them and understand why they are legends".[5] He didn't use the conventional biopic format.[6] He was inspired to create a film after listening to the radio about Savitri.[7] Ashwin said the film made him more responsible.[8]

text: Cast and crew[edit]

text: Soon after the announcement of the film, the media carried speculative reports that either Anushka Shetty, Sonakshi Sinha or Vidya Balan would be cast in the lead role. Denying the developments in August 2016, Ashwin stated he was still editing the script and intended to cast the lead actress with an audience poll.[9][10]

text: The team continued to hold talks with actresses for the lead role, and despite extended discussions with Nithya Menen, the actress did not agree to sign the film.[11][12] Samantha was the first lead actor to be cast in the film during December 2016 and revealed she would portray a character from whose point of view the story is narrated, rather than the role of Savitri.[13][14] Reports also suggested that she may be appearing as another actress from the era, Jamuna, a close friend of Savitri.[15] Subsequently, Keerthy Suresh was cast in the lead role of Savitri in early January 2017.[16][17] The film's first look poster featuring both actresses was released in March 2017 to coincide with International Women's Day.[18] For the role of Gemini Ganesan, Savitri's husband, Ashwin had considered Suriya or Madhavan, before finalising Dulquer Salmaan for the character.[19][20] Vijay Deverakonda, who had also earlier been considered for Gemini Ganesan's role, was then selected to portray another role in the film.[21][22] Keerthy and Savitri's daughter held conversations for working on the film.[23]

text: Filming[edit]

text: The principal shoot of the biopic began on 29 May 2017 at Ramakrishna Studios in Hyderabad, with the Telugu title confirmed as Mahanati and the Tamil title confirmed as Nadigaiyar Thilagam.[24][25] Actor Rajendra Prasad joined the cast during the first schedule, while Mickey J Meyer and Dani Sanchez-Lopez was confirmed as music composer and cinematographer respectively.[26] Ashwin revealed that Dulquer's portrayal of Gemini Ganesan would go beyond just the physical resemblance and the character's emotional side would also be depicted.[27] Keerthy Suresh was requested to put on weight for the film and shot for a vintage song in early June 2017 as a part of the first schedule.[28] Prakash Raj also joined the team in August 2017 to play the role of producer Aluri Chakrapani.[29] Deni Sanchez Lopez was the cinematographer.[30]

text: Music[edit]

text:
MahanatiSoundtrack album by Mickey J. MeyerReleased
1 May 2018 (2018-05-01)Recorded
2017Length
20:31Label
Aditya MusicProducer
Mickey J. MeyerMickey J. Meyer chronology

Mister(2017)

Mahanati(2018)

Srinivasa Kalyanam(2018)

text: All music composed by Mickey J. Meyer.

text: Telugu track listingNo.TitleLyricsSinger(s)Length1."Mooga Manasulu"Sirivennela Sitarama SastryShreya GhoshalAnurag Kulkarni4:192."Sada Nannu"Sirivennela Sitarama SastryCharulatha Mani3:373."Mahanati"Ramajogayya SastryAnurag Kulkarni4:444."Aagipo Baalyama"Sirivennela Sitarama SastryRamya Behara2:335."Chivaraku Migiledi"Sirivennela Sitarama SastrySunitha Upadrashta2:586."Gelupuleni Samaram"Sirivennela Sitarama SastryRamya Behara3:41Total length:20:31

text: All lyrics written by Madhan Karky.

text: Tamil track listingNo.TitleSinger(s)Length1."Mauna Mazhaiyilae"Nayana NairAnurag Kulkarni4:192."Thandhaay"Charulatha Mani3:373."Mahanadi"Anurag Kulkarni4:444."Kutti Rani"Nayana Nair2:335."Irudhiyil Enna Enjum"Nayana Nair2:586."Amudhai Pole Kidaithaai"Nayana Nair3:10Total length:20:01

text: Scroll.in review said, "soulful and dramatic, just like Savitri".[31]

text: Release[edit]

text: Mahanati and Nadigaiyar Thilagam were awarded a 'U' (universal) certificate by the Central Board of Film Certification, notwithstanding its long runtime of 176 minutes.[32] On 6 April and 10 April 2018 respectively, the first look posters featuring Akkineni and Devarakonda's characters were released, and received positive response.[33] On 14 April, the film's teaser and first look poster were released digitally.[33] They were positively received, and the teaser crossed 1 million views on YouTube within half a day.[33] 20 different looks of the 110 played by Keerthy in Mahanati were released by the makers through publicity posters, first looks and film stills.[34] The following week, a series of teasers revealed the various special appearances in the film.[35] Mahanati and Nadigaiyar Thilagam did not have theatrical trailers released, which was rare for an Indian film.[36]

text: Mahanati was released on 9 May 2018 worldwide,[37] while Nadigaiyar Thilagam was released on 11 May 2018.[38] The overseas distribution rights of Mahanati and all future films produced by Vyjayanthi Movies and Swapna Cinema were acquired by Nirvana Cinemas,[39] with the amount for the former being ₹45 million.[40] Chandrababu Naidu, then Chief Minister of Andhra Pradesh, was impressed with the film and offered to exempt Mahanati from taxes in Andhra Pradesh, but Ashwini Dutt politely declined it.[41] Mahanati was pirated on torrent websites from 17 May, taking a toll on its box office.[42] This was partly due to the inavailability of film tickets from holdovers of the year's previous releases Bharat Ane Nenu, Rangasthalam and Naa Peru Surya.[42] The number of screens were increased in the second week.[1] From the third week of June, Mahanati had free screenings conducted in old age homes.[43]

text: After Mahanati's release, Ganesan's first daughter Kamala Selvaraj alleged that the film was distorted and disagreed with his characterisation. Ashwin responded by stating that Selvaraj was made aware of the plot during its development.[44] Chamundeswari also defended the film, suggesting that Selvaraj may have watched the film through a different perspective.[45] The film's digital rights were acquired by Amazon Prime.[46] The television broadcast rights of Mahanati were acquired by Star India's subsidiary Star Maa; upon its initial run, the film registered a record target rating point (TRP) rating of 20.16.[47]

text: Reception[edit]

text: Critical response[edit]

text: Mahanati received positive reviews from critics.[48] Writing for The Times of India, Neeshita Nyayapati gave the film four stars out of five and commented, "To call Mahanati a celebration of Savitri’s life would be an understatement, because the film breathes her life in a span of almost 3 hours. There are few films in the Telugu film industry that make you feel the way this one does – empathize and fall head over heels in love with the protagonist even in her lowest of lows."[49] A review carried by Sify opines, "As a director, Nag Ashwin has told the story of Savitri in an emotional and poignant manner. His direction, his grip on emotions is outstanding". They found Keerthy "pitch perfect" as Savitri and felt Salmaan "gives the best performance".[50] Referring to the importance given to seemingly minor events, Janani K of India Today writes, "It is these little things that make Mahanati/Nadigaiyar Thilagam, one of the best biopics of all time".[51]

text: Manoj Kumar of The Indian Express gave the film four stars out of five, praising the performances, writing, "The four best things to happen to Nag's film are Dulquer Salmaan, Keerthy Suresh, Samantha Akkineni and the costume designers. These three actors are in top form and equally shoulder this film".[53] Priyanka Sundar of The Hindustan Times gave Mahanati 3.5 stars out of five and applauded Salmaan's performance "whether it is a frustrated and rude Ganesan when Savitri succumbs to coma or his cheesy Romeo avatar wooing Savitri".[54] Sangeetha Devi Dundoo of The Hindu writes, "The film belongs to Keerthy and Dulquer as they imbibe the mannerisms of the legendary actors they’re portraying but don’t end up as caricatures. Dulquer scores with his effortless ease, and Keerthy owns every moment on screen."[55]

text: Suresh Kavirayani, in his review for Deccan Chronicle, gave the film four stars out of five and praised the aspects of production, writing, "Avish deserves a pat on the back for creating the right atmosphere and old-world feel, and Mickey J. Mayer’s music and background score elevate the film".[56] Writing for Scroll.in, Archana Nathan particularly praised the execution: "The 176-minute movie is evenly paced, and the director ensures that the cameo-heavy screenplay is not reduced to a series of entries and exits."[57] Mridula Ramadugu of Firstpost felt the best part of the film was the lack of dramatising Savitri's life, opining, "It's hard not to be biased towards such a film".[58] Krishna Sripada of The News Minute writes, "[T]he story of Mahanati, probably south Indian film industry’s most revered female figure, is a once-in-a-lifetime epic. It is safe to say we are indebted now to Nag Ashwin, who has pulled off a stunner". She suggested that the film "will make you smile through the tears".[59]

text: Hemnath Kumar of Firstpost gave Mahanati three stars out of five; he felt that the film "leaves you craving for more drama", writing, "It is like reading a Wikipedia page of a famous person. You might get all that you are looking for but it is not the same as reading an in-depth book about the same person".[60] Akshara K. V. of Mathrubhumi also gave the film three stars out five and writes, "The length of the film could have been reduced because the second part after the interval gets lagging".[61] Reviewing Nadigaiyar Thilagam, Sudhir Srinivasan of Cinema Express was critical, calling it "a dubbed film in disguise" since, despite being promoted as a straight Tamil film, it retained many scenes from Mahanati without changing the lip-sync scenes.[62] Writing for Film Companion, Baradwaj Rangan said, "Mahanati, despite its numerous failings, cannot be easily dismissed – because it remembers a forgotten era, whose clips are lovingly recreated via scratchy prints and computerised colour".[63]

text: Box office[edit]

text: Mahanati and Nadigaiyar Thilagam became the highest-grossing South Indian film that started a woman as the primary character.[64] It became the fourth-best Telugu opener in the United States in 2018, earning more money than in Andhra Pradesh and Telangana combined on its first day.[65]