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Flamingo (disambiguation)

 

Flamingos or flamingoes /fləˈmɪŋɡoʊz/ are a type of wading bird in the genus Phoenicopterus, the only genus in the family Phoenicopteridae. There are four flamingo species in the Americas and two species in the Old World.

  

Etymology

 

Flamingo comes from Spanish flamenco, "with the colour of flame", in turn coming from Provençal flamenc from flama "flame" and Germanic-like suffix -ing, with a possible influence of words like Fleming. A similar etymology has the Latinate Greek term Phoenicopterus (from Greek: φοινικόπτερος phoinikopteros), literally "blood red-feathered".

  

Taxonomy and systematics

 

Traditionally, the long-legged Ciconiiformes, probably a paraphyletic assemblage, have been considered the flamingos' closest relatives and the family was included in the order. Usually the ibises and spoonbills of the Threskiornithidae were considered their closest relatives within this order. Earlier genetic studies, such as those of Charles Sibley and colleagues, also supported this relationship. Relationships to the waterfowl were considered as well, especially as flamingos are parasitized by feather lice of the genus Anaticola, which are otherwise exclusively found on ducks and geese. The peculiar presbyornithids were used to argue for a close relationship between flamingos, waterfowl, and waders. A 2002 paper concluded they are waterfowl, but a 2014 comprehensive study of bird orders found that flamingos and grebes are not waterfowl, but rather are part of Columbea along with doves, sandgrouse, and mesites.

  

Species

 

Six flamingo species are recognized by most sources, and these are generally placed in one genus. Two species, the Andean and the James's flamingo, are often placed in the genus Phoenicoparrus instead of Phoenicopterus.

 

Old World:

1, Greater flamingo (P. roseus) - Parts of Africa, S. Europe and S. and SW Asia (most widespread flamingo).

 

2, Lesser flamingo (P. minor) - Africa (e.g. Great Rift Valley) to NW India (most numerous flamingo).

 

New World:

1, Chilean flamingo (P. chilensis) - Temperate S. South America.

 

2, James's flamingo (P. jamesi) - High Andes in Peru, Chile, Bolivia and Argentina.

 

3, Andean flamingo (P. andinus) - High Andes in Peru, Chile, Bolivia and Argentina.

 

4, American flamingo (P. ruber) - Caribbean islands, Caribbean Mexico, Belize and Galapagos islands.

  

Prehistoric species of flamingo:

1, Phoenicopterus croizeti (Middle Oligocene – Middle Miocene of C Europe)

 

2, Phoenicopterus floridanus (Early Pliocene of Florida)

 

3, Phoenicopterus stocki (Middle Pliocene of Rincón, Mexico)

 

4, Phoenicopterus copei (Late Pleistocene of W North America and C Mexico)

 

5, Phoenicopterus minutus (Late Pleistocene of California, USA)

 

6, Phoenicopterus aethiopicus

 

7, Phoenicopterus eyrensis (Late Oligocene of South Australia)

 

8, Phoenicopterus novaehollandiae (Late Oligocene of South Australia)

  

Relationship with grebes

 

Recent molecular studies have suggested a relation with grebes, while morphological evidence also strongly supports a relationship between flamingos and grebes. They hold at least eleven morphological traits in common, which are not found in other birds. Many of these characteristics have been previously identified on flamingos, but not on grebes. The fossil palaelodids can be considered evolutionarily, and ecologically, intermediate between flamingos and grebes.

 

For the grebe-flamingo clade, the taxon Mirandornithes ("miraculous birds" due to their extreme divergence and apomorphies) has been proposed. Alternatively, they could be placed in one order, with Phoenocopteriformes taking priority.

  

Description

 

Flamingos often stand on one leg, the other leg tucked beneath the body. The reason for this behaviour is not fully understood. Recent research indicates that standing on one leg may allow the birds to conserve more body heat, given that they spend a significant amount of time wading in cold water. However, the behaviour also takes place in warm water. As well as standing in the water, flamingos may stamp their webbed feet in the mud to stir up food from the bottom.

 

Young flamingos hatch with greyish reddish plumage, but adults range from light pink to bright red due to aqueous bacteria and beta-Carotene obtained from their food supply. A well-fed, healthy flamingo is more vibrantly colored and thus a more desirable mate; a white or pale flamingo, however, is usually unhealthy or malnourished. Captive flamingos are a notable exception; many turn a pale pink as they are not fed carotene at levels comparable to the wild.

  

Behaviour and ecology

 

Feeding:

 

Flamingos filter-feed on brine shrimp and blue-green algae. Their beaks are specially adapted to separate mud and silt from the food they eat, and are uniquely used upside-down. The filtering of food items is assisted by hairy structures called lamellae which line the mandibles, and the large rough-surfaced tongue. The pink or reddish color of flamingos comes from carotenoids in their diet of animal and plant plankton. These carotenoids are broken down into pigments by liver enzymes. The source of this varies by species, and affects the saturation of color. Flamingos whose sole diet is blue-green algae are darker in color compared to those who get it second hand (e.g. from animals that have digested blue-green algae).

 

Lifecycle:

 

Flamingos are very social birds; they live in colonies whose population can number in the thousands. These large colonies are believed to serve three purposes for the flamingos: avoiding predators, maximizing food intake, and using scarce suitable nesting sites more efficiently. Before breeding, flamingo colonies split into breeding groups of between about 15 and 50 birds. Both males and females in these groups perform synchronized ritual displays. The members of a group stand together and display to each other by stretching their necks upwards, then uttering calls while head-flagging, and then flapping their wings. The displays do not seem to be directed towards an individual but instead occur randomly. These displays stimulate "synchronous nesting" (see below) and help pair up those birds who do not already have mates.

 

Flamingos form strong pair bonds although in larger colonies flamingos sometimes change mates, presumably because there are more mates to choose from. Flamingo pairs establish and defend nesting territories. They locate a suitable spot on the mudflat to build a nest (the spot is usually chosen by the female). It is during nest building that copulation usually occurs. Nest building is sometimes interrupted by another flamingo pair trying to commandeer the nesting site for their own use. Flamingos aggressively defend their nesting sites. Both the male and the female contribute to building the nest, and to defending the nest and egg.[citation needed] Occasional same-sex pairs have been reported.

 

After the chicks hatch, the only parental expense is feeding. Both the male and the female feed their chicks with a kind of crop milk, produced in glands lining the whole of the upper digestive tract (not just the crop). Production is stimulated by a hormone called prolactin. The milk contains fat, protein, and red and white blood cells. (Pigeons and doves—Columbidae—also produce a crop milk (just in the glands lining the crop), which contains less fat and more protein than flamingo crop milk.)

 

For the first six days after the chicks hatch, the adults and chicks stay in the nesting sites. At around seven to twelve days old, the chicks begin to move out of their nests and explore their surroundings. When they are two weeks old, the chicks congregate in groups, called "microcrèches", and their parents leave them alone. After a while, the microcrèches merge into "crèches" containing thousands of chicks. Chicks that do not stay in their crèches are vulnerable to predators.

  

Status and conservation

 

In captivity

 

The first flamingo hatched in a European zoo was a Chilean flamingo at Zoo Basel in Switzerland in 1958. Since then, over 389 flamingos have grown up in Basel and been distributed to other zoos around the globe.

 

An 83-year-old greater flamingo, believed to be the oldest in the world, died at the Adelaide Zoo in Australia in January, 2014.

  

[Credit: en.wikipedia.org]

Marbais, Belgium

A CLOSER LOOK AT CORVUS CORONE

  

LEGEND AND MYTHOLOGY

  

Crows appear in the Bible where Noah uses one to search for dry land and to check on the recession of the flood. Crows supposedly saved the prophet, Elijah, from famine and are an Inuit deity. Legend has it that England and its monarchy will end when there are no more crows in the Tower of London. And some believe that the crows went to the Tower attracted by the regular corpses following executions with written accounts of their presence at the executions of Anne Boleyn and Jane Gray.

  

In Welsh mythology, unfortunately Crows are seen as symbolic of evilness and black magic thanks to many references to witches transforming into crows or ravens and escaping. Indian legend tells of Kakabhusandi, a crow who sits on the branches of a wish-fulfilling tree called Kalpataru and a crow in Ramayana where Lord Rama blessed the crow with the power to foresee future events and communicate with the souls.

  

In Native American first nation legend the crow is sometimes considered to be something of a trickster, though they are also viewed positively by some tribes as messengers between this world and the next where they carry messages from the living to those deceased, and even carry healing medicines between both worlds. There is a belief that crows can foresee the future. The Klamath tribe in Oregon believe that when we die, we fly up to heaven as a crow. The Crow can also signify wisdom to some tribes who believe crows had the power to talk and were therefore considered to be one of the wisest of birds. Tribes with Crow Clans include the Chippewa (whose Crow Clan and its totem are called Aandeg), the Hopi (whose Crow Clan is called Angwusngyam or Ungwish-wungwa), the Menominee, the Caddo, the Tlingit, and the Pueblo tribes of New Mexico.

  

The crow features in the Nanissáanah (Ghost dance), popularized by Jerome Crow Dog, a Brulé Lakota sub-chief and warrior born at Horse Stealing Creek in Montana Territory in 1833, the crow symbolizing wisdom and the past, when the crow had became a guide and acted as a pathfinder during hunting. The Ghost dance movement was originally created in 1870 by Wodziwob, or Gray Hair, a prophet and medicine man of the Paiute tribe in an area that became known as Nevada. Ghost dancers wore crow and eagle feathers in their clothes and hair, and the fact that the Crow could talk placed it as one of the sages of the animal kingdom.

  

The five day dances seeking trance,prophecy and exhortations would eventually play a major part in the pathway towards the white man's broken treaties, the infamous battle at Wounded knee and the surrender of Matȟó Wanáȟtaka (Kicking Bear), after officials began to fear the ghost dancers and rituals which seemed to occur prior to battle.

  

Historically the Vikings are the group who made so many references to the crow, and Ragnarr Loðbrók and his sons used this species in his banner as well as appearances in many flags and coats of arms. Also, it had some kind of association with Odin, one of their main deities. Norse legend tells us that Odin is accompanied by two crows. Hugin, who symbolizes thought, and Munin, who represents a memory. These two crows were sent out each dawn to fly the entire world, returning at breakfast where they informed the Lord of the Nordic gods of everything that went on in their kingdoms. Odin was also referred to as Rafnagud (raven-god).

  

The raven appears in almost every skaldic poem describing warfare.Coins dating back to 940's minted by Olaf Cuaran depict the Viking war standard, the Raven and Viking war banners (Gonfalon) depicted the bird also.

  

In Scandinavian legends, crows are a representative of the Goddess of Death, known as Valkyrie (from old Norse 'Valkyrja'), one of the group of maidens who served the Norse deity Odin, visiting battlefields and sending him the souls of the slain worthy of a place in Valhalla. Odin ( also called Wodan, Woden, or Wotan), preferred that heroes be killed in battle and that the most valiant of souls be taken to Valhöll, the hall of slain warriors. It is the crow that provides the Valkyries with important information on who should go. In Hindu ceremonies that are associated to ancestors, the crow has an important place in Vedic rituals. They are seen as messengers of death in Indian culture too.

  

In Germanic legend, Crows are seen as psychonomes, meaning the act of guiding spirits to their final destination, and that the feathers of a crow could cure a victim who had been cursed. And yet, a lone black crow could symbolize impending death, whilst a group symbolizes a lucky omen! Vikings also saw good omens in the crow and would leave offerings of meat as a token.

  

The crow also has sacred and prophetic meaning within the Celtic civilization, where it stood for flesh ripped off due to combat and Morrighan, the warrior goddess, often appears in Celtic mythology as a raven or crow, or else is found to be in the company of the birds. Crow is sacred to Lugdnum, the Celtic god of creation who gave his name to the city of Lug

  

In Greek mythology according to Appolodorus, Apollo is supposedly responsible for the black feathers of the crow, turning them forever black from their pristine white original plumage as a punishment after they brought news that Κορωνις (Coronis) a princess of the Thessalian kingdom of Phlegyantis, Apollo's pregnant lover had left him to marry a mortal, Ischys. In one legend, Apollo burned the crows feathers and then burned Coronis to death, in another Coronis herself was turned into a black crow, and another that she was slain by the arrows of Αρτεμις (Artemis - twin to Apollo). Koronis was later set amongst the stars as the constellation Corvus ("the Crow"). Her name means "Curved One" from the Greek word korônis or "Crow" from the word korônê.

  

A similar Muslim legend allegedly tells of Muhammad, founder of Islam and the last prophet sent by God to Earth, who's secret location was given away by a white crow to his seekers, as he hid in caves. The crow shouted 'Ghar Ghar' (Cave, cave) and thus as punishment, Muhammad turned the crow black and cursed it for eternity to utter only one phrase, 'Ghar, ghar). Native Indian legend where the once rainbow coloured crows became forever black after shedding their colourful plumage over the other animals of the world.

  

In China the Crow is represented in art as a three legged bird on a solar disk, being a creature that helps the sun in its journey. In Japan there are myths of Crow Tengu who were priests who became vain, and turned into this spirit to serve as messengers until they learn the lesson of humility as well as a great Crow who takes part in Shinto creation stories.

  

In animal spirit guides there are general perceptions of what sightings of numbers of crows actually mean:

  

1 Crow Meaning: To carry a message from your near one who died recently.

 

2 Crows Meaning: Two crows sitting near your home signifies some good news is on your way.

 

3 Crows Meaning: An upcoming wedding in your family.

 

4 Crows Meaning: Symbolizes wealth and prosperity.

 

5 Crows Meaning: Diseases or pain.

 

6 Crows Meaning: A theft in your house!

 

7 Crows Meaning: Denotes travel or moving from your house.

 

8 Crows Meaning: Sorrowful events

  

Crows are generally seen as the symbolism when alive for doom bringing, misfortune and bad omens, and yet a dead crow symbolises potentially bringing good news and positive change to those who see it. This wonderful bird certainly gets a mixed bag of contradictory mythology and legend over the centuries and in modern days is often seen as a bit of a nuisance, attacking and killing the babies of other birds such as Starlings, Pigeons and House Sparrows as well as plucking the eyes out of lambs in the field, being loud and noisy and violently attacking poor victims in a 'crow court'....

  

There is even a classic horror film called 'THE CROW' released in 1994 by Miramax Films, directed by Alex Proyas and starring Brandon Lee in his final film appearance as Eric Draven, who is revived by a Crow tapping on his gravestone a year after he and his fiancée are murdered in Detroit by a street gang. The crow becomes his guide as he sets out to avenge the murders. The only son of martial arts expert Bruce Lee, Brandon lee suffered fatal injuries on the set of the film when the crew failed to remove the primer from a cartridge that hit Lee in the abdomen with the same force as a normal bullet. Lee died that day, March 31st 1993 aged 28.

  

The symbolism of the Crow resurrecting the dead star and accompanying him on his quest for revenge was powerful, and in some part based on the history of the carrion crow itself and the original film grossed more than $94 Million dollars with three subsequent sequels following.

  

TAKING A CLOSER LOOK

  

So let's move away from legend, mythology and stories passed down from our parents and grandparents and look at these amazing birds in isolation.

  

Carrion crow are passerines in the family Corvidae a group of Oscine passerine birds including Crows, Ravens, Rooks, Jackdaws, Jays, Magpies, Treepies, Choughs and Nutcrackers. Technically they are classed as Corvids, and the largest of passerine birds. Carrion crows are medium to large in size with rictal bristles and a single moult per year (most passerines moult twice). Carrion crow was one of the many species originally described by Swedish naturalist Carl Linnaeus (Carl Von Linne after his ennoblement) in his 1758 and 1759 editions of 'SYSTEMA NATURAE', and it still bears its original name of Corvus corone, derived from the Latin of Corvus, meaning Raven and the Greek κορώνη (korōnē), meaning crow.

  

Carrion crow are of the Animalia kingdom Phylum: Chordata Class: Aves Order: Passeriformes Family: Corvidae Genus: Corvus and Species: Corvus corone

  

Corvus corone can reach 45-47cm in length with a 93-104cm wingspan and weigh between 370-650g. They are protected under The Wildlife and Countryside Act 1981 in the United Kingdom with a Green UK conservation status which means they are of least concern with more than 1,000,000 territories. Breeding occurs in April with fledging of the chicks taking around twenty nine days following an incubation period of around twenty days with 3 to 4 eggs being the average norm.

  

They are abundant in the UK apart from Northwest Scotland and Ireland where the Hooded crow (Corvus cornix) was considered the same species until 2002. They have a lifespan of around four years, whilst Crow species can live to the age of Twenty years old, and the oldest known American crow in the wild was almost Thirty years old. The oldest documented captive crow died at age Fifty nine. They are smaller and have a shorter lifespan than the Raven, which again is used as a symbol in history to live life to the full and not waste a moment!

  

They are often mistaken for the Rook (Corvus frugilegus), a similar bird, though in the UK, the Rook is actually technically smaller than the Carrion crow averaging 44-46cm in length, 81-99cm wingspan and weighing up to 340g. Rooks have white beaks compared to the black beaks of Carrion crow, a more steeply raked ratio from head to beak, and longer straighter beaks as well as a different plumage pattern. There are documented cases in the UK of singular and grouped Rooks attacking and killing Carrion crows in their territory. Rooks nest in colonies unlike Carrion crows. Carrion crows have only a few natural enemies including powerful raptors such as the northern goshawk, the peregrine falcon, the Eurasian eagle-owl and the golden eagle which will all readily hunt them.

  

Regarded as one of the most intelligent birds, indeed creatures on the planet, studies suggest that Corvids cognitive abilities can rival that of primates such as chimpanzees and gorillas and even provide clues to understanding human intelligence. Crows have relatively large brains for their body size, compared to other animals. Their encephalization quotient (EQ) a ratio of brain to body size, adjusted for size because there isn’t a linear relationship is 4.1. That is remarkably close to chimps at 4.2 whilst humans are 8.1. Corvids also have a very high neuronal density, the number of neurons per gram of brain, factoring in the number of cortical neurons, neuron packing density, interneuronal distance and axonal conduction velocity shows that Corvids score high on this measure as well, with humans scoring the highest.

  

A corvid's pallium is packed with more neurons than a great ape's. Corvids have demonstrated the ability to use a combination of mental tools such as imagination, and anticipation of future events. They can craft tools from twigs and branches to hook grubs from deep recesses, they can solve puzzles and intricate methods of gaining access to food set by humans., and have even bent pieces of wire into hooks to obtain food. They have been proven to have a higher cognitive ability level than seven year old humans.

  

Communications wise, their repertoire of wraw-wraw's is not fully understood, but the intensity, rhythm, and duration of caws seems to form the basis of a possible language. They also remember the faces of humans who have hindered or hurt them and pass that information on to their offspring.

  

Aesop's fable of 'The Crow and the Pitcher, tells of a thirsty crow which drops stones into a water pitcher to raise the water level and enable it to take a drink. Scientists have conducted tests to see whether crows really are this intelligent. They placed floating treats in a deep tube and observed the crows indeed dropping dense objects carefully selected into the water until the treat floated within reach. They had the intelligence to pick up, weigh and discount objects that would float in the water, they also did not select ones that were too large for the container.

  

Pet crows develop a unique call for their owners, in effect actually naming them. They also know to sunbathe for a dose of vitamin D, regularly settling on wooden garden fences, opening their mouths and wings and raising their heads to the sun. In groups they warn of danger and communicate vocally. They store a cache of food for later if in abundance and are clever enough to move it if they feel it has been discovered. They leave markers for their cache. They have even learned to place walnuts and similar hard food items under car tyres at traffic lights as a means of cracking them!

  

Crows regularly gather around a dead fellow corvid, almost like a funeral, and it is thought they somehow learn from each death. They can even remember human faces for decades.Crows group together to attack larger predators and even steal their food, and they have different dialects in different areas, with the ability to mimic the dialect of the alpha males when they enter their territory!

  

They have a twenty year life span, the oldest on record reaching the age of Fifty nine. Crows can leave gifts for those who feed them such as buttons or bright shiny objects as a thank you, and they even kiss and make up after an argument, having mated for life.

  

In mythology they are associated with good and bad luck, being the bringers of omens and even witchcraft and are generally reviled for their attacks on baby birds and small mammals. They have an attack method of to stunning smaller birds before consuming them, tearing violently at smaller, less aggressive birds, which is simply down to the fact that they are so highly intelligent, and also the top of the food chain. Their diet includes over a thousand different items: Dead animals (as their name suggests), invertebrates, grain, as well as stealing eggs and chicks from other birds' nests, worms, insects, fruit, seeds, kitchen scraps. They are highly adaptable when food sources grow scarce. I absolutely love them, they are magnificent, bold, beautiful and incredibly interesting to watch and though at times it is hard to witness attacks made by them, I cannot help but adore them for so many other and more important reasons.

  

OBSERVATIONS ON THE PAIR IN MY GARDEN

  

Crows have been in the area for a while, but rarely had strayed into my garden, leaving the Magpies to own the territory. Things changed towards the end of May when a beautiful female Carrion crow appeared and began to take some of the food that I put down for the other birds. Within a few days she began to appear regularly, on occasions stocking up on food, whilst other times placing pieces in the birdbath to soften them. She would stand on the birdbath and eat and drink and come back over the course of the day to eat the softened food.

  

Shortly afterwards she brought along her mate, a tall and handsome fella, much larger than her who was also very vocal if he felt she was getting a little too close to me. By now I had moved from a seated position from the patio as an observer, to laying on a mat just five feet from the birdbath with my Nikon so that I could photograph the pair as they landed, scavenged and fed. She was now confident enough to let me be very close, and she even tolerated and recognized the clicking of the camera. At first I used silent mode to reduce the noise but this only allowed two shooting frame rates of single frame or continuous low frame which meant I was missing shots. I reverted back to normal continuous high frames and she soon got used to the whirring of the mechanisms as the mirror slapped back and forth.

  

The big fella would bark orders at her from the safety of the fence or the rear of the garden, whilst she rarely made a sound. That was until one day when in the sweltering heat she kept opening her beak and sunning on the grass, panting slightly in the heat. I placed the circular water sprayer nearby and had it rotating so that the birdbath and grass was bathed in gentle water droplets and she soon came back, landed and seemed to really like the cooling effect on offer. She then climbed onto the birdbath and opened her wings slightly and made some gentle purring, cooing noises....

  

I swear she was expressing happiness, joy....

  

On another blisteringly hot day when the sprayer was on, she came down, walked towards it and opened her wings up running into the water spray. Not once, but many times.

  

A further revelation into the unseen sides to these beautiful birds came with the male and female on the rear garden fence. They sat together, locked beaks like a kiss and then the male took his time gently preening her head feathers and the back of her neck as she made tiny happy sounds. They stayed together like that for several minutes, showing a gentle, softer side to their nature and demonstrating the deep bond between them. Into July and the pair started to bring their three youngsters to my garden, the nippers learning to use the birdbath for bathing and dipping food, the parents attentive as ever. Two of the youngsters headed off once large enough and strong enough.

  

I was privileged to be in close attendance as the last juvenile was brought down by the pair, taught to take food and then on a night in July, to soar and fly with it's mother in the evening sky as the light faded. She would swoop and twirl, and at regular intervals just touch the juvenile in flight with her wing tip feathers, as if to reassure it that she was close in attendance. What an amazing experience to view. A few days later, the juvenile, though now gaining independence and more than capable of tackling food scraps in the garden, was still on occasions demand feeding from it's mother who was now teaching him to take chicken breast, hotdogs or digestive biscuits and bury them in the garden beds for later delectation.

  

The juvenile also liked to gather up peanuts and bury them in the grass. On one occasion I witnessed a pair of rambunctious Pica Pica (Magpies), chasing the young crow on rooftops, leaping at him no matter how hard he tried to get away. He defended himself well and survived the attacks, much to my relief.

  

Into August and the last youngster remained with the adults, though now was very independent even though he still spent time with his parents on rooftops, and shared food gathering duties with his mum. Hotdog sausages were their favourite choice, followed by fish fingers and digestive biscuits which the adult male would gather up three at a time.

  

Corvus Corone.... magnificently misunderstood by some!

  

Paul Williams June 4th 2021

  

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Photograph taken at an altitude of Sixty two metres at 13:19pm on a cold and rainy afternoon on Wednesday 19th May 2021, off Hythe Avenue and Chessington Avenue in Bexleyheath, Kent.

  

Here we see a large adult Carrion crow (Corvus corone) , a passerine bird of the family Corvidae and the genus Raven (Higher classification: Corvus), which is native to western Europe and eastern Asia.

  

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Nikon D850 Focal length 600mm Shutter speed: 1/800s Electronic front-curtain Aperture f/6.3 iso250 Tripod mounted with Tamron VC Vibration Control set to position 3. Image area FX (36 x 24) NEF RAW L (4128 x 2752). JPeg basic (14 bit uncompressed) AF-C Priority Selection: Release. Nikon Back button focusing enabled. AF-S Priority selection: Focus. 3D Tracking watch area: Normal 55 Tracking points Exposure mode: Manual exposure mode Metering mode: Matrix metering White balance on: Auto1 (5160k) Colour space: RGB Picture control: Neutral (Sharpening +2)

  

Tamron SP 150-600mm F/5-6.3 Di VC USD G2. Nikon GP-1 GPS module. Lee SW150 MKII filter holder. Lee SW150 95mm screw in adapter ring. Lee SW150 circular polariser glass filter.Lee SW150 Filters field pouch. Hoodman HEYENRG round eyepiece oversized eyecup.Manfrotto MT057C3-G Carbon fiber Geared tripod 3 sections. Neewer Carbon Fiber Gimble tripod head 10088736 with Arca Swiss standard quick release plate. Neewer 9996 Arca Swiss release plate P860 x2.Jessops Tripod bag. Mcoplus professional MB-D850 multi function battery grip 6960.Two Nikon EN-EL15a batteries (Priority to battery in Battery grip). Black Rapid Curve Breathe strap. My Memory 128GB Class 10 SDXC 80MB/s card. Lowepro Flipside 400 AW camera bag.

     

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LATITUDE: N 51d 28m 27.99s

LONGITUDE: E 0d 8m 10.43s

ALTITUDE: 54.0m

  

RAW (TIFF) FILE: 130.00MB NEF FILE: 91.2MB

PROCESSED (JPeg) FILE: 38.20MB

   

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PROCESSING POWER:

  

Nikon D850 Firmware versions C 1.10 (9/05/2019) LD Distortion Data 2.018 (18/02/20) LF 1.00

  

HP 110-352na Desktop PC with AMD Quad-Core A6-5200 APU 64Bit processor. Radeon HD8400 graphics. 8 GB DDR3 Memory with 1TB Data storage. 64-bit Windows 10. Verbatim USB 2.0 1TB desktop hard drive. WD My Passport Ultra 1tb USB3 Portable hard drive. Nikon ViewNX-1 64bit Version 1.4.1 (18/02/2020). Nikon Capture NX-D 64bit Version 1.6.2 (18/02/2020). Nikon Picture Control Utility 2 (Version 2.4.5 (18/02/2020). Nikon Transfer 2 Version 2.13.5. Adobe photoshop Elements 8 Version 8.0 64bit.

   

Venus Optics Laowa 25mm f2.8 2.5-5x Ultra Macro of a Dark Chocolate Digestive @ 2.5x

Part of my Ultra Macro Sweets series

I always think of August as the mirror-image of February. There is often little let-up from the extremes of temperature that typically occur during the preceding month, but by the fourth week change is in the air. The season seems to have overstayed its welcome and one greets the coming adjustment. As the strength and duration of daylight rapidly increases during February, so in August it declines. The caterpillar-gnawed foliage of the roadside has lost its brilliance. Swallows assemble and look to the waning sun, feeling the draw of Afric's shores. Flocculent thistledown, suspended in the heavy air ...cont. p94.

Yes, here at Latitude 52° North the season is short and we must make hay while the sun shines. Without winter we wouldn't appreciate the summer so much. It's all so beautiful, yet one day we must take our leave of it. I think the crop on the right is what I call "sweet corn" ...grown in this country only as cattle feed. In America this grain, proof against all normal digestive processes, is regarded as fit for human consumption. My American-born wife insists that it is "corn", which I regard as a generic term covering all grain crops, citing Samuel Palmer as my authority. This is a long-running marital-linguistic dispute, equalled in its ferocity only by the zed/zee and aluminium/aluminum controversies. A film user-upper shot.

The Premier Biscuit of Britain.

Tahoe National Forest

 

Some birds have specialized digestive systems that enable them to eat winter foods that most other birds pass up. Yellow-rumped warblers migrate south from the coniferous forests of Alaska and Canada where they breed, but can spend their winter farther north than most other warblers because they can eat wax myrtle berries and bayberries. These fatty berries contain wax (colonial settlers used them to male candles), a high source of energy that other warblers can't digest.

 

Songbird Journeys - Miyoko Chu.

I have, over the years, gravitated to plain biscuits such as Digestives for my afters treat. However, come Christmas my son, who lives in Peterborough which has a wonderful British shop, gets me all kinds of goodies. This year I received a package of Marks & Spencer Milk Digestives. I also received a box of M&S tea, a Cadbury's Dairy Milk and Smarties (so much better than our versions here), Robertson's Shredless Marmalade, a jar of Pickled Onions and Imperial Leather Soap. It was all grand.

 

Funny story- kind of. I took these photos and whilst processing them I noticed it was the French side of the Christie Digestives box that was showing. (All products sold in stores in Canada have to have bilingual packaging by law). So back to my bedroom studio to shoot it all over again with the English side showing. It was a brain fart moment.

 

We're here looks at Biscuits today and I'm showing off a box of Christie Digestives and a half-eaten package of M&S Milk Chocolate Digestives. The bite out of the biscuit is courtesy of my teeth in case you were wondering. The rest magically disappeared at the end of the shoot.

Fripon après déjeuner...décontrasté dans le canapé...comme disait Garcimore le magicien de mon enfance.

500+ pieces of plastic debris that were found in the digestive tract of a dead albatross chick from the North Pacific Gyre.

 

About Mandy Barker from her website.......

  

"Mandy Barker is an international award-winning photographer whose work involving marine plastic debris for more than 12 years, has received global recognition. Working with scientists she aims to raise awareness about plastic pollution in the world's oceans, highlighting the harmful affect on marine life and ourselves - ultimately leading the viewer to take action.

 

Barker's work has been published in over 50 different countries including; National Geographic Magazine, TIME Magazine, The Guardian, The Financial Times, Smithsonian, The New Scientist, The Explorer's Journal, UNESCO, The British Journal of Photography, VOGUE, the World Wildlife Fund, and also to illustrate key academic and scientific research papers about current plastic research. Her work has been exhibited world-wide from MoMA Museum of Modern Art, and the United Nations headquarters in New York, the Victoria & Albert Museum London, and the Science & Technology Park Hong Kong. Barker was shortlisted for the Prix Pictet Award SPACE 2017, the world's leading photography award for sustainability, and nominated for the Magnum Foundation Fund, LOBA Award, and the Deutsche Börse Foundation Photography Prize 2020. She is a recipient of the 2018 National Geographic Society Grant for Research and Exploration. Her first book 'Beyond Drifting: Imperfectly Known Animals' was selected as one of the Ten Best Photography Books of 2017, by Smithsonian, and ‘Altered Ocean’ was chosen by The Royal Photographic Society as one of the most coveted titles and top 10 Photobooks of 2019. Barker is a member of the Union of Concerned Photographers UCP, which is dedicated to using the power of imagery to underline the urgency of environmental concerns.

 

In June 2019 Barker took part in the ‘Henderson Island Plastic Pollution Expedition’ which has been awarded the title of an ‘Explorers Club Flag Expedition’. With only 3 - 5 expeditions per year recognised in this way - previous others having included the Apollo 11 Space Mission, and the dive to Challenger Deep - this significant achievement included recording data as well as photographing marine plastic pollution, has now become part of the archives, accessible to other modern day explorers and scholars.

 

In 2012 she was awarded The Royal Photographic Society's Environmental Bursary enabling her to join scientists in a research expedition which sailed from Japan to Hawaii to examine the accumulation of marine plastic debris in the tsunami debris field in the Pacific Ocean. In June 2017 she was invited by Greenpeace to join the Beluga II Expedition which sailed around the remote and unique island locations of the Inner Hebrides, Scotland, to recover plastic debris in a commission for Greenpeace. Barker speaks internationally about her work to engage people with the plastic issue. She has been invited as a guest speaker at the National Geographic Photography Seminar 2018 Washington DC, Stanford University California, for the British Council, and on behalf of the British Embassy at the political festival Almedalen in Sweden, also as the opening keynote speaker for the EuroConference GlobalCapital Sustainable & Responsible Capital Markets Forum in Amsterdam, and on the panel for the Opening Session for EU Greenweek 2021 in Brussels with Ursula von der Leyen, President of the European Commission. Throughout the COP26 Climate Change Conference 2021 hosted in the UK, Barker’s images were screened in a short film made about her working process by First Move Productions, and presented by the British Council in the Pavilion, Glasgow. She also presented her work in 17 different countries during COP26 to engage the world with this critical climate change issue.

 

In 2019 Barker collaborated with Stanford University on the launch of the virtual reality experience, ‘Ripple: the unintended life of plastics in the sea’. Stanford’s Communication Programme in Journalism worked with several of Barker’s images to represent how ubiquitous plastic has now become part of our world, creating an experience available to everyone across all platforms, from a 360º headset to a mobile phone. Engaging the younger generation is an important part of Barker’s practise to inspire change, she has been committed to teaching workshops around the world for many years, with local communities, schools, and univeristies, from the Philippines to the Solomon Islands, and with individual initiatives for ‘World Oceans Days’. She was part of a youth mentoring programme with First Exposures, an organisation that empowers youth through photography in San Francisco and freely gives her time to mentor others.

 

"The aim of my work is to engage with and stimulate an emotional response in the viewer by combining a contradiction between initial aesthetic attraction along with the subsequent message of awareness. The research process is a vital part of my development as the images I make are based on scientific fact, essential to the integrity of my work. The impact of marine plastic is an area I have documented for more than 10 years and am committed to pursuing through visual interpretation, and in collaboration with science I hope it will ultimately lead to positive action in tackling this increasing environmental problem, which is currently of global concern".

 

“I hope your work does its job in raising an awareness of the cause we both care so much about.

With renewed wonder and best wishes”

Sir David Attenborough

(extract from personal handwritten letter)."

Villers-la-Ville, Belgium

www.desihealthtips.in/2018/10/digestive-system-prescripti...

Papadopoulos brand.

Ommatius coeraebus

Family: Asilidae

Order: Diptera

 

The robber flies in the family Asilidae are robust, bristly flies with a stout proboscis. They are predatory and the prey include a variety of arthropods. It uses its proboscis for stabbing its prey. it injects the prey with a mixture of digestive enzymes and powerful toxins, many of which are unique to robber flies. The liquified contents of the prey are then sucked out.

 

This robber fly species can be found all along the east coast of Australia. It is an opportunistic predator of a range of insects and is known to cannibalise its own species when opportunity arises.

 

I was pushing my luck on shutter speed to see how much I can rely on the optical stabilisation in the Sony lens. I took this shot at 1/80 using a 400mm lens on a monopod. The OS coped very well. (I tried it without OS and the results were not great). Aperture was F5.6. So by having the aperture wide open and the speed very slow I got the ISO down to 125 which allowed heavy cropping - which you need if you are using a long lens.

 

I focus stacked four shots using Boltnev and Kacher's excellent 'Focus Stacker' software. Distance to subject about 1.5m. Estimated size of subject 15-18mm.

  

DSC00884 _93 Robber Fly

I like this biscuits very much.

Sheffield is a city in South Yorkshire, England, whose name derives from the River Sheaf which runs through it. The city serves as the administrative centre of the City of Sheffield. It is historically part of the West Riding of Yorkshire and some of its southern suburbs were transferred from Derbyshire to the city council. It is the largest settlement in South Yorkshire.

 

The city is in the eastern foothills of the Pennines and the valleys of the River Don with its four tributaries: the Loxley, the Porter Brook, the Rivelin and the Sheaf. Sixty-one per cent of Sheffield's entire area is green space and a third of the city lies within the Peak District national park and is the fifth largest city in England. There are more than 250 parks, woodlands and gardens in the city, which is estimated to contain around 4.5 million trees. The city is 29 miles (47 km) south of Leeds and 32 miles (51 km) east of Manchester.

 

Sheffield played a crucial role in the Industrial Revolution, with many significant inventions and technologies having developed in the city. In the 19th century, the city saw a huge expansion of its traditional cutlery trade, when stainless steel and crucible steel were developed locally, fuelling an almost tenfold increase in the population. Sheffield received its municipal charter in 1843, becoming the City of Sheffield in 1893. International competition in iron and steel caused a decline in these industries in the 1970s and 1980s, coinciding with the collapse of coal mining in the area. The Yorkshire ridings became counties in their own right in 1889, the West Riding of Yorkshire county was disbanded in 1974. The city then became part of the county of South Yorkshire; this has been made up of separately-governed unitary authorities since 1986. The 21st century has seen extensive redevelopment in Sheffield, consistent with other British cities. Sheffield's gross value added (GVA) has increased by 60% since 1997, standing at £11.3 billion in 2015. The economy has experienced steady growth, averaging around 5% annually, which is greater than that of the broader region of Yorkshire and the Humber.

 

Sheffield had a population of 556,500 at the 2021 census, making it the second largest city in the Yorkshire and the Humber region. The Sheffield Built-up Area, of which the Sheffield sub-division is the largest part, had a population of 685,369 also including the town of Rotherham. The district borough, governed from the city, had a population of 554,401 at the mid-2019 estimate, making it the 7th most populous district in England. It is one of eleven British cities that make up the Core Cities Group. In 2011, the unparished area had a population of 490,070.

 

The city has a long sporting heritage and is home both to the world's oldest football club, Sheffield F.C., and the world's oldest football ground, Sandygate. Matches between the two professional clubs, Sheffield United and Sheffield Wednesday, are known as the Steel City derby. The city is also home to the World Snooker Championship and the Sheffield Steelers, the UK's first professional ice hockey team.

 

The history of Sheffield, a city in South Yorkshire, England, can be traced back to the founding of a settlement in a clearing beside the River Sheaf in the second half of the 1st millennium AD. The area now known as Sheffield had seen human occupation since at least the last ice age, but significant growth in the settlements that are now incorporated into the city did not occur until the Industrial Revolution.

 

Following the Norman conquest of England, Sheffield Castle was built to control the Saxon settlements and Sheffield developed into a small town, no larger than Sheffield City Centre. By the 14th century Sheffield was noted for the production of knives, and by 1600, overseen by the Company of Cutlers in Hallamshire, it had become the second centre of cutlery production in England after London. In the 1740s the crucible steel process was improved by Sheffield resident Benjamin Huntsman, allowing a much better production quality. At about the same time, Sheffield plate, a form of silver plating, was invented. The associated industries led to the rapid growth of Sheffield; the town was incorporated as a borough in 1843 and granted a city charter in 1893.

 

Sheffield remained a major industrial city throughout the first half of the 20th century, but the downturn in world trade following the 1973 oil crisis, technological improvements and economies of scale, and a wide-reaching restructuring of steel production throughout the European Economic Community led to the closure of many of the steelworks from the early 1970s onward. Urban and economic regeneration schemes began in the late 1980s to diversify the city's economy. Sheffield is now a centre for banking and insurance functions with HSBC, Santander and Aviva having regional offices in the city. The city has also attracted digital start-ups, with 25,000 now employed in the digital sector.

 

Early history

Photograph showing a moorland view. The moor is covered in heather of varying shades of brown. Stones are scattered across the moor. In the middle distance there is a rock outcrop atop a small hill. Behind it is a larger hill with a flat top.

Carl Wark, an Iron Age hill fort in southwest Sheffield.

The earliest known evidence of human occupation in the Sheffield area was found at Creswell Crags in Derbyshire to the east of the city. Artefacts and rock art found in caves at this site have been dated by archaeologists to the late Upper Palaeolithic period, at least 12,800 years ago. Other prehistoric remains found in Sheffield include a Mesolithic "house"—a circle of stones in the shape of a hut-base dating to around 8000 BC, found at Deepcar, in the northern part of the city. This has been ascribed to the Maglemosian culture. (grid reference SK 2920 9812). The site's culture has similarities to Star Carr in North Yorkshire, but gives its name to unique "Deepcar type assemblages" of microliths in the archaeology literature. A cup and ring-marked stone was discovered in Ecclesall Woods in 1981, and has been dated to the late Neolithic or Bronze Age periods. It, and an area around it of 2 m diameter, is a scheduled ancient monument.

 

During the Bronze Age (about 1500 BC) tribes sometimes called the Urn people started to settle in the area. They built numerous stone circles, examples of which can be found on Ash Cabin Flat, Froggatt Edge and Hordron Edge (Hordron Edge stone circle). Two Early Bronze Age urns were found at Crookes in 1887, and three Middle Bronze Age barrows found at Lodge Moor (both suburbs of the modern city).

 

Iron Age

During the British Iron Age the area became the southernmost territory of the Pennine tribe called the Brigantes. It is this tribe who in around 500 BC are thought to have constructed the hill fort that stands on the summit of a steep hill above the River Don at Wincobank, in what is now northeastern Sheffield. Other Iron Age hill forts in the area are Carl Wark on Hathersage Moor to the southwest of Sheffield, and one at Scholes Wood, near Rotherham. The rivers Sheaf and Don may have formed the boundary between the territory of the Brigantes and that of a rival tribe called the Corieltauvi who inhabited a large area of the northeastern Midlands.

 

Roman Britain

The Roman invasion of Britain began in AD 43. By 51 the Brigantes had submitted to the clientship of Rome, eventually being placed under direct rule in the early 70s. Few Roman remains have been found in the Sheffield area. A minor Roman road linking the Roman forts at Templeborough and Navio at Brough-on-Noe possibly ran through the centre of the area covered by the modern city, and Icknield Street is thought to have skirted its boundaries. The routes of these roads within this area are mostly unknown, although sections of the former were thought, by Hunter and Leader, be visible between Redmires and Stanage on an ancient road known as the Long Causeway. In recent years some scholars have cast doubt on this, with an initial survey of Barber Fields, Ringinglow, suggesting the Roman Road took a route over Burbage Edge. The remains of a Roman road, possibly linked to the latter, were discovered in Brinsworth in 1949.

 

In April 1761, tablets or diplomas dating from the Roman period were found in the Rivelin Valley south of Stannington, close to what was possibly the course of the Templeborough to Brough-on-Noe road. These tablets included a grant of citizenship and land or money to a retiring Roman auxiliary of the Sunuci tribe of Belgium.

 

To . . . . . . . . the son of Albanus, of the tribe of the Sunuci, late a foot soldier in the first cohort of the Sunuci commanded by M. Junius Claudianus.

 

In addition there have been finds dating from the Roman period on Walkley Bank Road, which leads onto the valley bottom.

 

There have been small finds of Roman coins throughout the Sheffield area, for example 30 to 40 Roman coins were found near the Old Great Dam at Crookesmoor, 19 coins were found near Meadowhall in 1891, 13 in Pitsmoor in 1906, and ten coins were found at a site alongside Eckington cemetery in December 2008. Roman burial urns were also found at Bank Street near Sheffield Cathedral, which, along with the name of the old lane behind the church (Campo Lane[n 2]), has led to speculation that there may have been a Roman camp at this site. It is unlikely that the settlement that grew into Sheffield existed at this time. In 2011 excavations revealed remains of a substantial 1st or 2nd century AD Roman rural estate centre, or 'villa' on what is believed to be a pre-existing Brigantian farmstead site at Whirlow Hall Farm in South-west Sheffield.

 

Following the departure of the Romans, the Sheffield area may have been the southern part of the Celtic kingdom of Elmet, with the rivers Sheaf and Don forming part of the boundary between this kingdom and the kingdom of Mercia. Gradually, Anglian settlers pushed west from the kingdom of Deira. The Britons of Elmet delayed this English expansion into the early part of the 7th century. An enduring Celtic presence within this area is evidenced by the settlements called Wales and Waleswood close to Sheffield—the word Wales derives from the Germanic word Walha, and was originally used by the Anglo-Saxons to refer to the native Britons.

 

The origins of Sheffield

The name Sheffield is Old English in origin. It derives from the River Sheaf, whose name is a corruption of shed or sheth, meaning to divide or separate. Field is a generic suffix deriving from the Old English feld, meaning a forest clearing. It is likely then that the origin of the present-day city of Sheffield is an Anglo-Saxon settlement in a clearing beside the confluence of the rivers Sheaf and Don founded between the arrival of the Anglo-Saxons in this region (roughly the 6th century) and the early 9th century.

 

The names of many of the other areas of Sheffield likely to have been established as settlements during this period end in ley, which signifies a clearing in the forest, or ton, which means an enclosed farmstead. These settlements include Heeley, Longley, Norton, Owlerton, Southey, Tinsley, Totley, Treeton, Wadsley, and Walkley.

 

The earliest evidence of this settlement is thought to be the shaft of a stone cross dating from the early 9th century that was found in Sheffield in the early 19th century. This shaft may be part of a cross removed from the church yard of the Sheffield parish church (now Sheffield Cathedral) in 1570. It is now kept in the British Museum.

 

A document from around the same time, an entry for the year 829 in the Anglo-Saxon Chronicle, refers to the submission of King Eanred of Northumbria to King Egbert of Wessex at the hamlet of Dore (now a suburb of Sheffield): "Egbert led an army against the Northumbrians as far as Dore, where they met him, and offered terms of obedience and subjection, on the acceptance of which they returned home". This event made Egbert the first Saxon to claim to be king of all of England.

 

The latter part of the 9th century saw a wave of Norse (Viking) settlers and the subsequent establishment of the Danelaw. The names of hamlets established by these settlers often end in thorpe, which means a farmstead. Examples of such settlements in the Sheffield area are Grimesthorpe, Hackenthorpe, Jordanthorpe, Netherthorpe, Upperthorpe, Waterthorpe, and Woodthorpe. By 918 the Danes south of the Humber had submitted to Edward the Elder, and by 926 Northumbria was under the control of King Æthelstan.

 

In 937 the combined armies of Olaf Guthfrithson, Viking king of Dublin, Constantine, king of Scotland and Owain ap Dyfnwal, king of the Cumbrians, invaded England. The invading force was met and defeated by an army from Wessex and Mercia led by King Æthelstan at the Battle of Brunanburh. The location of Brunanburh is unknown, but some historians have suggested a location between Tinsley in Sheffield and Brinsworth in Rotherham, on the slopes of White Hill. After the death of King Athelstan in 939 Olaf Guthfrithson invaded again and took control of Northumbria and part of Mercia. Subsequently, the Anglo-Saxons, under Edmund, re-conquered the Midlands, as far as Dore, in 942, and captured Northumbria in 944.

 

The Domesday Book of 1086, which was compiled following the Norman Conquest of 1066, contains the earliest known reference to the districts around Sheffield as the manor of "Hallun" (or Hallam). This manor retained its Saxon lord, Waltheof, for some years after the conquest. The Domesday Book was ordered written by William the Conqueror so that the value of the townships and manors of England could be assessed. The entries in the Domesday Book are written in a Latin shorthand; the extract for this area begins:

 

TERRA ROGERII DE BVSLI

M. hi Hallvn, cu XVI bereuvitis sunt. XXIX. carucate trae

Ad gld. Ibi hb Walleff com aula...

Translated it reads:

 

LANDS OF ROGER DE BUSLI

Photograph showing an old stone church with a short wide tower. The view is taken from a graveyard, there is a large tomb stone in the foreground and the church is surrounded by trees.

 

The remains of Beauchief Abbey.

In Hallam, one manor with its sixteen hamlets, there are twenty-nine carucates [~14 km2] to be taxed. There Earl Waltheof had an "Aula" [hall or court]. There may have been about twenty ploughs. This land Roger de Busli holds of the Countess Judith. He has himself there two carucates [~1 km2] and thirty-three villeins hold twelve carucates and a half [~6 km2]. There are eight acres [32,000 m2] of meadow, and a pasturable wood, four leuvae in length and four in breadth [~10 km2]. The whole manor is ten leuvae in length and eight broad [207 km2]. In the time of Edward the Confessor it was valued at eight marks of silver [£5.33]; now at forty shillings [£2.00].

In Attercliffe and Sheffield, two manors, Sweyn had five carucates of land [~2.4 km2] to be taxed. There may have been about three ploughs. This land is said to have been inland, demesne [domain] land of the manor of Hallam.

The reference is to Roger de Busli, tenant-in-chief in Domesday and one of the greatest of the new wave of Norman magnates. Waltheof, Earl of Northumbria had been executed in 1076 for his part in an uprising against William I. He was the last of the Anglo-Saxon earls still remaining in England a full decade after the Norman conquest. His lands had passed to his wife, Judith of Normandy, niece to William the Conqueror. The lands were held on her behalf by Roger de Busli.

 

The Domesday Book refers to Sheffield twice, first as Escafeld, then later as Scafeld. Sheffield historian S. O. Addy suggests that the second form, pronounced Shaffeld, is the truer form, as the spelling Sefeld is found in a deed issued less than one hundred years after the completion of the survey. Addy comments that the E in the first form may have been mistakenly added by the Norman scribe.

 

Roger de Busli died around the end of the 11th century, and was succeeded by a son, who died without an heir. The manor of Hallamshire passed to William de Lovetot, the grandson of a Norman baron who had come over to England with the Conqueror. William de Lovetot founded the parish churches of St Mary at Handsworth, St Nicholas at High Bradfield and St. Mary's at Ecclesfield at the start of the 12th century in addition to Sheffield's own parish church. He also built the original wooden Sheffield Castle, which stimulated the growth of the town.

 

Also dating from this time is Beauchief Abbey, which was founded by Robert FitzRanulf de Alfreton. The abbey was dedicated to Saint Mary and Saint Thomas Becket, who had been canonised in 1172. Thomas Tanner, writing in 1695, stated that it was founded in 1183. Samuel Pegge in his History of Beauchief Abbey notes that Albinas, the abbot of Derby, who was one of the witnesses to the charter of foundation, died in 1176, placing foundation before that date.

 

Medieval Sheffield

Following the death of William de Lovetot, the manor of Hallamshire passed to his son Richard de Lovetot and then his son William de Lovetot before being passed by marriage to Gerard de Furnival in about 1204. The de Furnivals held the manor for the next 180 years. The fourth Furnival lord, Thomas de Furnival, supported Simon de Montfort in the Second Barons' War. As a result of this, in 1266 a party of barons, led by John de Eyvill, marching from north Lincolnshire to Derbyshire passed through Sheffield and destroyed the town, burning the church and castle.

 

A new stone castle was constructed over the next four years and a new church was consecrated by William de Wickwane the Archbishop of York around 1280. In 1295 Thomas de Furnival's son (also Thomas) was the first lord of Hallamshire to be called to Parliament, thus taking the title Lord Furnivall. On 12 November 1296 Edward I granted a charter for a market to be held in Sheffield on Tuesday each week. This was followed on 10 August 1297 by a charter from Lord Furnival establishing Sheffield as a free borough.

 

The Sheffield Town Trust was established in the Charter to the Town of Sheffield, granted in 1297. De Furnival, granted land to the freeholders of Sheffield in return for an annual payment, and a Common Burgery administrated them. The Burgery originally consisted of public meetings of all the freeholders, who elected a Town Collector. Two more generations of Furnivals held Sheffield before it passed by marriage to Sir Thomas Nevil and then, in 1406, to John Talbot, the first Earl of Shrewsbury.

 

The Bishops' House.

In 1430 the 1280 Sheffield parish church building was pulled down and replaced. Parts of this new church still stand today and it is now Sheffield city centre's oldest surviving building, forming the core of Sheffield Cathedral. Other notable surviving buildings from this period include the Old Queen's Head pub in Pond Hill, which dates from around 1480, with its timber frame still intact, and Bishops' House and Broom Hall, both built around 1500.

 

Post-medieval Sheffield

The fourth Earl of Shrewsbury, George Talbot took up residence in Sheffield, building the Manor Lodge outside the town in about 1510 and adding a chapel to the Parish Church c1520 to hold the family vault. Memorials to the fourth and sixth Earls of Shrewsbury can still be seen in the church. In 1569 George Talbot, the sixth Earl of Shrewsbury, was given charge of Mary, Queen of Scots. Mary was regarded as a threat by Elizabeth I, and had been held captive since her arrival in England in 1568.

 

Talbot brought Mary to Sheffield in 1570, and she spent most of the next 14 years imprisoned in Sheffield Castle and its dependent buildings. The castle park extended beyond the present Manor Lane, where the remains of Manor Lodge are to be found. Beside them is the Turret House, an Elizabethan building, which may have been built to accommodate the captive queen. A room, believed to have been the queen's, has an elaborate plaster ceiling and overmantel, with heraldic decorations.[58] During the English Civil War, Sheffield changed hands several times, finally falling to the Parliamentarians, who demolished (slighted) the castle in 1648.

 

The Industrial Revolution brought large-scale steel making to Sheffield in the 18th century. Much of the medieval town was gradually replaced by a mix of Georgian and Victorian buildings. Large areas of Sheffield's city centre have been rebuilt in recent years, but among the modern buildings, some old buildings have been retained.

 

Industrial Sheffield

Sheffield developed after the industrial revolution because of its geography.

 

Fast-flowing rivers, such as the Sheaf, the Don and the Loxley, made it an ideal location for water-powered industries to develop. Raw materials, like coal, iron ore, ganister and millstone grit for grindstones, found in the nearby hills, were used in cutlery and blade production.

 

As early as the 14th century, Sheffield was noted for the production of knives:

 

Ay by his belt he baar a long panade,

And of a swerd ful trenchant was the blade.

A joly poppere baar he in his pouche;

Ther was no man, for peril, dorste hym touche.

A Sheffeld thwitel baar he in his hose.

Round was his face, and camus was his nose;

 

— Geoffrey Chaucer, The Reeve's Tale from The Canterbury Tales

 

By 1600 Sheffield was the main centre of cutlery production in England outside London, and in 1624 The Company of Cutlers in Hallamshire was formed to oversee the trade. Examples of water-powered blade and cutlery workshops from around this time can be seen at the Abbeydale Industrial Hamlet and Shepherd Wheel museums in Sheffield.

 

Around a century later, Daniel Defoe in his book A tour thro' the whole island of Great Britain, wrote:

 

This town of Sheffield is very populous and large, the streets narrow, and the houses dark and black, occasioned by the continued smoke of the forges, which are always at work: Here they make all sorts of cutlery-ware, but especially that of edged-tools, knives, razors, axes, &. and nails; and here the only mill of the sort, which was in use in England for some time was set up, (viz.) for turning their grindstones, though now 'tis grown more common. Here is a very spacious church, with a very handsome and high spire; and the town is said to have at least as many, if not more people in it than the city of York.

 

Sheffield area.

In the 1740s Benjamin Huntsman, a clock maker in Handsworth, invented a form of the crucible steel process for making a better quality of steel than had previously been available. At around the same time Thomas Boulsover invented a technique for fusing a thin sheet of silver onto a copper ingot producing a form of silver plating that became known as Sheffield plate. Originally hand-rolled Old Sheffield Plate was used for making silver buttons. Then in 1751 Joseph Hancock, previously apprenticed to Boulsover's friend Thomas Mitchell, first used it to make kitchen and tableware. This prospered and in 1762–65 Hancock built the water-powered Old Park Silver Mills at the confluence of the Loxley and the Don, one of the earliest factories solely producing an industrial semi-manufacture. Eventually Old Sheffield Plate was supplanted by cheaper electroplate in the 1840s. In 1773 Sheffield was given a silver assay office. In the late 18th century, Britannia metal, a pewter-based alloy similar in appearance to silver, was invented in the town.

 

Huntsman's process was only made obsolete in 1856 by Henry Bessemer's invention of the Bessemer converter, but production of crucible steel continued until well into the 20th century for special uses, as Bessemer's steel was not of the same quality, in the main replacing wrought iron for such applications as rails. Bessemer had tried to induce steelmakers to take up his improved system, but met with general rebuffs, and finally was driven to undertake the exploitation of the process himself. To this end he erected steelworks in Sheffield. Gradually the scale of production was enlarged until the competition became effective, and steel traders generally became aware that the firm of Henry Bessemer & Co. was underselling them to the extent of £20 a ton. One of Bessemer's converters can still be seen at Sheffield's Kelham Island Museum.

 

Stainless steel was discovered by Harry Brearley in 1912, at the Brown Firth Laboratories in Sheffield. His successor as manager at Brown Firth, Dr William Hatfield, continued Brealey's work. In 1924 he patented '18-8 stainless steel', which to this day is probably the most common alloy of this type.

 

These innovations helped Sheffield to gain a worldwide recognition for the production of cutlery; utensils such as the bowie knife were mass-produced and shipped to the United States. The population of the town increased rapidly. In 1736 Sheffield and its surrounding hamlets held about 7000 people, in 1801 there were 60,000, and by 1901, the population had grown to 451,195.

 

This growth spurred the reorganisation of the governance of the town. Prior to 1818, the town was run by a mixture of bodies. The Sheffield Town Trust and the Church Burgesses, for example, divided responsibility for the improvement of streets and bridges. By the 19th century both organisations lacked funds and struggled even to maintain existing infrastructure.[52] The Church Burgesses organised a public meeting on 27 May 1805 and proposed to apply to Parliament for an act to pave, light and clean the city's streets. The proposal was defeated.

 

The idea of a Commission was revived in 1810, and later in the decade Sheffield finally followed the model adopted by several other towns in petitioning for an Act to establish an Improvement Commission. This eventually led to the Sheffield Improvement Act 1818, which established the Commission and included several other provisions. In 1832 the town gained political representation with the formation of a Parliamentary borough. A municipal borough was formed by an Act of Incorporation in 1843, and this borough was granted the style and title of "City" by letters patent in 1893.

 

In 1832 an outbreak of cholera killed 402 people, including John Blake, the Master Cutler. Another 1,000 residents were infected by the disease. A memorial to the victims stands in Clay Wood where the victims of the outbreak are buried.

 

From the mid-18th century, a succession of public buildings were erected in the town. St Paul's Church, now demolished, was among the first, while the old Town Hall and the present Cutlers' Hall were among the major works of the 19th century. The town's water supply was improved by the Sheffield Waterworks Company, who built reservoirs around the town. Parts of Sheffield were devastated when, following a five-year construction project, the Dale Dyke dam collapsed on Friday 11 March 1864, resulting in the Great Sheffield Flood.

 

Sheffield's transport infrastructure was also improved. In the 18th century turnpike roads were built connecting Sheffield with Barnsley, Buxton, Chesterfield, Glossop, Intake, Penistone, Tickhill, and Worksop. In 1774 a 2-mile (3.2 km) wooden tramway was laid at the Duke of Norfolk's Nunnery Colliery. The tramway was destroyed by rioters, who saw it as part of a plan to raise the price of coal. A replacement tramway that used L-shaped rails was laid by John Curr in 1776 and was one of the earliest cast-iron railways. The Sheffield Canal opened in 1819 allowing the large-scale transport of freight.

 

This was followed by the Sheffield and Rotherham Railway in 1838, the Sheffield, Ashton-under-Lyne and Manchester Railway in 1845, and the Midland Railway in 1870. The Sheffield Tramway was started in 1873 with the construction of a horse tram route from Lady's Bridge to Attercliffe. This route was later extended to Brightside and Tinsley, and further routes were constructed to Hillsborough, Heeley, and Nether Edge. Due to the narrow medieval roads the tramways were initially banned from the town centre. An improvement scheme was passed in 1875; Pinstone Street and Leopold Street were constructed by 1879, and Fargate was widened in the 1880s. The 1875 plan also called for the widening of the High Street; disputes with property owners delayed this until 1895.

 

Steel production in the 19th century involved long working hours, in unpleasant conditions that offered little or no safety protection. Friedrich Engels in his The Condition of the Working Class in England in 1844 described the conditions prevalent in the city at that time:

 

In Sheffield wages are better, and the external state of the workers also. On the other hand, certain branches of work are to be noticed here, because of their extraordinarily injurious influence upon health. Certain operations require the constant pressure of tools against the chest, and engender consumption in many cases; others, file-cutting among them, retard the general development of the body and produce digestive disorders; bone-cutting for knife handles brings with it headache, biliousness, and among girls, of whom many are employed, anæmia. By far the most unwholesome work is the grinding of knife-blades and forks, which, especially when done with a dry stone, entails certain early death. The unwholesomeness of this work lies in part in the bent posture, in which chest and stomach are cramped; but especially in the quantity of sharp-edged metal dust particles freed in the cutting, which fill the atmosphere, and are necessarily inhaled. The dry grinders' average life is hardly thirty-five years, the wet grinders' rarely exceeds forty-five.

 

Sheffield became one of the main centres for trade union organisation and agitation in the UK. By the 1860s, the growing conflict between capital and labour provoked the so-called 'Sheffield Outrages', which culminated in a series of explosions and murders carried out by union militants. The Sheffield Trades Council organised a meeting in Sheffield in 1866 at which the United Kingdom Alliance of Organised Trades—a forerunner of the Trades Union Congress (TUC)—was founded.

 

The 20th century to the present

In 1914 Sheffield became a diocese of the Church of England, and the parish church became a cathedral. During the First World War the Sheffield City Battalion suffered heavy losses at the Somme and Sheffield itself was bombed by a German zeppelin.

 

The recession of the 1930s was only halted by the increasing tension as the Second World War loomed. The steel factories of Sheffield were set to work making weapons and ammunition for the war. As a result, once war was declared, the city once again became a target for bombing raids. In total there were 16 raids over Sheffield, but it was the heavy bombing over the nights of 12 and 15 December 1940 (now known as the Sheffield Blitz) when the most substantial damage occurred. More than 660 people died and numerous buildings were destroyed.

 

Following the war, the 1950s and 1960s saw many large scale developments in the city. The Sheffield Tramway was closed, and a new system of roads, including the Inner Ring Road, were laid out. Also at this time many of the old slums were cleared and replaced with housing schemes such as the Park Hill flats, and the Gleadless Valley estate.

 

In February 1962, the city was devastated by the Great Sheffield Gale. Extremely localised high winds across the city, reaching up to 97 mph (156 km/h), killed four people, injured more than 400, and damaged more than 150,000 houses across the city, leaving thousands homeless.

 

Sheffield's traditional manufacturing industries (along with those of many other areas in the UK), declined during the 20th century. In the 1980s, it was the setting for two films written by locally-born Barry Hines: Looks and Smiles, a 1981 film that portrayed the depression that the city was enduring, and Threads, a 1984 television film that simulated a nuclear winter in Sheffield after a warhead is dropped to the east of the city.

 

The building of the Meadowhall shopping centre on the site of a former steelworks in 1990 was a mixed blessing, creating much needed jobs but speeding the decline of the city centre. Attempts to regenerate the city were kick-started by the hosting of the 1991 World Student Games and the associated building of new sporting facilities such as the Sheffield Arena, Don Valley Stadium and the Ponds Forge complex. Sheffield began construction of a tram system in 1992, with the first section opening in 1994.

 

Starting in 1995, the Heart of the City Project has seen public works in the city centre: the Peace Gardens were renovated in 1998, the Millennium Gallery opened in April 2001, and a 1970s town hall extension was demolished in 2002 to make way for the Winter Garden, which opened on 22 May 2003. A series of other projects grouped under the title Sheffield One aim to regenerate the whole of the city centre.

 

Sheffield was particularly hard hit during the 2007 United Kingdom floods and the 2010 'Big Freeze'. The 2007 flooding on 25 June caused millions of pounds worth of damage to buildings in the city and led to the loss of two lives. Many landmark buildings such as Meadowhall and the Hillsborough Stadium flooded due to being close to rivers that flow through the city. In 2010, 5,000 properties in Sheffield were identified as still being at risk of flooding. In 2012 the city narrowly escaped another flood, despite extensive work by the Environment Agency to clear local river channels since the 2007 event. In 2014 Sheffield Council's cabinet approved plans to further reduce the possibility of flooding by adopting plans to increase water catchment on tributaries of the River Don. Another flood hit the city in 2019, resulting in shoppers being contained in Meadowhall Shopping Centre.

 

Between 2014 and 2018, there were disputes between the city council and residents over the fate of the city's 36,000 highway trees. Around 4,000 highway trees have since been felled as part of the ‘Streets Ahead’ Private Finance Initiative (PFI) contract signed in 2012 by the city council, Amey plc and the Department for Transport to maintain the city streets. The tree fellings have resulted in many arrests of residents and other protesters across the city even though most felled trees in the city have been replanted, including those historically felled and not previously replanted. The protests eventually stopped in 2018 after the council paused the tree felling programme as part of a new approach developed by the council for the maintenance of street trees in the city.

 

In July 2013 the Sevenstone project, which aimed to demolish and rebuild a large part of the city centre, and had been on hold since 2009, was further delayed and the company developing it was dropped. The city council is looking for partners to take a new version of the plan forwards. In April 2014 the council, together with Sheffield University, proposed a plan to reduce the blight of empty shops in the city centre by offering them free of charge to small businesses on a month-by-month basis.

 

In December 2022, thousands of homes in Hillsborough and Stannington were left without a gas supply for more than a week following a serious failure of the local network. Sheffield City Council declared a major incident as temperatures dropped below freezing in unheated homes, and aid was distributed to local residents.

"Digestive Walk"

 

Vallée de Zillertal (AUTRICHE 2015)

 

Website : www.fluidr.com/photos/pat21

 

"Copyright © – Patrick Bouchenard

The reproduction, publication, modification, transmission or exploitation of any work contained here in for any use, personal or commercial, without my prior written permission is strictly prohibited. All rights reserved

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Villers-la-Ville, Belgium

This was supposed to be a prymaid but the tin wasnt deep enough even though I tried to do it at an angle, its yummy it has apricots and nuts in it and chocolate orange on top

Buscando en los directorios encontre esta foto de un trabajo antiguo.

Corniche Kennedy, Marseille, France

Story and recipes Via delle rose

A big fat juicy beetle who just happened to get snagged in it's web to be followed by dessert of a little spider which has been enclosed in a silk 'coffin'. I think this is some kind of orbweaver. I watched it bite it's prey, wrap it in silk, waited for it to die, vomited over it and then it begin to eat. The process is the spider will literally vomit digestive fluid over the prey. The prey is then chewed with the "jaws" (chelicerae), and the fluid is sucked back into the mouth together with some liquefied "meat" from the prey. The spider repeats this process as often as necessary to digest, and ingest, all but the inedible hard parts. What is discarded afterwards is a small ball of residue......No wonder Miss Muffet was frightened away!!

 

Bronica | 400TX

Recipe and story on Un déjeuner de soleil

Villers-la-Ville, Belgium

ink on paper, 2017

Not my best pic by a long way but I thought my Flickr chums would be interested. This is a Portuguese-man-o-war. Not actually a jelly fish but a conglomerate of 4 polyps - the bladder at the top, the venomous tentacles (up to 65ft long) below, the digestive system and the reproductive system are all provided by separate organisms. Our guide spotted trhis while belting back to harbour and it was gone in a flash but a first sighting for me.

www.etsy.com/listing/76667574/anatomy-guts

Cuban Flamingo - Rare Species Centre, Kent, England - Sunday October 12th 2008.

Click here to see the Larger image

 

Click here to see My most interesting images

 

From Wikipedia, the free encyclopedia ~ Flamingos or flamingoes (pronunciation (help·info)) are gregarious wading birds in the genus Phoenicopterus and family Phoenicopteridae. They are found in both the Western Hemisphere and Eastern Hemisphere, but are more numerous in the former. There are four species in the Americas while two exist in the Old World. Two species, the Andean and the James's Flamingo, are often placed in the genus Phoenicoparrus instead of Phoenicopterus.

 

Description ~ Diet ~ Flamingos filter-feed on brine shrimp. Their oddly-shaped beaks are specially adapted to separate mud and silt from the food they eat, and are uniquely used upside-down. The filtering of food items is assisted by hairy structures called lamellae which line the mandibles, and the large rough-surfaced tongue. The flamingo's characteristic pink colouring is caused by the Beta carotene in their diet. The source of this varies by species, but shrimp and blue-green algae are common sources; zoo-fed flamingos may be given food with the additive canthaxanthin, which is often also given to farmed salmon. Flamingos produce a "milk" like pigeon milk due to the action of a hormone called prolactin (see Columbidae). It contains more fat and less protein than the latter does, and it is produced in glands lining the whole of the upper digestive tract, not just the crop. Both parents nurse their chick, and young flamingos feed on this milk, which also contains red and white blood cells, for about two months until their bills are developed enough to filter feed.

Appearance ~ Flamingos frequently stand on one leg. The reason for this behavior is not fully known. One common theory is that tucking one leg beneath the body may conserve body heat, but this has not been proven. It is often suggested that this is done in part to keep the legs from getting wet, in addition to conserving energy. As well as standing in the water, flamingos may stamp their webbed feet in the mud to stir up food from the bottom.

 

Young flamingos hatch with grey plumage, but adults range from light pink to bright red due to aqueous bacteria and beta carotene obtained from their food supply. A well-fed, healthy flamingo is more vibrantly coloured and thus a more desirable mate. A white or pale flamingo, however, is usually unhealthy or malnourished. Captive flamingos are a notable exception; many turn a pale pink as they are not fed carotene at levels comparable to the wild. This is changing as more zoos begin to add prawns and other supplements to the diets of their flamingos.

 

Culinary use ~ In Ancient Rome, flamingo tongues were considered a delicacy. Also, Andean miners have killed flamingos for their fat, believed to be a cure for tuberculosis.

 

Other ~ The Moche people of ancient Peru worshipped nature. They placed emphasis on animals and often depicted flamingos in their art.

A diagram of the human digestive system from the Washington Post.

www.washingtonpost.com/wp-dyn/content/graphic/2007/11/19/...

The digestive system

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4 shots of the Bearded Vulture (immature) that has been roaming round England since late June. Normally in mountains of Eurasia & parts of Africa & is red listed as critically endangered. This one believed to have come of a reintroduction scheme in the Alps. Diet is almost exclusively bones (for the bone marrow) & its digestive system can dissolve them easily.. I took loads of photos & several videos, one is uploaded today as well. I was lucky to get there early when spells of early sunshine were still about, but must have been 150 cars there before me, luckily I found a good spot with a view on my own & the 3000mm zoom of the P1000 was very usefull

mcvities milk chocolate & orange digestives

Villers-la-Ville, Belgium

A fungus (pl.: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the traditional eukaryotic kingdoms, along with Animalia, Plantae and either Protista or Protozoa and Chromista.

 

A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (i.e. they form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past mycology was regarded as a branch of botany, although it is now known that fungi are genetically more closely related to animals than to plants.

 

Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals, including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.

 

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven phyla, and ten subphyla.

 

Etymology

The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny. This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold').

 

The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by Robert Kaye Greville. In 1836 the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.

 

A group of all the fungi present in a particular region is known as mycobiota (plural noun, no singular). The term mycota is often used for this purpose, but many authors use it as a synonym of Fungi. The word funga has been proposed as a less ambiguous term morphologically similar to fauna and flora. The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.

 

Characteristics

 

Fungal hyphae cells

Hyphal wall

Septum

Mitochondrion

Vacuole

Ergosterol crystal

Ribosome

Nucleus

Endoplasmic reticulum

Lipid body

Plasma membrane

Spitzenkörper

Golgi apparatus

 

Fungal cell cycle showing Dikaryons typical of Higher Fungi

Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of mushrooms, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the Neoproterozoic Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:

 

With other eukaryotes: Fungal cells contain membrane-bound nuclei with chromosomes that contain DNA with noncoding regions called introns and coding regions called exons. Fungi have membrane-bound cytoplasmic organelles such as mitochondria, sterol-containing membranes, and ribosomes of the 80S type. They have a characteristic range of soluble carbohydrates and storage compounds, including sugar alcohols (e.g., mannitol), disaccharides, (e.g., trehalose), and polysaccharides (e.g., glycogen, which is also found in animals).

With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.

With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.

With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.

The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called the Spitzenkörper. Both fungi and oomycetes grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.

In common with some plant and animal species, more than one hundred fungal species display bioluminescence.

Unique features:

 

Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.

The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.

A whitish fan or funnel-shaped mushroom growing at the base of a tree.

Omphalotus nidiformis, a bioluminescent mushroom

Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants. Many of the plant and fungal enzymes that make these compounds differ from each other in sequence and other characteristics, which indicates separate origins and convergent evolution of these enzymes in the fungi and plants.

 

Diversity

Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.

 

As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. Species may also be distinguished by their biochemical and physiological characteristics, such as their ability to metabolize certain biochemicals, or their reaction to chemical tests. The biological species concept discriminates species based on their ability to mate. The application of molecular tools, such as DNA sequencing and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of genetic diversity within various taxonomic groups.

 

Mycology

Mycology is the branch of biology concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and psychotropic substances consumed for religious purposes, as well as their dangers, such as poisoning or infection. The field of phytopathology, the study of plant diseases, is closely related because many plant pathogens are fungi.

 

The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.

 

History

Mycology became a systematic science after the development of the microscope in the 17th century. Although fungal spores were first observed by Giambattista della Porta in 1588, the seminal work in the development of mycology is considered to be the publication of Pier Antonio Micheli's 1729 work Nova plantarum genera. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.

 

Morphology

Microscopic structures

Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.

An environmental isolate of Penicillium

Hypha

Conidiophore

Phialide

Conidia

Septa

Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells.

 

Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla,[63] and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.

 

Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.

 

Macroscopic structures

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called molds. Mycelia grown on solid agar media in laboratory petri dishes are usually referred to as colonies. These colonies can exhibit growth shapes and colors (due to spores or pigmentation) that can be used as diagnostic features in the identification of species or groups. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.

 

The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

 

Growth and physiology

Time-lapse photography sequence of a peach becoming progressively discolored and disfigured

Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form a structure called an appressorium that evolved to puncture plant tissues.[71] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[71] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.

 

The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular turgor by producing osmolytes such as glycerol. Adaptations such as these are complemented by hydrolytic enzymes secreted into the environment to digest large organic molecules—such as polysaccharides, proteins, and lipids—into smaller molecules that may then be absorbed as nutrients. The vast majority of filamentous fungi grow in a polar fashion (extending in one direction) by elongation at the tip (apex) of the hypha. Other forms of fungal growth include intercalary extension (longitudinal expansion of hyphal compartments that are below the apex) as in the case of some endophytic fungi, or growth by volume expansion during the development of mushroom stipes and other large organs. Growth of fungi as multicellular structures consisting of somatic and reproductive cells—a feature independently evolved in animals and plants—has several functions, including the development of fruit bodies for dissemination of sexual spores (see above) and biofilms for substrate colonization and intercellular communication.

 

Fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.

 

Reproduction

Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk

Polyporus squamosus

Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.

 

Asexual reproduction

Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.

 

Sexual reproduction

See also: Mating in fungi and Sexual selection in fungi

Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.

 

Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).

 

In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.

 

Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).

 

In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming a gametangium, a specialized cell structure that becomes a fertile gamete-producing cell. The gametangium develops into a zygospore, a thick-walled spore formed by the union of gametes. When the zygospore germinates, it undergoes meiosis, generating new haploid hyphae, which may then form asexual sporangiospores. These sporangiospores allow the fungus to rapidly disperse and germinate into new genetically identical haploid fungal mycelia.

 

Spore dispersal

The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or hydrophobic spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.

 

Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.

 

Homothallism

In homothallic sexual reproduction, two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an Aspergillus-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus Cochliobolus, and the ascomycete Pneumocystis jirovecii. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.

 

Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.

 

Evolution

In contrast to plants and animals, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal fruiting bodies, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble extant fungi. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.

 

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.

 

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).

 

Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415 Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90 Ma.

 

Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.

 

Sixty-five million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".

 

Taxonomy

Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts. Analyses using molecular phylogenetics support a monophyletic origin of fungi. The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.

 

There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).

 

The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.

 

The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. As of 2019, nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycotina, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.

 

Phylogenetic analysis has demonstrated that the Microsporidia, unicellular parasites of animals and protists, are fairly recent and highly derived endobiotic fungi (living within the tissue of another species). Previously considered to be "primitive" protozoa, they are now thought to be either a basal branch of the Fungi, or a sister group–each other's closest evolutionary relative.

 

The Chytridiomycota are commonly known as chytrids. These fungi are distributed worldwide. Chytrids and their close relatives Neocallimastigomycota and Blastocladiomycota (below) are the only fungi with active motility, producing zoospores that are capable of active movement through aqueous phases with a single flagellum, leading early taxonomists to classify them as protists. Molecular phylogenies, inferred from rRNA sequences in ribosomes, suggest that the Chytrids are a basal group divergent from the other fungal phyla, consisting of four major clades with suggestive evidence for paraphyly or possibly polyphyly.

 

The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. Molecular data and ultrastructural characteristics, however, place the Blastocladiomycota as a sister clade to the Zygomycota, Glomeromycota, and Dikarya (Ascomycota and Basidiomycota). The blastocladiomycetes are saprotrophs, feeding on decomposing organic matter, and they are parasites of all eukaryotic groups. Unlike their close relatives, the chytrids, most of which exhibit zygotic meiosis, the blastocladiomycetes undergo sporic meiosis.

 

The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. Members of this small phylum are anaerobic organisms, living in the digestive system of larger herbivorous mammals and in other terrestrial and aquatic environments enriched in cellulose (e.g., domestic waste landfill sites). They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.

 

Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres

Arbuscular mycorrhiza seen under microscope. Flax root cortical cells containing paired arbuscules.

Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)

Diagram of an apothecium (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.

Members of the Glomeromycota form arbuscular mycorrhizae, a form of mutualist symbiosis wherein fungal hyphae invade plant root cells and both species benefit from the resulting increased supply of nutrients. All known Glomeromycota species reproduce asexually. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. Formerly part of the Zygomycota (commonly known as 'sugar' and 'pin' molds), the Glomeromycota were elevated to phylum status in 2001 and now replace the older phylum Zygomycota. Fungi that were placed in the Zygomycota are now being reassigned to the Glomeromycota, or the subphyla incertae sedis Mucoromycotina, Kickxellomycotina, the Zoopagomycotina and the Entomophthoromycotina. Some well-known examples of fungi formerly in the Zygomycota include black bread mold (Rhizopus stolonifer), and Pilobolus species, capable of ejecting spores several meters through the air. Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

 

The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. These fungi form meiotic spores called ascospores, which are enclosed in a special sac-like structure called an ascus. This phylum includes morels, a few mushrooms and truffles, unicellular yeasts (e.g., of the genera Saccharomyces, Kluyveromyces, Pichia, and Candida), and many filamentous fungi living as saprotrophs, parasites, and mutualistic symbionts (e.g. lichens). Prominent and important genera of filamentous ascomycetes include Aspergillus, Penicillium, Fusarium, and Claviceps. Many ascomycete species have only been observed undergoing asexual reproduction (called anamorphic species), but analysis of molecular data has often been able to identify their closest teleomorphs in the Ascomycota. Because the products of meiosis are retained within the sac-like ascus, ascomycetes have been used for elucidating principles of genetics and heredity (e.g., Neurospora crassa).

 

Members of the Basidiomycota, commonly known as the club fungi or basidiomycetes, produce meiospores called basidiospores on club-like stalks called basidia. Most common mushrooms belong to this group, as well as rust and smut fungi, which are major pathogens of grains. Other important basidiomycetes include the maize pathogen Ustilago maydis, human commensal species of the genus Malassezia, and the opportunistic human pathogen, Cryptococcus neoformans.

 

Fungus-like organisms

Because of similarities in morphology and lifestyle, the slime molds (mycetozoans, plasmodiophorids, acrasids, Fonticula and labyrinthulids, now in Amoebozoa, Rhizaria, Excavata, Opisthokonta and Stramenopiles, respectively), water molds (oomycetes) and hyphochytrids (both Stramenopiles) were formerly classified in the kingdom Fungi, in groups like Mastigomycotina, Gymnomycota and Phycomycetes. The slime molds were studied also as protozoans, leading to an ambiregnal, duplicated taxonomy.

 

Unlike true fungi, the cell walls of oomycetes contain cellulose and lack chitin. Hyphochytrids have both chitin and cellulose. Slime molds lack a cell wall during the assimilative phase (except labyrinthulids, which have a wall of scales), and take in nutrients by ingestion (phagocytosis, except labyrinthulids) rather than absorption (osmotrophy, as fungi, labyrinthulids, oomycetes and hyphochytrids). Neither water molds nor slime molds are closely related to the true fungi, and, therefore, taxonomists no longer group them in the kingdom Fungi. Nonetheless, studies of the oomycetes and myxomycetes are still often included in mycology textbooks and primary research literature.

 

The Eccrinales and Amoebidiales are opisthokont protists, previously thought to be zygomycete fungi. Other groups now in Opisthokonta (e.g., Corallochytrium, Ichthyosporea) were also at given time classified as fungi. The genus Blastocystis, now in Stramenopiles, was originally classified as a yeast. Ellobiopsis, now in Alveolata, was considered a chytrid. The bacteria were also included in fungi in some classifications, as the group Schizomycetes.

 

The Rozellida clade, including the "ex-chytrid" Rozella, is a genetically disparate group known mostly from environmental DNA sequences that is a sister group to fungi. Members of the group that have been isolated lack the chitinous cell wall that is characteristic of fungi. Alternatively, Rozella can be classified as a basal fungal group.

 

The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinomycetota), a group with many filamentous bacteria, were also long believed to be fungi.

 

Ecology

Although often inconspicuous, fungi occur in every environment on Earth and play very important roles in most ecosystems. Along with bacteria, fungi are the major decomposers in most terrestrial (and some aquatic) ecosystems, and therefore play a critical role in biogeochemical cycles and in many food webs. As decomposers, they play an essential role in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms.

 

Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all kingdoms. These interactions can be mutualistic or antagonistic in nature, or in the case of commensal fungi are of no apparent benefit or detriment to the host.

 

With plants

Mycorrhizal symbiosis between plants and fungi is one of the most well-known plant–fungus associations and is of significant importance for plant growth and persistence in many ecosystems; over 90% of all plant species engage in mycorrhizal relationships with fungi and are dependent upon this relationship for survival.

 

A microscopic view of blue-stained cells, some with dark wavy lines in them

The dark filaments are hyphae of the endophytic fungus Epichloë coenophiala in the intercellular spaces of tall fescue leaf sheath tissue

The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. It often increases the plant's uptake of inorganic compounds, such as nitrate and phosphate from soils having low concentrations of these key plant nutrients. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. Similar to mycorrhiza, endophytic colonization by fungi may benefit both symbionts; for example, endophytes of grasses impart to their host increased resistance to herbivores and other environmental stresses and receive food and shelter from the plant in return.

 

With algae and cyanobacteria

A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface

The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species

Lichens are a symbiotic relationship between fungi and photosynthetic algae or cyanobacteria. The photosynthetic partner in the relationship is referred to in lichen terminology as a "photobiont". The fungal part of the relationship is composed mostly of various species of ascomycetes and a few basidiomycetes. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession, and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves. As in mycorrhizas, the photobiont provides sugars and other carbohydrates via photosynthesis to the fungus, while the fungus provides minerals and water to the photobiont. The functions of both symbiotic organisms are so closely intertwined that they function almost as a single organism; in most cases the resulting organism differs greatly from the individual components. Lichenization is a common mode of nutrition for fungi; around 27% of known fungi—more than 19,400 species—are lichenized. Characteristics common to most lichens include obtaining organic carbon by photosynthesis, slow growth, small size, long life, long-lasting (seasonal) vegetative reproductive structures, mineral nutrition obtained largely from airborne sources, and greater tolerance of desiccation than most other photosynthetic organisms in the same habitat.

 

With insects

Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. Likewise, females of several wood wasp species (genus Sirex) inject their eggs together with spores of the wood-rotting fungus Amylostereum areolatum into the sapwood of pine trees; the growth of the fungus provides ideal nutritional conditions for the development of the wasp larvae. At least one species of stingless bee has a relationship with a fungus in the genus Monascus, where the larvae consume and depend on fungus transferred from old to new nests. Termites on the African savannah are also known to cultivate fungi, and yeasts of the genera Candida and Lachancea inhabit the gut of a wide range of insects, including neuropterans, beetles, and cockroaches; it is not known whether these fungi benefit their hosts. Fungi growing in dead wood are essential for xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. The larvae of many families of fungicolous flies, particularly those within the superfamily Sciaroidea such as the Mycetophilidae and some Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae.

 

A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.

The plant pathogen Puccinia magellanicum (calafate rust) causes the defect known as witch's broom, seen here on a barberry shrub in Chile.

 

Gram stain of Candida albicans from a vaginal swab from a woman with candidiasis, showing hyphae, and chlamydospores, which are 2–4 µm in diameter.

Many fungi are parasites on plants, animals (including humans), and other fungi. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the rice blast fungus Magnaporthe oryzae, tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease, Cryphonectria parasitica responsible for chestnut blight, and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. Some carnivorous fungi, like Paecilomyces lilacinus, are predators of nematodes, which they capture using an array of specialized structures such as constricting rings or adhesive nets. Many fungi that are plant pathogens, such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead.

 

Some fungi can cause serious diseases in humans, several of which may be fatal if untreated. These include aspergillosis, candidiasis, coccidioidomycosis, cryptococcosis, histoplasmosis, mycetomas, and paracoccidioidomycosis. Furthermore, persons with immuno-deficiencies are particularly susceptible to disease by genera such as Aspergillus, Candida, Cryptoccocus, Histoplasma, and Pneumocystis. Other fungi can attack eyes, nails, hair, and especially skin, the so-called dermatophytic and keratinophilic fungi, and cause local infections such as ringworm and athlete's foot. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.

 

As targets of mycoparasites

Organisms that parasitize fungi are known as mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.

 

Communication

Main article: Mycorrhizal networks

There appears to be electrical communication between fungi in word-like components according to spiking characteristics.

 

Possible impact on climate

According to a study published in the academic journal Current Biology, fungi can soak from the atmosphere around 36% of global fossil fuel greenhouse gas emissions.

 

Mycotoxins

(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide

Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations

Many fungi produce biologically active compounds, several of which are toxic to animals or plants and are therefore called mycotoxins. Of particular relevance to humans are mycotoxins produced by molds causing food spoilage, and poisonous mushrooms (see above). Particularly infamous are the lethal amatoxins in some Amanita mushrooms, and ergot alkaloids, which have a long history of causing serious epidemics of ergotism (St Anthony's Fire) in people consuming rye or related cereals contaminated with sclerotia of the ergot fungus, Claviceps purpurea. Other notable mycotoxins include the aflatoxins, which are insidious liver toxins and highly carcinogenic metabolites produced by certain Aspergillus species often growing in or on grains and nuts consumed by humans, ochratoxins, patulin, and trichothecenes (e.g., T-2 mycotoxin) and fumonisins, which have significant impact on human food supplies or animal livestock.

 

Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.

 

Pathogenic mechanisms

Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as U. maydis. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species at the site of the attempted invasion. U. maydis can respond to the oxidative burst with an oxidative stress response, regulated by the gene YAP1. The response protects U. maydis from the host defense, and is necessary for the pathogen's virulence. Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.

 

Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. C. neoformans usually infects the lungs, where it is phagocytosed by alveolar macrophages. Some C. neoformans can survive inside macrophages, which appears to be the basis for latency, disseminated disease, and resistance to antifungal agents. One mechanism by which C. neoformans survives the hostile macrophage environment is by up-regulating the expression of genes involved in the oxidative stress response. Another mechanism involves meiosis. The majority of C. neoformans are mating "type a". Filaments of mating "type a" ordinarily have haploid nuclei, but they can become diploid (perhaps by endoduplication or by stimulated nuclear fusion) to form blastospores. The diploid nuclei of blastospores can undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed. This process is referred to as monokaryotic fruiting. This process requires a gene called DMC1, which is a conserved homologue of genes recA in bacteria and RAD51 in eukaryotes, that mediates homologous chromosome pairing during meiosis and repair of DNA double-strand breaks. Thus, C. neoformans can undergo a meiosis, monokaryotic fruiting, that promotes recombinational repair in the oxidative, DNA damaging environment of the host macrophage, and the repair capability may contribute to its virulence.

 

Human use

See also: Human interactions with fungi

Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres

Saccharomyces cerevisiae cells shown with DIC microscopy

The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history. Mushroom farming and mushroom gathering are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as ethnomycology. Because of the capacity of this group to produce an enormous range of natural products with antimicrobial or other biological activities, many species have long been used or are being developed for industrial production of antibiotics, vitamins, and anti-cancer and cholesterol-lowering drugs. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing bioeconomy, with applications under research and development including use for textiles, meat substitution and general fungal biotechnology.

 

Therapeutic uses

Modern chemotherapeutics

Many species produce metabolites that are major sources of pharmacologically active drugs.

 

Antibiotics

Particularly important are the antibiotics, including the penicillins, a structurally related group of β-lactam antibiotics that are synthesized from small peptides. Although naturally occurring penicillins such as penicillin G (produced by Penicillium chrysogenum) have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by chemical modification of the natural penicillins. Modern penicillins are semisynthetic compounds, obtained initially from fermentation cultures, but then structurally altered for specific desirable properties. Other antibiotics produced by fungi include: ciclosporin, commonly used as an immunosuppressant during transplant surgery; and fusidic acid, used to help control infection from methicillin-resistant Staphylococcus aureus bacteria. Widespread use of antibiotics for the treatment of bacterial diseases, such as tuberculosis, syphilis, leprosy, and others began in the early 20th century and continues to date. In nature, antibiotics of fungal or bacterial origin appear to play a dual role: at high concentrations they act as chemical defense against competition with other microorganisms in species-rich environments, such as the rhizosphere, and at low concentrations as quorum-sensing molecules for intra- or interspecies signaling.

 

Other

Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections, and statins (HMG-CoA reductase inhibitors), used to inhibit cholesterol synthesis. Examples of statins found in fungi include mevastatin from Penicillium citrinum and lovastatin from Aspergillus terreus and the oyster mushroom. Psilocybin from fungi is investigated for therapeutic use and appears to cause global increases in brain network integration. Fungi produce compounds that inhibit viruses and cancer cells. Specific metabolites, such as polysaccharide-K, ergotamine, and β-lactam antibiotics, are routinely used in clinical medicine. The shiitake mushroom is a source of lentinan, a clinical drug approved for use in cancer treatments in several countries, including Japan. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.

 

Traditional medicine

Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance

Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end.

The fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right) are used in traditional medicine practices

Certain mushrooms are used as supposed therapeutics in folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include Agaricus subrufescens, Ganoderma lucidum, and Ophiocordyceps sinensis.

 

Cultured foods

Baker's yeast or Saccharomyces cerevisiae, a unicellular fungus, is used to make bread and other wheat-based products, such as pizza dough and dumplings. Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso while Rhizopus species are used for making tempeh. Several of these fungi are domesticated species that were bred or selected according to their capacity to ferment food without producing harmful mycotoxins (see below), which are produced by very closely related Aspergilli. Quorn, a meat substitute, is made from Fusarium venenatum.

Villers-la-Ville, Belgium