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Die Böhmische Schweiz (Bohemian Switzerland) liegt im Nordwesten von Tschechien und ist Teil des Elbsandsteingebirges. Sonnenaufgang im Tal, in der näheren Umgebung des Dorfes Hohen Leipa (Vysoká Lípa).
The Photo was taken in the North of the Czech Republic . The Region is called in czech "Ceske Svycarsko" (engl. "Bohemian Switzerland"). It is the czech part of the Elbe Sandstone Mountains in northern Bohemia.
© www.borais.com | Do not use without my permission. Image is under full copyright.
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under acrylic glass (US and UK) under: Bohemian Switzerland Print
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Das Bild kann online als professioneller Print (aufgezogen) bestellt und geliefert werden: Böhmische Schweiz Abzug
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Khajuraho, a UNESCO World Heritage Site, is a famous tourist and archaeological site known for its sculptured temples dedicated to Shiva, Vishnu, Jain patriarchs, and others. Khajuraho was one of the capitals of the Chandela kings, who from the 9th to the 12th century CE developed a large realm, which at its height included almost all of what is now Madhya Pradesh state. Khajuraho extended over 21 sq. km and contained about 85 temples, built by multiple rulers mostly between 950 to 1050. In the 12th century the Chandelas, in a period of chaos and decline, abandoned Khajuraho and moved to hill forts elsewhere. Khajuraho puttered along until the 14th century (Ibn Batuta was impressed by its temples) but was afterwards largely forgotten and overtaken by jungles, which probably saved it from the desecration that Muslim conquerors sometimes inflicted on Hindu monuments. In 1838 a British army captain, TS Burt, employed by the Asiatic Society in Calcutta, came upon information that led him to the rediscovery of the complex of temples in the jungle in Khajuraho.
Of the 85 original temples—most constructed of sandstone—about 20 are still reasonably well preserved. Both internally and externally the temples are richly carved with excellent sculptures that are frequently sensual and often sexually explicit. The temples are divided into three complexes—the western is the largest and best known, containing the magnificent Shaivite temple Kandariya Mahadev, a 31m high agglomeration of porches and turrets culminating in a spire. Khajuraho's name derives from the prevalence of khajur, or date palms, in the area.
There are many theories explaining the real reason for incorporating erotic art with the temples. One of the most popular theories is that they were meant to provide education about earthly desires. During the medieval era, young boys were sent to hermitage and practice bramhacharya till reaching maturity. The sculptures were meant to educate them about grahasthahram.
Yet another theory states that they were meant to represent kama (desire) as the third purushartha (aim of life). The sculptures are also accredited to the rise of the Tantric cults during that time. However, several experts reject these theories.
However, the most credible explanations for the erotic sculptures at Khajuraho are as follows:
1. Sign of happiness, prosperity and auspiciousness: During the medieval era there was a common belief that having erotic sculptures were considered alankaras or decorative motifs, protective and auspicious. This hypothesis is based on the authoritative religious texts like the Shilpashastras and the Brihat Samhita. According to the Brihat Samhita, mithunas (couples), goblins, creepers and erotic sculptures were meant to be carved on the temple door to bring in good luck and as a sign of auspiciousness.
2. Mock the Ascetics: Yet another theory explaining the old erotic art at Khajuraho states that the coital couples represent ascetics as well as people from the royal class. It is also believed that the sculptures of couples participating in the orgies are scenes imagined by the artists. Also, it is also said that the artists used ascetics in sensuous and passionate moods as a way to mock the extreme Tantric sects that rose during that era.
3. Code Language: According to experts, the erotic figures were used as a code language to convey Tantric doctrines and non-communicable experiences. For e.g. A sculpture wherein a washerwoman clings to an ascetic may look erotic and sensuous for the layman. However, in the Tantric language, the washerwoman represents the Kundalini energy that has ascended up to the chakras i.e. the neck of the ascetic. Thus, the erotic sculptures have a deeper meaning related to the Tantric cult.
4. Conceal the Magico-Propitiory Yantra: According to the architectural text of the Shilpa Prakasha, every temple must have a Kamakala Yantra strategically placed to protect it from evil spirits and natural calamities. However, the Yantra, which is basically a set of lines drawn symmetrically, must not be visible to the lay man. Thus, a few erotic sculptures with the head down posture were made corresponding to the lines of the Yantra and superimposed on it.
5. Non-duality: It is possible that the erotic sculptures symbolically represent the union of two opposing forces or energies like, inhalation and exhalation, in a timeless state of non-duality.
The erotic art at Khajuraho is considered to be the pinnacle of love and passion. However, during the period between 900 – 1300 AD most Hindu, Jain and Buddhist temples across western and southern India comprised of erotic art.
However, at other temples, these sculptures were carved at the plinth level, below the eye level and thus did not get noticed. It is only at Khajuraho that these sculptures were so prominently displayed on the main wall of the temples
The blowhole of the humpback whale like all baleen whales have two openings unlike the toothed whales which have only one.
The first sign of whales in an area is usually the 'blow'. Air leaves the blowhole at more than 400 kilometres an hour. When tie whales surface, the exhalation of warm air produces a distinctive cloud, formed by condensation of vapour excelled from the lungs under great pressure. Adult humpbacks have two lungs, each the size of a small car, which they can empty and refill in less than two seconds.
Whale breath is quite fishy to smell, and if a blow happens to drift on to an unsuspecting photographer they will find their equipment will have a light smear of oil on it.
Note: Mother and Baby Calf swimming north towards Queensland...
++++++++++++++++++++++++++++++++++++++++++++++++++++++++++
Caught between a window and a face-palm, the little ectoparasite spilled her guts in a death slide.
Mosquitoes are attracted to CO₂, and they are able to home in on humans by detecting the trace gas in our exhalations, so watch out for that heavy breathing around mosquitoes, and smash them when you can.
I watched this one bouncing against the door window for several minutes before I went out and smashed it. I don't know why mosquitoes do that; maybe they can smell us through the windows. Normally, I don't pursue them outside, and give chase only when they invade my space.
However, I should add I've killed many mosquitoes outdoors while watching and photographing birds in line with my general strategy: kill all mosquitoes!
Khajuraho, a UNESCO World Heritage Site, is a famous tourist and archaeological site known for its sculptured temples dedicated to Shiva, Vishnu, Jain patriarchs, and others. Khajuraho was one of the capitals of the Chandela kings, who from the 9th to the 12th century CE developed a large realm, which at its height included almost all of what is now Madhya Pradesh state. Khajuraho extended over 21 sq. km and contained about 85 temples, built by multiple rulers mostly between 950 to 1050. In the 12th century the Chandelas, in a period of chaos and decline, abandoned Khajuraho and moved to hill forts elsewhere. Khajuraho puttered along until the 14th century (Ibn Batuta was impressed by its temples) but was afterwards largely forgotten and overtaken by jungles, which probably saved it from the desecration that Muslim conquerors sometimes inflicted on Hindu monuments. In 1838 a British army captain, TS Burt, employed by the Asiatic Society in Calcutta, came upon information that led him to the rediscovery of the complex of temples in the jungle in Khajuraho.
Of the 85 original temples—most constructed of sandstone—about 20 are still reasonably well preserved. Both internally and externally the temples are richly carved with excellent sculptures that are frequently sensual and often sexually explicit. The temples are divided into three complexes—the western is the largest and best known, containing the magnificent Shaivite temple Kandariya Mahadev, a 31m high agglomeration of porches and turrets culminating in a spire. Khajuraho's name derives from the prevalence of khajur, or date palms, in the area.
There are many theories explaining the real reason for incorporating erotic art with the temples. One of the most popular theories is that they were meant to provide education about earthly desires. During the medieval era, young boys were sent to hermitage and practice bramhacharya till reaching maturity. The sculptures were meant to educate them about grahasthahram.
Yet another theory states that they were meant to represent kama (desire) as the third purushartha (aim of life). The sculptures are also accredited to the rise of the Tantric cults during that time. However, several experts reject these theories.
However, the most credible explanations for the erotic sculptures at Khajuraho are as follows:
1. Sign of happiness, prosperity and auspiciousness: During the medieval era there was a common belief that having erotic sculptures were considered alankaras or decorative motifs, protective and auspicious. This hypothesis is based on the authoritative religious texts like the Shilpashastras and the Brihat Samhita. According to the Brihat Samhita, mithunas (couples), goblins, creepers and erotic sculptures were meant to be carved on the temple door to bring in good luck and as a sign of auspiciousness.
2. Mock the Ascetics: Yet another theory explaining the old erotic art at Khajuraho states that the coital couples represent ascetics as well as people from the royal class. It is also believed that the sculptures of couples participating in the orgies are scenes imagined by the artists. Also, it is also said that the artists used ascetics in sensuous and passionate moods as a way to mock the extreme Tantric sects that rose during that era.
3. Code Language: According to experts, the erotic figures were used as a code language to convey Tantric doctrines and non-communicable experiences. For e.g. A sculpture wherein a washerwoman clings to an ascetic may look erotic and sensuous for the layman. However, in the Tantric language, the washerwoman represents the Kundalini energy that has ascended up to the chakras i.e. the neck of the ascetic. Thus, the erotic sculptures have a deeper meaning related to the Tantric cult.
4. Conceal the Magico-Propitiory Yantra: According to the architectural text of the Shilpa Prakasha, every temple must have a Kamakala Yantra strategically placed to protect it from evil spirits and natural calamities. However, the Yantra, which is basically a set of lines drawn symmetrically, must not be visible to the lay man. Thus, a few erotic sculptures with the head down posture were made corresponding to the lines of the Yantra and superimposed on it.
5. Non-duality: It is possible that the erotic sculptures symbolically represent the union of two opposing forces or energies like, inhalation and exhalation, in a timeless state of non-duality.
The erotic art at Khajuraho is considered to be the pinnacle of love and passion. However, during the period between 900 – 1300 AD most Hindu, Jain and Buddhist temples across western and southern India comprised of erotic art.
However, at other temples, these sculptures were carved at the plinth level, below the eye level and thus did not get noticed. It is only at Khajuraho that these sculptures were so prominently displayed on the main wall of the temples
I've got words to speak for me, that's what carries the distance. They say that we're supposed to speak with actions, but actions are silent, unless you make noise, then in that case they're noisy. But written words are silent as well, so am I really saying anything by writing? I am, yes, to a seeing person, even to a deaf person. But to a blind person, no, although I would like to believe that blind people have magical powers and could somehow decipher what I was writing if they were sitting beside me in the dark and listening to the sound that my fingers were making while hitting each individual keystroke. But maybe each keystroke sounds exactly the same as the next, in which case it'd be a pointless exercise.
But I like to believe that such things are possible, because I don't like to admit that there are limits to the human condition. I tend to think that the only real conditions of the human condition are those that we place on ourselves. I'm not even sure what those phrases mean that go on and on about the meaning of life and so forth. Maybe we're just trying to figure out how we can make an impact when all that really justifies our actions is ourselves. Do I really believe that? I don't know, maybe you do. I do know that when it all comes full circle, it's dust to dust and ash to ash, and only the ends justify the ends.
It's a big temptation to end a thought with a word like "end", but I'm not done thinking yet. Sometimes we let our minds get twisted around a three act narrative as if that's the way life really works. I figure that we're either living countless, endless acts or our lives are just one big sprawling act until the curtain falls on the time we lived. I hesitate to use artistic metaphors to describe life like we're in a fictional story, narrowing our lives down to the lowest common denominator as if we're writing something more real than reality. But nobody would ever read something as slow as life. I've seen it happen all around me. Eight straight hours of sleep, the slowest chapter of every novel. Page after page with descriptions of exhalation, tossing and turning, maybe some snoring to spice things up. By morning we'd be hanging on the edge of our seats for an indecipherable dream or two. Talk about a thrill ride. Fiction is better with nocturnal omissions intact.
Have you taken a look at your life lately and realized how badly paced it is? There's a lot of in-between time, a lot of staring out a window or into a screen. The average film can depict inaction for about a minute before losing viewer interest, but is it really any better from the other side? It's almost like we can't tell anymore, we're so focused on seeing ourselves from an external perspective. If it wouldn't be interesting to watch, we're not sure if we want to do it anymore. What audience are we playing to? Take a look in the mirror - do you even know what you don't look like anymore? Do you remember what it was like when you didn't know more things? Your lack of knowledge was a sense of freedom that made it so that you didn't care about appearance for appearance's sake, and what about when discovery was all that seemed important?
Isn't discovery all that really drives us in the end? "What does that sensation mean?" we ask. Touch it, hear it, taste it, feel it. Your eyes take you half the way there, let your body bring you closer. Find more love, dive deeper. Seek more pain, reach further. We outgrow discovery and settle for familiarity, until the only thing we want is what we're already used to. Again, do it again. Sing that same song, eat that same meal, swallow down your entertainment like the pornography it is and make love to yourself and everyone else in the same way as yesterday. Where's the hollowness gone? Nowhere at all, you know better. It's gnawing at the core of you, rodent teeth tearing through your tendons and tail flicking against your innards. Discovery is your savior, peace and action, wild and alive. Take your body and awaken your brain. Reach your soul with your lightning mind. Seek a sense of wonder, be loved, then FEEL loved. You'll find it all, and more, into your body and through the red door.
"Va a lavorare in miniera!" è una esortazione verso gli sfaccendati che, almeno in Italia, ha praticamente perso valore perché i numerosi giacimenti che esistevano nel passato sono stati tutti chiusi ad eccezione di alcuni a cielo aperto, come le cave di pietra o di marmo o quelle di salgemma. Ma in cosa consisteva questa minaccia?
Lavorare in miniera significava infilarsi in un buco ogni giorno e rimanerci per dieci, dodici ore; significava martellare sulle pietre con picconi o martelli pneumatici fino ad assordarsi; respirare le polveri sottili dei minerali frantumati e finire per ammalarsi polmoni di silicosi. E poi rischiare la vita per l'esalazione di gas striscianti e inodori come il grisù oppure per i crolli delle gallerie.
Foto tratta dalla cartellonistica del Parco Minerario di Gavorrano (Toscana).
Miner
"Goes to work at the mine!" is an exhortation to the idlers that, almost in Italy, has virtually lost value because the numerous mines existing in the past were all closed except a few quarries of stone or marble or those of rock salt. But what the threat was?
Work in the mines meant going into a hole and stay there every day for ten to twelve hours; it meant hammering on the rocks with pickaxes or pneumatic drills until deafened by this noise; breathing fine dust of ground ore and end up getting sick lungs of silicosis. And then risk their lives for the exhalation of gas creeping and odorless as firedamp or for the collapses of tunnels.
Photo taken from the Mining Park of Gavorrano (Tuscany)
Use without permission is illegal.
Please don't use this image on websites, blogs or other media without my explicit permission.
"What we commonly call fire. It is not really fire, for fire is an excess of heat and a sort of ebullition; but in reality, of what we call air, the part surrounding the earth is moist and warm, because it contains both vapour and a dry exhalation from the earth." - Fire Element (wikipedia.com)
Note:
Fire element interpretation. All ambient lighting.
Credits: Shiella M. & Sarah U.
Can't say if some days take my breath away, or I give it. Either way, I'm happy to lose it, to feel that big, wide open exhalation, making space for exultation. I stayed till there was nothing left, blown back in a sea of green and rolling grasses, encapsulating a dead oasis. Like some desert island isolation, that one palm tree traded for a single family spire. Perched between pitiful picket fences, the loneliness is incredible, like fighting a battle to find soldiers on all sides. I can't tell you how it tears my heart out, when the last sun breaks on black clouds, and I'm standing like a bastion between bright light darkness. It's a metaphor for my mindset, tip-toed to the teetering edge, looking in on a memory not far from forgotten. Good words for an ardent explorer, try to treat myself well while I sift through the shadows.
November 6, 2020
Bayard Family Cemetery
South Farmington, Nova Scotia
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My prayer is that everyone will have free access to clean and healthy water. We must heed Leonardo Da Vinci's words and realize that everything is made up of water. Every exhalation against a mirror reminds us that we are sustained generously in such mysterious ways. Today thousands of people all around the world are sending healing energy and gratitude to all forms of water.
***Looking forward to catching up with all of you
soon*****************
Added: January 28, 2015
*****A MUST SEE*****
"Flow for the Love of Water" documentary (Irena Salina)
It will wake you up, break your heart and refresh your understanding and relationship with water. It's getting harder to find but here is a link:
An "electric chariot" group are assemble in Plaza San Luis.
WHITE KNIGHT TOURS
For the first time in Philippine history, an Electric Chariots tour of Intramuros has been launched by White Knight tours at Hotel Intramuros fronting San Agustin Chruch. The business started in 2012.
The Electric Chariots are 2 wheel equipments with automatic balancing electronic gadget that moves forward if your body leans forward and stops or move backward as you lean backwards.
The guided tours using electric chariots amplify the most recent tourism slogan of "It's more fun in the Philippines" for indeed, the exhalation as well. White Knight offers 30 minutes and 1 hour guided tours of Intramuros with price ranging from USD 13.00 for 30minutes and USD 25.00 for 1 hour.
Unlike riding the horse driver Chariots tours which provide passive involvements for passengers and tourist, the Electric Chariots reinvigorates the user with its interactive role of balancing and inspires a sense of adrenaline rush.
It's an experience in a life time that one should never miss!
Sheep (pl.: sheep) or domestic sheep (Ovis aries) are a domesticated, ruminant mammal typically kept as livestock. Although the term sheep can apply to other species in the genus Ovis, in everyday usage it almost always refers to domesticated sheep. Like all ruminants, sheep are members of the order Artiodactyla, the even-toed ungulates. Numbering a little over one billion, domestic sheep are also the most numerous species of sheep. An adult female is referred to as a ewe (/juː/ yoo), an intact male as a ram, occasionally a tup, a castrated male as a wether, and a young sheep as a lamb.
Sheep are most likely descended from the wild mouflon of Europe and Asia, with Iran being a geographic envelope of the domestication center. One of the earliest animals to be domesticated for agricultural purposes, sheep are raised for fleeces, meat (lamb, hogget or mutton) and milk. A sheep's wool is the most widely used animal fiber, and is usually harvested by shearing. In Commonwealth countries, ovine meat is called lamb when from younger animals and mutton when from older ones; in the United States, meat from both older and younger animals is usually called lamb. Sheep continue to be important for wool and meat today, and are also occasionally raised for pelts, as dairy animals, or as model organisms for science.
Sheep husbandry is practised throughout the majority of the inhabited world, and has been fundamental to many civilizations. In the modern era, Australia, New Zealand, the southern and central South American nations, and the British Isles are most closely associated with sheep production.
There is a large lexicon of unique terms for sheep husbandry which vary considerably by region and dialect. Use of the word sheep began in Middle English as a derivation of the Old English word scēap. A group of sheep is called a flock. Many other specific terms for the various life stages of sheep exist, generally related to lambing, shearing, and age.
Being a key animal in the history of farming, sheep have a deeply entrenched place in human culture, and are represented in much modern language and symbolism. As livestock, sheep are most often associated with pastoral, Arcadian imagery. Sheep figure in many mythologies—such as the Golden Fleece—and major religions, especially the Abrahamic traditions. In both ancient and modern religious ritual, sheep are used as sacrificial animals.
History
Main article: History of the domestic sheep
The exact line of descent from wild ancestors to domestic sheep is unclear. The most common hypothesis states that Ovis aries is descended from the Asiatic (O. gmelini) species of mouflon; the European mouflon (Ovis aries musimon) is a direct descendant of this population. Sheep were among the first animals to be domesticated by humankind (although the domestication of dogs probably took place 10 to 20 thousand years earlier); the domestication date is estimated to fall between 11,000 and 9000 B.C in Mesopotamia and possibly around 7000 BC in Mehrgarh in the Indus Valley. The rearing of sheep for secondary products, and the resulting breed development, began in either southwest Asia or western Europe. Initially, sheep were kept solely for meat, milk and skins. Archaeological evidence from statuary found at sites in Iran suggests that selection for woolly sheep may have begun around 6000 BC, and the earliest woven wool garments have been dated to two to three thousand years later.
Sheep husbandry spread quickly in Europe. Excavations show that in about 6000 BC, during the Neolithic period of prehistory, the Castelnovien people, living around Châteauneuf-les-Martigues near present-day Marseille in the south of France, were among the first in Europe to keep domestic sheep. Practically from its inception, ancient Greek civilization relied on sheep as primary livestock, and were even said to name individual animals. Ancient Romans kept sheep on a wide scale, and were an important agent in the spread of sheep raising. Pliny the Elder, in his Natural History (Naturalis Historia), speaks at length about sheep and wool. European colonists spread the practice to the New World from 1493 onwards.
Characteristics
Domestic sheep are relatively small ruminants, usually with a crimped hair called wool and often with horns forming a lateral spiral. They differ from their wild relatives and ancestors in several respects, having become uniquely neotenic as a result of selective breeding by humans. A few primitive breeds of sheep retain some of the characteristics of their wild cousins, such as short tails. Depending on breed, domestic sheep may have no horns at all (i.e. polled), or horns in both sexes, or in males only. Most horned breeds have a single pair, but a few breeds may have several.
Sheep in Turkmenistan
Another trait unique to domestic sheep as compared to wild ovines is their wide variation in color. Wild sheep are largely variations of brown hues, and variation within species is extremely limited. Colors of domestic sheep range from pure white to dark chocolate brown, and even spotted or piebald. Sheep keepers also sometimes artificially paint "smit marks" onto their sheep in any pattern or color for identification. Selection for easily dyeable white fleeces began early in sheep domestication, and as white wool is a dominant trait it spread quickly. However, colored sheep do appear in many modern breeds, and may even appear as a recessive trait in white flocks. While white wool is desirable for large commercial markets, there is a niche market for colored fleeces, mostly for handspinning. The nature of the fleece varies widely among the breeds, from dense and highly crimped, to long and hairlike. There is variation of wool type and quality even among members of the same flock, so wool classing is a step in the commercial processing of the fibre.
Suffolks are a medium wool, black-faced breed of meat sheep that make up 60% of the sheep population in the U.S.
Depending on breed, sheep show a range of heights and weights. Their rate of growth and mature weight is a heritable trait that is often selected for in breeding. Ewes typically weigh between 45 and 100 kilograms (100 and 220 lb), and rams between 45 and 160 kilograms (100 and 350 lb). When all deciduous teeth have erupted, the sheep has 20 teeth. Mature sheep have 32 teeth. As with other ruminants, the front teeth in the lower jaw bite against a hard, toothless pad in the upper jaw. These are used to pick off vegetation, then the rear teeth grind it before it is swallowed. There are eight lower front teeth in ruminants, but there is some disagreement as to whether these are eight incisors, or six incisors and two incisor-shaped canines. This means that the dental formula for sheep is either
0.0.3.3
4.0.3.3
or
0.0.3.3
3.1.3.3
There is a large diastema between the incisors and the molars.
In the first few years of life one can calculate the age of sheep from their front teeth, as a pair of milk teeth is replaced by larger adult teeth each year, the full set of eight adult front teeth being complete at about four years of age. The front teeth are then gradually lost as sheep age, making it harder for them to feed and hindering the health and productivity of the animal. For this reason, domestic sheep on normal pasture begin to slowly decline from four years on, and the life expectancy of a sheep is 10 to 12 years, though some sheep may live as long as 20 years.
Skull
Sheep have good hearing, and are sensitive to noise when being handled. Sheep have horizontal slit-shaped pupils, with excellent peripheral vision; with visual fields of about 270° to 320°, sheep can see behind themselves without turning their heads. Many breeds have only short hair on the face, and some have facial wool (if any) confined to the poll and or the area of the mandibular angle; the wide angles of peripheral vision apply to these breeds. A few breeds tend to have considerable wool on the face; for some individuals of these breeds, peripheral vision may be greatly reduced by "wool blindness", unless recently shorn about the face. Sheep have poor depth perception; shadows and dips in the ground may cause sheep to baulk. In general, sheep have a tendency to move out of the dark and into well-lit areas, and prefer to move uphill when disturbed. Sheep also have an excellent sense of smell, and, like all species of their genus, have scent glands just in front of the eyes, and interdigitally on the feet. The purpose of these glands is uncertain, but those on the face may be used in breeding behaviors. The foot glands might also be related to reproduction, but alternative functions, such as secretion of a waste product or a scent marker to help lost sheep find their flock, have also been proposed.
Comparison with goats
Sheep and goats are closely related: both are in the subfamily Caprinae. However, they are separate species, so hybrids rarely occur and are always infertile. A hybrid of a ewe and a buck (a male goat) is called a sheep-goat hybrid, known as geep. Visual differences between sheep and goats include the beard of goats and divided upper lip of sheep. Sheep tails also hang down, even when short or docked, while the short tails of goats are held upwards. Also, sheep breeds are often naturally polled (either in both sexes or just in the female), while naturally polled goats are rare (though many are polled artificially). Males of the two species differ in that buck goats acquire a unique and strong odor during the rut, whereas rams do not.
Breeds
The domestic sheep is a multi-purpose animal, and the more than 200 breeds now in existence were created to serve these diverse purposes. Some sources give a count of a thousand or more breeds, but these numbers cannot be verified, according to some sources. However, several hundred breeds of sheep have been identified by the Food and Agriculture Organization of the UN (FAO), with the estimated number varying somewhat from time to time: e.g. 863 breeds as of 1993, 1314 breeds as of 1995 and 1229 breeds as of 2006. (These numbers exclude extinct breeds, which are also tallied by the FAO.) For the purpose of such tallies, the FAO definition of a breed is "either a subspecific group of domestic livestock with definable and identifiable external characteristics that enable it to be separated by visual appraisal from other similarly defined groups within the same species or a group for which geographical and/or cultural separation from phenotypically similar groups has led to acceptance of its separate identity." Almost all sheep are classified as being best suited to furnishing a certain product: wool, meat, milk, hides, or a combination in a dual-purpose breed. Other features used when classifying sheep include face color (generally white or black), tail length, presence or lack of horns, and the topography for which the breed has been developed. This last point is especially stressed in the UK, where breeds are described as either upland (hill or mountain) or lowland breeds. A sheep may also be of a fat-tailed type, which is a dual-purpose sheep common in Africa and Asia with larger deposits of fat within and around its tail.
Breeds are often categorized by the type of their wool. Fine wool breeds are those that have wool of great crimp and density, which are preferred for textiles. Most of these were derived from Merino sheep, and the breed continues to dominate the world sheep industry. Downs breeds have wool between the extremes, and are typically fast-growing meat and ram breeds with dark faces. Some major medium wool breeds, such as the Corriedale, are dual-purpose crosses of long and fine-wooled breeds and were created for high-production commercial flocks. Long wool breeds are the largest of sheep, with long wool and a slow rate of growth. Long wool sheep are most valued for crossbreeding to improve the attributes of other sheep types. For example: the American Columbia breed was developed by crossing Lincoln rams (a long wool breed) with fine-wooled Rambouillet ewes.
Coarse or carpet wool sheep are those with a medium to long length wool of characteristic coarseness. Breeds traditionally used for carpet wool show great variability, but the chief requirement is a wool that will not break down under heavy use (as would that of the finer breeds). As the demand for carpet-quality wool declines, some breeders of this type of sheep are attempting to use a few of these traditional breeds for alternative purposes. Others have always been primarily meat-class sheep.
A minor class of sheep are the dairy breeds. Dual-purpose breeds that may primarily be meat or wool sheep are often used secondarily as milking animals, but there are a few breeds that are predominantly used for milking. These sheep produce a higher quantity of milk and have slightly longer lactation curves. In the quality of their milk, the fat and protein content percentages of dairy sheep vary from non-dairy breeds, but lactose content does not.
A last group of sheep breeds is that of fur or hair sheep, which do not grow wool at all. Hair sheep are similar to the early domesticated sheep kept before woolly breeds were developed, and are raised for meat and pelts. Some modern breeds of hair sheep, such as the Dorper, result from crosses between wool and hair breeds. For meat and hide producers, hair sheep are cheaper to keep, as they do not need shearing. Hair sheep are also more resistant to parasites and hot weather.
With the modern rise of corporate agribusiness and the decline of localized family farms, many breeds of sheep are in danger of extinction. The Rare Breeds Survival Trust of the UK lists 22 native breeds as having only 3,000 registered animals (each), and The Livestock Conservancy lists 14 as either "critical" or "threatened". Preferences for breeds with uniform characteristics and fast growth have pushed heritage (or heirloom) breeds to the margins of the sheep industry. Those that remain are maintained through the efforts of conservation organizations, breed registries, and individual farmers dedicated to their preservation.
Diet
Sheep are herbivorous mammals. Most breeds prefer to graze on grass and other short roughage, avoiding the taller woody parts of plants that goats readily consume. Both sheep and goats use their lips and tongues to select parts of the plant that are easier to digest or higher in nutrition. Sheep, however, graze well in monoculture pastures where most goats fare poorly.
Ruminant system of a sheep
Like all ruminants, sheep have a complex digestive system composed of four chambers, allowing them to break down cellulose from stems, leaves, and seed hulls into simpler carbohydrates. When sheep graze, vegetation is chewed into a mass called a bolus, which is then passed into the rumen, via the reticulum. The rumen is a 19- to 38-liter (5 to 10 gallon) organ in which feed is fermented. The fermenting organisms include bacteria, fungi, and protozoa. (Other important rumen organisms include some archaea, which produce methane from carbon dioxide.) The bolus is periodically regurgitated back to the mouth as cud for additional chewing and salivation. After fermentation in the rumen, feed passes into the reticulum and the omasum; special feeds such as grains may bypass the rumen altogether. After the first three chambers, food moves into the abomasum for final digestion before processing by the intestines. The abomasum is the only one of the four chambers analogous to the human stomach, and is sometimes called the "true stomach".
Other than forage, the other staple feed for sheep is hay, often during the winter months. The ability to thrive solely on pasture (even without hay) varies with breed, but all sheep can survive on this diet. Also included in some sheep's diets are minerals, either in a trace mix or in licks. Feed provided to sheep must be specially formulated, as most cattle, poultry, pig, and even some goat feeds contain levels of copper that are lethal to sheep. The same danger applies to mineral supplements such as salt licks.
Grazing behavior
Sheep follow a diurnal pattern of activity, feeding from dawn to dusk, stopping sporadically to rest and chew their cud. Ideal pasture for sheep is not lawnlike grass, but an array of grasses, legumes and forbs. Types of land where sheep are raised vary widely, from pastures that are seeded and improved intentionally to rough, native lands. Common plants toxic to sheep are present in most of the world, and include (but are not limited to) cherry, some oaks and acorns, tomato, yew, rhubarb, potato, and rhododendron.
Effects on pasture
Sheep are largely grazing herbivores, unlike browsing animals such as goats and deer that prefer taller foliage. With a much narrower face, sheep crop plants very close to the ground and can overgraze a pasture much faster than cattle. For this reason, many shepherds use managed intensive rotational grazing, where a flock is rotated through multiple pastures, giving plants time to recover. Paradoxically, sheep can both cause and solve the spread of invasive plant species. By disturbing the natural state of pasture, sheep and other livestock can pave the way for invasive plants. However, sheep also prefer to eat invasives such as cheatgrass, leafy spurge, kudzu and spotted knapweed over native species such as sagebrush, making grazing sheep effective for conservation grazing. Research conducted in Imperial County, California compared lamb grazing with herbicides for weed control in seedling alfalfa fields. Three trials demonstrated that grazing lambs were just as effective as herbicides in controlling winter weeds. Entomologists also compared grazing lambs to insecticides for insect control in winter alfalfa. In this trial, lambs provided insect control as effectively as insecticides.
Behavior
Sheep are flock animals and strongly gregarious; much sheep behavior can be understood on the basis of these tendencies. The dominance hierarchy of sheep and their natural inclination to follow a leader to new pastures were the pivotal factors in sheep being one of the first domesticated livestock species. Furthermore, in contrast to the red deer and gazelle (two other ungulates of primary importance to meat production in prehistoric times), sheep do not defend territories although they do form home ranges. All sheep have a tendency to congregate close to other members of a flock, although this behavior varies with breed, and sheep can become stressed when separated from their flock members. During flocking, sheep have a strong tendency to follow, and a leader may simply be the first individual to move. Relationships in flocks tend to be closest among related sheep: in mixed-breed flocks, subgroups of the same breed tend to form, and a ewe and her direct descendants often move as a unit within large flocks. Sheep can become hefted to one particular local pasture (heft) so they do not roam freely in unfenced landscapes. Lambs learn the heft from ewes and if whole flocks are culled it must be retaught to the replacement animals.
Flock behaviour in sheep is generally only exhibited in groups of four or more sheep; fewer sheep may not react as expected when alone or with few other sheep. Being a prey species, the primary defense mechanism of sheep is to flee from danger when their flight zone is entered. Cornered sheep may charge and butt, or threaten by hoof stamping and adopting an aggressive posture. This is particularly true for ewes with newborn lambs.
In regions where sheep have no natural predators, none of the native breeds of sheep exhibit a strong flocking behavior.
Herding
Farmers exploit flocking behavior to keep sheep together on unfenced pastures such as hill farming, and to move them more easily. For this purpose shepherds may use herding dogs in this effort, with a highly bred herding ability. Sheep are food-oriented, and association of humans with regular feeding often results in sheep soliciting people for food. Those who are moving sheep may exploit this behavior by leading sheep with buckets of feed.
Dominance hierarchy
Sheep establish a dominance hierarchy through fighting, threats and competitiveness. Dominant animals are inclined to be more aggressive with other sheep, and usually feed first at troughs. Primarily among rams, horn size is a factor in the flock hierarchy. Rams with different size horns may be less inclined to fight to establish the dominance order, while rams with similarly sized horns are more so. Merinos have an almost linear hierarchy whereas there is a less rigid structure in Border Leicesters when a competitive feeding situation arises.
In sheep, position in a moving flock is highly correlated with social dominance, but there is no definitive study to show consistent voluntary leadership by an individual sheep.
Intelligence and learning ability
Sheep are frequently thought of as unintelligent animals. Their flocking behavior and quickness to flee and panic can make shepherding a difficult endeavor for the uninitiated. Despite these perceptions, a University of Illinois monograph on sheep reported their intelligence to be just below that of pigs and on par with that of cattle. Sheep can recognize individual human and ovine faces and remember them for years; they can remember 50 other different sheep faces for over two years; they can recognize and are attracted to individual sheep and humans by their faces, as they possess similar specialized neural systems in the temporal and frontal lobes of their brains to humans and have a greater involvement of the right brain hemisphere. In addition to long-term facial recognition of individuals, sheep can also differentiate emotional states through facial characteristics.[68][69] If worked with patiently, sheep may learn their names, and many sheep are trained to be led by halter for showing and other purposes. Sheep have also responded well to clicker training. Sheep have been used as pack animals; Tibetan nomads distribute baggage equally throughout a flock as it is herded between living sites.
It has been reported that some sheep have apparently shown problem-solving abilities; a flock in West Yorkshire, England allegedly found a way to get over cattle grids by rolling on their backs, although documentation of this has relied on anecdotal accounts.
Vocalisations
Sounds made by domestic sheep include bleats, grunts, rumbles and snorts. Bleating ("baaing") is used mostly for contact communication, especially between dam and lambs, but also at times between other flock members. The bleats of individual sheep are distinctive, enabling the ewe and her lambs to recognize each other's vocalizations. Vocal communication between lambs and their dam declines to a very low level within several weeks after parturition. A variety of bleats may be heard, depending on sheep age and circumstances. Apart from contact communication, bleating may signal distress, frustration or impatience; however, sheep are usually silent when in pain. Isolation commonly prompts bleating by sheep. Pregnant ewes may grunt when in labor. Rumbling sounds are made by the ram during courting; somewhat similar rumbling sounds may be made by the ewe, especially when with her neonate lambs. A snort (explosive exhalation through the nostrils) may signal aggression or a warning, and is often elicited from startled sheep.
Lamb
In sheep breeds lacking facial wool, the visual field is wide. In 10 sheep (Cambridge, Lleyn and Welsh Mountain breeds, which lack facial wool), the visual field ranged from 298° to 325°, averaging 313.1°, with binocular overlap ranging from 44.5° to 74°, averaging 61.7°. In some breeds, unshorn facial wool can limit the visual field; in some individuals, this may be enough to cause "wool blindness". In 60 Merinos, visual fields ranged from 219.1° to 303.0°, averaging 269.9°, and the binocular field ranged from 8.9° to 77.7°, averaging 47.5°; 36% of the measurements were limited by wool, although photographs of the experiments indicate that only limited facial wool regrowth had occurred since shearing. In addition to facial wool (in some breeds), visual field limitations can include ears and (in some breeds) horns, so the visual field can be extended by tilting the head. Sheep eyes exhibit very low hyperopia and little astigmatism. Such visual characteristics are likely to produce a well-focused retinal image of objects in both the middle and long distance. Because sheep eyes have no accommodation, one might expect the image of very near objects to be blurred, but a rather clear near image could be provided by the tapetum and large retinal image of the sheep's eye, and adequate close vision may occur at muzzle length. Good depth perception, inferred from the sheep's sure-footedness, was confirmed in "visual cliff" experiments; behavioral responses indicating depth perception are seen in lambs at one day old. Sheep are thought to have colour vision, and can distinguish between a variety of colours: black, red, brown, green, yellow and white. Sight is a vital part of sheep communication, and when grazing, they maintain visual contact with each other. Each sheep lifts its head upwards to check the position of other sheep in the flock. This constant monitoring is probably what keeps the sheep in a flock as they move along grazing. Sheep become stressed when isolated; this stress is reduced if they are provided with a mirror, indicating that the sight of other sheep reduces stress.
Taste is the most important sense in sheep, establishing forage preferences, with sweet and sour plants being preferred and bitter plants being more commonly rejected. Touch and sight are also important in relation to specific plant characteristics, such as succulence and growth form.
The ram uses his vomeronasal organ (sometimes called the Jacobson's organ) to sense the pheromones of ewes and detect when they are in estrus. The ewe uses her vomeronasal organ for early recognition of her neonate lamb.
Reproduction
Sheep follow a similar reproductive strategy to other herd animals. A group of ewes is generally mated by a single ram, who has either been chosen by a breeder or (in feral populations) has established dominance through physical contest with other rams. Most sheep are seasonal breeders, although some are able to breed year-round. Ewes generally reach sexual maturity at six to eight months old, and rams generally at four to six months. However, there are exceptions. For example, Finnsheep ewe lambs may reach puberty as early as 3 to 4 months, and Merino ewes sometimes reach puberty at 18 to 20 months. Ewes have estrus cycles about every 17 days, during which they emit a scent and indicate readiness through physical displays towards rams.
In feral sheep, rams may fight during the rut to determine which individuals may mate with ewes. Rams, especially unfamiliar ones, will also fight outside the breeding period to establish dominance; rams can kill one another if allowed to mix freely. During the rut, even usually friendly rams may become aggressive towards humans due to increases in their hormone levels.
After mating, sheep have a gestation period of about five months, and normal labor takes one to three hours. Although some breeds regularly throw larger litters of lambs, most produce single or twin lambs. During or soon after labor, ewes and lambs may be confined to small lambing jugs, small pens designed to aid both careful observation of ewes and to cement the bond between them and their lambs.
A lamb's first steps
Ovine obstetrics can be problematic. By selectively breeding ewes that produce multiple offspring with higher birth weights for generations, sheep producers have inadvertently caused some domestic sheep to have difficulty lambing; balancing ease of lambing with high productivity is one of the dilemmas of sheep breeding. In the case of any such problems, those present at lambing may assist the ewe by extracting or repositioning lambs. After the birth, ewes ideally break the amniotic sac (if it is not broken during labor), and begin licking clean the lamb. Most lambs will begin standing within an hour of birth. In normal situations, lambs nurse after standing, receiving vital colostrum milk. Lambs that either fail to nurse or are rejected by the ewe require help to survive, such as bottle-feeding or fostering by another ewe.
Most lambs begin life being born outdoors. After lambs are several weeks old, lamb marking (ear tagging, docking, mulesing, and castrating) is carried out. Vaccinations are usually carried out at this point as well. Ear tags with numbers are attached, or ear marks are applied, for ease of later identification of sheep. Docking and castration are commonly done after 24 hours (to avoid interference with maternal bonding and consumption of colostrum) and are often done not later than one week after birth, to minimize pain, stress, recovery time and complications. The first course of vaccinations (commonly anti-clostridial) is commonly given at an age of about 10 to 12 weeks; i.e. when the concentration of maternal antibodies passively acquired via colostrum is expected to have fallen low enough to permit development of active immunity. Ewes are often revaccinated annually about 3 weeks before lambing, to provide high antibody concentrations in colostrum during the first several hours after lambing. Ram lambs that will either be slaughtered or separated from ewes before sexual maturity are not usually castrated. Objections to all these procedures have been raised by animal rights groups, but farmers defend them by saying they save money, and inflict only temporary pain.
Homosexuality
Sheep are the only species of mammal except for humans which exhibits exclusive homosexual behavior. About 10% of rams refuse to mate with ewes but readily mate with other rams, and thirty percent of all rams demonstrate at least some homosexual behavior. Additionally, a small number of females that were accompanied by a male fetus in utero (i.e. as fraternal twins) are freemartins (female animals that are behaviorally masculine and lack functioning ovaries).
Health
Sheep may fall victim to poisons, infectious diseases, and physical injuries. As a prey species, a sheep's system is adapted to hide the obvious signs of illness, to prevent being targeted by predators. However, some signs of ill health are obvious, with sick sheep eating little, vocalizing excessively, and being generally listless. Throughout history, much of the money and labor of sheep husbandry has aimed to prevent sheep ailments. Historically, shepherds often created remedies by experimentation on the farm. In some developed countries, including the United States, sheep lack the economic importance for drug companies to perform expensive clinical trials required to approve more than a relatively limited number of drugs for ovine use. However, extra-label drug use in sheep production is permitted in many jurisdictions, subject to certain restrictions. In the US, for example, regulations governing extra-label drug use in animals are found in 21 CFR (Code of Federal Regulations) Part 530. In the 20th and 21st centuries, a minority of sheep owners have turned to alternative treatments such as homeopathy, herbalism and even traditional Chinese medicine to treat sheep veterinary problems. Despite some favorable anecdotal evidence, the effectiveness of alternative veterinary medicine has been met with skepticism in scientific journals. The need for traditional anti-parasite drugs and antibiotics is widespread, and is the main impediment to certified organic farming with sheep.
Many breeders take a variety of preventive measures to ward off problems. The first is to ensure all sheep are healthy when purchased. Many buyers avoid outlets known to be clearing houses for animals culled from healthy flocks as either sick or simply inferior. This can also mean maintaining a closed flock, and quarantining new sheep for a month. Two fundamental preventive programs are maintaining good nutrition and reducing stress in the sheep. Restraint, isolation, loud noises, novel situations, pain, heat, extreme cold, fatigue and other stressors can lead to secretion of cortisol, a stress hormone, in amounts that may indicate welfare problems. Excessive stress can compromise the immune system. "Shipping fever" (pneumonic mannheimiosis, formerly called pasteurellosis) is a disease of particular concern, that can occur as a result of stress, notably during transport and (or) handling. Pain, fear and several other stressors can cause secretion of epinephrine (adrenaline). Considerable epinephrine secretion in the final days before slaughter can adversely affect meat quality (by causing glycogenolysis, removing the substrate for normal post-slaughter acidification of meat) and result in meat becoming more susceptible to colonization by spoilage bacteria. Because of such issues, low-stress handling is essential in sheep management. Avoiding poisoning is also important; common poisons are pesticide sprays, inorganic fertilizer, motor oil, as well as radiator coolant containing ethylene glycol.
Common forms of preventive medication for sheep are vaccinations and treatments for parasites. Both external and internal parasites are the most prevalent malady in sheep, and are either fatal, or reduce the productivity of flocks. Worms are the most common internal parasites. They are ingested during grazing, incubate within the sheep, and are expelled through the digestive system (beginning the cycle again). Oral anti-parasitic medicines, known as drenches, are given to a flock to treat worms, sometimes after worm eggs in the feces has been counted to assess infestation levels. Afterwards, sheep may be moved to a new pasture to avoid ingesting the same parasites. External sheep parasites include: lice (for different parts of the body), sheep keds, nose bots, sheep itch mites, and maggots. Keds are blood-sucking parasites that cause general malnutrition and decreased productivity, but are not fatal. Maggots are those of the bot fly and the blow-fly, commonly Lucilia sericata or its relative L. cuprina. Fly maggots cause the extremely destructive condition of flystrike. Flies lay their eggs in wounds or wet, manure-soiled wool; when the maggots hatch they burrow into a sheep's flesh, eventually causing death if untreated. In addition to other treatments, crutching (shearing wool from a sheep's rump) is a common preventive method. Some countries allow mulesing, a practice that involves stripping away the skin on the rump to prevent fly-strike, normally performed when the sheep is a lamb. Nose bots are fly larvae that inhabit a sheep's sinuses, causing breathing difficulties and discomfort. Common signs are a discharge from the nasal passage, sneezing, and frantic movement such as head shaking. External parasites may be controlled through the use of backliners, sprays or immersive sheep dips.
A wide array of bacterial and viral diseases affect sheep. Diseases of the hoof, such as foot rot and foot scald may occur, and are treated with footbaths and other remedies. Foot rot is present in over 97% of flocks in the UK. These painful conditions cause lameness and hinder feeding. Ovine Johne's disease is a wasting disease that affects young sheep. Bluetongue disease is an insect-borne illness causing fever and inflammation of the mucous membranes. Ovine rinderpest (or peste des petits ruminants) is a highly contagious and often fatal viral disease affecting sheep and goats. Sheep may also be affected by primary or secondary photosensitization. Tetanus can also afflict sheep through wounds from shearing, docking, castration, or vaccination. The organism also can be introduced into the reproductive tract by unsanitary humans who assist ewes during lambing.
A few sheep conditions are transmissible to humans. Orf (also known as scabby mouth, contagious ecthyma or soremouth) is a skin disease leaving lesions that is transmitted through skin-to-skin contact. Cutaneous anthrax is also called woolsorter's disease, as the spores can be transmitted in unwashed wool. More seriously, the organisms that can cause spontaneous enzootic abortion in sheep are easily transmitted to pregnant women. Also of concern are the prion disease scrapie and the virus that causes foot-and-mouth disease (FMD), as both can devastate flocks. The latter poses a slight risk to humans. During the 2001 FMD pandemic in the UK, hundreds of sheep were culled and some rare British breeds were at risk of extinction due to this.
Of the 600,300 sheep lost to the US economy in 2004, 37.3% were lost to predators, while 26.5% were lost to some form of disease. Poisoning accounted for 1.7% of non-productive deaths.
Predators
A lamb being attacked by coyotes with a bite to the throat
Other than parasites and disease, predation is a threat to sheep and the profitability of sheep raising. Sheep have little ability to defend themselves, compared with other species kept as livestock. Even if sheep survive an attack, they may die from their injuries or simply from panic. However, the impact of predation varies dramatically with region. In Africa, Australia, the Americas, and parts of Europe and Asia predators are a serious problem. In the United States, for instance, over one third of sheep deaths in 2004 were caused by predation. In contrast, other nations are virtually devoid of sheep predators, particularly islands known for extensive sheep husbandry. Worldwide, canids—including the domestic dog—are responsible for most sheep deaths. Other animals that occasionally prey on sheep include: felines, bears, birds of prey, ravens and feral hogs.
Sheep producers have used a wide variety of measures to combat predation. Pre-modern shepherds used their own presence, livestock guardian dogs, and protective structures such as barns and fencing. Fencing (both regular and electric), penning sheep at night and lambing indoors all continue to be widely used. More modern shepherds used guns, traps, and poisons to kill predators, causing significant decreases in predator populations. In the wake of the environmental and conservation movements, the use of these methods now usually falls under the purview of specially designated government agencies in most developed countries.
The 1970s saw a resurgence in the use of livestock guardian dogs and the development of new methods of predator control by sheep producers, many of them non-lethal. Donkeys and guard llamas have been used since the 1980s in sheep operations, using the same basic principle as livestock guardian dogs. Interspecific pasturing, usually with larger livestock such as cattle or horses, may help to deter predators, even if such species do not actively guard sheep. In addition to animal guardians, contemporary sheep operations may use non-lethal predator deterrents such as motion-activated lights and noisy alarms.
Economic importance
Main article: Agricultural economics
Global sheep stock
in 2019
Number in millions
1. China163.5 (13.19%)
2. India74.3 (5.99%)
3. Australia65.8 (5.31%)
4. Nigeria46.9 (3.78%)
5. Iran41.3 (3.33%)
6. Sudan40.9 (3.3%)
7. Chad35.9 (2.9%)
8. Turkey35.2 (2.84%)
9. United Kingdom33.6 (2.71%)
10. Mongolia32.3 (2.61%)
World total1,239.8
Source: UN Food and Agriculture Organization
Sheep are an important part of the global agricultural economy. However, their once vital status has been largely replaced by other livestock species, especially the pig, chicken, and cow. China, Australia, India, and Iran have the largest modern flocks, and serve both local and exportation needs for wool and mutton. Other countries such as New Zealand have smaller flocks but retain a large international economic impact due to their export of sheep products. Sheep also play a major role in many local economies, which may be niche markets focused on organic or sustainable agriculture and local food customers. Especially in developing countries, such flocks may be a part of subsistence agriculture rather than a system of trade. Sheep themselves may be a medium of trade in barter economies.
Domestic sheep provide a wide array of raw materials. Wool was one of the first textiles, although in the late 20th century wool prices began to fall dramatically as the result of the popularity and cheap prices for synthetic fabrics. For many sheep owners, the cost of shearing is greater than the possible profit from the fleece, making subsisting on wool production alone practically impossible without farm subsidies. Fleeces are used as material in making alternative products such as wool insulation. In the 21st century, the sale of meat is the most profitable enterprise in the sheep industry, even though far less sheep meat is consumed than chicken, pork or beef.
Sheepskin is likewise used for making clothes, footwear, rugs, and other products. Byproducts from the slaughter of sheep are also of value: sheep tallow can be used in candle and soap making, sheep bone and cartilage has been used to furnish carved items such as dice and buttons as well as rendered glue and gelatin. Sheep intestine can be formed into sausage casings, and lamb intestine has been formed into surgical sutures, as well as strings for musical instruments and tennis rackets. Sheep droppings, which are high in cellulose, have even been sterilized and mixed with traditional pulp materials to make paper. Of all sheep byproducts, perhaps the most valuable is lanolin: the waterproof, fatty substance found naturally in sheep's wool and used as a base for innumerable cosmetics and other products.
Some farmers who keep sheep also make a profit from live sheep. Providing lambs for youth programs such as 4-H and competition at agricultural shows is often a dependable avenue for the sale of sheep. Farmers may also choose to focus on a particular breed of sheep in order to sell registered purebred animals, as well as provide a ram rental service for breeding. A new option for deriving profit from live sheep is the rental of flocks for grazing; these "mowing services" are hired in order to keep unwanted vegetation down in public spaces and to lessen fire hazard.
Despite the falling demand and price for sheep products in many markets, sheep have distinct economic advantages when compared with other livestock. They do not require expensive housing, such as that used in the intensive farming of chickens or pigs. They are an efficient use of land; roughly six sheep can be kept on the amount that would suffice for a single cow or horse. Sheep can also consume plants, such as noxious weeds, that most other animals will not touch, and produce more young at a faster rate. Also, in contrast to most livestock species, the cost of raising sheep is not necessarily tied to the price of feed crops such as grain, soybeans and corn. Combined with the lower cost of quality sheep, all these factors combine to equal a lower overhead for sheep producers, thus entailing a higher profitability potential for the small farmer. Sheep are especially beneficial for independent producers, including family farms with limited resources, as the sheep industry is one of the few types of animal agriculture that has not been vertically integrated by agribusiness. However, small flocks, from 10 to 50 ewes, often are not profitable because they tend to be poorly managed. The primary reason is that mechanization is not feasible, so return per hour of labor is not maximized. Small farm flocks generally are used simply to control weeds on irrigation ditches or maintained as a hobby.
Shoulder of lamb
Sheep meat and milk were one of the earliest staple proteins consumed by human civilization after the transition from hunting and gathering to agriculture. Sheep meat prepared for food is known as either mutton or lamb, and approximately 540 million sheep are slaughtered each year for meat worldwide. "Mutton" is derived from the Old French moton, which was the word for sheep used by the Anglo-Norman rulers of much of the British Isles in the Middle Ages. This became the name for sheep meat in English, while the Old English word sceap was kept for the live animal. Throughout modern history, "mutton" has been limited to the meat of mature sheep usually at least two years of age; "lamb" is used for that of immature sheep less than a year.
In the 21st century, the nations with the highest consumption of sheep meat are the Arab states of the Persian Gulf, New Zealand, Australia, Greece, Uruguay, the United Kingdom and Ireland. These countries eat 14–40 lbs (3–18 kg) of sheep meat per capita, per annum. Sheep meat is also popular in France, Africa (especially the Arab world), the Caribbean, the rest of the Middle East, India, and parts of China. This often reflects a history of sheep production. In these countries in particular, dishes comprising alternative cuts and offal may be popular or traditional. Sheep testicles—called animelles or lamb fries—are considered a delicacy in many parts of the world. Perhaps the most unusual dish of sheep meat is the Scottish haggis, composed of various sheep innards cooked along with oatmeal and chopped onions inside its stomach. In comparison, countries such as the U.S. consume only a pound or less (under 0.5 kg), with Americans eating 50 pounds (22 kg) of pork and 65 pounds (29 kg) of beef. In addition, such countries rarely eat mutton, and may favor the more expensive cuts of lamb: mostly lamb chops and leg of lamb.
Though sheep's milk may be drunk rarely in fresh form, today it is used predominantly in cheese and yogurt making. Sheep have only two teats, and produce a far smaller volume of milk than cows. However, as sheep's milk contains far more fat, solids, and minerals than cow's milk, it is ideal for the cheese-making process. It also resists contamination during cooling better because of its much higher calcium content. Well-known cheeses made from sheep milk include the feta of Bulgaria and Greece, Roquefort of France, Manchego from Spain, the pecorino romano (the Italian word for "sheep" is pecore) and ricotta of Italy. Yogurts, especially some forms of strained yogurt, may also be made from sheep milk. Many of these products are now often made with cow's milk, especially when produced outside their country of origin. Sheep milk contains 4.8% lactose, which may affect those who are intolerant.
As with other domestic animals, the meat of uncastrated males is inferior in quality, especially as they grow. A "bucky" lamb is a lamb which was not castrated early enough, or which was castrated improperly (resulting in one testicle being retained). These lambs are worth less at market.
In science
Sheep are generally too large and reproduce too slowly to make ideal research subjects, and thus are not a common model organism. They have, however, played an influential role in some fields of science. In particular, the Roslin Institute of Edinburgh, Scotland used sheep for genetics research that produced groundbreaking results. In 1995, two ewes named Megan and Morag were the first mammals cloned from differentiated cells, also referred to as gynomerogony. A year later, a Finnish Dorset sheep named Dolly, dubbed "the world's most famous sheep" in Scientific American, was the first mammal to be cloned from an adult somatic cell. Following this, Polly and Molly were the first mammals to be simultaneously cloned and transgenic.
As of 2008, the sheep genome has not been fully sequenced, although a detailed genetic map has been published, and a draft version of the complete genome produced by assembling sheep DNA sequences using information given by the genomes of other mammals. In 2012, a transgenic sheep named "Peng Peng" was cloned by Chinese scientists, who spliced his genes with that of a roundworm (C. elegans) in order to increase production of fats healthier for human consumption.
In the study of natural selection, the population of Soay sheep that remain on the island of Hirta have been used to explore the relation of body size and coloration to reproductive success. Soay sheep come in several colors, and researchers investigated why the larger, darker sheep were in decline; this occurrence contradicted the rule of thumb that larger members of a population tend to be more successful reproductively. The feral Soays on Hirta are especially useful subjects because they are isolated.
Domestic sheep are sometimes used in medical research, particularly for researching cardiovascular physiology, in areas such as hypertension and heart failure. Pregnant sheep are also a useful model for human pregnancy, and have been used to investigate the effects on fetal development of malnutrition and hypoxia. In behavioral sciences, sheep have been used in isolated cases for the study of facial recognition, as their mental process of recognition is qualitatively similar to humans.
Cultural impact
Sheep have had a strong presence in many cultures, especially in areas where they form the most common type of livestock. In the English language, to call someone a sheep or ovine may allude that they are timid and easily led. In contradiction to this image, male sheep are often used as symbols of virility and power; the logos of the Los Angeles Rams football team and the Dodge Ram pickup truck allude to males of the bighorn sheep, Ovis canadensis.
Counting sheep is popularly said to be an aid to sleep, and some ancient systems of counting sheep persist today. Sheep also enter in colloquial sayings and idiom frequently with such phrases as "black sheep". To call an individual a black sheep implies that they are an odd or disreputable member of a group. This usage derives from the recessive trait that causes an occasional black lamb to be born into an entirely white flock. These black sheep were considered undesirable by shepherds, as black wool is not as commercially viable as white wool. Citizens who accept overbearing governments have been referred to by the Portmanteau neologism of sheeple. Somewhat differently, the adjective "sheepish" is also used to describe embarrassment.
In heraldry
In British heraldry, sheep appear in the form of rams, sheep proper and lambs. These are distinguished by the ram being depicted with horns and a tail, the sheep with neither and the lamb with its tail only. A further variant of the lamb, termed the Paschal lamb, is depicted as carrying a Christian cross and with a halo over its head. Rams' heads, portrayed without a neck and facing the viewer, are also found in British armories. The fleece, depicted as an entire sheepskin carried by a ring around its midsection, originally became known through its use in the arms of the Order of the Golden Fleece and was later adopted by towns and individuals with connections to the wool industry. A sheep on a blue field is depicted on the greater/royal arms of the king of Denmark to represent the Faroe Islands. In 2004 a modernized arms has been adopted by the Faroe Islands, which based on a 15th century coat of arms.
Religion and folklore
In antiquity, symbolism involving sheep cropped up in religions in the ancient Near East, the Mideast, and the Mediterranean area: Çatalhöyük, ancient Egyptian religion, the Cana'anite and Phoenician tradition, Judaism, Greek religion, and others. Religious symbolism and ritual involving sheep began with some of the first known faiths: Skulls of rams (along with bulls) occupied central placement in shrines at the Çatalhöyük settlement in 8,000 BCE. In Ancient Egyptian religion, the ram was the symbol of several gods: Khnum, Heryshaf and Amun (in his incarnation as a god of fertility). Other deities occasionally shown with ram features include the goddess Ishtar, the Phoenician god Baal-Hamon, and the Babylonian god Ea-Oannes. In Madagascar, sheep were not eaten as they were believed to be incarnations of the souls of ancestors.
There are many ancient Greek references to sheep: that of Chrysomallos, the golden-fleeced ram, continuing to be told through into the modern era. Astrologically, Aries, the ram, is the first sign of the classical Greek zodiac, and the sheep is the eighth of the twelve animals associated with the 12-year cycle of in the Chinese zodiac, related to the Chinese calendar. It is said in Chinese traditions that Hou ji sacrificed sheep. Mongolia, shagai are an ancient form of dice made from the cuboid bones of sheep that are often used for fortunetelling purposes.
Sheep play an important role in all the Abrahamic faiths; Abraham, Isaac, Jacob, Moses, and King David were all shepherds. According to the Biblical story of the Binding of Isaac, a ram is sacrificed as a substitute for Isaac after an angel stays Abraham's hand (in the Islamic tradition, Abraham was about to sacrifice Ishmael). Eid al-Adha is a major annual festival in Islam in which sheep (or other animals) are sacrificed in remembrance of this act. Sheep are occasionally sacrificed to commemorate important secular events in Islamic cultures. Greeks and Romans sacrificed sheep regularly in religious practice, and Judaism once sacrificed sheep as a Korban (sacrifice), such as the Passover lamb. Ovine symbols—such as the ceremonial blowing of a shofar—still find a presence in modern Judaic traditions.
Collectively, followers of Christianity are often referred to as a flock, with Christ as the Good Shepherd, and sheep are an element in the Christian iconography of the birth of Jesus. Some Christian saints are considered patrons of shepherds, and even of sheep themselves. Christ is also portrayed as the Sacrificial lamb of God (Agnus Dei) and Easter celebrations in Greece and Romania traditionally feature a meal of Paschal lamb. A church leader is often called the pastor, which is derived from the Latin word for shepherd. In many western Christian traditions bishops carry a staff, which also serves as a symbol of the episcopal office, known as a crosier, which is modeled on the shepherd's crook.
Sheep are key symbols in fables and nursery rhymes like The Wolf in Sheep's Clothing, Little Bo Peep, Baa, Baa, Black Sheep, and Mary Had a Little Lamb; novels such as George Orwell's Animal Farm and Haruki Murakami's A Wild Sheep Chase; songs such as Bach's Sheep may safely graze (Schafe können sicher weiden) and Pink Floyd's "Sheep", and poems like William Blake's "The Lamb".
18x28 giclee on canvas
Escape From Heaven is based on the Legend of the Hindu Apsara (angel) Urvashi.
Urvashi found the atmosphere among the Gods stifling. Everything in heaven seemed too perfect. The colors were too bright, the flowers smelled too sweet. Nothing seemed real to Urvashi. Wanting a taste of reality, Urvashi often snuck away to the Earth at night with her fellow Apsaras to feel the wet dew under her feet and the soft breeze against her body. It was during one of these many earthly excursions that she met King Pururava.
King Pururava envied the Gods. He was a regular invitee to Indra's court and dreamed at night of the grandeur he saw there. He would often take his chariot above the clouds and hurtle through the skies at break-neck speed. It was on such an occasion that the two met.
Urvashi was returning to heaven just before dawn with the other apsaras, when she was abducted by a demon . Pururava saw the abduction and chased the demon on his chariot and freed Urvashi from his clutches. The brief period their bodies touched changed their lives forever. For the first time Urvashi experienced the warm flesh of a mortal, for the first time she heard blood pounding in veins and for the first time she heard the inhalation and exhalation of breath. Pururavas had seen Urvashi in Indra’s court before and to actually be close to the most beautiful woman in heaven exhilarated him. Pururava left Urvashi with her friends but when they parted each was madly in love with the other.
Eventually the Urvashi became the wife of King Pururava and remained on earth with him for the rest of his life. The two had many sons.
Copyright 2007 Christopher Beikmann. All Rights Reserved
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Text by Ánxel Vázquez de la Cruz
"THE LAND’S REPLY"
First there was land:
Its supply of darkness,
moisture, and silence.
It took in the seed:
a whole life in two embracing spirals,
precarious reserves for the hazardous wait.
And after rains,
suns, mists…
the silent exhalation of the land:
That symmetry of colour and peace.
The poet put away its river melody.
And, later, he told the land:
- We don’t deserve it! We don´t deserve this!
And he heard the land’s silence:
- You are right: I know; but it´s just the way I am.
Ánxel Vázquez de la Cruz
____________________
"A RESPOSTA DA TERRA"
Texto de Ánxel Vázquez de la Cruz
Primeiro foi a terra:
A súa oferta de penumbra,
humidade e silencio.
Acolleu un día a semente:
a vida toda en dúas espirais que se abrazan,
precarias reservas para a azarosa espera.
E despois de chuvias,
soles, orballos…
a silandeira exhalación da terra:
esa simetría de cor e paz.
O poeta gardou a súa melodía fluvial.
E, despois, díxolle á terra:
- Non o merecemos:¡ Non merecemos isto!
E oíu o silencio da terra:
- Tes razón: xa o sei; pero é a miña forma de ser.
Ánxel Vázquez de la Cruz
www.xerais.es/cgigeneral/newFichaProducto.pl?obrcod=25200...
www.laopinioncoruna.es/cultura/2010/08/03/angel-vazquez-c...
www.farodevigo.es/comarcas/2010/10/06/medico-tudense-ange...
www.lavozdegalicia.es/coruna/2010/08/03/0003_8647038.htm
ver a Horta de Carnati:
www.xoanpinon.com/galerias/colecciones/colecs01/galego/co...
www.xoanpinon.com/galerias/colecciones/colecs02/galego/co...
At 21:47 GMT, the equinox happened, and so from then on, light is destined to win over darkness. Which meant, of course, that the day before then was the shortest "day", or amount of daylight.
This is the end of the year, the build up and excitement before Christmas, and at the same time, looking back at the year, and what has happened in the previous 50 or so weeks. So, a time of mixed emotions, good and bad, happy and sad.
But I was on vacation, or not going to work.
I am not up to date, but I did all the tasks I was supposed to do, threw a few electronic grenades over the walls, and was now happy not to think of that shit for two whole weeks.
For Jools, however, there was half a day to do, and then her employers paid for all those employed at the factory to go to a fancy place in Folkestone for lunch, drinks at the bar and a bottle of wine between four folks.
It was, in short, a time for celebration. Something I realise has not happened in my job since I left operational quality, to be happy and give thanks to those we work with. And be recognised for the good job we do.
So, I was to take Jools to work, and have the car for the day.
Jools was conscious that my plan for the day involved driving to the far west of Kent, so realised I needed an early start, and not dropping her off in Hythe at seven.
We left after coffee just after six, driving through Dover and Folkestone on the main road and motorway before turning over the downs into Hythe. I dropped her off in the town, so she could get some walking in. She always didn't walk, as waves of showers swept over the town, and me as I drove back home for breakfast and do all the chores before leaving on a mini-churchcrawl.
So, back home for breakfast, more coffee, wash up, do the bird feeders and with postcodes, set out for points in the extreme west. Now, Kent is not a big county, not say, Texas big, but it takes some time to get to some parts of the west of the county. Main roads run mainly from London to the coast, so going cross-country or cross-county would take time.
At first it was as per normal up the A20 then onto the motorway to Ashford then to Maidstone until the junction before the M26 starts. One of the reasons for going later was to avoid rush hours in and around Maidstone, Tonbridge and Tunbridge Wells.
As it was, after turning down the A road, things were fine until I got to Mereworth, but from there the road began to twist and turn until it lead me into Tonbridge. Once upon a time, this was a sleepy village or small town. The the railways came and it became a major junction. The road to Penshurt took me though the one way system, then down the wide High Street, over the river Medway and up the hill the other side.
Two more turns took me to my target, through what were once called stockbroker mansions, then down a hill, with the village laid out before me just visible through the trees.
The village was built around the outskirts of Penshurst Place, home to the Sidney family since Tudor times. Just about everything is named the Leicester something, the village having its own Leicester Square, though with no cinemas, and all timber framed houses and painfully picturesque.
The church lays behind the houses, the tower in golden sandstone topped with four spirelets.
I parked the car, and armed with two cameras, several lenses and a photographer's eye, walked to the church.
The reason for coming was I can only remember a little about my previous visit, but the Leicester name thing triggered in my head the thought the memorials and tombs might be worth a revisit.
So there I was.
Gilbert Scott was very busy here, so there is little of anything prior to the 19th century, but the memorials are there. Including one which features the heads of the children of Robert Sidney (d1702) in a cloud. Including the eldest son who died, apparently, so young he wasn't named, and is recorded as being the first born.
This is in the Sidney Chapel where the great and good are buried and remembered, it has a colourful roof, or roof beams, and heraldic shields. It has a 15th century font, which, sadly, has been brightly painted so is gaudy in the extreme.
I go around getting my shots, leave a fiver for the church. Go back to the car and program Speldhurst into the sat nav.
Its just a ten minute drive, but there is no place to park anywhere near the church. I could see from my slow drive-by the porch doors closed, and I convinced myself they were locked and not worth checking out.
I went on to Groombridge, where there is a small chapel with fabulous glass. I had been here before too, but wanted to redo my shots.
It was by now pouring with rain, and as dark as twilight, I missed the church on first pass, went to the mini-roundabout only to discover that it and the other church in the village were in Sussex. I turned round, the church looked dark and was almost certainly locked. I told myself.
I didn't stop here either, so instead of going to the final village church, I went straigh to Tunbridge Wells where there was another church to revisit.
I drove into the town, over the man road and to the car park with no waiting in traffic, how odd, I thought.
It was hard to find a parking space, but high up in the parking house there were finally spaced. I parked near the stairs down, grabbed my cameras and went down.
I guess I could have parked nearer the church, but once done it would be easier to leave the town as the road back home went past the exit.
I ambled down the hill leading to the station, over the bridge and down the narrow streets, all lined with shops. I think its fair to say that it is a richer town than Dover because on one street there were three stores offering beposke designer kitchens.
The church is across the road from the Georgian square known at The Pantiles, but it was the church I was here to visit.
I go in, and there is a service underway. I decide to sit at the back and observe.
And pray.
I did not take communion, though. The only one there who didn't.
About eight elderly parishioners did, though.
I was here to photograph the ceiling, and then the other details I failed to record when we were last here over a decade ago.
I was quizzed strongly by a warden as to why I was doing this. I had no answer other than I enjoyed it, and for me that is enough.
After getting my shots, I leave and begin the slog back up to the car, but on the way keeping my promise to a young man selling the Big Issue that I would come back and buy a copy. I did better than that in that I gave him a fiver and didn't take a copy.
He nearly burst into tears. I said, there is kindness in the world, and some of us do keep our promises.
By the time I got to the car park, it was raining hard again. I had two and a half hours to get to Folkestone to pick up Jools after her meal.
Traffic into Tunbridge Wells from this was was crazy, miles and miles of queues, so I was more than happy going the other way.
I get back to the M20, cruise down to Ashford, stopping at Stop 24 services for a coffee and something to eat. I had 90 minutes to kill, so eat, drink and scroll Twitter as I had posted yet more stuff that morning. In other news: nothing changed, sadly.
At quarter past four I went to pick up Jools, stopping outside the restaurant. When she got in she declared she had been drinking piña coladas. Just two, but she was bubby and jabbering away all the way home.
With Jools having eaten out, and with snacks I had, no dinner was needed, so when suppertime came round, we dined on cheese and crackers, followed by a large slice of Christmas cake.
She was now done for Christmas too.
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The red brick church stands on a busy junction at the end of the Pantiles whose patrons it was built to serve in 1678. Within thirty years it had been extended on two occasions to more or less reach its present size. The ceiling bears the date 1678 and is rather domestic in character, based on deep circular domes with putti, palms and swags. The stained glass in the east window is based on a picture by Alex Ender and was designed by Heaton, Butler and Bayne in 1901. There is an excellent window under the north gallery designed by Lawrence Lee in 1969. The church was sympathetically restored by Ewan Christian in 1882, when the shallow chancel was added. The woodwork it contains was brought from one of Wren's City of London churches. Outside the west wall of the church, set into the footpath, is a boundary marker to show the former parish boundaries of Tonbridge and Speldhurst.
www.kentchurches.info/church.asp?p=Tunbridge+Wells+1
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The large and populous hamlet or village of TUNBRIDGE-WELLS is situated at the south-east boundary of this parish; part of it only is in Speldhurst, another part in the parish of Tunbridge, and the remainder in that of Fant, in the county of Suffex. It consists of four smaller districts, named from the hills on which they stand, Mount Ephraim, Mount Pleasant, and Mount Sion; the other is called The Wells, from their being within it, which altogether form a considerable town; but the last is the centre of business and pleasure, for there, besides the Wells themselves, are the market, public parades, assembly rooms, taverns, shops, &c. Near the Wells is the chapel, which stands remarkably in the three parishes above mentioned—the pulpit in Speldhurst, the altar in Tunbridge, and the vestry in Fant, and the stream, which parted the two counties of Kent and Suffex, formerly ran underneath it, but is now turned to a further distance from it. The right of patronage is claimed by the rector of Speldhurst, though he has never yet possessed the chapel or presented to it; the value of it is about two hundred pounds per annum, which sum is raised by voluntary subscription; divine service is performed in it every day in summer, and three times a week in winter. Adjoining to it is a charity school, for upwards of fifty poor boys and girls, which is supported by a contribution, collected at the chapel doors, two or three times a year.
The trade of Tunbridge-wells is similar to that of Spa, in Germany, and consists chiefly in a variety of toys, made of wood, commonly called Tunbridge ware, which employs a great number of hands. The wood principally used for this purpose is beech and sycamore, with yew and holly inlaid, and beautifully polished. To the market of this place is brought, in great plenty, from the South downs, in Sussex, the little bird, called the wheatear, which, from its delicacy, is usually called the English ortolan. It is not bigger in size than a lark; it is almost a lump of fat, and of a very delicious taste; it is in season only in the midst of summer, when the heat of the weather, and the fatness of it, prevents its being sent to London, which otherwise would, in all likelihood, monopolize every one of them. On the other or Suffex side of the Medway, above a mile from the Wells, are the rocks, which consist of a great number of rude eminences, adjoining to each other, several of which are seventy feet in height; in several places there are cliffs and chasms which lead quite through the midst of them, by narrow gloomy passages, which strike the beholder with astonishment.
THESE MEDICINAL WATERS, commonly called TUNBRIDGE-WELLS, lie so near to the county of Suffex that part of them are within it, for which reason they were for some time called Fant-wells, as being within that parish. (fn. 1) Their efficacy is reported to have been accidentally found out by Dudley lord North, in the beginning of the reign of king James I. Whilst he resided at Eridge-house for his health, lord Abergavenny's seat, in this neighbourhood, and that he was entirely cured of the lingering consumptive disorder he laboured under by the use of them.
The springs, which were then discovered, seem to have been seven in number, two of the principal of which were some time afterwards, by lord Abergavenny's care, inclosed, and were afterwards much resorted to by many of the middling and lower sort, whose ill health had real occasion for the use of them. In which state they continued till queen Henrietta Maria, wife of king Charles I. having been sent hither by her physicians, in the year 1630, for the reestablishment of her health, soon brought these waters into fashion, and occasioned a great resort to them from that time. In compliment to her doctor, Lewis Rowzee, in his treatise on them, calls these springs the Queen's-wells; but this name lasted but a small time, and they were soon afterwards universally known by that of Tunbridge-wells, which names they acquired from the company usually residing at Tunbridge town, when they came into these parts for the benefit of drinking the waters.
The town of Tunbridge being five miles distant from the wells, occasioned some few houses to be built in the hamlets of Southborough and Rusthall, for the accommodation of the company resorting hither, and this place now becoming fashionable, was visited by numbers for the sake of pleasure and dissipation, as well as for the cure of their infirmities; and soon after the Restoration every kind of building, for public amusements, was erected at the two hamlets above mentioned, lodgings and other buildings were built at and near the wells, the springs themselves were secured, and other conveniencies added to them. In 1664, the queen came here by the advice of her physicians, in hopes of reinstating her health, which was greatly impaired by a dangerous fever, and her success, in being perfectly cured by these waters, greatly raised the reputation of them, and the company increasing yearly, it induced the inhabitants to make every accommodation for them adjoining to the Wells, so that both Rusthall and Southborough became ruinous and deserted by all but their native inhabitants. The duke of York, with his duchess, and the two princesses their daughters, visited Tunbridge-wells in the year 1670, which brought much more company than usual to them, and raised their reputation still higher; and the annual increase continuing, it induced the lord of the manor to think of improving this humour of visiting the wells to his own profit as well as the better accommodation of the company. To effect which, he entered into an agreement with his tenants, and hired of them the herbage of the waste of the manor for the term of fifty years, at the yearly rent of ten shillings to each tenant, and then erected shops and houses on and near the walks and springs, in every convenient spot for that purpose; by which means Tunbridge wells became a populous and flourishing village, well inhabited, for whose convenience, and the company resorting thither, a chapel was likewise built, in 1684, by subscription, on some ground given by the lady viscountess Purbeck, which was, about twelve years afterwards, enlarged by an additional subscription, amounting together to near twenty-three hundred pounds.
About the year 1726, the building lease, which had been granted by the lord of the manor of Rusthall, in which this hamlet is situated, expiring, the tenants of the manor claimed a share in the buildings, as a compensation for the loss of the herbage, which was covered by his houses. This occasioned a long and very expensive law suit between them, which was at last determined in favour of the tenants, who were adjudged to have a right to a third part of the buildings then erected on the estate, in lieu of their right to the herbage; upon which all the shops and houses, which had been built on the manor waste, were divided into three lots, of which the tenants were to draw one, and the other two were to remain to the lord of the manor; the lot which the tenants drew was the middle one, which included the assembly room on the public walk, which has since turned out much the most advantageous of the three. After which long articles of agreement, in 1739, were entered into between Maurice Conyers, esq. then lord of the manor of Rusthall, and the above mentioned tenants of it, in which, among many other matters, he agreed to permit the public walks and wells, and divers other premises there, to be made use of for the public benefit of the nobility and gentry resorting thereto, and several regulations were made in them concerning the walks, wells, and wastes of the manor, and for the restraining buildings on the waste, between the lord and his tenants, according to a plan therein specified; all which were confirmed and established by an act of parliament, passed in 1740. Since which several of the royal family have honoured these wells with their presence, and numbers of the nobility and persons of rank and fashion yearly resortto them, so that this place is now in a most flourishing state, having great numbers of good houses built for lodgings, and every other necessary accommodation for the company. Its customs are settled; the employment of the dippers regulated; (fn. 2) its pleasures regulated; its markets well and plentifully supplied, at a reasonable rate, with sowl, fish, meat, every other kind of food, and every convenience added that can contribute to give health and pleasure.
¶The whole neighbourhood of Tunbridge-wells abounds with springs of mineral water, but as the properties of all are nearly the same, only those two, which at the first discovery of them were adjudged the best, are held in any particular estimation. These two wells are enclosed with a handsome triangular stone wall; over the springs are placed two convenient basons of Portland stone, with perforations at the bottom; one of them being given by queen Anne, and the other by the lord of the manor; through which they receive the water, which at the spring is extremely clear and bright. Its taste is steely, but not disagreeable; it has hardly any smell, though sometimes, in a dense air, its ferruginous exhalations are very distinguishable. In point of heat it is invariably temperate, the spring lying so deep in the earth, that neither the heat of summer, nor the cold of winter, affects it. When this water is first taken up in a large glass, its particles continue at rest till it is warmed to nearly the heat of the atmosphere, then a few airy globules begin to separate themselves, and adhere to the sides of the glass, and in a few hours a light copper coloured scum begins to float on the surface, after which an ochreous sediment settles at the bottom. Long continued rains sometimes give the water a milky appearance, but do not otherwise sensibly affect it. From the experiments of different physicians, it appears that the component parts of this water are, steely particles, marine salts, an oily matter, an ochreous substance, simple water, and a volatile vitriolic spirit, too subtile for any chemical analysis. In weight it is, in seven ounces and a quarter, four grains lighter than the German Spa (to which it is preferable on that account) and ten grains lighter than common water; with syrup of violets this water gives a deep green, as vitriols do. (fn. 3) It requires five drops of oleum sulphuris, or elixir of vitriol, to a quart of water, to preserve its virtues to a distance from the spring.
This water is said to be an impregnation of rain in some of the neighbouring eminences, which abound in iron mineral, where it is further enriched with the marine salts and all the valuable ingredients, which constitute it a light and pure chalybeate, which instantly searches the most remote recesses of the human frame, warms and invigorates the relaxed constitution, restores the weakened fibres to their due tone and elasticity, removes those obstructions to which the minuter vessels of the body are liable, and is consequently adapted to most cold chronical disorders, lowness of spirits, weak digestions, and nervous complaints. Dr. Lodowick Rowzee, of Ashford, in this county, wrote a Treatise of the Nature and Virtues of these Waters, printed in 12mo. 1671; and Dr. Patrick Madan wrote a Philosophical and Medical Essay on them, in 1687, in quarto.
"...and silent bells in doors
of Childhood's friends; among recorded
sounds of unanimous inscriptions, and aura's
perfumes of oldies tapes; among erased
by needles vinyl records with Opera's fragments,
assembled and uniformly distributed
collective exhalations of Oceanic depth
from Whales inspired by songs of own wedding's ..."
As the sun sets over Sebastian Inlet, the light catches an exhalation from Catalog #3450. The right whale and her calf swam into the inlet on the morning of Feb. 8, 2016 and spent most of their day about ¾ mile from the ocean. Biologists don't know why the whales entered the inlet, but it was likely accidental. The pair appear healthy.
Photo Taken: Feb. 8, 2016
Photo Credit (full credit required for use):
Florida Fish and Wildlife Conservation Commission (FWC)
Khajuraho, a UNESCO World Heritage Site, is a famous tourist and archaeological site known for its sculptured temples dedicated to Shiva, Vishnu, Jain patriarchs, and others. Khajuraho was one of the capitals of the Chandela kings, who from the 9th to the 12th century CE developed a large realm, which at its height included almost all of what is now Madhya Pradesh state. Khajuraho extended over 21 sq. km and contained about 85 temples, built by multiple rulers mostly between 950 to 1050. In the 12th century the Chandelas, in a period of chaos and decline, abandoned Khajuraho and moved to hill forts elsewhere. Khajuraho puttered along until the 14th century (Ibn Batuta was impressed by its temples) but was afterwards largely forgotten and overtaken by jungles, which probably saved it from the desecration that Muslim conquerors sometimes inflicted on Hindu monuments. In 1838 a British army captain, TS Burt, employed by the Asiatic Society in Calcutta, came upon information that led him to the rediscovery of the complex of temples in the jungle in Khajuraho.
Of the 85 original temples—most constructed of sandstone—about 20 are still reasonably well preserved. Both internally and externally the temples are richly carved with excellent sculptures that are frequently sensual and often sexually explicit. The temples are divided into three complexes—the western is the largest and best known, containing the magnificent Shaivite temple Kandariya Mahadev, a 31m high agglomeration of porches and turrets culminating in a spire. Khajuraho's name derives from the prevalence of khajur, or date palms, in the area.
There are many theories explaining the real reason for incorporating erotic art with the temples. One of the most popular theories is that they were meant to provide education about earthly desires. During the medieval era, young boys were sent to hermitage and practice bramhacharya till reaching maturity. The sculptures were meant to educate them about grahasthahram.
Yet another theory states that they were meant to represent kama (desire) as the third purushartha (aim of life). The sculptures are also accredited to the rise of the Tantric cults during that time. However, several experts reject these theories.
However, the most credible explanations for the erotic sculptures at Khajuraho are as follows:
1. Sign of happiness, prosperity and auspiciousness: During the medieval era there was a common belief that having erotic sculptures were considered alankaras or decorative motifs, protective and auspicious. This hypothesis is based on the authoritative religious texts like the Shilpashastras and the Brihat Samhita. According to the Brihat Samhita, mithunas (couples), goblins, creepers and erotic sculptures were meant to be carved on the temple door to bring in good luck and as a sign of auspiciousness.
2. Mock the Ascetics: Yet another theory explaining the old erotic art at Khajuraho states that the coital couples represent ascetics as well as people from the royal class. It is also believed that the sculptures of couples participating in the orgies are scenes imagined by the artists. Also, it is also said that the artists used ascetics in sensuous and passionate moods as a way to mock the extreme Tantric sects that rose during that era.
3. Code Language: According to experts, the erotic figures were used as a code language to convey Tantric doctrines and non-communicable experiences. For e.g. A sculpture wherein a washerwoman clings to an ascetic may look erotic and sensuous for the layman. However, in the Tantric language, the washerwoman represents the Kundalini energy that has ascended up to the chakras i.e. the neck of the ascetic. Thus, the erotic sculptures have a deeper meaning related to the Tantric cult.
4. Conceal the Magico-Propitiory Yantra: According to the architectural text of the Shilpa Prakasha, every temple must have a Kamakala Yantra strategically placed to protect it from evil spirits and natural calamities. However, the Yantra, which is basically a set of lines drawn symmetrically, must not be visible to the lay man. Thus, a few erotic sculptures with the head down posture were made corresponding to the lines of the Yantra and superimposed on it.
5. Non-duality: It is possible that the erotic sculptures symbolically represent the union of two opposing forces or energies like, inhalation and exhalation, in a timeless state of non-duality.
The erotic art at Khajuraho is considered to be the pinnacle of love and passion. However, during the period between 900 – 1300 AD most Hindu, Jain and Buddhist temples across western and southern India comprised of erotic art.
However, at other temples, these sculptures were carved at the plinth level, below the eye level and thus did not get noticed. It is only at Khajuraho that these sculptures were so prominently displayed on the main wall of the temples
We stopped at the barachois in Gaskiers-PointLa Haye on the way to St. Vincent's Beach, looking across to the Point La Haye Light. The wheeling herring gulls, black backed gulls and northern gannets meant there were capelin in the area for their annual spawning on the shoreline. And capelin drawn humpback whales who stay in these waters to feed on capelin all summer. Sure enough we saw a plume of mist, the whale's exhalation . . . and then the sight of the dorsal fin as the humpback whale rolled. There were two in this area and others at St. Vincent's Beach which is well known for its whale watching from the shore.
I absolutely love photography. It's not the end result necessarily, so much as the process itself. Walking about, camera in hand, the whole world becomes sharper, clearer, imbued with life, a life of its own. The Self opens its eyes and begins to see, the Self raises the camera, collects the shot, exhales its Self out into nothingness, the Ego vanishes, the shutter clicks, the viewfinder goes black. Everything suddenly disappears- and for that brief instant, you are totally free. And so you keep doing it, over and over again... and something of that seeing, that exhalation, and that vanishing remains with you, like a remembered movement, retaining that feeling when you see your friend, or do your work, or whatever else you may do.
Enlightenment is for the film, not the operator. The finger that clicks the shutter is limited in purpose, driven by will, and focused on a pinpoint- but the film is open, empty, and receptive.
That's what I long to be like, some day.
Thanks to all of you my friends, who are supporting me in what is basically a brief time of absence from here- I'm not online much at all at the moment, and everything is in transition. I miss you guys, and I miss your photos and the daily inspiration they give me, and the deeper appreciation for the world we live in, as it is seen by friends and comrades, and the sense of community and sharing we have... it is of immense personal value to me, and I hope to be back here soon! Winter has passed, summer is beginning, and all is well, if a little unpredictable... and that is usually fertile ground for new blessings. My very best wishes go with you all!
At 21:47 GMT, the equinox happened, and so from then on, light is destined to win over darkness. Which meant, of course, that the day before then was the shortest "day", or amount of daylight.
This is the end of the year, the build up and excitement before Christmas, and at the same time, looking back at the year, and what has happened in the previous 50 or so weeks. So, a time of mixed emotions, good and bad, happy and sad.
But I was on vacation, or not going to work.
I am not up to date, but I did all the tasks I was supposed to do, threw a few electronic grenades over the walls, and was now happy not to think of that shit for two whole weeks.
For Jools, however, there was half a day to do, and then her employers paid for all those employed at the factory to go to a fancy place in Folkestone for lunch, drinks at the bar and a bottle of wine between four folks.
It was, in short, a time for celebration. Something I realise has not happened in my job since I left operational quality, to be happy and give thanks to those we work with. And be recognised for the good job we do.
So, I was to take Jools to work, and have the car for the day.
Jools was conscious that my plan for the day involved driving to the far west of Kent, so realised I needed an early start, and not dropping her off in Hythe at seven.
We left after coffee just after six, driving through Dover and Folkestone on the main road and motorway before turning over the downs into Hythe. I dropped her off in the town, so she could get some walking in. She always didn't walk, as waves of showers swept over the town, and me as I drove back home for breakfast and do all the chores before leaving on a mini-churchcrawl.
So, back home for breakfast, more coffee, wash up, do the bird feeders and with postcodes, set out for points in the extreme west. Now, Kent is not a big county, not say, Texas big, but it takes some time to get to some parts of the west of the county. Main roads run mainly from London to the coast, so going cross-country or cross-county would take time.
At first it was as per normal up the A20 then onto the motorway to Ashford then to Maidstone until the junction before the M26 starts. One of the reasons for going later was to avoid rush hours in and around Maidstone, Tonbridge and Tunbridge Wells.
As it was, after turning down the A road, things were fine until I got to Mereworth, but from there the road began to twist and turn until it lead me into Tonbridge. Once upon a time, this was a sleepy village or small town. The the railways came and it became a major junction. The road to Penshurt took me though the one way system, then down the wide High Street, over the river Medway and up the hill the other side.
Two more turns took me to my target, through what were once called stockbroker mansions, then down a hill, with the village laid out before me just visible through the trees.
The village was built around the outskirts of Penshurst Place, home to the Sidney family since Tudor times. Just about everything is named the Leicester something, the village having its own Leicester Square, though with no cinemas, and all timber framed houses and painfully picturesque.
The church lays behind the houses, the tower in golden sandstone topped with four spirelets.
I parked the car, and armed with two cameras, several lenses and a photographer's eye, walked to the church.
The reason for coming was I can only remember a little about my previous visit, but the Leicester name thing triggered in my head the thought the memorials and tombs might be worth a revisit.
So there I was.
Gilbert Scott was very busy here, so there is little of anything prior to the 19th century, but the memorials are there. Including one which features the heads of the children of Robert Sidney (d1702) in a cloud. Including the eldest son who died, apparently, so young he wasn't named, and is recorded as being the first born.
This is in the Sidney Chapel where the great and good are buried and remembered, it has a colourful roof, or roof beams, and heraldic shields. It has a 15th century font, which, sadly, has been brightly painted so is gaudy in the extreme.
I go around getting my shots, leave a fiver for the church. Go back to the car and program Speldhurst into the sat nav.
Its just a ten minute drive, but there is no place to park anywhere near the church. I could see from my slow drive-by the porch doors closed, and I convinced myself they were locked and not worth checking out.
I went on to Groombridge, where there is a small chapel with fabulous glass. I had been here before too, but wanted to redo my shots.
It was by now pouring with rain, and as dark as twilight, I missed the church on first pass, went to the mini-roundabout only to discover that it and the other church in the village were in Sussex. I turned round, the church looked dark and was almost certainly locked. I told myself.
I didn't stop here either, so instead of going to the final village church, I went straigh to Tunbridge Wells where there was another church to revisit.
I drove into the town, over the man road and to the car park with no waiting in traffic, how odd, I thought.
It was hard to find a parking space, but high up in the parking house there were finally spaced. I parked near the stairs down, grabbed my cameras and went down.
I guess I could have parked nearer the church, but once done it would be easier to leave the town as the road back home went past the exit.
I ambled down the hill leading to the station, over the bridge and down the narrow streets, all lined with shops. I think its fair to say that it is a richer town than Dover because on one street there were three stores offering beposke designer kitchens.
The church is across the road from the Georgian square known at The Pantiles, but it was the church I was here to visit.
I go in, and there is a service underway. I decide to sit at the back and observe.
And pray.
I did not take communion, though. The only one there who didn't.
About eight elderly parishioners did, though.
I was here to photograph the ceiling, and then the other details I failed to record when we were last here over a decade ago.
I was quizzed strongly by a warden as to why I was doing this. I had no answer other than I enjoyed it, and for me that is enough.
After getting my shots, I leave and begin the slog back up to the car, but on the way keeping my promise to a young man selling the Big Issue that I would come back and buy a copy. I did better than that in that I gave him a fiver and didn't take a copy.
He nearly burst into tears. I said, there is kindness in the world, and some of us do keep our promises.
By the time I got to the car park, it was raining hard again. I had two and a half hours to get to Folkestone to pick up Jools after her meal.
Traffic into Tunbridge Wells from this was was crazy, miles and miles of queues, so I was more than happy going the other way.
I get back to the M20, cruise down to Ashford, stopping at Stop 24 services for a coffee and something to eat. I had 90 minutes to kill, so eat, drink and scroll Twitter as I had posted yet more stuff that morning. In other news: nothing changed, sadly.
At quarter past four I went to pick up Jools, stopping outside the restaurant. When she got in she declared she had been drinking piña coladas. Just two, but she was bubby and jabbering away all the way home.
With Jools having eaten out, and with snacks I had, no dinner was needed, so when suppertime came round, we dined on cheese and crackers, followed by a large slice of Christmas cake.
She was now done for Christmas too.
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The red brick church stands on a busy junction at the end of the Pantiles whose patrons it was built to serve in 1678. Within thirty years it had been extended on two occasions to more or less reach its present size. The ceiling bears the date 1678 and is rather domestic in character, based on deep circular domes with putti, palms and swags. The stained glass in the east window is based on a picture by Alex Ender and was designed by Heaton, Butler and Bayne in 1901. There is an excellent window under the north gallery designed by Lawrence Lee in 1969. The church was sympathetically restored by Ewan Christian in 1882, when the shallow chancel was added. The woodwork it contains was brought from one of Wren's City of London churches. Outside the west wall of the church, set into the footpath, is a boundary marker to show the former parish boundaries of Tonbridge and Speldhurst.
www.kentchurches.info/church.asp?p=Tunbridge+Wells+1
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The large and populous hamlet or village of TUNBRIDGE-WELLS is situated at the south-east boundary of this parish; part of it only is in Speldhurst, another part in the parish of Tunbridge, and the remainder in that of Fant, in the county of Suffex. It consists of four smaller districts, named from the hills on which they stand, Mount Ephraim, Mount Pleasant, and Mount Sion; the other is called The Wells, from their being within it, which altogether form a considerable town; but the last is the centre of business and pleasure, for there, besides the Wells themselves, are the market, public parades, assembly rooms, taverns, shops, &c. Near the Wells is the chapel, which stands remarkably in the three parishes above mentioned—the pulpit in Speldhurst, the altar in Tunbridge, and the vestry in Fant, and the stream, which parted the two counties of Kent and Suffex, formerly ran underneath it, but is now turned to a further distance from it. The right of patronage is claimed by the rector of Speldhurst, though he has never yet possessed the chapel or presented to it; the value of it is about two hundred pounds per annum, which sum is raised by voluntary subscription; divine service is performed in it every day in summer, and three times a week in winter. Adjoining to it is a charity school, for upwards of fifty poor boys and girls, which is supported by a contribution, collected at the chapel doors, two or three times a year.
The trade of Tunbridge-wells is similar to that of Spa, in Germany, and consists chiefly in a variety of toys, made of wood, commonly called Tunbridge ware, which employs a great number of hands. The wood principally used for this purpose is beech and sycamore, with yew and holly inlaid, and beautifully polished. To the market of this place is brought, in great plenty, from the South downs, in Sussex, the little bird, called the wheatear, which, from its delicacy, is usually called the English ortolan. It is not bigger in size than a lark; it is almost a lump of fat, and of a very delicious taste; it is in season only in the midst of summer, when the heat of the weather, and the fatness of it, prevents its being sent to London, which otherwise would, in all likelihood, monopolize every one of them. On the other or Suffex side of the Medway, above a mile from the Wells, are the rocks, which consist of a great number of rude eminences, adjoining to each other, several of which are seventy feet in height; in several places there are cliffs and chasms which lead quite through the midst of them, by narrow gloomy passages, which strike the beholder with astonishment.
THESE MEDICINAL WATERS, commonly called TUNBRIDGE-WELLS, lie so near to the county of Suffex that part of them are within it, for which reason they were for some time called Fant-wells, as being within that parish. (fn. 1) Their efficacy is reported to have been accidentally found out by Dudley lord North, in the beginning of the reign of king James I. Whilst he resided at Eridge-house for his health, lord Abergavenny's seat, in this neighbourhood, and that he was entirely cured of the lingering consumptive disorder he laboured under by the use of them.
The springs, which were then discovered, seem to have been seven in number, two of the principal of which were some time afterwards, by lord Abergavenny's care, inclosed, and were afterwards much resorted to by many of the middling and lower sort, whose ill health had real occasion for the use of them. In which state they continued till queen Henrietta Maria, wife of king Charles I. having been sent hither by her physicians, in the year 1630, for the reestablishment of her health, soon brought these waters into fashion, and occasioned a great resort to them from that time. In compliment to her doctor, Lewis Rowzee, in his treatise on them, calls these springs the Queen's-wells; but this name lasted but a small time, and they were soon afterwards universally known by that of Tunbridge-wells, which names they acquired from the company usually residing at Tunbridge town, when they came into these parts for the benefit of drinking the waters.
The town of Tunbridge being five miles distant from the wells, occasioned some few houses to be built in the hamlets of Southborough and Rusthall, for the accommodation of the company resorting hither, and this place now becoming fashionable, was visited by numbers for the sake of pleasure and dissipation, as well as for the cure of their infirmities; and soon after the Restoration every kind of building, for public amusements, was erected at the two hamlets above mentioned, lodgings and other buildings were built at and near the wells, the springs themselves were secured, and other conveniencies added to them. In 1664, the queen came here by the advice of her physicians, in hopes of reinstating her health, which was greatly impaired by a dangerous fever, and her success, in being perfectly cured by these waters, greatly raised the reputation of them, and the company increasing yearly, it induced the inhabitants to make every accommodation for them adjoining to the Wells, so that both Rusthall and Southborough became ruinous and deserted by all but their native inhabitants. The duke of York, with his duchess, and the two princesses their daughters, visited Tunbridge-wells in the year 1670, which brought much more company than usual to them, and raised their reputation still higher; and the annual increase continuing, it induced the lord of the manor to think of improving this humour of visiting the wells to his own profit as well as the better accommodation of the company. To effect which, he entered into an agreement with his tenants, and hired of them the herbage of the waste of the manor for the term of fifty years, at the yearly rent of ten shillings to each tenant, and then erected shops and houses on and near the walks and springs, in every convenient spot for that purpose; by which means Tunbridge wells became a populous and flourishing village, well inhabited, for whose convenience, and the company resorting thither, a chapel was likewise built, in 1684, by subscription, on some ground given by the lady viscountess Purbeck, which was, about twelve years afterwards, enlarged by an additional subscription, amounting together to near twenty-three hundred pounds.
About the year 1726, the building lease, which had been granted by the lord of the manor of Rusthall, in which this hamlet is situated, expiring, the tenants of the manor claimed a share in the buildings, as a compensation for the loss of the herbage, which was covered by his houses. This occasioned a long and very expensive law suit between them, which was at last determined in favour of the tenants, who were adjudged to have a right to a third part of the buildings then erected on the estate, in lieu of their right to the herbage; upon which all the shops and houses, which had been built on the manor waste, were divided into three lots, of which the tenants were to draw one, and the other two were to remain to the lord of the manor; the lot which the tenants drew was the middle one, which included the assembly room on the public walk, which has since turned out much the most advantageous of the three. After which long articles of agreement, in 1739, were entered into between Maurice Conyers, esq. then lord of the manor of Rusthall, and the above mentioned tenants of it, in which, among many other matters, he agreed to permit the public walks and wells, and divers other premises there, to be made use of for the public benefit of the nobility and gentry resorting thereto, and several regulations were made in them concerning the walks, wells, and wastes of the manor, and for the restraining buildings on the waste, between the lord and his tenants, according to a plan therein specified; all which were confirmed and established by an act of parliament, passed in 1740. Since which several of the royal family have honoured these wells with their presence, and numbers of the nobility and persons of rank and fashion yearly resortto them, so that this place is now in a most flourishing state, having great numbers of good houses built for lodgings, and every other necessary accommodation for the company. Its customs are settled; the employment of the dippers regulated; (fn. 2) its pleasures regulated; its markets well and plentifully supplied, at a reasonable rate, with sowl, fish, meat, every other kind of food, and every convenience added that can contribute to give health and pleasure.
¶The whole neighbourhood of Tunbridge-wells abounds with springs of mineral water, but as the properties of all are nearly the same, only those two, which at the first discovery of them were adjudged the best, are held in any particular estimation. These two wells are enclosed with a handsome triangular stone wall; over the springs are placed two convenient basons of Portland stone, with perforations at the bottom; one of them being given by queen Anne, and the other by the lord of the manor; through which they receive the water, which at the spring is extremely clear and bright. Its taste is steely, but not disagreeable; it has hardly any smell, though sometimes, in a dense air, its ferruginous exhalations are very distinguishable. In point of heat it is invariably temperate, the spring lying so deep in the earth, that neither the heat of summer, nor the cold of winter, affects it. When this water is first taken up in a large glass, its particles continue at rest till it is warmed to nearly the heat of the atmosphere, then a few airy globules begin to separate themselves, and adhere to the sides of the glass, and in a few hours a light copper coloured scum begins to float on the surface, after which an ochreous sediment settles at the bottom. Long continued rains sometimes give the water a milky appearance, but do not otherwise sensibly affect it. From the experiments of different physicians, it appears that the component parts of this water are, steely particles, marine salts, an oily matter, an ochreous substance, simple water, and a volatile vitriolic spirit, too subtile for any chemical analysis. In weight it is, in seven ounces and a quarter, four grains lighter than the German Spa (to which it is preferable on that account) and ten grains lighter than common water; with syrup of violets this water gives a deep green, as vitriols do. (fn. 3) It requires five drops of oleum sulphuris, or elixir of vitriol, to a quart of water, to preserve its virtues to a distance from the spring.
This water is said to be an impregnation of rain in some of the neighbouring eminences, which abound in iron mineral, where it is further enriched with the marine salts and all the valuable ingredients, which constitute it a light and pure chalybeate, which instantly searches the most remote recesses of the human frame, warms and invigorates the relaxed constitution, restores the weakened fibres to their due tone and elasticity, removes those obstructions to which the minuter vessels of the body are liable, and is consequently adapted to most cold chronical disorders, lowness of spirits, weak digestions, and nervous complaints. Dr. Lodowick Rowzee, of Ashford, in this county, wrote a Treatise of the Nature and Virtues of these Waters, printed in 12mo. 1671; and Dr. Patrick Madan wrote a Philosophical and Medical Essay on them, in 1687, in quarto.
Persephone supervising Sisyphus in the Underworld, Attica black-figure amphora (vase), c. 530 BC, Staatliche Antikensammlungen[1]
In Greek mythology Sisyphus (/ˈsɪsᵻfəs/;[2] Greek: Σίσυφος, Sísuphos) was the king of Ephyra (now known as Corinth). He was punished for his self-aggrandizing craftiness and deceitfulness by being forced to roll an immense boulder up a hill, only to watch it come back to hit him, repeating this action for eternity. Through the classical influence on modern culture, tasks that are both laborious and futile are therefore described as Sisyphean (/ˌsɪsᵻˈfiːən/).
Sisyphus was the son of King Aeolus of Thessaly and Enarete. He was the founder and first king of Ephyra (supposedly the original name of Corinth).He was the father of Glaucus, Ornytion, Almus, and Thersander by the nymph Merope, the brother of Salmoneus, and the grandfather of Bellerophon through Glaucus.King Sisyphus promoted navigation and commerce but was avaricious and deceitful. He also killed travellers and guests, a violation of xenia, which fell under Zeus's domain. He took pleasure in these killings because they allowed him to maintain his iron-fisted rule.
Sisyphus and his brother Salmoneus were known to hate each other, and Sisyphus consulted with the Oracle of Delphi on just how to kill Salmoneus without incurring any severe consequences for himself. From Homer onward, Sisyphus was famed as the craftiest of men. He seduced Salmoneus's daughter Tyro in one of his plots to kill Salmoneus, only for Tyro to slay the children she bore by him when she discovered that Sisyphus was planning on eventually using them to dethrone her father.
King Sisyphus also betrayed one of Zeus' secrets by revealing the whereabouts of Aegina (an Asopides who was taken away by Zeus) to her father (the river god Asopus) in return for causing a spring to flow on the Corinthian acropolis.[citation needed]
Zeus then ordered Thanatos, Death, to chain King Sisyphus down below in Tartarus. Sisyphus was curious as to why Hermes, whose job it was to guide souls to the Underworld, had not appeared on this occasion. King Sisyphus slyly asked Thanatos to demonstrate how the chains worked. As Thanatos was granting him his wish, Sisyphus seized the opportunity and trapped Thanatos in the chains instead. Once Thanatos was bound by the strong chains, no one died on earth. This caused an uproar especially for Ares (who was annoyed that his battles had lost their fun because his opponents would not die), and so he intervened. The exasperated Ares freed Thanatos and turned King Sisyphus over to Thanatos.[9]
In another version, Hades was sent to chain Sisyphus and was chained himself. As long as Hades was tied up, nobody could die. Because of this, sacrifices could not be made to the gods, and those that were old and sick were suffering. The gods finally threatened to make life so miserable for Sisyphus that he would wish he were dead. He then had no choice but to release Hades.[10]
Before King Sisyphus died, he had told his wife to throw his naked body into the middle of the public square (purportedly as a test of his wife's love for him). This caused King Sisyphus to end up on the shores of the river Styx. Then, complaining to Persephone that this was a sign of his wife's disrespect for him, King Sisyphus persuaded her to allow him to return to the upper world. Once back in Corinth, the spirit of King Sisyphus scolded his wife for not burying his body and giving it a proper funeral (as a loving wife should). When King Sisyphus refused to return to the Underworld, he was forcibly dragged back there by Hermes.[citation needed] In another version of the myth, Persephone, goddess of the Underworld, was tricked by Sisyphus that he had been conducted to Tartarus by mistake, and so she ordered that he be released.
In Philoctetes by Sophocles, there is a reference to the father of Odysseus (rumoured to have been Sisyphus, and not Laërtes, whom we know as the father in the Odyssey) upon having returned from the dead.
As a punishment for his trickery, King Sisyphus was made to endlessly roll a huge boulder up a steep hill. The maddening nature of the punishment was reserved for King Sisyphus due to his hubristic belief that his cleverness surpassed that of Zeus himself. Zeus accordingly displayed his own cleverness by enchanting the boulder into rolling away from King Sisyphus before he reached the top, which ended up consigning Sisyphus to an eternity of useless efforts and unending frustration. Thus it came to pass that pointless or interminable activities are sometimes described as Sisyphean. King Sisyphus was a common subject for ancient writers and was depicted by the painter Polygnotus on the walls of the Lesche at Delphi.
Interpretations
According to the solar theory, King Sisyphus is the disk of the sun that rises every day in the east and then sinks into the west.[15] Other scholars regard him as a personification of waves rising and falling, or of the treacherous sea.[15] The 1st-century BC Epicurean philosopher Lucretius interprets the myth of Sisyphus as personifying politicians aspiring for political office who are constantly defeated, with the quest for power, in itself an "empty thing", being likened to rolling the boulder up the hill.[16] Søren Kierkegaard saw the myth as pertaining to anything a person loves too much: "It is comic that a mentally disordered man picks up any piece of granite and carries it around because he thinks it is money, and in the same way it is comic that Don Juan has 1,003 mistresses, for the number simply indicates that they have no value. Therefore, one should stay within one’s means in the use of the word “love."[17] Friedrich Welcker suggested that he symbolises the vain struggle of man in the pursuit of knowledge, and Salomon Reinach[18] that his punishment is based on a picture in which Sisyphus was represented rolling a huge stone Acrocorinthus, symbolic of the labour and skill involved in the building of the Sisypheum. Albert Camus, in his 1942 essay The Myth of Sisyphus, saw Sisyphus as personifying the absurdity of human life, but Camus concludes "one must imagine Sisyphus happy" as "The struggle itself towards the heights is enough to fill a man's heart." More recently, J. Nigro Sansonese,[19] building on the work of Georges Dumézil, speculates that the origin of the name "Sisyphos" is onomatopoetic of the continual back-and-forth, susurrant sound ("siss phuss") made by the breath in the nasal passages, situating the mythology of Sisyphus in a far larger context of archaic (see Proto-Indo-European religion) trance-inducing techniques related to breath control. The repetitive inhalation–exhalation cycle is described esoterically in the myth as an up–down motion of Sisyphus and his boulder on a hill.
In experiments that test how workers respond when the meaning of their task is diminished, the test condition is referred to as the Sisyphusian condition. The two main conclusions of the experiment are that people work harder when their work seems more meaningful, and that people underestimate the relationship between meaning and motivation.
Literary interpretations.
Sisyphys (1548–49) by Titian, Prado Museum, Madrid, Spain
Sisypher dwarves by Tomasz Moczek (pl) in Wrocław
Homer describes Sisyphus in both Book VI of the Iliad and Book XI of the Odyssey.
Ovid, the Roman poet, makes reference to Sisyphus in the story of Orpheus and Eurydice. When Orpheus descends and confronts Hades and Persephone, he sings a song so that they will grant his wish to bring Eurydice back from the dead. After this song is sung, Ovid shows how moving it was by noting that Sisyphus, emotionally affected, for just a moment, stops his eternal task and sits on his rock, the Latin wording being inque tuo sedisti, Sisyphe, saxo ("you sat upon your rock, Sisyphus").
Though purported to be one of the dialogues of Greek philosopher Plato, the Sisyphus is generally believed to be apocryphal, possibly written by one of his pupils. In his Apology, Socrates considers Sisyphus to be a wise man he may meet in the afterlife.
Albert Camus, the French absurdist, wrote an essay entitled The Myth of Sisyphus, in which he elevates Sisyphus to the status of absurd hero. Franz Kafka repeatedly referred to Sisyphus as a bachelor; Kafkaesque for him were those qualities that brought out the Sisyphus-like qualities in himself. According to Frederick Karl: "The man who struggled to reach the heights only to be thrown down to the depths embodied all of Kafka's aspirations; and he remained himself, alone, solitary." The philosopher Richard Taylor uses the myth of Sisyphus as a representation of a life made meaningless because it consists of bare repetition. James Clement van Pelt, co-founder of Yale's Initiative in Religion, Science & Technology, suggests that Sisyphus also personifies humanity and its disastrous pursuit of perfection by any means necessary, in which the great rock repeatedly rushing down the mount symbolizes the accelerating pace of unsustainable civilization toward cataclysmic collapse and cultural oblivion that ends each historical age and restarts the sisyphean cycle. The Myth of Sisyphus, saw Sisyphus as personifying the absurdity of human life, but Camus concludes "one must imagine Sisyphus happy"
The American alligator (Alligator mississippiensis), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the genus Alligator, and is larger than the only other living alligator species, the Chinese alligator.
Adult male American alligators measure 3.4 to 4.6 m (11.2 to 15.1 ft) in length, and can weigh up to 500 kg (1,100 lb), with unverified sizes of up to 5.84 m (19.2 ft) and weights of 1,000 kg (2,200 lb) making it the second largest member of the family Alligatoridae, after the black caiman. Females are smaller, measuring 2.6 to 3 m (8.5 to 9.8 ft) in length. The American alligator inhabits subtropical and tropical freshwater wetlands, such as marshes and cypress swamps, from southern Texas to North Carolina. It is distinguished from the sympatric American crocodile by its broader snout, with overlapping jaws and darker coloration, and is less tolerant of saltwater but more tolerant of cooler climates than the American crocodile, which is found only in tropical and warm subtropical climates.
American alligators are apex predators and consume fish, amphibians, reptiles, birds, and mammals. Hatchlings feed mostly on invertebrates. They play an important role as ecosystem engineers in wetland ecosystems through the creation of alligator holes, which provide both wet and dry habitats for other organisms. Throughout the year (in particular during the breeding season), American alligators bellow to declare territory, and locate suitable mates. Male American alligators use infrasound to attract females. Eggs are laid in a nest of vegetation, sticks, leaves, and mud in a sheltered spot in or near the water. Young are born with yellow bands around their bodies and are protected by their mother for up to one year.[
The conservation status of the American alligator is listed as Least Concern by the International Union for Conservation of Nature. Historically, hunting had decimated their population, and the American alligator was listed as an endangered species by the Endangered Species Act of 1973. Subsequent conservation efforts have allowed their numbers to increase and the species was removed from endangered status in 1987. The species is the official state reptile of three states: Florida, Louisiana, and Mississippi.
Taxonomy
The American alligator was first classified by French zoologist François Marie Daudin as Crocodilus mississipiensis in 1801. In 1807, Georges Cuvier created the genus Alligator; the American alligator and the Chinese alligator are the only extant species in the genus. They are grouped in the family Alligatoridae with the caimans. The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the American alligator than to either the Nile crocodile (Crocodylus niloticus) or the gharial (Gavialis gangeticus).
Phylogeny
Members of this superfamily first arose in the late Cretaceous, about 100–65 million years ago (Mya). Leidyosuchus of Alberta is the earliest known fossil, from the Campanian era 83 to 72 Mya. Fossil alligatoroids have been found throughout Eurasia, because bridges across both the North Atlantic and the Bering Strait connected North America to Eurasia about 66 to 23 Mya.
Alligators and caimans split in North America during the late Cretaceous, and the caimans reached South America by the Paleogene, before the closure of the Isthmus of Panama during the Neogene period, from about 23 to 2.58 Mya. The Chinese alligator likely descended from a lineage that crossed the Bering land bridge during the Neogene. Fossils identical to the existing American alligator are found throughout the Pleistocene, from 2.5 million to 11.7 thousand years ago. In 2016, a Miocene (about 23 to 5.3 Mya) fossil skull of an alligator was found at Marion County, Florida. Unlike the other extinct alligator species of the same genus, the fossil skull was virtually indistinguishable from that of the modern American alligator. This alligator and the American alligator are now considered to be sister taxa, meaning that the A. mississippiensis lineage has existed in North America for over 8 million years.
The alligator's full mitochondrial genome was sequenced in the 1990s, and it suggests the animal evolved at a rate similar to mammals and greater than birds and most cold-blooded vertebrates. However, the full genome, published in 2014, suggests that the alligator evolved much more slowly than mammals and birds.
Characteristics
American alligator skull
Domestic American alligators range from long and slender to short and robust, possibly in response to variations in factors such as growth rate, diet, and climate.
Size
The American alligator is a relatively large species of crocodilian. On average, it is the largest species in the family Alligatoridae, with only the black caiman being possibly larger. Weight varies considerably depending on length, age, health, season, and available food sources. Similar to many other reptiles that range expansively into temperate zones, American alligators from the northern end of their range, such as southern Arkansas, Alabama, and northern North Carolina, tend to reach smaller sizes. Large adult American alligators tend to be relatively robust and bulky compared to other similar-length crocodilians; for example, captive males measuring 3 to 4 m (9 ft 10 in to 13 ft 1 in) were found to weigh 200 to 350 kg (440 to 770 lb), although captive specimens may outweigh wild specimens due to lack of hunting behavior and other stressors.
Large male American alligators reach an expected maximum size up to 4.6 m (15 ft 1 in) in length and weighing up to 500 kg (1,100 lb), while females reach an expected maximum of 3 m (9 ft 10 in). However, the largest free-ranging female had a total length of 3.22 m (10 ft 7 in) and weighed 170 kg (370 lb). On rare occasions, a large, old male may grow to an even greater length.
Largest
During the 19th and 20th centuries, larger males reaching 5 to 6 m (16 ft 5 in to 19 ft 8 in) were reported. The largest reported individual size was a male killed in 1890 on Marsh Island, Louisiana, and reportedly measured at 5.84 m (19 ft 2 in) in length, but no voucher specimen was available, since the American alligator was left on a muddy bank after having been measured due to having been too massive to relocate. If the size of this animal was correct, it would have weighed about 1,000 kg (2,200 lb). In Arkansas, a man killed an American alligator that was 4.04 m (13 ft 3 in) and 626 kg (1,380 lb). The largest American alligator ever killed in Florida was 5.31 m (17 ft 5 in), as reported by the Everglades National Park, although this record is unverified. The largest American alligator scientifically verified in Florida for the period from 1977 to 1993 was reportedly 4.23 m (13 ft 11 in) and weighed 473 kg (1,043 lb), although another specimen (size estimated from skull) may have measured 4.54 m (14 ft 11 in). A specimen that was 4.5 m (14 ft 9 in) long and weighed 458.8 kg (1,011.5 lb) is the largest American alligator killed in Alabama and has been declared the SCI world record in 2014.
Average
American alligators do not normally reach such extreme sizes. In mature males, most specimens grow up to about 3.4 m (11 ft 2 in) in length, and weigh up to 360 kg (790 lb), while in females, the mature size is normally around 2.6 m (8 ft 6 in), with a body weight up to 91 kg (201 lb). In Newnans Lake, Florida, adult males averaged 73.2 kg (161 lb) in weight and 2.47 m (8 ft 1 in) in length, while adult females averaged 55.1 kg (121 lb) and measured 2.22 m (7 ft 3 in). In Lake Griffin State Park, Florida, adults weighed on average 57.9 kg (128 lb). Weight at sexual maturity per one study was stated as averaging 30 kg (66 lb) while adult weight was claimed as 160 kg (350 lb).
Relation to age
There is a common belief stated throughout reptilian literature that crocodilians, including the American alligator, exhibit indeterminate growth, meaning the animal continues to grow for the duration of its life. However, these claims are largely based on assumptions and observations of juvenile and young adult crocodilians, and recent studies are beginning to contradict this claim. For example, one long-term mark-recapture study (1979–2015) done at the Tom Yawkey Wildlife Center in South Carolina found evidence to support patterns of determinate growth, with growth ceasing upon reaching a certain age (43 years for males and 31 years for females).
Sexual dimorphism
While noticeable in very mature specimens, the sexual dimorphism in size of the American alligator is relatively modest among crocodilians. For contrast, the sexual dimorphism of saltwater crocodiles is much more extreme, with mature males nearly twice as long as and at least four times as heavy as female saltwater crocodiles. Given that female American alligators have relatively higher survival rates at an early age and a large percentage of given populations consists of immature or young breeding American alligators, relatively few large mature males of the expected mature length of 3.4 m (11 ft 2 in) or more are typically seen.
Color
Dorsally, adult American alligators may be olive, brown, gray, or black. However, they are on average one of the most darkly colored modern crocodilians (although other alligatorid family members are also fairly dark), and can be reliably be distinguished by color via their more blackish dorsal scales against crocodiles. Meanwhile, their undersides are cream-colored. Some American alligators are missing or have an inhibited gene for melanin, which makes them albino. These American alligators are extremely rare and almost impossible to find in the wild. They could only survive in captivity, as they are very vulnerable to the sun and predators.
Jaws, teeth, and snout
American alligators have 74–80 teeth. As they grow and develop, the morphology of their teeth and jaws change significantly. Juveniles have small, needle-like teeth that become much more robust and narrow snouts that become broader as the individuals develop. These morphological changes correspond to shifts in the American alligators' diets, from smaller prey items such as fish and insects to larger prey items such as turtles, birds, and other large vertebrates. American alligators have broad snouts, especially in captive individuals. When the jaws are closed, the edges of the upper jaws cover the lower teeth, which fit into the jaws' hollows. Like the spectacled caiman, this species has a bony nasal ridge, though it is less prominent. American alligators are often mistaken for a similar animal: the American crocodile. An easy characteristic to distinguish the two is the fourth tooth. Whenever an American alligator's mouth is closed, the fourth tooth is no longer visible. It is enclosed in a pocket in the upper jaw.
Bite
Adult American alligators held the record as having the strongest laboratory-measured bite of any living animal, measured at up to 13,172 N (1,343.2 kgf; 2,961 lbf). This experiment had not been, at the time of the paper published, replicated in any other crocodilians, and the same laboratory was able to measure a greater bite force of 16,414 N (1,673.8 kgf; 3,690 lbf) in saltwater crocodiles; notwithstanding this very high biting force, the muscles opening the American alligator's jaw are quite weak, and the jaws can be held closed by hand or tape when an American alligator is captured. No significant difference is noted between the bite forces of male and female American alligators of equal size. Another study noted that as the American alligator increases in size, the force of its bite also increases.
Movement
When on land, an American alligator moves either by sprawling or walking, the latter involving the reptile lifting its belly off the ground. The sprawling of American alligators and other crocodylians is not similar to that of salamanders and lizards, being similar to walking. Therefore, the two forms of land locomotion can be termed the "low walk" and the "high walk". Unlike most other land vertebrates, American alligators increase their speed through the distal rather than proximal ends of their limbs. In the water, American alligators swim like fish, moving their pelvic regions and tails from side to side. During respiration, air flow is unidirectional, looping through the lungs during inhalation and exhalation; the American alligator's abdominal muscles can alter the position of the lungs within the torso, thus shifting the center of buoyancy, which allows the American alligator to dive, rise, and roll within the water.
Distribution
American alligators, being native both to the Nearctic and Neotropical realms, are found in the wild in the Southeastern United States, from the Lowcountry in South Carolina, south to Everglades National Park in Florida, and west to the southeastern region of Texas. They are found in parts of North Carolina, South Carolina, Georgia, Florida, Louisiana, Alabama, Mississippi, Arkansas, Oklahoma and Texas. Some of these locations appear to be relatively recent introductions, with often small but reproductive populations. Louisiana has the largest American alligator population of any U.S. state. In the future, possible American alligator populations may be found in areas of Mexico adjacent to the Texas border. American alligators have been naturally expanding their range into Tennessee, and have established a small population in the southwestern part of that state via inland waterways, according to the state's wildlife agency. They have been extirpated from Virginia, and occasional vagrants from North Carolina wander into the Great Dismal Swamp.
Conservation status
American alligators are currently listed as least concern by the IUCN Red List, even though from the 1800s to the mid-1900s, they were being hunted and poached by humans unsustainably.
Historically, hunting and habitat loss have severely affected American alligator populations throughout their range, and whether the species would survive was in doubt. In 1967, the American alligator was listed as an endangered species (under a law that was the precursor Endangered Species Act of 1973), since it was believed to be in danger of extinction throughout all or a significant portion of its range.
Both the United States Fish and Wildlife Service (USFWS) and state wildlife agencies in the South contributed to the American alligator's recovery. Protection under the Endangered Species Act allowed the species to recuperate in many areas where it had been depleted. States began monitoring their American alligator populations to ensure that they would continue to grow. In 1987, the USFWS removed the animal from the endangered species list, as it was considered to be fully recovered. The USFWS still regulates the legal trade in American alligators and their products to protect still endangered crocodilians that may be passed off as American alligators during trafficking.
American alligators are listed under Appendix II of the Convention on International Trade in Endangered Species (CITES) meaning that international trade in the species (including parts and derivatives) is regulated.
Habitat
They inhabit swamps, streams, rivers, ponds, and lakes. A lone American alligator was spotted for over 10 years living in a river north of Atlanta, Georgia. Females and juveniles are also found in Carolina Bays and other seasonal wetlands. While they prefer fresh water, American alligators may sometimes wander into brackish water, but are less tolerant of salt water than American crocodiles, as the salt glands on their tongues do not function. One study of American alligators in north-central Florida found the males preferred open lake water during the spring, while females used both swampy and open-water areas. During summer, males still preferred open water, while females remained in the swamps to construct their nests and lay their eggs. Both sexes may den underneath banks or clumps of trees during the winter.
In some areas of their range, American alligators are an unusual example of urban wildlife; golf courses are often favored by the species due to an abundance of water and a frequent supply of prey animals such as fish and birds.
Cold tolerance
American alligators are less vulnerable to cold than American crocodiles. Unlike an American crocodile, which would immediately succumb to the cold and drown in water at 45 °F (7 °C) or less, an American alligator can survive in such temperatures for some time without displaying any signs of discomfort. This adaptiveness is thought to be why American alligators are widespread further north than the American crocodile. In fact, the American alligator is found farther from the equator and is more equipped to handle cooler conditions than any other crocodilian. When the water begins to freeze, American alligators go into a period of brumation; they stick their snouts through the surface, which allows them to breathe above the ice, and they can remain in this state for several days.
Ecology and behavior
Basking
American alligators primarily bask on shore, but also climb into and perch on tree limbs to bask if no shoreline is available. This is not often seen, since if disturbed, they quickly retreat back into the water by jumping from their perch.
Holes
American alligators modify wetland habitats, most dramatically in flat areas such as the Everglades, by constructing small ponds known as alligator holes. This behavior has qualified the American alligator to be considered a keystone species. Alligator holes retain water during the dry season and provide a refuge for aquatic organisms, which survive the dry season by seeking refuge in alligator holes, so are a source of future populations. The construction of nests along the periphery of alligator holes, as well as a buildup of soils during the excavation process, provides drier areas for other reptiles to nest and a place for plants that are intolerant of inundation to colonize. Alligator holes are an oasis during the Everglades dry season, so are consequently important foraging sites for other organisms. In the limestone depressions of cypress swamps, alligator holes tend to be large and deep, while those in marl prairies and rocky glades are usually small and shallow, and those in peat depressions of ridge and slough wetlands are more variable.
Bite and mastication
The teeth of the American alligator are designed to grip prey, but cannot rip or chew flesh like teeth of some other predators (such as canids and felids), and depend on their gizzard, instead, to masticate their food. The American alligator is capable of biting through a turtle's shell or a moderately sized mammal bone.
Tool use
American alligators have been documented using lures to hunt prey such as birds. This means they are among the first reptiles recorded to use tools. By balancing sticks and branches on their heads, American alligators are able to lure birds looking for suitable nesting material to kill and consume. This strategy, which is shared by the mugger crocodile, is particularly effective during the nesting season, in which birds are more likely to gather appropriate nesting materials. This strategy has been documented in two Florida zoos occurring multiple times a day in peak nesting season and in some parks in Louisiana. The use of tools was documented primarily during the peak rookery season when birds were primarily looking for sticks.
Aquatic vs terrestrial
Fish and other aquatic prey taken in the water or at the water's edge form the major part of American alligator's diet and may be eaten at any time of the day or night. Adult American alligators also spend considerable time hunting on land, up to 160 feet (50 m) from water, ambushing terrestrial animals on trailsides and road shoulders. Usually, terrestrial hunting occurs on nights with warm temperatures. When hunting terrestrial prey, American alligators may also ambush them from the edge of the water by grabbing them and pulling the prey into the water, the preferred method of predation of larger crocodiles.
An American alligator in the process of consuming a box turtle
Additionally, American alligators have recently been filmed and documented killing and eating sharks and rays; four incidents documented indicated that bonnetheads, lemon sharks, Atlantic stingrays, and nurse sharks are components of the animal's diet. Sharks are also known to prey on American alligators, in turn, indicating that encounters between the two predators are common.
Common
American alligators are considered an apex predator throughout their range. They are opportunists and their diet is determined largely by both their size and age and the size and availability of prey. Most American alligators eat a wide variety of animals, including invertebrates, fish, birds, turtles, snakes, amphibians, and mammals. Hatchlings mostly feed on invertebrates such as insects, insect larvae, snails, spiders, and worms, as well as small fish and frogs. As they grow, American alligators gradually expand to larger prey. Once an American alligator reaches full size and power in adulthood, any animal living in the water or coming to the water to drink is potential prey. Most animals captured by American alligators are considerably smaller than itself. A few examples of animals consumed are largemouth bass, spotted gar, freshwater pearl mussels, American green tree frogs, yellow mud turtles, cottonmouths, common moorhens, and feral wild boars. Stomach contents show, among native mammals, muskrats and raccoons are some of the most commonly eaten species. In Louisiana, where introduced nutria are common, they are perhaps the most regular prey for adult American alligators, although only larger adults commonly eat this species. It has also been reported that large American alligators prey on medium-sized American alligators, which had preyed on hatchlings and smaller juveniles.
If an American alligator's primary food resource is not available, it will sometimes feed on carrion and non-prey items such as rocks and artificial objects, like bottle caps. These items help the American alligator in the process of digestion by crushing up the meat and bones of animals, especially animals with shells.
Large animals
Other animals may occasionally be eaten, even large deer or feral wild boars, but these are not normally part of the diet. American alligators occasionally prey on large mammals, such as deer, but usually do so when fish and smaller prey levels go down. Rarely, American alligators have been observed killing and eating bobcats, but such events are not common and have little effect on bobcat populations. Although American alligators have been listed as predators of the Nilgai and the West Indian manatees, very little evidence exists of such predation. In the 2000s, when invasive Burmese pythons first occupied the Everglades, American alligators have been recorded preying on sizable snakes, possibly controlling populations and preventing the invasive species from spreading northwards. However, the python is also known to occasionally prey on alligators, a form of both competition and predation. American alligator predation on Florida panthers is rare, but has been documented. Such incidents usually involve a panther trying to cross a waterway or coming down to a swamp or river to get a drink. American alligator predation on American black bears has also been recorded.
Domestic animals
Occasionally, domestic animals, including dogs, cats, and calves, are taken as available, but are secondary to wild and feral prey. Other prey, including snakes, lizards, and various invertebrates, are eaten occasionally by adults.
Birds
Water birds, such as herons, egrets, storks, waterfowl and large dabbling rails such as gallinules or coots, are taken when possible. Occasionally, unwary adult birds are grabbed and eaten by American alligators, but most predation on bird species occurs with unsteady fledgling birds in late summer, as fledgling birds attempt to make their first flights near the water's edge.
Fruit
In 2013, American alligators and other crocodilians were reported to also eat fruit. Such behavior has been witnessed, as well as documented from stomach contents, with the American alligators eating such fruit as wild grapes, elderberries, and citrus fruits directly from the trees. Thirty-four families and 46 genera of plants were represented among seeds and fruits found in the stomach contents of alligators. The discovery of this unexpected part of the American alligator diet further reveals that they may be responsible for spreading seeds from the fruit they consume across their habitat.
Cooperative hunting
Additionally, American alligators engage in what seems to be cooperative hunting. One observation of cooperative hunting techniques was where there are pushing American alligators and catching American alligators and they were observed taking turns in each position. Another observation said that about 60 American alligators gathered in an area and would form a semicircle with about half of them and would push the fish closer to the bank. Once one of the American alligators caught a fish another one would enter into its spot, and it would take the fish to the resting area. This was reported to have occurred two days in a row.
In Florida and East Texas
The diet of adult American alligators from central Florida lakes is dominated by fish, but the species is highly opportunistically based upon local availability. In Lake Griffin, fish made up 54% of the diet by weight, with catfish being most commonly consumed, while in Lake Apopka, fish made up 90% of the food and mostly shad were taken; in Lake Woodruff, the diet was 84% fish and largely consists of bass and sunfish. Unusually in these regions, reptiles and amphibians were the most important nonpiscivore prey, mostly turtles and water snakes. In southern Louisiana, crustaceans (largely crawfish and crabs) were found to be present in the southeastern American alligators, but largely absent in southwestern American alligators, which consumed a relatively high proportion of reptiles, although fish were the most recorded prey for adults, and adult males consumed a large portion of mammals.
In East Texas, diets were diverse and adult American alligators took mammals, reptiles, amphibians, and invertebrates (e.g. snails) in often equal measure as they did fish.
Vocalizations
Mechanism
An American alligator is able to abduct and adduct the vocal folds of its larynx, but not to elongate or shorten them; yet in spite of this, it can modulate fundamental frequency very well. Their vocal folds consists of epithelium, lamina propria and muscle. Sounds ranged from 50 to 1200 Hz. In one experiment conducted on the larynx, the fundamental frequency depended on both the glottal gap and stiffness of the larynx tissues. As the frequency increases, there's high tension and large strains. The fundamental frequency has been influenced by the glottal gap size and subglottal pressure and when the phonation threshold pressure has been exceeded, there will be vocal fold vibration.
Calls
Crocodilians are the most vocal of all non-avian reptiles and have a variety of different calls depending on the age, size, and sex of the animal. The American alligator can perform specific vocalizations to declare territory, signal distress, threaten competitors, and locate suitable mates. Juveniles can perform a high-pitched hatchling call (a "yelping" trait common to many crocodilian species' hatchling young)[118] to alert their mothers when they are ready to emerge from the nest. Juveniles also make a distress call to alert their mothers if they are being threatened. Adult American alligators can growl, hiss, or cough to threaten others and declare territory.
Bellowing
Both males and females bellow loudly by sucking air into their lungs and blowing it out in intermittent, deep-toned roars to attract mates and declare territory. Males are known to use infrasound during mating bellows. Bellowing is performed in a "head oblique, tail arched" posture. Infrasonic waves from a bellowing male can cause the surface of the water directly over and to either side of his back to literally "sprinkle", in what is commonly called the "water dance". Large bellowing "choruses" of American alligators during the breeding season are commonly initiated by females and perpetuated by males. Observers of large bellowing choruses have noted they are often felt more than they are heard due to the intense infrasound emitted by males. American alligators bellow in B flat (specifically "B♭1", defined as an audio frequency of 58.27 Hz), and bellowing choruses can be induced by tuba players, sonic booms, and large aircraft.
Lifespan
American alligators typically live to the age of 50, and possibly over 70 years old. Males reach sexual maturity at around 11.6 years, and females at around 15.8 years. Although it was originally thought that American alligators never stop growing, studies have now found that males stop growing at around the age of 43 years, and females stop growing at around the age of 31 years.
Reproduction
Breeding season
The breeding season begins in the spring. On spring nights, American alligators gather in large numbers for group courtship, in the aforementioned "water dances". The female builds a nest of vegetation, sticks, leaves, and mud in a sheltered spot in or near the water.
Eggs
After the female lays her 20 to 50 white eggs, about the size of a goose egg, she covers them with more vegetation, which heats as it decays, helping to keep the eggs warm. This differs from Nile crocodiles, which lay their eggs in pits. The temperature at which American alligator eggs develop determines their sex (see temperature-dependent sex determination). Studies have found that eggs hatched at a temperature below 88.7 °F or a temperature above 94.1 °F will produce female offspring, while those at a temperature between 90.5 °F to 92.3 °F will produce male offspring. The nests built on levees are warmer, thus produce males, while the cooler nests of wet marsh produce females. The female remains near the nest throughout the 65-day incubation period, protecting it from intruders. When the young begin to hatch — their "yelping" calls can sometimes even be heard just before hatching commences — the mother quickly digs them out and carries them to the water in her mouth, as some other crocodilian species are known to do.
Young
The young are tiny replicas of adults, with a series of yellow bands around their bodies that serve as camouflage. Hatchlings gather into pods and are guarded by their mother and keep in contact with her through their "yelping" vocalizations. Young American alligators eat small fish, frogs, crayfish, and insects. They are preyed on by large fish, birds, raccoons, Florida panthers, and adult American alligators. Mother American alligators eventually become more aggressive towards their young, which encourages them to disperse. Young American alligators grow 3–8 in (7.6–20.3 cm) a year and reach adulthood at 6 ft (1.8 m).
Parasites
American alligators are commonly infected with parasites. In a 2016 Texas study, 100% of the specimens collected were infected with parasites, and by at least 20 different species of parasites, including lung pentastomids, gastric nematodes, intestinal helminths. When compared to American alligators from different states there was no significant difference in prevalence.
Nutria were introduced into coastal marshes from South America in the mid-20th century, and their population has since exploded into the millions. They cause serious damage to coastal marshes and may dig burrows in levees. Hence, Louisiana has had a bounty to try to reduce nutria numbers. Large American alligators feed heavily on nutria, so American alligators may not only control nutria populations in Louisiana, but also prevent them spreading east into the Everglades. Since hunting and trapping preferentially take the large American alligators that are the most important in eating nutria, some changes in harvesting may be needed to capitalize on their ability to control nutria.
Recently, a population of Burmese pythons became established in Everglades National Park. Substantial American alligator populations in the Everglades might be a contributing factor, as a competitor, in keeping the python populations low, preventing the spread of the species north. While events of predation by Burmese pythons on any sizable American alligators have been observed, no evidence of a net negative effect has been seen on overall American alligator populations.
American alligators play an important role in the restoration of the Everglades as biological indicators of restoration success. American alligators are highly sensitive to changes in the hydrology, salinity, and productivity of their ecosystems; all are factors that are expected to change with Everglades restoration. American alligators also may control the long-term vegetation dynamics in wetlands by reducing the population of small mammals, particularly nutria, which may otherwise overgraze marsh vegetation. In this way, the vital ecological service they provide may be important in reducing rates of coastal wetland losses in Louisiana. They may provide a protection service for water birds nesting on islands in freshwater wetlands. American alligators prevent predatory mammals from reaching island-based rookeries and in return eat spilled food and birds that fall from their nests. Wading birds appear to be attracted to areas with American alligators and have been known to nest at heavily trafficked tourist attractions with large numbers of American alligators, such as the St. Augustine Alligator Farm in St. Augustine, Florida.
Relationship with humans
Attacks on humans
Main article: List of fatal alligator attacks in the United States
American alligators are capable of killing humans, but fatal attacks are rare. Mistaken identity leading to an attack is always possible, especially in or near cloudy waters. American alligators are often less aggressive towards humans than larger crocodile species, a few of which (mainly the Nile and saltwater crocodiles) may prey on humans with some regularity. Alligator bites are serious injuries, due to the reptile's sheer bite force and risk of infection. Even with medical treatment, an American alligator bite may still result in a fatal infection.
As human populations increase, and as they build houses in low-lying areas, or fish or hunt near water, incidents are inevitable where humans intrude on American alligators and their habitats. Since 1948, 257 documented attacks on humans in Florida (about five incidents per year) have been reported, of which an estimated 23 resulted in death. Only nine fatal attacks occurred in the United States throughout the 1970s–1990s, but American alligators killed 12 people between 2001 and 2007. An additional report of alligator attacks showed a total of 376 injuries and 15 deaths recorded all from 1948 to 2004, leading this to an increase of the alligator population. In May 2006, American alligators killed three Floridians in less than a week. At least 28 fatal attacks by American alligators have occurred in the United States since 1970.
Wrestling
Main article: Alligator wrestling
Since the late 1880s, alligator wrestling has been a source of entertainment for some. Created by the Miccosukee and Seminole tribes prior to the arrival of Europeans, this tourism tradition remains popular despite criticism from animal-rights activists.
Farming
Main article: Alligator farm
Today, alligator farming is a large, growing industry in Georgia, Florida, Texas, and Louisiana. These states produce a combined annual total of some 45,000 alligator hides. Alligator hides bring good prices and hides in the 6- to 7-ft range have sold for $300 each. The market for alligator meat is growing, and about 300,000 pounds (140,000 kg) of meat are produced annually. According to the Florida Department of Agriculture and Consumer Services, raw alligator meat contains roughly 200 Calories (840 kJ) per 3-oz (85-g) portion, of which 27 Calories (130 kJ) come from fat.
The American alligator is the official state reptile of Florida, Louisiana, and Mississippi. Several organizations and products from Florida have been named after the animal.
"Gators" has been the nickname of the University of Florida's sports teams since 1911. In 1908, a printer made a spur-of-the-moment decision to print an alligator emblem on a shipment of the school's football pennants. The mascot stuck, and was made official in 1911, perhaps because the team captain's nickname was Gator. Allegheny College and San Francisco State University both have Gators as their mascots, as well.
The Gator Bowl is a college football game held in Jacksonville annually since 1946, with Gator Bowl Stadium hosting the event until the 1993 edition. The Gatornationals is a NHRA drag race held at the Gainesville Raceway in Gainesville since 1970.
When in water, the body is more swollen, and both shell and body are more translucent, lighter and brighter in colour, and less reflective, than when in air.
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Preface
[EDIT July 2021: specimens illustrated in this account which were supplied to Amgueddfa Cymru (the Natural History Museum, Wales) were sequenced by Ben Rowson who found no difference in the DNA of M. myosotis and M. denticulata and concluded that they were a single species; Myosotella myosotis. This has now been accepted by WoRMS; see www.marinespecies.org/aphia.php?p=taxdetails&id=139672 ]
The World Register of Marine Species (WoRMS) accepts Myosotella myosotis and M. denticulata as valid species, but those identified as such in Britain may be distinct ecotypes of a single species. A possibility, raised by Martins (2013), is that the true M. myosotis (Draparnaud, 1801) occurs in the Mediterranean and that both British shell forms are ecotypes of M. denticulata (Montagu, 1803). This account treats them separately as, whichever status is determined by planned DNA sequencing, they have distinct apertural sculptures associated with different habitats.
Because of its special habitat intermediate between terrestrial and marine, this species, and its Myosotella and Leucophytia relatives in the family Ellobiidae, are omitted from some identification guides, while variously appearing in others devoted solely to either terrestrial, marine or even freshwater mollusca.
Myosotella myosotis (Draparnaud, 1801)
Synonyms: Auricula myosotis Draparnoud, 1801; Ovatella myosotis (Draparnaud, 1801); Alexia myosotis (Draparnaud, 1801); Phytia myosotis (Draparnaud, 1801); Conovulus denticulatus var. myosotis in Forbes & Hanley (1853); Melampus myosotis in Jeffreys (1869);
Vernacular Probably also applied to M. denticulata: Mouse-eared Alexia, Mouse ear(ed) snail (English); Clust llygoden (Welsh); Evesnegl (Danish); Muizenoortje (Dutch); Ovatelle naine des vases (French); Stranddvärgsnäcka (Swedish); Mäuseöhrchen (German);
Applied to just this species/ecotype: Estuarine mouse-ear (English); Gewoon muizenoortje (Dutch);
Description
When in water, the body is more swollen, and both shell and body are more translucent, lighter and brighter in colour, and less reflective, than when in air 1Mm flic.kr/p/2drL1Pw . The following shell description is of specimens in air.
Shell
Juvenile shell usually less than 6.5mm high. Adult shell usually up to 8mm high and 3.5mm broad, exceptionally 10mm high and 5mm broad; ridge often within palatal (outer) lip 2Mm flic.kr/p/2exLLRc . Fusiform shell, width 45% to 50% of height 3Mm flic.kr/p/23Wmsve . Small spire with sharp apex; body whorl c. 73% of height of 7.5mm adult; 77% of 5.8mm juvenile; 80% of 4.4mm juvenile. Apex slightly twisted due to change from sinistral protoconch to dextral teleoconch. Shell-wall thin, opaque or slightly translucent, with a silky sheen when clean 2Mm flic.kr/p/2exLLRc . Up to 8 moderately convex whorls separated by distinct shallow sutures. On juveniles, the periostracum is drawn into a row of bristles below the sutures 4Mm flic.kr/p/23Wmsjc , but they are worn off over time; a few bristles may survive on adult shells. Earliest juveniles with three or fewer whorls lack periostracum and bristles; their shells are white-translucent with punctate spiral lines which may persist for a time as the shell grows 5Mm flic.kr/p/23Wms8v ; other later whorls may have them concealed under the periostracum. Very fine, closely spaced, costal lines sometimes visible on adults, especially on spire whorls 6Mm flic.kr/p/23Wms6g . Growth lines sometimes emphasised by change of shell colour 7Mm flic.kr/p/2drL14d . Usually no umbilicus except for an umbilicus-like slit in the apex caused by the change from the sinistral larval shell (protoconch) to a dextral shell 5Mm flic.kr/p/23Wms8v . Within the shell, when it reaches 2½ whorls, the columella and septa between the spire whorls are resorbed by the mantle, leaving an open space except for the columella and septum of the body whorl 8Mm flic.kr/p/23WmrU4 & 9Mm flic.kr/p/2drKZRQ . Aperture about 50% of adult shell height, 65% of 4.6mm juvenile; shaped like a narrow ear with a rounded base and a sharp adapical angle 3Mm flic.kr/p/23Wmsve & 10Mm flic.kr/p/QMuYsB . Palatal (outer) lip of juveniles (under c.6mm shell-height) is thin without protrusions (folds/teeth/denticles); lip sometimes weakly reflected 4Mm flic.kr/p/23Wmsjc on adults (over c.6mm high) often with a pale calcareous ridge within the aperture near the palatal rim. The ridge sometimes contains a single raised white denticle that is often only weakly developed. The columellar/parietal lip (inner lip of aperture) has two or three protrusions . The parietal lip consists of a wide glazed area on the body whorl. For a clear view of the features within the aperture, including, sometimes, a far-back palatal ridge formed at a previous pause in growth, the animal may need a prod with a small brush to make it withdraw, and the shell requires tilting at different angles 11Mm flic.kr/p/2drKZKC . There is no operculum 12Mm flic.kr/p/QMuYge . Exterior colour varies from pale yellowish brown to dark reddish brown or, sometimes, purplish brown 2Mm flic.kr/p/2exLLRc . Sometimes the shade of brown changes at growth lines 7Mm flic.kr/p/2drL14d , and, frequently, the spire is darker than the body whorl 3Mm flic.kr/p/23Wmsve . The protoconch and juvenile shell up to 1.4mm height are white 13Mm flic.kr/p/2drKZr1 , and are retained as a white apex on the adult 2Mm flic.kr/p/2exLLRc . On adults, the pale ridge within the aperture may show as a pale band on the exterior of the slightly translucent shell 11Mm flic.kr/p/2drKZKC . Shells on saltings are often coated with mud 14Mm flic.kr/p/QMuY3i . On dead stranded shells the periostracum often peels off and the colour bleaches to whitish 15Mm flic.kr/p/2drKZdL .
Body
Upper parts of head and body that are exposed by normal extension are various shades of grey including whitish-grey, steel-grey and brownish-grey to nearly black 6Mm flic.kr/p/23Wms6g & 16Mm flic.kr/p/QMuXMt , but rarely, if ever, pure white; colour on an individual varies with degree of extension and whether in air or water, and its intensity may increase with age. The colour is arranged in transverse bands across the dorsum 7Mm flic.kr/p/2drL14d , and as a mosaic of tessellating blotches on the sides of the head 16Mm flic.kr/p/QMuXMt . Sides of foot are greyish white or a paler grey than the dorsum 17Mm flic.kr/p/2drKYVb . Body parts normally concealed by the shell have less pigment and show the white of the internal oesophagus with flanking salivary glands, reproductive organs and retractor muscles when the body is extended to its maximum 7Mm flic.kr/p/2drL14d & 18Mm flic.kr/p/QMuXBi . When immersed, the body absorbs water and swells, the body colour becomes paler, and translucency of the shell and body increases 19Mm flic.kr/p/2drKYKG . The mantle sometimes projects a short way beyond the aperture rim of the palatal lip, but is not reflected onto it 20Mm flic.kr/p/2da1VEX . The parietal lip on the body whorl is a glaze formed by the mantle extending onto it. Within the shell, the mantle is very thin, semi-transparent and colourless apart from a faint, fine, pale-grey speckling. Where it roofs the mantle cavity 21Mm flic.kr/p/2da1VyK , it contains many haemolymph vessels which are occasionally visible through translucent juvenile shells 22Mm flic.kr/p/2drKYpG or on dissected mantle. The mantle cavity, which functions as a lung for respiration, is sealed off from the exterior by a thick, white or brownish-white, membranous mantle-collar which fits closely round the body as it extends or retracts 23Mm flic.kr/p/QMuSQV & 24Mm flic.kr/p/2drKYky . The collar has a pneumostome which, when in air, can be opened and closed 25Mm flic.kr/p/2exLJ2K for respiration and humidity control but, when immersed, does not effectively retain air or exclude water 26Mm flic.kr/p/2drKY7h . The rectum and part of the intestine, visible through translucent shells in water 22Mm flic.kr/p/2drKYpG , runs along the rear edge of the roof of the mantle cavity 21Mm flic.kr/p/2da1VyK to the anus which opens to the exterior in a folded part of the mantle-collar 27Mm flic.kr/p/2da1UTX in the adapical angle of the aperture close to the pneumostome. The head has two cephalic tentacles; nearly linear with a bluntly pointed tip (subulate) when dry, and conical and paler when swollen with water 16Mm flic.kr/p/QMuXMt . When not extended, they are contracted into crumpled stumps 28Mm flic.kr/p/2drKY1q , not retracted by inversion into the body. Both when extended or contracted, the tentacles widely diverge from their bases near the midline of the head 28Mm flic.kr/p/2drKY1q & 29Mm flic.kr/p/2da1UNr . The distal half of the tentacles, sometimes slightly bulbous, is brownish and contains sensory chemoreceptor cells (Wondrak, 1984) 29Mm flic.kr/p/2da1UNr . There is an internal black eye within the posteromesial base of each tentacle 29Mm flic.kr/p/2da1UNr & 30Mm flic.kr/p/2drKXUJ . Distinctness of eyes varies with intensity of body pigment and amount of flesh they are viewed through. The head in front of the tentacles forms a broad, slightly bilobed “muzzle” (Forbes & Hanley,1853) which can be variably configured, but not cylindrically to form a snout like that of many marine gastropods. When not feeding, it forms a shallow curve over the mouth in anterior view, and a steep anterior in profile view 28Mm flic.kr/p/2drKY1q . Near the anterior edge of the muzzle are two button-like, faintly-brownish grey pads (“fungiform bodies” of Wondrak, 1984) 28Mm flic.kr/p/2drKY1q which contain sensory cells. Ventrally, the mouth is protected by white outer-lip lobes. When feeding, the ventrally translucent-white muzzle is spread out flat on the substrate and the outer lips moved aside 31Mm flic.kr/p/SpE2GS to expose the mouth edged anteriorly by the rim of the red-brown jaw, and to allow the extension of the anterior of the radula covered in thousands of white teeth. When translucent, the muzzle may reveal dorsally the oral tube leading from the mouth to the buccal mass, and the oesophagus passing from it towards the stomach 18Mm flic.kr/p/QMuXBi . The stomach is a large sac, partly surrounded by digestive gland. On specimens removed from the shell, the transparent lateral surface showing the stomach contents is prominently visible on the left of the visceral mass 32Mm flic.kr/p/2exLHtF , and the gizzard-like girdle of strong, white, folded muscle fibre surrounding it may be seen if the visceral mass is opened 33Mm flic.kr/p/2da1Uvc & 34Mm flic.kr/p/2exLHiF . Unlike the spiral viscera of most gastropods, the viscera of M. myosotis forms a non-spiracular, approximate cone 35Mm flic.kr/p/2da1Uo8 to fill the spire after the columella and septa are resorbed 8Mm flic.kr/p/23WmrU4 & 9Mm flic.kr/p/2drKZRQ , with a small colourless tip formed in the shell's early pre-resorption stage. On weakly pigmented, translucent specimens (most often juveniles) the dumbbell-shaped, dorsal part of the nerve ring with two cerebral ganglia may be visible 24Mm flic.kr/p/2drKYky . The ring encircles the oesophagus. It and its ganglia that innervate organs on the head are the nearest approximation in gastropods to a centralised brain, but other ganglia distributed on nerve cords around the body innervate other organs 36Mm flic.kr/p/2exLH8v . The anterior edge of the translucent white sole is broad and almost straight, sometimes with a slightly indented middle, and tapers to a rounded posterior 25Mm flic.kr/p/2exLJ2K . M. myosotis is a protandrous hermaphrodite with a penis shaped as a stout cone with the vas deferens running up the centre. The penis is normally inverted within the body, and everted for mating by hydrostatic pressure of haemolymph from an opening on the right side of the head to the rear of the tentacle 37Mm flic.kr/p/2da1Uac .
Key identification features
Features 1 to 4, below, accord with Forbes & Hanley (1853) and Gittenberger (2004). The former aggregated M. myosotis sensu stricto with M. denticulata but “scrupulously kept apart their description.” Many currently used identification guides aggregate them and their features under M. myosotis sensu lato. Consequently, distribution maps on GBIF and NBN include many M. denticulata occurrences under “M. myosotis”, and the M. denticulata maps have under-representation of its occurrence. If making a differentiated record, 'sensu stricto' should be added to the record to enable it to be distinguished from lumped records.
To observe aperture sculpture the animal must be well withdrawn, and the shell tilted at different angles. Sometimes the outer (palatal) lip sculpture of an earlier growth stage is visible deep into the aperture and should be used if the sculpture on new growth has not yet developed. It is advisable to examine several specimens of different sizes from a site; sometimes both are present.
Myosotella myosotis
1. Live shell brown 2Mm flic.kr/p/2exLLRc (beachworn shells may be dull whitish 15Mm flic.kr/p/2drKZdL ). Usual adult height 6.5mm to 8mm, exceptionally 10mm .
2. Inner (columellar/parietal) lip has only 2 or 3 apertural protrusions 3Mm flic.kr/p/23Wmsve .
3. Outer (palatal) lip has a single apertural denticle or none 3Mm flic.kr/p/23Wmsve . Some have a pale apertural ridge running close to the lip.
4. Flesh colour of normally extended dorsal body is grey 1Mm flic.kr/p/2drL1Pw . Shade and intensity varies with age, extension and whether in air or water, but not pure white when adult.
5. Habitat: among vegetation, often under driftwood, on low salinity estuarine saltings 38Mm flic.kr/p/2exLH2t and Saltmarsh-grass sward by tidal rivers 39Mm flic.kr/p/2da1TMi a little above and below EHWS. Locally abundant. (May occur with M. denticulata under stones on/near saltings 40Mm flic.kr/p/2exLGc2 .)
Similar species/ecotype
Myosotella denticulata
(full account flic.kr/s/aHskQdY4cp)
1. Live shell brown (beachworn shells may be dull whitish). Usual adult height 3.5 mm to 7.5 mm, exceptionally 10 mm 45Mm flic.kr/p/QMuTi8 .
2. Inner (columellar/parietal) lip has 3 to 5 apertural protrusions 45Mm flic.kr/p/QMuTi8 .
3.Outer (palatal) lip has 2 to 7 (or more) apertural protrusions 45Mm flic.kr/p/QMuTi8 , sometimes set into a pale ridge which occasionally submerges them. [If no protrusions, check further back in aperture for protrusions on earlier lip position; may be visible from exterior through translucent shell , with or without connecting streaks.]
4. In its typical non-salting habitat, the flesh colour of normally extended dorsal body is white or very pale whitish grey, with darker grey tentacles 46Mm flic.kr/p/2da1NUc . But when it occurs in muddier conditions, it may be as dark as M. myosotis 47Mm flic.kr/p/QMuSYv .
5. Habitat: typically under slightly embedded stones at Extreme High Water Spring level and above (supralittoral) on sheltered coast without salting vegetation at fully marine salinity. Occasionally under stones on landward edge of Saltmarsh-grass sward by tidal rivers with low salinity 40Mm flic.kr/p/2exLGc2 .
Leucophytia bidentata (Montagu, 1808).
(Full account flic.kr/s/aHsmwhDvaL )
Features 1 to 4 conform with Montagu's original description and image.
1. Live shell slightly-translucent ivory-white; yellow viscera may show through spire 48Mm flic.kr/p/2da1NEe . Usual adult height to 5 mm, occasionally to 7 mm. Sutures shallower and whorls less rounded than on M. myosotis 49Mm flic.kr/p/2da21sH .
2. Inner (columellar/parietal) lip has 2 protrusions within the aperture; not more 49Mm flic.kr/p/2da21sH .
3. Outer (palatal) lip has no protrusions or rib (sometimes in a photo, a strong growth line might be mistaken for a rib 49Mm flic.kr/p/2da21sH ).
4. Flesh colour of normally extended dorsal body is amost pure white 48Mm flic.kr/p/2da1NEe , but when contracted into body-whorl colour saturation gives it a cream appearance.
5. Lives in deep, silty, rock crevices between High Water Neap level and Low Water Spring level. Also under stones embedded into soil-like substrate at Extreme High Water Spring level and a little above on sheltered coast where it is often with M. denticulata.
Habits and ecology
M. myosotis lives in the upper littoral fringe at, and a little below, the level of EHWS tides at the base of halophyte vascular plants on estuarine saltings 41Mm flic.kr/p/QMuUAP and in Saltmarsh-grass sward (Puccinellia maritima) along tidal rivers 39Mm flic.kr/p/2da1TMi & 42Mm flic.kr/p/2exLEJH . This zone can be extensive on large, flat, estuarine saltings 38Mm flic.kr/p/2exLH2t or as little as a metre wide on steep river banks 39Mm flic.kr/p/2da1TMi . At its upper limits, at or a little above EHWS, M. myosotis intermingles with some terrestrial invertebrates. It does not live in permanently submerged in pools, but can survive and be active for the short period of immersion (c. 1 hour) that occurs on 2 to 6 days per month. It lives semi-subterraneanly under driftwood 43Mm flic.kr/p/QMuTCB or large debris, or in the groundcover matt of vegetation and debris which, for moving through, its spindle shaped shell is well adapted. When moving, the foot is cushioned on a layer of watery mucus and the shell has a thicker layer between it and the substrate 17Mm flic.kr/p/2drKYVb . Under large pieces of wood or debris it favours the central darkest part while Assiminea grayana, if present, is found near the periphery. After immersion or during rain, it may be active in the open when its tentacles wave in the air to detect odours and, in the absence of anterior tentacles found on most pulmonates, its oral tube tests wet substances on the substrate (Wondrak, 1984). The two button-like, faintly-brownish grey pads (“fungiform bodies” of Wondrak, 1984) 28Mm flic.kr/p/2drKY1q near the anterior edge of the muzzle are also sensory.
It is a euryhaline species capable of surviving immersion in water from 0 p.p.t to full marine salinity or more, but individuals require time to adapt to changes in salinity and may become inactive/moribund when abruptly immersed in water they are unaccustomed to.
Respiration is of atmospheric air in the mantle cavity which is sealed by a white collar of thickened mantle 23Mm flic.kr/p/QMuSQV that firmly embraces the body but allows it to extend-from/retract-into the shell 24Mm flic.kr/p/2drKYky . A pneumostome (respiratory pore) in the collar 25Mm flic.kr/p/2exLJ2K can be opened for inhalation/exhalation of air or closed to seal the cavity against dehydration.The roof of the mantle cavity contains a network of haemolymph vessels 21Mm flic.kr/p/2da1VyK and is very thin, enabling oxygen from inhaled air to diffuse into the vessels and for carbon dioxide to leave with the exhaled air. When immersed, air escapes from the mantle cavity 26Mm flic.kr/p/2drKY7h and water enters 24Mm flic.kr/p/2drKYky . M. myosotis can survive immersion for at least three days with no access to atmospheric air. Long submersion with water in the mantle cavity is well tolerated if the water's oxygen content is high 22Mm flic.kr/p/2drKYpG (Seelmann, 1968, in Gittenberger, 2004).
When feeding, the muzzle is spread out on the substrate and the radula is extended 31Mm flic.kr/p/SpE2GS to gather, with the red jaw as a backstop, decaying vegetation, diatoms (Wiese & Richling, 2008) and sediment rich in organic material which are bound into food boli with mucus from the supra pedal gland brought to the mouth along a median groove. The boli travel along the oesophagus 36Mm flic.kr/p/2exLH8v to the stomach 32Mm flic.kr/p/2exLHtF & 34Mm flic.kr/p/2exLHiF where strong muscular contractions of the internally-folded gizzard triturate them and squeeze out semi-fluid nutrient which passes into the diverticula of the digestive gland for digestion. The residual mass is squeezed as faecal boli into the intestine by muscular contractions 34Mm flic.kr/p/2exLHiF and passes to and through the rectum 21Mm flic.kr/p/2da1VyK . Unlike marine prosobranch gastropods, which defecate into a mantle cavity that is cleared by water currents, M. myosotis has a rectum that opens to the exterior through an anus in the mantle collar, near to, but separate from, the pneumostome 27Mm flic.kr/p/2da1UTX so that faeces are expelled without fouling the respiratory mantle-cavity. The soft faeces, wet and loosely bound with mucus when fresh, dry to thin granular threads consisting mostly of fine mineral particles 44Mm flic.kr/p/2da1Pyi .
Reproduction: in NW Germany, copulation is in all months with peaks in April/May and August/September (Schultes,2014). Ova are laid when the temperature is above 15°C from late spring to late summer (Gittenberger, 2004). It is a protandrous hermaphrodite which changes its sexual function in the wild when 1½ to 2 years old, so younger, 1 to 1½ years, fully mature males mate with older, over 1½ years, females (Schultes, 2014) using the stout, conical penis everted from the side near the posterior of the right tentacle. Sometimes, a chain of three individuals mate, with the middle one acting as both male and female. Under optimal laboratory conditions, newly hatched animals can reach a shell length of 5 mm and start laying eggs in 8 weeks (Wiese & Richling, 2008). Each month of the breeding season, a female deposits 15 to 80 egg capsules (Schultes, 2014) in a small, yellow or white, frog-spawn-like mass (Morton, 1954 and Gittenberger, 2004). Each ovoid capsule contains a single ovum. The cases are attached to each other in a loosely convoluted chain by a filament (chalaziform process) at each end. The closely packed cases with intervening clear fluid are contained in a tough binding membrane which is attached to stones, plant stems, wood etc in moist situations (Morton, 1954) or in soil cavities in clusters from several females (Schultes, 2014).There is a larval veliger stage, with sinistral shell, which is passed entirely within the ovum (Morton, 1954). At 10 °C to 20 °C and 18 p.p.t. salinity, crawling juveniles emerge after about two and a half weeks. At less favourable salinities, less than 18 p.p.t or more than 54p.p.t., development takes several weeks longer (Gittenberger, 2004), though the eggs are moist but not immersed for most of the time. Juveniles in northern Germany hatch after 3-7 weeks, and 12-15 days in France (Schultes, 2014). Individuals can live to 3 or 4 years of age (Wiese & Richling, 2008).
Distribution and status
Europe from Orkney, Scotland and southern Denmark to Mediterranean and Azores; majority of records are from Britain and Ireland. Locally abundant in suitable habitat in Britain. Occurs in German Baltic saltmarshes, but habitat threatened in Germany (Wiese & Richling, 2008). Assumed to have been introduced to temperate coasts of Australia and North America (Atlantic and Pacific) GBIF map www.gbif.org/species/2297460 , but comparison of soft part morphology suggests that many different species have been aggregated because of similar shell morphology (Martins, 2013).
Widespread around Britain but non-estuarine records are likely to be the species/ecotype M. denticulata NBN map
species.nbnatlas.org/species/NHMSYS0001702112#tab_mapView
Irish distribution, National biodiversity data centre, in Mollusc Ireland: www.habitas.org.uk/molluscireland/species.asp?ID=121
Acknowledgements
I gratefully thank Ben Rowson of the National Museum of Wales/ Amgueddfa Cymru for his help with the account, but any errors or omissions are mine.
Links and references
Anderson, R. MolluscIreland, accessed January 2019. www.habitas.org.uk/molluscireland/species.asp?ID=121
Forbes, E. & Hanley S. 1849-53. A history of the British mollusca and their shells. vol. 4 (1853), 190 – 197 & plate CXXV. London, van Voorst. (AsConovulus denticulatus var. myosotis.)
Free pdf at archive.org/details/historyofbritish04forbe/page/190
plate at archive.org/details/historyofbritish04forbe/page/n565
Fretter, V. & Peake, J. 1975. Pulmonates functional anatomy and physiology. Vol.1. London. Academic Press.
Gittenberger, E. et al. 2004. De Nederlandse zoetwatermollusken. Leiden, Netherlands, Nationaal Natuurhistorisch Museum Naturalis.
Heller J. 2015. Marine Ancestors of most Land Snails: Pulmonates. In: Sea Snails. Springer, Cham. link.springer.com/chapter/10.1007%2F978-3-319-15452-7_10
Jeffreys, J.G. 1862-69. British conchology. vol. 5 (1869). London, van Voorst. (As Melampus myosotis (including var. ringens = Myosotella denticulata); Free pdf at archive.org/stream/britishconcholog05jeffr#page/106/mode/2up . Use slide at base of page to select pp.106-109.)
Martins, A.M. de F. 1996. Anatomy and systematics of the western Atlantic Ellobidae (Gastropoda: Pulmonata). Malacologia 37(2): 163 – 332.
www.biodiversitylibrary.org/page/13113594#page/179/mode/1up
Martins, A.M. de F. & Mendes, A.R.M. 2013. Do cosmopolitans speciate? Anatomical diversity of Myosotella in Azores. Centro de Investigação em Biodiversidade e Recursos Genéticos. Ponta Delgada, Açores, Portugal. Poster for World Congress of Malacology 2013 in pdf: www.researchgate.net/publication/264339925_Do_cosmopolita... .
Montagu, G. 1808. Supplement to: 1803 Testacea Britannica, or, Natural history of British shells, marine, land, and fresh-water, including the most minute : systematically arranged and embellished with figures. London, J. White.
Description of Leucophtia bidentata as Voluta bidentata pp. 100-101.
www.biodiversitylibrary.org/page/24430722#page/806/mode/1up
Plate 30, fig.2:
www.biodiversitylibrary.org/page/24430722#page/917/mode/1up
Morton, J. E. 1955. The functional morphology of the British Ellobiidae (Gastropoda Pulmonata) with special reference to the digestive and reproductive systems. Phil. Trans. R. Soc. Ser. B .
239, No. 661: 89-160 www.jstor.org/stable/92507
Schultes, F.W. 2014. Species summary for Ovatella myosotis (Draparnoud, 1801). AnimalBase. SUB Göttingen. www.animalbase.uni-goettingen.de/zooweb/servlet/AnimalBas... Accessed January 2019.
Watson, H. I943. Notes on a list of the British non-marine Mollusca. J. Conch. 22: 13 - 22.
Wiese, V. & Richling, I. 2008. Das Mäuseöhrchen Myosotella myosotis (Draparnaud 1801). Arbeitskreis Mollusken NRW.
www.mollusken-nrw.de/weichtier_des_jahres/weichtier2008.htm
Wondrak, G. 1984. Ultrastructure of the sensory epithelia of oral tube, fungiform sensory bodies, and terminal knobs of tentacles of Ovatella
myosotis. Draparnaud (Archaeopulmonata, Gastropoda) J. Morphol. 181: 333-347 .
onlinelibrary.wiley.com/doi/pdf/10.1002/jmor.1051810307
Current taxonomy:
www.marinespecies.org/aphia.php?p=taxdetails&id=139673
Glossary
adapical angle = angle at which outer lip meets body-whorl.
boli = (sing. bolus) small rounded masses, especially of triturated food material.
cerebral = to do with integration of sensory and neural functions to initiate and coordinate body activity.
chalaziform = resembling the two spiral bands (chalazae) in a bird's egg that attach the yolk to opposite ends of the lining membrane.
columella = solid or hollow axial “little column” around which gastropod shell spirals; hidden inside shell, except on final whorl next to lower part of inner lip of aperture where hollow ones may end in an umbilicus or siphonal canal.
columellar = (adj.) of or near central axis of coiled gastropod.
columellar lip = lower (abapical) part of inner lip of aperture.
costa (pl. costae) = rib running across a whorl of a gastropod shell at approximately right-angles to direction of coiling and any spiral striae.
costal (adj.) = of, or arranged like, costae.
dextral = (of gastropod shell) in apertural view with spire uppermost, the aperture is on the right. Most gastropod species adults have dextral shells.
distal = away from centre of body or from point of attachment.
diverticula = (for digestion) blind ended tubules in the digestive gland that receive nutrients for digestion.
EHWS = extreme high water spring tide.
euryhaline = able to tolerate a wide variation in salinty.
fusiform = slender, spindle-shaped, tapering almost equally towards both ends.
ganglia = (sing. ganglion) knots on a nerve cord containing sensory cell bodies that conduct impulses to (innervate) organs of the body.
haemolymph = circulating fluid in molluscs that carries nutrients, waste and hormones. Analagous to vertebrate blood, but most molluscs have copper-based haemocyanin in it instead of red haemoglobin to carry oxygen. It may be tinged blue when oxygenated; colourless when depleted of oxygen.
halophyte = plant tolerant of saline soil and periodic tidal immersion, usually on saltmarshes, estuarine shores and sides of tidal rivers.
mantle = sheet of tissue that secretes the shell, covers the viscera and forms a cavity in gastropods. In terrestrial gastropods ('pulmonates') the cavity roof contains a network of haemolymph ('blood') vessels enabling the cavity to act like a lung.
mesial = on or facing towards the midline of the body.
operculum = plate of horny conchiolin, rarely calcareous, used to close shell aperture of prosobranch gastropods.
palatal lip = outer lip of gastropod aperture.
parietal lip ( or parietal wall) = upper part of inner side of gastropod aperture, often lacking clear lip structure with just a glaze on side of whorl adapically of columellar lip.
periostracum = thin horny layer of proteinaceous material often coating shells.
posteromesial = at the rear facing towards the midline of the body.
prosobranch = member of Prosobranchia, one of three subclasses into which the class Gastropoda (slugs and snails) was divided during the 20th Century (other two were Pulmonata and Opisthobranchia). This classification is no longer used by scientists, but prosobranch is a useful informal term to signify (mainly marine) snails breathing with a ctenidium (comblike gill inside mantle cavity), an operculum, and a shell which can accommodate the whole body.
protandrous hermaphrodite = each individual starts mature life as a functioning male, later changing to female function.
protoconch = apical whorls produced during embryonic and larval stages of gastropod; often different in form from other whorls (teleoconch).
protrusions = teeth, denticles, folds, lamellae or cogs (terms used by various authors).
punctate = with pinprick-like depressions.
resorb = absorb what was previously secreted; break it down into component materials and disperse into the circulation.
resorption = the process of absorbing what was previously secreted by breaking it down into component materials and dispersal into the circulation.
salting = area of salt tolerant vascular plants rooted in sediment between mean high water mark (MHW) and extreme high water of spring tides (EHWS). [Preferred synonym for “saltmarsh” as much of salting not marshy.]
septa = plural of septum; internal partition separating two chambers/ shell-whorls of a gastropod.
septum = internal partition separating two chambers/ shell-whorls of a gastropod.
sinistral = (of gastropod shell) in apertural view with spire uppermost, the aperture is on the left. Most gastropod species adults have dextral shells.
subsutural = close below the suture when shell positioned with apex uppermost.
subulate = slender and tapering to a point like onion leaf or awl.
suture = groove or line where whorls of gastropod shell adjoin.
teleoconch = entire gastropod shell other than the apical, embryonic & larval stage protoconch.
triturate = reduce to small particles.
vascular plants = plants that have vascular tissues to transport water and nutrients through the plant. Include all seed-bearing plants, ferns and horsetails. Usually terrestrial or in freshwater or brackish water; a few, such as Zostera, live in fully marine salinity water.
The American alligator (Alligator mississippiensis), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the genus Alligator, and is larger than the only other living alligator species, the Chinese alligator.
Adult male American alligators measure 3.4 to 4.6 m (11.2 to 15.1 ft) in length, and can weigh up to 500 kg (1,100 lb), with unverified sizes of up to 5.84 m (19.2 ft) and weights of 1,000 kg (2,200 lb) making it the second largest member of the family Alligatoridae, after the black caiman. Females are smaller, measuring 2.6 to 3 m (8.5 to 9.8 ft) in length. The American alligator inhabits subtropical and tropical freshwater wetlands, such as marshes and cypress swamps, from southern Texas to North Carolina. It is distinguished from the sympatric American crocodile by its broader snout, with overlapping jaws and darker coloration, and is less tolerant of saltwater but more tolerant of cooler climates than the American crocodile, which is found only in tropical and warm subtropical climates.
American alligators are apex predators and consume fish, amphibians, reptiles, birds, and mammals. Hatchlings feed mostly on invertebrates. They play an important role as ecosystem engineers in wetland ecosystems through the creation of alligator holes, which provide both wet and dry habitats for other organisms. Throughout the year (in particular during the breeding season), American alligators bellow to declare territory, and locate suitable mates. Male American alligators use infrasound to attract females. Eggs are laid in a nest of vegetation, sticks, leaves, and mud in a sheltered spot in or near the water. Young are born with yellow bands around their bodies and are protected by their mother for up to one year.[
The conservation status of the American alligator is listed as Least Concern by the International Union for Conservation of Nature. Historically, hunting had decimated their population, and the American alligator was listed as an endangered species by the Endangered Species Act of 1973. Subsequent conservation efforts have allowed their numbers to increase and the species was removed from endangered status in 1987. The species is the official state reptile of three states: Florida, Louisiana, and Mississippi.
Taxonomy
The American alligator was first classified by French zoologist François Marie Daudin as Crocodilus mississipiensis in 1801. In 1807, Georges Cuvier created the genus Alligator; the American alligator and the Chinese alligator are the only extant species in the genus. They are grouped in the family Alligatoridae with the caimans. The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the American alligator than to either the Nile crocodile (Crocodylus niloticus) or the gharial (Gavialis gangeticus).
Phylogeny
Members of this superfamily first arose in the late Cretaceous, about 100–65 million years ago (Mya). Leidyosuchus of Alberta is the earliest known fossil, from the Campanian era 83 to 72 Mya. Fossil alligatoroids have been found throughout Eurasia, because bridges across both the North Atlantic and the Bering Strait connected North America to Eurasia about 66 to 23 Mya.
Alligators and caimans split in North America during the late Cretaceous, and the caimans reached South America by the Paleogene, before the closure of the Isthmus of Panama during the Neogene period, from about 23 to 2.58 Mya. The Chinese alligator likely descended from a lineage that crossed the Bering land bridge during the Neogene. Fossils identical to the existing American alligator are found throughout the Pleistocene, from 2.5 million to 11.7 thousand years ago. In 2016, a Miocene (about 23 to 5.3 Mya) fossil skull of an alligator was found at Marion County, Florida. Unlike the other extinct alligator species of the same genus, the fossil skull was virtually indistinguishable from that of the modern American alligator. This alligator and the American alligator are now considered to be sister taxa, meaning that the A. mississippiensis lineage has existed in North America for over 8 million years.
The alligator's full mitochondrial genome was sequenced in the 1990s, and it suggests the animal evolved at a rate similar to mammals and greater than birds and most cold-blooded vertebrates. However, the full genome, published in 2014, suggests that the alligator evolved much more slowly than mammals and birds.
Characteristics
American alligator skull
Domestic American alligators range from long and slender to short and robust, possibly in response to variations in factors such as growth rate, diet, and climate.
Size
The American alligator is a relatively large species of crocodilian. On average, it is the largest species in the family Alligatoridae, with only the black caiman being possibly larger. Weight varies considerably depending on length, age, health, season, and available food sources. Similar to many other reptiles that range expansively into temperate zones, American alligators from the northern end of their range, such as southern Arkansas, Alabama, and northern North Carolina, tend to reach smaller sizes. Large adult American alligators tend to be relatively robust and bulky compared to other similar-length crocodilians; for example, captive males measuring 3 to 4 m (9 ft 10 in to 13 ft 1 in) were found to weigh 200 to 350 kg (440 to 770 lb), although captive specimens may outweigh wild specimens due to lack of hunting behavior and other stressors.
Large male American alligators reach an expected maximum size up to 4.6 m (15 ft 1 in) in length and weighing up to 500 kg (1,100 lb), while females reach an expected maximum of 3 m (9 ft 10 in). However, the largest free-ranging female had a total length of 3.22 m (10 ft 7 in) and weighed 170 kg (370 lb). On rare occasions, a large, old male may grow to an even greater length.
Largest
During the 19th and 20th centuries, larger males reaching 5 to 6 m (16 ft 5 in to 19 ft 8 in) were reported. The largest reported individual size was a male killed in 1890 on Marsh Island, Louisiana, and reportedly measured at 5.84 m (19 ft 2 in) in length, but no voucher specimen was available, since the American alligator was left on a muddy bank after having been measured due to having been too massive to relocate. If the size of this animal was correct, it would have weighed about 1,000 kg (2,200 lb). In Arkansas, a man killed an American alligator that was 4.04 m (13 ft 3 in) and 626 kg (1,380 lb). The largest American alligator ever killed in Florida was 5.31 m (17 ft 5 in), as reported by the Everglades National Park, although this record is unverified. The largest American alligator scientifically verified in Florida for the period from 1977 to 1993 was reportedly 4.23 m (13 ft 11 in) and weighed 473 kg (1,043 lb), although another specimen (size estimated from skull) may have measured 4.54 m (14 ft 11 in). A specimen that was 4.5 m (14 ft 9 in) long and weighed 458.8 kg (1,011.5 lb) is the largest American alligator killed in Alabama and has been declared the SCI world record in 2014.
Average
American alligators do not normally reach such extreme sizes. In mature males, most specimens grow up to about 3.4 m (11 ft 2 in) in length, and weigh up to 360 kg (790 lb), while in females, the mature size is normally around 2.6 m (8 ft 6 in), with a body weight up to 91 kg (201 lb). In Newnans Lake, Florida, adult males averaged 73.2 kg (161 lb) in weight and 2.47 m (8 ft 1 in) in length, while adult females averaged 55.1 kg (121 lb) and measured 2.22 m (7 ft 3 in). In Lake Griffin State Park, Florida, adults weighed on average 57.9 kg (128 lb). Weight at sexual maturity per one study was stated as averaging 30 kg (66 lb) while adult weight was claimed as 160 kg (350 lb).
Relation to age
There is a common belief stated throughout reptilian literature that crocodilians, including the American alligator, exhibit indeterminate growth, meaning the animal continues to grow for the duration of its life. However, these claims are largely based on assumptions and observations of juvenile and young adult crocodilians, and recent studies are beginning to contradict this claim. For example, one long-term mark-recapture study (1979–2015) done at the Tom Yawkey Wildlife Center in South Carolina found evidence to support patterns of determinate growth, with growth ceasing upon reaching a certain age (43 years for males and 31 years for females).
Sexual dimorphism
While noticeable in very mature specimens, the sexual dimorphism in size of the American alligator is relatively modest among crocodilians. For contrast, the sexual dimorphism of saltwater crocodiles is much more extreme, with mature males nearly twice as long as and at least four times as heavy as female saltwater crocodiles. Given that female American alligators have relatively higher survival rates at an early age and a large percentage of given populations consists of immature or young breeding American alligators, relatively few large mature males of the expected mature length of 3.4 m (11 ft 2 in) or more are typically seen.
Color
Dorsally, adult American alligators may be olive, brown, gray, or black. However, they are on average one of the most darkly colored modern crocodilians (although other alligatorid family members are also fairly dark), and can be reliably be distinguished by color via their more blackish dorsal scales against crocodiles. Meanwhile, their undersides are cream-colored. Some American alligators are missing or have an inhibited gene for melanin, which makes them albino. These American alligators are extremely rare and almost impossible to find in the wild. They could only survive in captivity, as they are very vulnerable to the sun and predators.
Jaws, teeth, and snout
American alligators have 74–80 teeth. As they grow and develop, the morphology of their teeth and jaws change significantly. Juveniles have small, needle-like teeth that become much more robust and narrow snouts that become broader as the individuals develop. These morphological changes correspond to shifts in the American alligators' diets, from smaller prey items such as fish and insects to larger prey items such as turtles, birds, and other large vertebrates. American alligators have broad snouts, especially in captive individuals. When the jaws are closed, the edges of the upper jaws cover the lower teeth, which fit into the jaws' hollows. Like the spectacled caiman, this species has a bony nasal ridge, though it is less prominent. American alligators are often mistaken for a similar animal: the American crocodile. An easy characteristic to distinguish the two is the fourth tooth. Whenever an American alligator's mouth is closed, the fourth tooth is no longer visible. It is enclosed in a pocket in the upper jaw.
Bite
Adult American alligators held the record as having the strongest laboratory-measured bite of any living animal, measured at up to 13,172 N (1,343.2 kgf; 2,961 lbf). This experiment had not been, at the time of the paper published, replicated in any other crocodilians, and the same laboratory was able to measure a greater bite force of 16,414 N (1,673.8 kgf; 3,690 lbf) in saltwater crocodiles; notwithstanding this very high biting force, the muscles opening the American alligator's jaw are quite weak, and the jaws can be held closed by hand or tape when an American alligator is captured. No significant difference is noted between the bite forces of male and female American alligators of equal size. Another study noted that as the American alligator increases in size, the force of its bite also increases.
Movement
When on land, an American alligator moves either by sprawling or walking, the latter involving the reptile lifting its belly off the ground. The sprawling of American alligators and other crocodylians is not similar to that of salamanders and lizards, being similar to walking. Therefore, the two forms of land locomotion can be termed the "low walk" and the "high walk". Unlike most other land vertebrates, American alligators increase their speed through the distal rather than proximal ends of their limbs. In the water, American alligators swim like fish, moving their pelvic regions and tails from side to side. During respiration, air flow is unidirectional, looping through the lungs during inhalation and exhalation; the American alligator's abdominal muscles can alter the position of the lungs within the torso, thus shifting the center of buoyancy, which allows the American alligator to dive, rise, and roll within the water.
Distribution
American alligators, being native both to the Nearctic and Neotropical realms, are found in the wild in the Southeastern United States, from the Lowcountry in South Carolina, south to Everglades National Park in Florida, and west to the southeastern region of Texas. They are found in parts of North Carolina, South Carolina, Georgia, Florida, Louisiana, Alabama, Mississippi, Arkansas, Oklahoma and Texas. Some of these locations appear to be relatively recent introductions, with often small but reproductive populations. Louisiana has the largest American alligator population of any U.S. state. In the future, possible American alligator populations may be found in areas of Mexico adjacent to the Texas border. American alligators have been naturally expanding their range into Tennessee, and have established a small population in the southwestern part of that state via inland waterways, according to the state's wildlife agency. They have been extirpated from Virginia, and occasional vagrants from North Carolina wander into the Great Dismal Swamp.
Conservation status
American alligators are currently listed as least concern by the IUCN Red List, even though from the 1800s to the mid-1900s, they were being hunted and poached by humans unsustainably.
Historically, hunting and habitat loss have severely affected American alligator populations throughout their range, and whether the species would survive was in doubt. In 1967, the American alligator was listed as an endangered species (under a law that was the precursor Endangered Species Act of 1973), since it was believed to be in danger of extinction throughout all or a significant portion of its range.
Both the United States Fish and Wildlife Service (USFWS) and state wildlife agencies in the South contributed to the American alligator's recovery. Protection under the Endangered Species Act allowed the species to recuperate in many areas where it had been depleted. States began monitoring their American alligator populations to ensure that they would continue to grow. In 1987, the USFWS removed the animal from the endangered species list, as it was considered to be fully recovered. The USFWS still regulates the legal trade in American alligators and their products to protect still endangered crocodilians that may be passed off as American alligators during trafficking.
American alligators are listed under Appendix II of the Convention on International Trade in Endangered Species (CITES) meaning that international trade in the species (including parts and derivatives) is regulated.
Habitat
They inhabit swamps, streams, rivers, ponds, and lakes. A lone American alligator was spotted for over 10 years living in a river north of Atlanta, Georgia. Females and juveniles are also found in Carolina Bays and other seasonal wetlands. While they prefer fresh water, American alligators may sometimes wander into brackish water, but are less tolerant of salt water than American crocodiles, as the salt glands on their tongues do not function. One study of American alligators in north-central Florida found the males preferred open lake water during the spring, while females used both swampy and open-water areas. During summer, males still preferred open water, while females remained in the swamps to construct their nests and lay their eggs. Both sexes may den underneath banks or clumps of trees during the winter.
In some areas of their range, American alligators are an unusual example of urban wildlife; golf courses are often favored by the species due to an abundance of water and a frequent supply of prey animals such as fish and birds.
Cold tolerance
American alligators are less vulnerable to cold than American crocodiles. Unlike an American crocodile, which would immediately succumb to the cold and drown in water at 45 °F (7 °C) or less, an American alligator can survive in such temperatures for some time without displaying any signs of discomfort. This adaptiveness is thought to be why American alligators are widespread further north than the American crocodile. In fact, the American alligator is found farther from the equator and is more equipped to handle cooler conditions than any other crocodilian. When the water begins to freeze, American alligators go into a period of brumation; they stick their snouts through the surface, which allows them to breathe above the ice, and they can remain in this state for several days.
Ecology and behavior
Basking
American alligators primarily bask on shore, but also climb into and perch on tree limbs to bask if no shoreline is available. This is not often seen, since if disturbed, they quickly retreat back into the water by jumping from their perch.
Holes
American alligators modify wetland habitats, most dramatically in flat areas such as the Everglades, by constructing small ponds known as alligator holes. This behavior has qualified the American alligator to be considered a keystone species. Alligator holes retain water during the dry season and provide a refuge for aquatic organisms, which survive the dry season by seeking refuge in alligator holes, so are a source of future populations. The construction of nests along the periphery of alligator holes, as well as a buildup of soils during the excavation process, provides drier areas for other reptiles to nest and a place for plants that are intolerant of inundation to colonize. Alligator holes are an oasis during the Everglades dry season, so are consequently important foraging sites for other organisms. In the limestone depressions of cypress swamps, alligator holes tend to be large and deep, while those in marl prairies and rocky glades are usually small and shallow, and those in peat depressions of ridge and slough wetlands are more variable.
Bite and mastication
The teeth of the American alligator are designed to grip prey, but cannot rip or chew flesh like teeth of some other predators (such as canids and felids), and depend on their gizzard, instead, to masticate their food. The American alligator is capable of biting through a turtle's shell or a moderately sized mammal bone.
Tool use
American alligators have been documented using lures to hunt prey such as birds. This means they are among the first reptiles recorded to use tools. By balancing sticks and branches on their heads, American alligators are able to lure birds looking for suitable nesting material to kill and consume. This strategy, which is shared by the mugger crocodile, is particularly effective during the nesting season, in which birds are more likely to gather appropriate nesting materials. This strategy has been documented in two Florida zoos occurring multiple times a day in peak nesting season and in some parks in Louisiana. The use of tools was documented primarily during the peak rookery season when birds were primarily looking for sticks.
Aquatic vs terrestrial
Fish and other aquatic prey taken in the water or at the water's edge form the major part of American alligator's diet and may be eaten at any time of the day or night. Adult American alligators also spend considerable time hunting on land, up to 160 feet (50 m) from water, ambushing terrestrial animals on trailsides and road shoulders. Usually, terrestrial hunting occurs on nights with warm temperatures. When hunting terrestrial prey, American alligators may also ambush them from the edge of the water by grabbing them and pulling the prey into the water, the preferred method of predation of larger crocodiles.
An American alligator in the process of consuming a box turtle
Additionally, American alligators have recently been filmed and documented killing and eating sharks and rays; four incidents documented indicated that bonnetheads, lemon sharks, Atlantic stingrays, and nurse sharks are components of the animal's diet. Sharks are also known to prey on American alligators, in turn, indicating that encounters between the two predators are common.
Common
American alligators are considered an apex predator throughout their range. They are opportunists and their diet is determined largely by both their size and age and the size and availability of prey. Most American alligators eat a wide variety of animals, including invertebrates, fish, birds, turtles, snakes, amphibians, and mammals. Hatchlings mostly feed on invertebrates such as insects, insect larvae, snails, spiders, and worms, as well as small fish and frogs. As they grow, American alligators gradually expand to larger prey. Once an American alligator reaches full size and power in adulthood, any animal living in the water or coming to the water to drink is potential prey. Most animals captured by American alligators are considerably smaller than itself. A few examples of animals consumed are largemouth bass, spotted gar, freshwater pearl mussels, American green tree frogs, yellow mud turtles, cottonmouths, common moorhens, and feral wild boars. Stomach contents show, among native mammals, muskrats and raccoons are some of the most commonly eaten species. In Louisiana, where introduced nutria are common, they are perhaps the most regular prey for adult American alligators, although only larger adults commonly eat this species. It has also been reported that large American alligators prey on medium-sized American alligators, which had preyed on hatchlings and smaller juveniles.
If an American alligator's primary food resource is not available, it will sometimes feed on carrion and non-prey items such as rocks and artificial objects, like bottle caps. These items help the American alligator in the process of digestion by crushing up the meat and bones of animals, especially animals with shells.
Large animals
Other animals may occasionally be eaten, even large deer or feral wild boars, but these are not normally part of the diet. American alligators occasionally prey on large mammals, such as deer, but usually do so when fish and smaller prey levels go down. Rarely, American alligators have been observed killing and eating bobcats, but such events are not common and have little effect on bobcat populations. Although American alligators have been listed as predators of the Nilgai and the West Indian manatees, very little evidence exists of such predation. In the 2000s, when invasive Burmese pythons first occupied the Everglades, American alligators have been recorded preying on sizable snakes, possibly controlling populations and preventing the invasive species from spreading northwards. However, the python is also known to occasionally prey on alligators, a form of both competition and predation. American alligator predation on Florida panthers is rare, but has been documented. Such incidents usually involve a panther trying to cross a waterway or coming down to a swamp or river to get a drink. American alligator predation on American black bears has also been recorded.
Domestic animals
Occasionally, domestic animals, including dogs, cats, and calves, are taken as available, but are secondary to wild and feral prey. Other prey, including snakes, lizards, and various invertebrates, are eaten occasionally by adults.
Birds
Water birds, such as herons, egrets, storks, waterfowl and large dabbling rails such as gallinules or coots, are taken when possible. Occasionally, unwary adult birds are grabbed and eaten by American alligators, but most predation on bird species occurs with unsteady fledgling birds in late summer, as fledgling birds attempt to make their first flights near the water's edge.
Fruit
In 2013, American alligators and other crocodilians were reported to also eat fruit. Such behavior has been witnessed, as well as documented from stomach contents, with the American alligators eating such fruit as wild grapes, elderberries, and citrus fruits directly from the trees. Thirty-four families and 46 genera of plants were represented among seeds and fruits found in the stomach contents of alligators. The discovery of this unexpected part of the American alligator diet further reveals that they may be responsible for spreading seeds from the fruit they consume across their habitat.
Cooperative hunting
Additionally, American alligators engage in what seems to be cooperative hunting. One observation of cooperative hunting techniques was where there are pushing American alligators and catching American alligators and they were observed taking turns in each position. Another observation said that about 60 American alligators gathered in an area and would form a semicircle with about half of them and would push the fish closer to the bank. Once one of the American alligators caught a fish another one would enter into its spot, and it would take the fish to the resting area. This was reported to have occurred two days in a row.
In Florida and East Texas
The diet of adult American alligators from central Florida lakes is dominated by fish, but the species is highly opportunistically based upon local availability. In Lake Griffin, fish made up 54% of the diet by weight, with catfish being most commonly consumed, while in Lake Apopka, fish made up 90% of the food and mostly shad were taken; in Lake Woodruff, the diet was 84% fish and largely consists of bass and sunfish. Unusually in these regions, reptiles and amphibians were the most important nonpiscivore prey, mostly turtles and water snakes. In southern Louisiana, crustaceans (largely crawfish and crabs) were found to be present in the southeastern American alligators, but largely absent in southwestern American alligators, which consumed a relatively high proportion of reptiles, although fish were the most recorded prey for adults, and adult males consumed a large portion of mammals.
In East Texas, diets were diverse and adult American alligators took mammals, reptiles, amphibians, and invertebrates (e.g. snails) in often equal measure as they did fish.
Vocalizations
Mechanism
An American alligator is able to abduct and adduct the vocal folds of its larynx, but not to elongate or shorten them; yet in spite of this, it can modulate fundamental frequency very well. Their vocal folds consists of epithelium, lamina propria and muscle. Sounds ranged from 50 to 1200 Hz. In one experiment conducted on the larynx, the fundamental frequency depended on both the glottal gap and stiffness of the larynx tissues. As the frequency increases, there's high tension and large strains. The fundamental frequency has been influenced by the glottal gap size and subglottal pressure and when the phonation threshold pressure has been exceeded, there will be vocal fold vibration.
Calls
Crocodilians are the most vocal of all non-avian reptiles and have a variety of different calls depending on the age, size, and sex of the animal. The American alligator can perform specific vocalizations to declare territory, signal distress, threaten competitors, and locate suitable mates. Juveniles can perform a high-pitched hatchling call (a "yelping" trait common to many crocodilian species' hatchling young)[118] to alert their mothers when they are ready to emerge from the nest. Juveniles also make a distress call to alert their mothers if they are being threatened. Adult American alligators can growl, hiss, or cough to threaten others and declare territory.
Bellowing
Both males and females bellow loudly by sucking air into their lungs and blowing it out in intermittent, deep-toned roars to attract mates and declare territory. Males are known to use infrasound during mating bellows. Bellowing is performed in a "head oblique, tail arched" posture. Infrasonic waves from a bellowing male can cause the surface of the water directly over and to either side of his back to literally "sprinkle", in what is commonly called the "water dance". Large bellowing "choruses" of American alligators during the breeding season are commonly initiated by females and perpetuated by males. Observers of large bellowing choruses have noted they are often felt more than they are heard due to the intense infrasound emitted by males. American alligators bellow in B flat (specifically "B♭1", defined as an audio frequency of 58.27 Hz), and bellowing choruses can be induced by tuba players, sonic booms, and large aircraft.
Lifespan
American alligators typically live to the age of 50, and possibly over 70 years old. Males reach sexual maturity at around 11.6 years, and females at around 15.8 years. Although it was originally thought that American alligators never stop growing, studies have now found that males stop growing at around the age of 43 years, and females stop growing at around the age of 31 years.
Reproduction
Breeding season
The breeding season begins in the spring. On spring nights, American alligators gather in large numbers for group courtship, in the aforementioned "water dances". The female builds a nest of vegetation, sticks, leaves, and mud in a sheltered spot in or near the water.
Eggs
After the female lays her 20 to 50 white eggs, about the size of a goose egg, she covers them with more vegetation, which heats as it decays, helping to keep the eggs warm. This differs from Nile crocodiles, which lay their eggs in pits. The temperature at which American alligator eggs develop determines their sex (see temperature-dependent sex determination). Studies have found that eggs hatched at a temperature below 88.7 °F or a temperature above 94.1 °F will produce female offspring, while those at a temperature between 90.5 °F to 92.3 °F will produce male offspring. The nests built on levees are warmer, thus produce males, while the cooler nests of wet marsh produce females. The female remains near the nest throughout the 65-day incubation period, protecting it from intruders. When the young begin to hatch — their "yelping" calls can sometimes even be heard just before hatching commences — the mother quickly digs them out and carries them to the water in her mouth, as some other crocodilian species are known to do.
Young
The young are tiny replicas of adults, with a series of yellow bands around their bodies that serve as camouflage. Hatchlings gather into pods and are guarded by their mother and keep in contact with her through their "yelping" vocalizations. Young American alligators eat small fish, frogs, crayfish, and insects. They are preyed on by large fish, birds, raccoons, Florida panthers, and adult American alligators. Mother American alligators eventually become more aggressive towards their young, which encourages them to disperse. Young American alligators grow 3–8 in (7.6–20.3 cm) a year and reach adulthood at 6 ft (1.8 m).
Parasites
American alligators are commonly infected with parasites. In a 2016 Texas study, 100% of the specimens collected were infected with parasites, and by at least 20 different species of parasites, including lung pentastomids, gastric nematodes, intestinal helminths. When compared to American alligators from different states there was no significant difference in prevalence.
Nutria were introduced into coastal marshes from South America in the mid-20th century, and their population has since exploded into the millions. They cause serious damage to coastal marshes and may dig burrows in levees. Hence, Louisiana has had a bounty to try to reduce nutria numbers. Large American alligators feed heavily on nutria, so American alligators may not only control nutria populations in Louisiana, but also prevent them spreading east into the Everglades. Since hunting and trapping preferentially take the large American alligators that are the most important in eating nutria, some changes in harvesting may be needed to capitalize on their ability to control nutria.
Recently, a population of Burmese pythons became established in Everglades National Park. Substantial American alligator populations in the Everglades might be a contributing factor, as a competitor, in keeping the python populations low, preventing the spread of the species north. While events of predation by Burmese pythons on any sizable American alligators have been observed, no evidence of a net negative effect has been seen on overall American alligator populations.
American alligators play an important role in the restoration of the Everglades as biological indicators of restoration success. American alligators are highly sensitive to changes in the hydrology, salinity, and productivity of their ecosystems; all are factors that are expected to change with Everglades restoration. American alligators also may control the long-term vegetation dynamics in wetlands by reducing the population of small mammals, particularly nutria, which may otherwise overgraze marsh vegetation. In this way, the vital ecological service they provide may be important in reducing rates of coastal wetland losses in Louisiana. They may provide a protection service for water birds nesting on islands in freshwater wetlands. American alligators prevent predatory mammals from reaching island-based rookeries and in return eat spilled food and birds that fall from their nests. Wading birds appear to be attracted to areas with American alligators and have been known to nest at heavily trafficked tourist attractions with large numbers of American alligators, such as the St. Augustine Alligator Farm in St. Augustine, Florida.
Relationship with humans
Attacks on humans
Main article: List of fatal alligator attacks in the United States
American alligators are capable of killing humans, but fatal attacks are rare. Mistaken identity leading to an attack is always possible, especially in or near cloudy waters. American alligators are often less aggressive towards humans than larger crocodile species, a few of which (mainly the Nile and saltwater crocodiles) may prey on humans with some regularity. Alligator bites are serious injuries, due to the reptile's sheer bite force and risk of infection. Even with medical treatment, an American alligator bite may still result in a fatal infection.
As human populations increase, and as they build houses in low-lying areas, or fish or hunt near water, incidents are inevitable where humans intrude on American alligators and their habitats. Since 1948, 257 documented attacks on humans in Florida (about five incidents per year) have been reported, of which an estimated 23 resulted in death. Only nine fatal attacks occurred in the United States throughout the 1970s–1990s, but American alligators killed 12 people between 2001 and 2007. An additional report of alligator attacks showed a total of 376 injuries and 15 deaths recorded all from 1948 to 2004, leading this to an increase of the alligator population. In May 2006, American alligators killed three Floridians in less than a week. At least 28 fatal attacks by American alligators have occurred in the United States since 1970.
Wrestling
Main article: Alligator wrestling
Since the late 1880s, alligator wrestling has been a source of entertainment for some. Created by the Miccosukee and Seminole tribes prior to the arrival of Europeans, this tourism tradition remains popular despite criticism from animal-rights activists.
Farming
Main article: Alligator farm
Today, alligator farming is a large, growing industry in Georgia, Florida, Texas, and Louisiana. These states produce a combined annual total of some 45,000 alligator hides. Alligator hides bring good prices and hides in the 6- to 7-ft range have sold for $300 each. The market for alligator meat is growing, and about 300,000 pounds (140,000 kg) of meat are produced annually. According to the Florida Department of Agriculture and Consumer Services, raw alligator meat contains roughly 200 Calories (840 kJ) per 3-oz (85-g) portion, of which 27 Calories (130 kJ) come from fat.
The American alligator is the official state reptile of Florida, Louisiana, and Mississippi. Several organizations and products from Florida have been named after the animal.
"Gators" has been the nickname of the University of Florida's sports teams since 1911. In 1908, a printer made a spur-of-the-moment decision to print an alligator emblem on a shipment of the school's football pennants. The mascot stuck, and was made official in 1911, perhaps because the team captain's nickname was Gator. Allegheny College and San Francisco State University both have Gators as their mascots, as well.
The Gator Bowl is a college football game held in Jacksonville annually since 1946, with Gator Bowl Stadium hosting the event until the 1993 edition. The Gatornationals is a NHRA drag race held at the Gainesville Raceway in Gainesville since 1970.
© BlueFunambulist 2014 | Spain
Deep inhalations, deep exhalations.
The plain land with it greenish, brown and bright red colours surrounded the road. A greyish way that ate the time faster than the actual speed of the wheels and the blades of the windmills. Halfway there, the monkeys of Gibraltar made smoke signals to welcome me back to the region, more than a decade later, as so did the cows with their tails. The spotless white buildings and houses started to appear far away and the road changed into one of those yellow and untreated paths. But it wasn't until when the car engine had stopped and I opened the door when the smell of the summers of my childhood suddenly embraced me. I have been remembering that smell for years and I thought I would not be able to feel it never again. I was wrong. Still as fresh and salty, still a real pleasure for my lungs.
The soft and warm sand said «run» and later on my body intensely felt it in some kind of extrasensory combination with a light-skinned one. My mind said «peace» and my heart said «be». Unceasingly I stared at the transparent glances from his yellow eyes. The sudden goosebumps were embraced as hard as I did with my surfboard, in a mutual protection attitude. My feets drew semi-circles and my fingers drew joy. The laughs were might listened by the fishes from all over the coast.
The red light of the cars indicators turned into a blurry view, and a license plate told me that 7772.
Listening to: Pánico Práctico, Izal.
I've come to accept that she's generally anxious about time
A function of matronly love perhaps
Harrying her lazy guests to hurry up, hands motioning in sweeping gestures
Shooing away dust motes and hopefully my lethargy, as I'm almost always
The last out the door, struggling with my five toed shoes
Much to my chagrin, she was amazingly blase when bringing us to the airport
Dropping hints by the truckloads
The furtive looks to my phone to check the large digital clock
Deep breaths and not-so-silent exhalations, paired with a tannic frown
I think I almost balked when she stopped in the underground street mall passageway
To take photos of the travelling band, a matsuri pre-cursor
My snarky mind instantly labelled as created to make me miss my flight
But make it we did, a dash of brisk walking
Parting as always, with grains of sand in our eyes
Hearts trapped in cat-cradles filled with flotsam
>>
In calmer times, Haneda
The Christmas lights everywhere made decent target practice for creating bokeh balls
Not too shabby for old AIS lenses
Sheep (pl.: sheep) or domestic sheep (Ovis aries) are a domesticated, ruminant mammal typically kept as livestock. Although the term sheep can apply to other species in the genus Ovis, in everyday usage it almost always refers to domesticated sheep. Like all ruminants, sheep are members of the order Artiodactyla, the even-toed ungulates. Numbering a little over one billion, domestic sheep are also the most numerous species of sheep. An adult female is referred to as a ewe (/juː/ yoo), an intact male as a ram, occasionally a tup, a castrated male as a wether, and a young sheep as a lamb.
Sheep are most likely descended from the wild mouflon of Europe and Asia, with Iran being a geographic envelope of the domestication center. One of the earliest animals to be domesticated for agricultural purposes, sheep are raised for fleeces, meat (lamb, hogget or mutton) and milk. A sheep's wool is the most widely used animal fiber, and is usually harvested by shearing. In Commonwealth countries, ovine meat is called lamb when from younger animals and mutton when from older ones; in the United States, meat from both older and younger animals is usually called lamb. Sheep continue to be important for wool and meat today, and are also occasionally raised for pelts, as dairy animals, or as model organisms for science.
Sheep husbandry is practised throughout the majority of the inhabited world, and has been fundamental to many civilizations. In the modern era, Australia, New Zealand, the southern and central South American nations, and the British Isles are most closely associated with sheep production.
There is a large lexicon of unique terms for sheep husbandry which vary considerably by region and dialect. Use of the word sheep began in Middle English as a derivation of the Old English word scēap. A group of sheep is called a flock. Many other specific terms for the various life stages of sheep exist, generally related to lambing, shearing, and age.
Being a key animal in the history of farming, sheep have a deeply entrenched place in human culture, and are represented in much modern language and symbolism. As livestock, sheep are most often associated with pastoral, Arcadian imagery. Sheep figure in many mythologies—such as the Golden Fleece—and major religions, especially the Abrahamic traditions. In both ancient and modern religious ritual, sheep are used as sacrificial animals.
History
Main article: History of the domestic sheep
The exact line of descent from wild ancestors to domestic sheep is unclear. The most common hypothesis states that Ovis aries is descended from the Asiatic (O. gmelini) species of mouflon; the European mouflon (Ovis aries musimon) is a direct descendant of this population. Sheep were among the first animals to be domesticated by humankind (although the domestication of dogs probably took place 10 to 20 thousand years earlier); the domestication date is estimated to fall between 11,000 and 9000 B.C in Mesopotamia and possibly around 7000 BC in Mehrgarh in the Indus Valley. The rearing of sheep for secondary products, and the resulting breed development, began in either southwest Asia or western Europe. Initially, sheep were kept solely for meat, milk and skins. Archaeological evidence from statuary found at sites in Iran suggests that selection for woolly sheep may have begun around 6000 BC, and the earliest woven wool garments have been dated to two to three thousand years later.
Sheep husbandry spread quickly in Europe. Excavations show that in about 6000 BC, during the Neolithic period of prehistory, the Castelnovien people, living around Châteauneuf-les-Martigues near present-day Marseille in the south of France, were among the first in Europe to keep domestic sheep. Practically from its inception, ancient Greek civilization relied on sheep as primary livestock, and were even said to name individual animals. Ancient Romans kept sheep on a wide scale, and were an important agent in the spread of sheep raising. Pliny the Elder, in his Natural History (Naturalis Historia), speaks at length about sheep and wool. European colonists spread the practice to the New World from 1493 onwards.
Characteristics
Domestic sheep are relatively small ruminants, usually with a crimped hair called wool and often with horns forming a lateral spiral. They differ from their wild relatives and ancestors in several respects, having become uniquely neotenic as a result of selective breeding by humans. A few primitive breeds of sheep retain some of the characteristics of their wild cousins, such as short tails. Depending on breed, domestic sheep may have no horns at all (i.e. polled), or horns in both sexes, or in males only. Most horned breeds have a single pair, but a few breeds may have several.
Sheep in Turkmenistan
Another trait unique to domestic sheep as compared to wild ovines is their wide variation in color. Wild sheep are largely variations of brown hues, and variation within species is extremely limited. Colors of domestic sheep range from pure white to dark chocolate brown, and even spotted or piebald. Sheep keepers also sometimes artificially paint "smit marks" onto their sheep in any pattern or color for identification. Selection for easily dyeable white fleeces began early in sheep domestication, and as white wool is a dominant trait it spread quickly. However, colored sheep do appear in many modern breeds, and may even appear as a recessive trait in white flocks. While white wool is desirable for large commercial markets, there is a niche market for colored fleeces, mostly for handspinning. The nature of the fleece varies widely among the breeds, from dense and highly crimped, to long and hairlike. There is variation of wool type and quality even among members of the same flock, so wool classing is a step in the commercial processing of the fibre.
Suffolks are a medium wool, black-faced breed of meat sheep that make up 60% of the sheep population in the U.S.
Depending on breed, sheep show a range of heights and weights. Their rate of growth and mature weight is a heritable trait that is often selected for in breeding. Ewes typically weigh between 45 and 100 kilograms (100 and 220 lb), and rams between 45 and 160 kilograms (100 and 350 lb). When all deciduous teeth have erupted, the sheep has 20 teeth. Mature sheep have 32 teeth. As with other ruminants, the front teeth in the lower jaw bite against a hard, toothless pad in the upper jaw. These are used to pick off vegetation, then the rear teeth grind it before it is swallowed. There are eight lower front teeth in ruminants, but there is some disagreement as to whether these are eight incisors, or six incisors and two incisor-shaped canines. This means that the dental formula for sheep is either
0.0.3.3
4.0.3.3
or
0.0.3.3
3.1.3.3
There is a large diastema between the incisors and the molars.
In the first few years of life one can calculate the age of sheep from their front teeth, as a pair of milk teeth is replaced by larger adult teeth each year, the full set of eight adult front teeth being complete at about four years of age. The front teeth are then gradually lost as sheep age, making it harder for them to feed and hindering the health and productivity of the animal. For this reason, domestic sheep on normal pasture begin to slowly decline from four years on, and the life expectancy of a sheep is 10 to 12 years, though some sheep may live as long as 20 years.
Skull
Sheep have good hearing, and are sensitive to noise when being handled. Sheep have horizontal slit-shaped pupils, with excellent peripheral vision; with visual fields of about 270° to 320°, sheep can see behind themselves without turning their heads. Many breeds have only short hair on the face, and some have facial wool (if any) confined to the poll and or the area of the mandibular angle; the wide angles of peripheral vision apply to these breeds. A few breeds tend to have considerable wool on the face; for some individuals of these breeds, peripheral vision may be greatly reduced by "wool blindness", unless recently shorn about the face. Sheep have poor depth perception; shadows and dips in the ground may cause sheep to baulk. In general, sheep have a tendency to move out of the dark and into well-lit areas, and prefer to move uphill when disturbed. Sheep also have an excellent sense of smell, and, like all species of their genus, have scent glands just in front of the eyes, and interdigitally on the feet. The purpose of these glands is uncertain, but those on the face may be used in breeding behaviors. The foot glands might also be related to reproduction, but alternative functions, such as secretion of a waste product or a scent marker to help lost sheep find their flock, have also been proposed.
Comparison with goats
Sheep and goats are closely related: both are in the subfamily Caprinae. However, they are separate species, so hybrids rarely occur and are always infertile. A hybrid of a ewe and a buck (a male goat) is called a sheep-goat hybrid, known as geep. Visual differences between sheep and goats include the beard of goats and divided upper lip of sheep. Sheep tails also hang down, even when short or docked, while the short tails of goats are held upwards. Also, sheep breeds are often naturally polled (either in both sexes or just in the female), while naturally polled goats are rare (though many are polled artificially). Males of the two species differ in that buck goats acquire a unique and strong odor during the rut, whereas rams do not.
Breeds
The domestic sheep is a multi-purpose animal, and the more than 200 breeds now in existence were created to serve these diverse purposes. Some sources give a count of a thousand or more breeds, but these numbers cannot be verified, according to some sources. However, several hundred breeds of sheep have been identified by the Food and Agriculture Organization of the UN (FAO), with the estimated number varying somewhat from time to time: e.g. 863 breeds as of 1993, 1314 breeds as of 1995 and 1229 breeds as of 2006. (These numbers exclude extinct breeds, which are also tallied by the FAO.) For the purpose of such tallies, the FAO definition of a breed is "either a subspecific group of domestic livestock with definable and identifiable external characteristics that enable it to be separated by visual appraisal from other similarly defined groups within the same species or a group for which geographical and/or cultural separation from phenotypically similar groups has led to acceptance of its separate identity." Almost all sheep are classified as being best suited to furnishing a certain product: wool, meat, milk, hides, or a combination in a dual-purpose breed. Other features used when classifying sheep include face color (generally white or black), tail length, presence or lack of horns, and the topography for which the breed has been developed. This last point is especially stressed in the UK, where breeds are described as either upland (hill or mountain) or lowland breeds. A sheep may also be of a fat-tailed type, which is a dual-purpose sheep common in Africa and Asia with larger deposits of fat within and around its tail.
Breeds are often categorized by the type of their wool. Fine wool breeds are those that have wool of great crimp and density, which are preferred for textiles. Most of these were derived from Merino sheep, and the breed continues to dominate the world sheep industry. Downs breeds have wool between the extremes, and are typically fast-growing meat and ram breeds with dark faces. Some major medium wool breeds, such as the Corriedale, are dual-purpose crosses of long and fine-wooled breeds and were created for high-production commercial flocks. Long wool breeds are the largest of sheep, with long wool and a slow rate of growth. Long wool sheep are most valued for crossbreeding to improve the attributes of other sheep types. For example: the American Columbia breed was developed by crossing Lincoln rams (a long wool breed) with fine-wooled Rambouillet ewes.
Coarse or carpet wool sheep are those with a medium to long length wool of characteristic coarseness. Breeds traditionally used for carpet wool show great variability, but the chief requirement is a wool that will not break down under heavy use (as would that of the finer breeds). As the demand for carpet-quality wool declines, some breeders of this type of sheep are attempting to use a few of these traditional breeds for alternative purposes. Others have always been primarily meat-class sheep.
A minor class of sheep are the dairy breeds. Dual-purpose breeds that may primarily be meat or wool sheep are often used secondarily as milking animals, but there are a few breeds that are predominantly used for milking. These sheep produce a higher quantity of milk and have slightly longer lactation curves. In the quality of their milk, the fat and protein content percentages of dairy sheep vary from non-dairy breeds, but lactose content does not.
A last group of sheep breeds is that of fur or hair sheep, which do not grow wool at all. Hair sheep are similar to the early domesticated sheep kept before woolly breeds were developed, and are raised for meat and pelts. Some modern breeds of hair sheep, such as the Dorper, result from crosses between wool and hair breeds. For meat and hide producers, hair sheep are cheaper to keep, as they do not need shearing. Hair sheep are also more resistant to parasites and hot weather.
With the modern rise of corporate agribusiness and the decline of localized family farms, many breeds of sheep are in danger of extinction. The Rare Breeds Survival Trust of the UK lists 22 native breeds as having only 3,000 registered animals (each), and The Livestock Conservancy lists 14 as either "critical" or "threatened". Preferences for breeds with uniform characteristics and fast growth have pushed heritage (or heirloom) breeds to the margins of the sheep industry. Those that remain are maintained through the efforts of conservation organizations, breed registries, and individual farmers dedicated to their preservation.
Diet
Sheep are herbivorous mammals. Most breeds prefer to graze on grass and other short roughage, avoiding the taller woody parts of plants that goats readily consume. Both sheep and goats use their lips and tongues to select parts of the plant that are easier to digest or higher in nutrition. Sheep, however, graze well in monoculture pastures where most goats fare poorly.
Ruminant system of a sheep
Like all ruminants, sheep have a complex digestive system composed of four chambers, allowing them to break down cellulose from stems, leaves, and seed hulls into simpler carbohydrates. When sheep graze, vegetation is chewed into a mass called a bolus, which is then passed into the rumen, via the reticulum. The rumen is a 19- to 38-liter (5 to 10 gallon) organ in which feed is fermented. The fermenting organisms include bacteria, fungi, and protozoa. (Other important rumen organisms include some archaea, which produce methane from carbon dioxide.) The bolus is periodically regurgitated back to the mouth as cud for additional chewing and salivation. After fermentation in the rumen, feed passes into the reticulum and the omasum; special feeds such as grains may bypass the rumen altogether. After the first three chambers, food moves into the abomasum for final digestion before processing by the intestines. The abomasum is the only one of the four chambers analogous to the human stomach, and is sometimes called the "true stomach".
Other than forage, the other staple feed for sheep is hay, often during the winter months. The ability to thrive solely on pasture (even without hay) varies with breed, but all sheep can survive on this diet. Also included in some sheep's diets are minerals, either in a trace mix or in licks. Feed provided to sheep must be specially formulated, as most cattle, poultry, pig, and even some goat feeds contain levels of copper that are lethal to sheep. The same danger applies to mineral supplements such as salt licks.
Grazing behavior
Sheep follow a diurnal pattern of activity, feeding from dawn to dusk, stopping sporadically to rest and chew their cud. Ideal pasture for sheep is not lawnlike grass, but an array of grasses, legumes and forbs. Types of land where sheep are raised vary widely, from pastures that are seeded and improved intentionally to rough, native lands. Common plants toxic to sheep are present in most of the world, and include (but are not limited to) cherry, some oaks and acorns, tomato, yew, rhubarb, potato, and rhododendron.
Effects on pasture
Sheep are largely grazing herbivores, unlike browsing animals such as goats and deer that prefer taller foliage. With a much narrower face, sheep crop plants very close to the ground and can overgraze a pasture much faster than cattle. For this reason, many shepherds use managed intensive rotational grazing, where a flock is rotated through multiple pastures, giving plants time to recover. Paradoxically, sheep can both cause and solve the spread of invasive plant species. By disturbing the natural state of pasture, sheep and other livestock can pave the way for invasive plants. However, sheep also prefer to eat invasives such as cheatgrass, leafy spurge, kudzu and spotted knapweed over native species such as sagebrush, making grazing sheep effective for conservation grazing. Research conducted in Imperial County, California compared lamb grazing with herbicides for weed control in seedling alfalfa fields. Three trials demonstrated that grazing lambs were just as effective as herbicides in controlling winter weeds. Entomologists also compared grazing lambs to insecticides for insect control in winter alfalfa. In this trial, lambs provided insect control as effectively as insecticides.
Behavior
Sheep are flock animals and strongly gregarious; much sheep behavior can be understood on the basis of these tendencies. The dominance hierarchy of sheep and their natural inclination to follow a leader to new pastures were the pivotal factors in sheep being one of the first domesticated livestock species. Furthermore, in contrast to the red deer and gazelle (two other ungulates of primary importance to meat production in prehistoric times), sheep do not defend territories although they do form home ranges. All sheep have a tendency to congregate close to other members of a flock, although this behavior varies with breed, and sheep can become stressed when separated from their flock members. During flocking, sheep have a strong tendency to follow, and a leader may simply be the first individual to move. Relationships in flocks tend to be closest among related sheep: in mixed-breed flocks, subgroups of the same breed tend to form, and a ewe and her direct descendants often move as a unit within large flocks. Sheep can become hefted to one particular local pasture (heft) so they do not roam freely in unfenced landscapes. Lambs learn the heft from ewes and if whole flocks are culled it must be retaught to the replacement animals.
Flock behaviour in sheep is generally only exhibited in groups of four or more sheep; fewer sheep may not react as expected when alone or with few other sheep. Being a prey species, the primary defense mechanism of sheep is to flee from danger when their flight zone is entered. Cornered sheep may charge and butt, or threaten by hoof stamping and adopting an aggressive posture. This is particularly true for ewes with newborn lambs.
In regions where sheep have no natural predators, none of the native breeds of sheep exhibit a strong flocking behavior.
Herding
Farmers exploit flocking behavior to keep sheep together on unfenced pastures such as hill farming, and to move them more easily. For this purpose shepherds may use herding dogs in this effort, with a highly bred herding ability. Sheep are food-oriented, and association of humans with regular feeding often results in sheep soliciting people for food. Those who are moving sheep may exploit this behavior by leading sheep with buckets of feed.
Dominance hierarchy
Sheep establish a dominance hierarchy through fighting, threats and competitiveness. Dominant animals are inclined to be more aggressive with other sheep, and usually feed first at troughs. Primarily among rams, horn size is a factor in the flock hierarchy. Rams with different size horns may be less inclined to fight to establish the dominance order, while rams with similarly sized horns are more so. Merinos have an almost linear hierarchy whereas there is a less rigid structure in Border Leicesters when a competitive feeding situation arises.
In sheep, position in a moving flock is highly correlated with social dominance, but there is no definitive study to show consistent voluntary leadership by an individual sheep.
Intelligence and learning ability
Sheep are frequently thought of as unintelligent animals. Their flocking behavior and quickness to flee and panic can make shepherding a difficult endeavor for the uninitiated. Despite these perceptions, a University of Illinois monograph on sheep reported their intelligence to be just below that of pigs and on par with that of cattle. Sheep can recognize individual human and ovine faces and remember them for years; they can remember 50 other different sheep faces for over two years; they can recognize and are attracted to individual sheep and humans by their faces, as they possess similar specialized neural systems in the temporal and frontal lobes of their brains to humans and have a greater involvement of the right brain hemisphere. In addition to long-term facial recognition of individuals, sheep can also differentiate emotional states through facial characteristics.[68][69] If worked with patiently, sheep may learn their names, and many sheep are trained to be led by halter for showing and other purposes. Sheep have also responded well to clicker training. Sheep have been used as pack animals; Tibetan nomads distribute baggage equally throughout a flock as it is herded between living sites.
It has been reported that some sheep have apparently shown problem-solving abilities; a flock in West Yorkshire, England allegedly found a way to get over cattle grids by rolling on their backs, although documentation of this has relied on anecdotal accounts.
Vocalisations
Sounds made by domestic sheep include bleats, grunts, rumbles and snorts. Bleating ("baaing") is used mostly for contact communication, especially between dam and lambs, but also at times between other flock members. The bleats of individual sheep are distinctive, enabling the ewe and her lambs to recognize each other's vocalizations. Vocal communication between lambs and their dam declines to a very low level within several weeks after parturition. A variety of bleats may be heard, depending on sheep age and circumstances. Apart from contact communication, bleating may signal distress, frustration or impatience; however, sheep are usually silent when in pain. Isolation commonly prompts bleating by sheep. Pregnant ewes may grunt when in labor. Rumbling sounds are made by the ram during courting; somewhat similar rumbling sounds may be made by the ewe, especially when with her neonate lambs. A snort (explosive exhalation through the nostrils) may signal aggression or a warning, and is often elicited from startled sheep.
Lamb
In sheep breeds lacking facial wool, the visual field is wide. In 10 sheep (Cambridge, Lleyn and Welsh Mountain breeds, which lack facial wool), the visual field ranged from 298° to 325°, averaging 313.1°, with binocular overlap ranging from 44.5° to 74°, averaging 61.7°. In some breeds, unshorn facial wool can limit the visual field; in some individuals, this may be enough to cause "wool blindness". In 60 Merinos, visual fields ranged from 219.1° to 303.0°, averaging 269.9°, and the binocular field ranged from 8.9° to 77.7°, averaging 47.5°; 36% of the measurements were limited by wool, although photographs of the experiments indicate that only limited facial wool regrowth had occurred since shearing. In addition to facial wool (in some breeds), visual field limitations can include ears and (in some breeds) horns, so the visual field can be extended by tilting the head. Sheep eyes exhibit very low hyperopia and little astigmatism. Such visual characteristics are likely to produce a well-focused retinal image of objects in both the middle and long distance. Because sheep eyes have no accommodation, one might expect the image of very near objects to be blurred, but a rather clear near image could be provided by the tapetum and large retinal image of the sheep's eye, and adequate close vision may occur at muzzle length. Good depth perception, inferred from the sheep's sure-footedness, was confirmed in "visual cliff" experiments; behavioral responses indicating depth perception are seen in lambs at one day old. Sheep are thought to have colour vision, and can distinguish between a variety of colours: black, red, brown, green, yellow and white. Sight is a vital part of sheep communication, and when grazing, they maintain visual contact with each other. Each sheep lifts its head upwards to check the position of other sheep in the flock. This constant monitoring is probably what keeps the sheep in a flock as they move along grazing. Sheep become stressed when isolated; this stress is reduced if they are provided with a mirror, indicating that the sight of other sheep reduces stress.
Taste is the most important sense in sheep, establishing forage preferences, with sweet and sour plants being preferred and bitter plants being more commonly rejected. Touch and sight are also important in relation to specific plant characteristics, such as succulence and growth form.
The ram uses his vomeronasal organ (sometimes called the Jacobson's organ) to sense the pheromones of ewes and detect when they are in estrus. The ewe uses her vomeronasal organ for early recognition of her neonate lamb.
Reproduction
Sheep follow a similar reproductive strategy to other herd animals. A group of ewes is generally mated by a single ram, who has either been chosen by a breeder or (in feral populations) has established dominance through physical contest with other rams. Most sheep are seasonal breeders, although some are able to breed year-round. Ewes generally reach sexual maturity at six to eight months old, and rams generally at four to six months. However, there are exceptions. For example, Finnsheep ewe lambs may reach puberty as early as 3 to 4 months, and Merino ewes sometimes reach puberty at 18 to 20 months. Ewes have estrus cycles about every 17 days, during which they emit a scent and indicate readiness through physical displays towards rams.
In feral sheep, rams may fight during the rut to determine which individuals may mate with ewes. Rams, especially unfamiliar ones, will also fight outside the breeding period to establish dominance; rams can kill one another if allowed to mix freely. During the rut, even usually friendly rams may become aggressive towards humans due to increases in their hormone levels.
After mating, sheep have a gestation period of about five months, and normal labor takes one to three hours. Although some breeds regularly throw larger litters of lambs, most produce single or twin lambs. During or soon after labor, ewes and lambs may be confined to small lambing jugs, small pens designed to aid both careful observation of ewes and to cement the bond between them and their lambs.
A lamb's first steps
Ovine obstetrics can be problematic. By selectively breeding ewes that produce multiple offspring with higher birth weights for generations, sheep producers have inadvertently caused some domestic sheep to have difficulty lambing; balancing ease of lambing with high productivity is one of the dilemmas of sheep breeding. In the case of any such problems, those present at lambing may assist the ewe by extracting or repositioning lambs. After the birth, ewes ideally break the amniotic sac (if it is not broken during labor), and begin licking clean the lamb. Most lambs will begin standing within an hour of birth. In normal situations, lambs nurse after standing, receiving vital colostrum milk. Lambs that either fail to nurse or are rejected by the ewe require help to survive, such as bottle-feeding or fostering by another ewe.
Most lambs begin life being born outdoors. After lambs are several weeks old, lamb marking (ear tagging, docking, mulesing, and castrating) is carried out. Vaccinations are usually carried out at this point as well. Ear tags with numbers are attached, or ear marks are applied, for ease of later identification of sheep. Docking and castration are commonly done after 24 hours (to avoid interference with maternal bonding and consumption of colostrum) and are often done not later than one week after birth, to minimize pain, stress, recovery time and complications. The first course of vaccinations (commonly anti-clostridial) is commonly given at an age of about 10 to 12 weeks; i.e. when the concentration of maternal antibodies passively acquired via colostrum is expected to have fallen low enough to permit development of active immunity. Ewes are often revaccinated annually about 3 weeks before lambing, to provide high antibody concentrations in colostrum during the first several hours after lambing. Ram lambs that will either be slaughtered or separated from ewes before sexual maturity are not usually castrated. Objections to all these procedures have been raised by animal rights groups, but farmers defend them by saying they save money, and inflict only temporary pain.
Homosexuality
Sheep are the only species of mammal except for humans which exhibits exclusive homosexual behavior. About 10% of rams refuse to mate with ewes but readily mate with other rams, and thirty percent of all rams demonstrate at least some homosexual behavior. Additionally, a small number of females that were accompanied by a male fetus in utero (i.e. as fraternal twins) are freemartins (female animals that are behaviorally masculine and lack functioning ovaries).
Health
Sheep may fall victim to poisons, infectious diseases, and physical injuries. As a prey species, a sheep's system is adapted to hide the obvious signs of illness, to prevent being targeted by predators. However, some signs of ill health are obvious, with sick sheep eating little, vocalizing excessively, and being generally listless. Throughout history, much of the money and labor of sheep husbandry has aimed to prevent sheep ailments. Historically, shepherds often created remedies by experimentation on the farm. In some developed countries, including the United States, sheep lack the economic importance for drug companies to perform expensive clinical trials required to approve more than a relatively limited number of drugs for ovine use. However, extra-label drug use in sheep production is permitted in many jurisdictions, subject to certain restrictions. In the US, for example, regulations governing extra-label drug use in animals are found in 21 CFR (Code of Federal Regulations) Part 530. In the 20th and 21st centuries, a minority of sheep owners have turned to alternative treatments such as homeopathy, herbalism and even traditional Chinese medicine to treat sheep veterinary problems. Despite some favorable anecdotal evidence, the effectiveness of alternative veterinary medicine has been met with skepticism in scientific journals. The need for traditional anti-parasite drugs and antibiotics is widespread, and is the main impediment to certified organic farming with sheep.
Many breeders take a variety of preventive measures to ward off problems. The first is to ensure all sheep are healthy when purchased. Many buyers avoid outlets known to be clearing houses for animals culled from healthy flocks as either sick or simply inferior. This can also mean maintaining a closed flock, and quarantining new sheep for a month. Two fundamental preventive programs are maintaining good nutrition and reducing stress in the sheep. Restraint, isolation, loud noises, novel situations, pain, heat, extreme cold, fatigue and other stressors can lead to secretion of cortisol, a stress hormone, in amounts that may indicate welfare problems. Excessive stress can compromise the immune system. "Shipping fever" (pneumonic mannheimiosis, formerly called pasteurellosis) is a disease of particular concern, that can occur as a result of stress, notably during transport and (or) handling. Pain, fear and several other stressors can cause secretion of epinephrine (adrenaline). Considerable epinephrine secretion in the final days before slaughter can adversely affect meat quality (by causing glycogenolysis, removing the substrate for normal post-slaughter acidification of meat) and result in meat becoming more susceptible to colonization by spoilage bacteria. Because of such issues, low-stress handling is essential in sheep management. Avoiding poisoning is also important; common poisons are pesticide sprays, inorganic fertilizer, motor oil, as well as radiator coolant containing ethylene glycol.
Common forms of preventive medication for sheep are vaccinations and treatments for parasites. Both external and internal parasites are the most prevalent malady in sheep, and are either fatal, or reduce the productivity of flocks. Worms are the most common internal parasites. They are ingested during grazing, incubate within the sheep, and are expelled through the digestive system (beginning the cycle again). Oral anti-parasitic medicines, known as drenches, are given to a flock to treat worms, sometimes after worm eggs in the feces has been counted to assess infestation levels. Afterwards, sheep may be moved to a new pasture to avoid ingesting the same parasites. External sheep parasites include: lice (for different parts of the body), sheep keds, nose bots, sheep itch mites, and maggots. Keds are blood-sucking parasites that cause general malnutrition and decreased productivity, but are not fatal. Maggots are those of the bot fly and the blow-fly, commonly Lucilia sericata or its relative L. cuprina. Fly maggots cause the extremely destructive condition of flystrike. Flies lay their eggs in wounds or wet, manure-soiled wool; when the maggots hatch they burrow into a sheep's flesh, eventually causing death if untreated. In addition to other treatments, crutching (shearing wool from a sheep's rump) is a common preventive method. Some countries allow mulesing, a practice that involves stripping away the skin on the rump to prevent fly-strike, normally performed when the sheep is a lamb. Nose bots are fly larvae that inhabit a sheep's sinuses, causing breathing difficulties and discomfort. Common signs are a discharge from the nasal passage, sneezing, and frantic movement such as head shaking. External parasites may be controlled through the use of backliners, sprays or immersive sheep dips.
A wide array of bacterial and viral diseases affect sheep. Diseases of the hoof, such as foot rot and foot scald may occur, and are treated with footbaths and other remedies. Foot rot is present in over 97% of flocks in the UK. These painful conditions cause lameness and hinder feeding. Ovine Johne's disease is a wasting disease that affects young sheep. Bluetongue disease is an insect-borne illness causing fever and inflammation of the mucous membranes. Ovine rinderpest (or peste des petits ruminants) is a highly contagious and often fatal viral disease affecting sheep and goats. Sheep may also be affected by primary or secondary photosensitization. Tetanus can also afflict sheep through wounds from shearing, docking, castration, or vaccination. The organism also can be introduced into the reproductive tract by unsanitary humans who assist ewes during lambing.
A few sheep conditions are transmissible to humans. Orf (also known as scabby mouth, contagious ecthyma or soremouth) is a skin disease leaving lesions that is transmitted through skin-to-skin contact. Cutaneous anthrax is also called woolsorter's disease, as the spores can be transmitted in unwashed wool. More seriously, the organisms that can cause spontaneous enzootic abortion in sheep are easily transmitted to pregnant women. Also of concern are the prion disease scrapie and the virus that causes foot-and-mouth disease (FMD), as both can devastate flocks. The latter poses a slight risk to humans. During the 2001 FMD pandemic in the UK, hundreds of sheep were culled and some rare British breeds were at risk of extinction due to this.
Of the 600,300 sheep lost to the US economy in 2004, 37.3% were lost to predators, while 26.5% were lost to some form of disease. Poisoning accounted for 1.7% of non-productive deaths.
Predators
A lamb being attacked by coyotes with a bite to the throat
Other than parasites and disease, predation is a threat to sheep and the profitability of sheep raising. Sheep have little ability to defend themselves, compared with other species kept as livestock. Even if sheep survive an attack, they may die from their injuries or simply from panic. However, the impact of predation varies dramatically with region. In Africa, Australia, the Americas, and parts of Europe and Asia predators are a serious problem. In the United States, for instance, over one third of sheep deaths in 2004 were caused by predation. In contrast, other nations are virtually devoid of sheep predators, particularly islands known for extensive sheep husbandry. Worldwide, canids—including the domestic dog—are responsible for most sheep deaths. Other animals that occasionally prey on sheep include: felines, bears, birds of prey, ravens and feral hogs.
Sheep producers have used a wide variety of measures to combat predation. Pre-modern shepherds used their own presence, livestock guardian dogs, and protective structures such as barns and fencing. Fencing (both regular and electric), penning sheep at night and lambing indoors all continue to be widely used. More modern shepherds used guns, traps, and poisons to kill predators, causing significant decreases in predator populations. In the wake of the environmental and conservation movements, the use of these methods now usually falls under the purview of specially designated government agencies in most developed countries.
The 1970s saw a resurgence in the use of livestock guardian dogs and the development of new methods of predator control by sheep producers, many of them non-lethal. Donkeys and guard llamas have been used since the 1980s in sheep operations, using the same basic principle as livestock guardian dogs. Interspecific pasturing, usually with larger livestock such as cattle or horses, may help to deter predators, even if such species do not actively guard sheep. In addition to animal guardians, contemporary sheep operations may use non-lethal predator deterrents such as motion-activated lights and noisy alarms.
Economic importance
Main article: Agricultural economics
Global sheep stock
in 2019
Number in millions
1. China163.5 (13.19%)
2. India74.3 (5.99%)
3. Australia65.8 (5.31%)
4. Nigeria46.9 (3.78%)
5. Iran41.3 (3.33%)
6. Sudan40.9 (3.3%)
7. Chad35.9 (2.9%)
8. Turkey35.2 (2.84%)
9. United Kingdom33.6 (2.71%)
10. Mongolia32.3 (2.61%)
World total1,239.8
Source: UN Food and Agriculture Organization
Sheep are an important part of the global agricultural economy. However, their once vital status has been largely replaced by other livestock species, especially the pig, chicken, and cow. China, Australia, India, and Iran have the largest modern flocks, and serve both local and exportation needs for wool and mutton. Other countries such as New Zealand have smaller flocks but retain a large international economic impact due to their export of sheep products. Sheep also play a major role in many local economies, which may be niche markets focused on organic or sustainable agriculture and local food customers. Especially in developing countries, such flocks may be a part of subsistence agriculture rather than a system of trade. Sheep themselves may be a medium of trade in barter economies.
Domestic sheep provide a wide array of raw materials. Wool was one of the first textiles, although in the late 20th century wool prices began to fall dramatically as the result of the popularity and cheap prices for synthetic fabrics. For many sheep owners, the cost of shearing is greater than the possible profit from the fleece, making subsisting on wool production alone practically impossible without farm subsidies. Fleeces are used as material in making alternative products such as wool insulation. In the 21st century, the sale of meat is the most profitable enterprise in the sheep industry, even though far less sheep meat is consumed than chicken, pork or beef.
Sheepskin is likewise used for making clothes, footwear, rugs, and other products. Byproducts from the slaughter of sheep are also of value: sheep tallow can be used in candle and soap making, sheep bone and cartilage has been used to furnish carved items such as dice and buttons as well as rendered glue and gelatin. Sheep intestine can be formed into sausage casings, and lamb intestine has been formed into surgical sutures, as well as strings for musical instruments and tennis rackets. Sheep droppings, which are high in cellulose, have even been sterilized and mixed with traditional pulp materials to make paper. Of all sheep byproducts, perhaps the most valuable is lanolin: the waterproof, fatty substance found naturally in sheep's wool and used as a base for innumerable cosmetics and other products.
Some farmers who keep sheep also make a profit from live sheep. Providing lambs for youth programs such as 4-H and competition at agricultural shows is often a dependable avenue for the sale of sheep. Farmers may also choose to focus on a particular breed of sheep in order to sell registered purebred animals, as well as provide a ram rental service for breeding. A new option for deriving profit from live sheep is the rental of flocks for grazing; these "mowing services" are hired in order to keep unwanted vegetation down in public spaces and to lessen fire hazard.
Despite the falling demand and price for sheep products in many markets, sheep have distinct economic advantages when compared with other livestock. They do not require expensive housing, such as that used in the intensive farming of chickens or pigs. They are an efficient use of land; roughly six sheep can be kept on the amount that would suffice for a single cow or horse. Sheep can also consume plants, such as noxious weeds, that most other animals will not touch, and produce more young at a faster rate. Also, in contrast to most livestock species, the cost of raising sheep is not necessarily tied to the price of feed crops such as grain, soybeans and corn. Combined with the lower cost of quality sheep, all these factors combine to equal a lower overhead for sheep producers, thus entailing a higher profitability potential for the small farmer. Sheep are especially beneficial for independent producers, including family farms with limited resources, as the sheep industry is one of the few types of animal agriculture that has not been vertically integrated by agribusiness. However, small flocks, from 10 to 50 ewes, often are not profitable because they tend to be poorly managed. The primary reason is that mechanization is not feasible, so return per hour of labor is not maximized. Small farm flocks generally are used simply to control weeds on irrigation ditches or maintained as a hobby.
Shoulder of lamb
Sheep meat and milk were one of the earliest staple proteins consumed by human civilization after the transition from hunting and gathering to agriculture. Sheep meat prepared for food is known as either mutton or lamb, and approximately 540 million sheep are slaughtered each year for meat worldwide. "Mutton" is derived from the Old French moton, which was the word for sheep used by the Anglo-Norman rulers of much of the British Isles in the Middle Ages. This became the name for sheep meat in English, while the Old English word sceap was kept for the live animal. Throughout modern history, "mutton" has been limited to the meat of mature sheep usually at least two years of age; "lamb" is used for that of immature sheep less than a year.
In the 21st century, the nations with the highest consumption of sheep meat are the Arab states of the Persian Gulf, New Zealand, Australia, Greece, Uruguay, the United Kingdom and Ireland. These countries eat 14–40 lbs (3–18 kg) of sheep meat per capita, per annum. Sheep meat is also popular in France, Africa (especially the Arab world), the Caribbean, the rest of the Middle East, India, and parts of China. This often reflects a history of sheep production. In these countries in particular, dishes comprising alternative cuts and offal may be popular or traditional. Sheep testicles—called animelles or lamb fries—are considered a delicacy in many parts of the world. Perhaps the most unusual dish of sheep meat is the Scottish haggis, composed of various sheep innards cooked along with oatmeal and chopped onions inside its stomach. In comparison, countries such as the U.S. consume only a pound or less (under 0.5 kg), with Americans eating 50 pounds (22 kg) of pork and 65 pounds (29 kg) of beef. In addition, such countries rarely eat mutton, and may favor the more expensive cuts of lamb: mostly lamb chops and leg of lamb.
Though sheep's milk may be drunk rarely in fresh form, today it is used predominantly in cheese and yogurt making. Sheep have only two teats, and produce a far smaller volume of milk than cows. However, as sheep's milk contains far more fat, solids, and minerals than cow's milk, it is ideal for the cheese-making process. It also resists contamination during cooling better because of its much higher calcium content. Well-known cheeses made from sheep milk include the feta of Bulgaria and Greece, Roquefort of France, Manchego from Spain, the pecorino romano (the Italian word for "sheep" is pecore) and ricotta of Italy. Yogurts, especially some forms of strained yogurt, may also be made from sheep milk. Many of these products are now often made with cow's milk, especially when produced outside their country of origin. Sheep milk contains 4.8% lactose, which may affect those who are intolerant.
As with other domestic animals, the meat of uncastrated males is inferior in quality, especially as they grow. A "bucky" lamb is a lamb which was not castrated early enough, or which was castrated improperly (resulting in one testicle being retained). These lambs are worth less at market.
In science
Sheep are generally too large and reproduce too slowly to make ideal research subjects, and thus are not a common model organism. They have, however, played an influential role in some fields of science. In particular, the Roslin Institute of Edinburgh, Scotland used sheep for genetics research that produced groundbreaking results. In 1995, two ewes named Megan and Morag were the first mammals cloned from differentiated cells, also referred to as gynomerogony. A year later, a Finnish Dorset sheep named Dolly, dubbed "the world's most famous sheep" in Scientific American, was the first mammal to be cloned from an adult somatic cell. Following this, Polly and Molly were the first mammals to be simultaneously cloned and transgenic.
As of 2008, the sheep genome has not been fully sequenced, although a detailed genetic map has been published, and a draft version of the complete genome produced by assembling sheep DNA sequences using information given by the genomes of other mammals. In 2012, a transgenic sheep named "Peng Peng" was cloned by Chinese scientists, who spliced his genes with that of a roundworm (C. elegans) in order to increase production of fats healthier for human consumption.
In the study of natural selection, the population of Soay sheep that remain on the island of Hirta have been used to explore the relation of body size and coloration to reproductive success. Soay sheep come in several colors, and researchers investigated why the larger, darker sheep were in decline; this occurrence contradicted the rule of thumb that larger members of a population tend to be more successful reproductively. The feral Soays on Hirta are especially useful subjects because they are isolated.
Domestic sheep are sometimes used in medical research, particularly for researching cardiovascular physiology, in areas such as hypertension and heart failure. Pregnant sheep are also a useful model for human pregnancy, and have been used to investigate the effects on fetal development of malnutrition and hypoxia. In behavioral sciences, sheep have been used in isolated cases for the study of facial recognition, as their mental process of recognition is qualitatively similar to humans.
Cultural impact
Sheep have had a strong presence in many cultures, especially in areas where they form the most common type of livestock. In the English language, to call someone a sheep or ovine may allude that they are timid and easily led. In contradiction to this image, male sheep are often used as symbols of virility and power; the logos of the Los Angeles Rams football team and the Dodge Ram pickup truck allude to males of the bighorn sheep, Ovis canadensis.
Counting sheep is popularly said to be an aid to sleep, and some ancient systems of counting sheep persist today. Sheep also enter in colloquial sayings and idiom frequently with such phrases as "black sheep". To call an individual a black sheep implies that they are an odd or disreputable member of a group. This usage derives from the recessive trait that causes an occasional black lamb to be born into an entirely white flock. These black sheep were considered undesirable by shepherds, as black wool is not as commercially viable as white wool. Citizens who accept overbearing governments have been referred to by the Portmanteau neologism of sheeple. Somewhat differently, the adjective "sheepish" is also used to describe embarrassment.
In heraldry
In British heraldry, sheep appear in the form of rams, sheep proper and lambs. These are distinguished by the ram being depicted with horns and a tail, the sheep with neither and the lamb with its tail only. A further variant of the lamb, termed the Paschal lamb, is depicted as carrying a Christian cross and with a halo over its head. Rams' heads, portrayed without a neck and facing the viewer, are also found in British armories. The fleece, depicted as an entire sheepskin carried by a ring around its midsection, originally became known through its use in the arms of the Order of the Golden Fleece and was later adopted by towns and individuals with connections to the wool industry. A sheep on a blue field is depicted on the greater/royal arms of the king of Denmark to represent the Faroe Islands. In 2004 a modernized arms has been adopted by the Faroe Islands, which based on a 15th century coat of arms.
Religion and folklore
In antiquity, symbolism involving sheep cropped up in religions in the ancient Near East, the Mideast, and the Mediterranean area: Çatalhöyük, ancient Egyptian religion, the Cana'anite and Phoenician tradition, Judaism, Greek religion, and others. Religious symbolism and ritual involving sheep began with some of the first known faiths: Skulls of rams (along with bulls) occupied central placement in shrines at the Çatalhöyük settlement in 8,000 BCE. In Ancient Egyptian religion, the ram was the symbol of several gods: Khnum, Heryshaf and Amun (in his incarnation as a god of fertility). Other deities occasionally shown with ram features include the goddess Ishtar, the Phoenician god Baal-Hamon, and the Babylonian god Ea-Oannes. In Madagascar, sheep were not eaten as they were believed to be incarnations of the souls of ancestors.
There are many ancient Greek references to sheep: that of Chrysomallos, the golden-fleeced ram, continuing to be told through into the modern era. Astrologically, Aries, the ram, is the first sign of the classical Greek zodiac, and the sheep is the eighth of the twelve animals associated with the 12-year cycle of in the Chinese zodiac, related to the Chinese calendar. It is said in Chinese traditions that Hou ji sacrificed sheep. Mongolia, shagai are an ancient form of dice made from the cuboid bones of sheep that are often used for fortunetelling purposes.
Sheep play an important role in all the Abrahamic faiths; Abraham, Isaac, Jacob, Moses, and King David were all shepherds. According to the Biblical story of the Binding of Isaac, a ram is sacrificed as a substitute for Isaac after an angel stays Abraham's hand (in the Islamic tradition, Abraham was about to sacrifice Ishmael). Eid al-Adha is a major annual festival in Islam in which sheep (or other animals) are sacrificed in remembrance of this act. Sheep are occasionally sacrificed to commemorate important secular events in Islamic cultures. Greeks and Romans sacrificed sheep regularly in religious practice, and Judaism once sacrificed sheep as a Korban (sacrifice), such as the Passover lamb. Ovine symbols—such as the ceremonial blowing of a shofar—still find a presence in modern Judaic traditions.
Collectively, followers of Christianity are often referred to as a flock, with Christ as the Good Shepherd, and sheep are an element in the Christian iconography of the birth of Jesus. Some Christian saints are considered patrons of shepherds, and even of sheep themselves. Christ is also portrayed as the Sacrificial lamb of God (Agnus Dei) and Easter celebrations in Greece and Romania traditionally feature a meal of Paschal lamb. A church leader is often called the pastor, which is derived from the Latin word for shepherd. In many western Christian traditions bishops carry a staff, which also serves as a symbol of the episcopal office, known as a crosier, which is modeled on the shepherd's crook.
Sheep are key symbols in fables and nursery rhymes like The Wolf in Sheep's Clothing, Little Bo Peep, Baa, Baa, Black Sheep, and Mary Had a Little Lamb; novels such as George Orwell's Animal Farm and Haruki Murakami's A Wild Sheep Chase; songs such as Bach's Sheep may safely graze (Schafe können sicher weiden) and Pink Floyd's "Sheep", and poems like William Blake's "The Lamb".
Sheep (pl.: sheep) or domestic sheep (Ovis aries) are a domesticated, ruminant mammal typically kept as livestock. Although the term sheep can apply to other species in the genus Ovis, in everyday usage it almost always refers to domesticated sheep. Like all ruminants, sheep are members of the order Artiodactyla, the even-toed ungulates. Numbering a little over one billion, domestic sheep are also the most numerous species of sheep. An adult female is referred to as a ewe (/juː/ yoo), an intact male as a ram, occasionally a tup, a castrated male as a wether, and a young sheep as a lamb.
Sheep are most likely descended from the wild mouflon of Europe and Asia, with Iran being a geographic envelope of the domestication center. One of the earliest animals to be domesticated for agricultural purposes, sheep are raised for fleeces, meat (lamb, hogget or mutton) and milk. A sheep's wool is the most widely used animal fiber, and is usually harvested by shearing. In Commonwealth countries, ovine meat is called lamb when from younger animals and mutton when from older ones; in the United States, meat from both older and younger animals is usually called lamb. Sheep continue to be important for wool and meat today, and are also occasionally raised for pelts, as dairy animals, or as model organisms for science.
Sheep husbandry is practised throughout the majority of the inhabited world, and has been fundamental to many civilizations. In the modern era, Australia, New Zealand, the southern and central South American nations, and the British Isles are most closely associated with sheep production.
There is a large lexicon of unique terms for sheep husbandry which vary considerably by region and dialect. Use of the word sheep began in Middle English as a derivation of the Old English word scēap. A group of sheep is called a flock. Many other specific terms for the various life stages of sheep exist, generally related to lambing, shearing, and age.
Being a key animal in the history of farming, sheep have a deeply entrenched place in human culture, and are represented in much modern language and symbolism. As livestock, sheep are most often associated with pastoral, Arcadian imagery. Sheep figure in many mythologies—such as the Golden Fleece—and major religions, especially the Abrahamic traditions. In both ancient and modern religious ritual, sheep are used as sacrificial animals.
History
Main article: History of the domestic sheep
The exact line of descent from wild ancestors to domestic sheep is unclear. The most common hypothesis states that Ovis aries is descended from the Asiatic (O. gmelini) species of mouflon; the European mouflon (Ovis aries musimon) is a direct descendant of this population. Sheep were among the first animals to be domesticated by humankind (although the domestication of dogs probably took place 10 to 20 thousand years earlier); the domestication date is estimated to fall between 11,000 and 9000 B.C in Mesopotamia and possibly around 7000 BC in Mehrgarh in the Indus Valley. The rearing of sheep for secondary products, and the resulting breed development, began in either southwest Asia or western Europe. Initially, sheep were kept solely for meat, milk and skins. Archaeological evidence from statuary found at sites in Iran suggests that selection for woolly sheep may have begun around 6000 BC, and the earliest woven wool garments have been dated to two to three thousand years later.
Sheep husbandry spread quickly in Europe. Excavations show that in about 6000 BC, during the Neolithic period of prehistory, the Castelnovien people, living around Châteauneuf-les-Martigues near present-day Marseille in the south of France, were among the first in Europe to keep domestic sheep. Practically from its inception, ancient Greek civilization relied on sheep as primary livestock, and were even said to name individual animals. Ancient Romans kept sheep on a wide scale, and were an important agent in the spread of sheep raising. Pliny the Elder, in his Natural History (Naturalis Historia), speaks at length about sheep and wool. European colonists spread the practice to the New World from 1493 onwards.
Characteristics
Domestic sheep are relatively small ruminants, usually with a crimped hair called wool and often with horns forming a lateral spiral. They differ from their wild relatives and ancestors in several respects, having become uniquely neotenic as a result of selective breeding by humans. A few primitive breeds of sheep retain some of the characteristics of their wild cousins, such as short tails. Depending on breed, domestic sheep may have no horns at all (i.e. polled), or horns in both sexes, or in males only. Most horned breeds have a single pair, but a few breeds may have several.
Sheep in Turkmenistan
Another trait unique to domestic sheep as compared to wild ovines is their wide variation in color. Wild sheep are largely variations of brown hues, and variation within species is extremely limited. Colors of domestic sheep range from pure white to dark chocolate brown, and even spotted or piebald. Sheep keepers also sometimes artificially paint "smit marks" onto their sheep in any pattern or color for identification. Selection for easily dyeable white fleeces began early in sheep domestication, and as white wool is a dominant trait it spread quickly. However, colored sheep do appear in many modern breeds, and may even appear as a recessive trait in white flocks. While white wool is desirable for large commercial markets, there is a niche market for colored fleeces, mostly for handspinning. The nature of the fleece varies widely among the breeds, from dense and highly crimped, to long and hairlike. There is variation of wool type and quality even among members of the same flock, so wool classing is a step in the commercial processing of the fibre.
Suffolks are a medium wool, black-faced breed of meat sheep that make up 60% of the sheep population in the U.S.
Depending on breed, sheep show a range of heights and weights. Their rate of growth and mature weight is a heritable trait that is often selected for in breeding. Ewes typically weigh between 45 and 100 kilograms (100 and 220 lb), and rams between 45 and 160 kilograms (100 and 350 lb). When all deciduous teeth have erupted, the sheep has 20 teeth. Mature sheep have 32 teeth. As with other ruminants, the front teeth in the lower jaw bite against a hard, toothless pad in the upper jaw. These are used to pick off vegetation, then the rear teeth grind it before it is swallowed. There are eight lower front teeth in ruminants, but there is some disagreement as to whether these are eight incisors, or six incisors and two incisor-shaped canines. This means that the dental formula for sheep is either
0.0.3.3
4.0.3.3
or
0.0.3.3
3.1.3.3
There is a large diastema between the incisors and the molars.
In the first few years of life one can calculate the age of sheep from their front teeth, as a pair of milk teeth is replaced by larger adult teeth each year, the full set of eight adult front teeth being complete at about four years of age. The front teeth are then gradually lost as sheep age, making it harder for them to feed and hindering the health and productivity of the animal. For this reason, domestic sheep on normal pasture begin to slowly decline from four years on, and the life expectancy of a sheep is 10 to 12 years, though some sheep may live as long as 20 years.
Skull
Sheep have good hearing, and are sensitive to noise when being handled. Sheep have horizontal slit-shaped pupils, with excellent peripheral vision; with visual fields of about 270° to 320°, sheep can see behind themselves without turning their heads. Many breeds have only short hair on the face, and some have facial wool (if any) confined to the poll and or the area of the mandibular angle; the wide angles of peripheral vision apply to these breeds. A few breeds tend to have considerable wool on the face; for some individuals of these breeds, peripheral vision may be greatly reduced by "wool blindness", unless recently shorn about the face. Sheep have poor depth perception; shadows and dips in the ground may cause sheep to baulk. In general, sheep have a tendency to move out of the dark and into well-lit areas, and prefer to move uphill when disturbed. Sheep also have an excellent sense of smell, and, like all species of their genus, have scent glands just in front of the eyes, and interdigitally on the feet. The purpose of these glands is uncertain, but those on the face may be used in breeding behaviors. The foot glands might also be related to reproduction, but alternative functions, such as secretion of a waste product or a scent marker to help lost sheep find their flock, have also been proposed.
Comparison with goats
Sheep and goats are closely related: both are in the subfamily Caprinae. However, they are separate species, so hybrids rarely occur and are always infertile. A hybrid of a ewe and a buck (a male goat) is called a sheep-goat hybrid, known as geep. Visual differences between sheep and goats include the beard of goats and divided upper lip of sheep. Sheep tails also hang down, even when short or docked, while the short tails of goats are held upwards. Also, sheep breeds are often naturally polled (either in both sexes or just in the female), while naturally polled goats are rare (though many are polled artificially). Males of the two species differ in that buck goats acquire a unique and strong odor during the rut, whereas rams do not.
Breeds
The domestic sheep is a multi-purpose animal, and the more than 200 breeds now in existence were created to serve these diverse purposes. Some sources give a count of a thousand or more breeds, but these numbers cannot be verified, according to some sources. However, several hundred breeds of sheep have been identified by the Food and Agriculture Organization of the UN (FAO), with the estimated number varying somewhat from time to time: e.g. 863 breeds as of 1993, 1314 breeds as of 1995 and 1229 breeds as of 2006. (These numbers exclude extinct breeds, which are also tallied by the FAO.) For the purpose of such tallies, the FAO definition of a breed is "either a subspecific group of domestic livestock with definable and identifiable external characteristics that enable it to be separated by visual appraisal from other similarly defined groups within the same species or a group for which geographical and/or cultural separation from phenotypically similar groups has led to acceptance of its separate identity." Almost all sheep are classified as being best suited to furnishing a certain product: wool, meat, milk, hides, or a combination in a dual-purpose breed. Other features used when classifying sheep include face color (generally white or black), tail length, presence or lack of horns, and the topography for which the breed has been developed. This last point is especially stressed in the UK, where breeds are described as either upland (hill or mountain) or lowland breeds. A sheep may also be of a fat-tailed type, which is a dual-purpose sheep common in Africa and Asia with larger deposits of fat within and around its tail.
Breeds are often categorized by the type of their wool. Fine wool breeds are those that have wool of great crimp and density, which are preferred for textiles. Most of these were derived from Merino sheep, and the breed continues to dominate the world sheep industry. Downs breeds have wool between the extremes, and are typically fast-growing meat and ram breeds with dark faces. Some major medium wool breeds, such as the Corriedale, are dual-purpose crosses of long and fine-wooled breeds and were created for high-production commercial flocks. Long wool breeds are the largest of sheep, with long wool and a slow rate of growth. Long wool sheep are most valued for crossbreeding to improve the attributes of other sheep types. For example: the American Columbia breed was developed by crossing Lincoln rams (a long wool breed) with fine-wooled Rambouillet ewes.
Coarse or carpet wool sheep are those with a medium to long length wool of characteristic coarseness. Breeds traditionally used for carpet wool show great variability, but the chief requirement is a wool that will not break down under heavy use (as would that of the finer breeds). As the demand for carpet-quality wool declines, some breeders of this type of sheep are attempting to use a few of these traditional breeds for alternative purposes. Others have always been primarily meat-class sheep.
A minor class of sheep are the dairy breeds. Dual-purpose breeds that may primarily be meat or wool sheep are often used secondarily as milking animals, but there are a few breeds that are predominantly used for milking. These sheep produce a higher quantity of milk and have slightly longer lactation curves. In the quality of their milk, the fat and protein content percentages of dairy sheep vary from non-dairy breeds, but lactose content does not.
A last group of sheep breeds is that of fur or hair sheep, which do not grow wool at all. Hair sheep are similar to the early domesticated sheep kept before woolly breeds were developed, and are raised for meat and pelts. Some modern breeds of hair sheep, such as the Dorper, result from crosses between wool and hair breeds. For meat and hide producers, hair sheep are cheaper to keep, as they do not need shearing. Hair sheep are also more resistant to parasites and hot weather.
With the modern rise of corporate agribusiness and the decline of localized family farms, many breeds of sheep are in danger of extinction. The Rare Breeds Survival Trust of the UK lists 22 native breeds as having only 3,000 registered animals (each), and The Livestock Conservancy lists 14 as either "critical" or "threatened". Preferences for breeds with uniform characteristics and fast growth have pushed heritage (or heirloom) breeds to the margins of the sheep industry. Those that remain are maintained through the efforts of conservation organizations, breed registries, and individual farmers dedicated to their preservation.
Diet
Sheep are herbivorous mammals. Most breeds prefer to graze on grass and other short roughage, avoiding the taller woody parts of plants that goats readily consume. Both sheep and goats use their lips and tongues to select parts of the plant that are easier to digest or higher in nutrition. Sheep, however, graze well in monoculture pastures where most goats fare poorly.
Ruminant system of a sheep
Like all ruminants, sheep have a complex digestive system composed of four chambers, allowing them to break down cellulose from stems, leaves, and seed hulls into simpler carbohydrates. When sheep graze, vegetation is chewed into a mass called a bolus, which is then passed into the rumen, via the reticulum. The rumen is a 19- to 38-liter (5 to 10 gallon) organ in which feed is fermented. The fermenting organisms include bacteria, fungi, and protozoa. (Other important rumen organisms include some archaea, which produce methane from carbon dioxide.) The bolus is periodically regurgitated back to the mouth as cud for additional chewing and salivation. After fermentation in the rumen, feed passes into the reticulum and the omasum; special feeds such as grains may bypass the rumen altogether. After the first three chambers, food moves into the abomasum for final digestion before processing by the intestines. The abomasum is the only one of the four chambers analogous to the human stomach, and is sometimes called the "true stomach".
Other than forage, the other staple feed for sheep is hay, often during the winter months. The ability to thrive solely on pasture (even without hay) varies with breed, but all sheep can survive on this diet. Also included in some sheep's diets are minerals, either in a trace mix or in licks. Feed provided to sheep must be specially formulated, as most cattle, poultry, pig, and even some goat feeds contain levels of copper that are lethal to sheep. The same danger applies to mineral supplements such as salt licks.
Grazing behavior
Sheep follow a diurnal pattern of activity, feeding from dawn to dusk, stopping sporadically to rest and chew their cud. Ideal pasture for sheep is not lawnlike grass, but an array of grasses, legumes and forbs. Types of land where sheep are raised vary widely, from pastures that are seeded and improved intentionally to rough, native lands. Common plants toxic to sheep are present in most of the world, and include (but are not limited to) cherry, some oaks and acorns, tomato, yew, rhubarb, potato, and rhododendron.
Effects on pasture
Sheep are largely grazing herbivores, unlike browsing animals such as goats and deer that prefer taller foliage. With a much narrower face, sheep crop plants very close to the ground and can overgraze a pasture much faster than cattle. For this reason, many shepherds use managed intensive rotational grazing, where a flock is rotated through multiple pastures, giving plants time to recover. Paradoxically, sheep can both cause and solve the spread of invasive plant species. By disturbing the natural state of pasture, sheep and other livestock can pave the way for invasive plants. However, sheep also prefer to eat invasives such as cheatgrass, leafy spurge, kudzu and spotted knapweed over native species such as sagebrush, making grazing sheep effective for conservation grazing. Research conducted in Imperial County, California compared lamb grazing with herbicides for weed control in seedling alfalfa fields. Three trials demonstrated that grazing lambs were just as effective as herbicides in controlling winter weeds. Entomologists also compared grazing lambs to insecticides for insect control in winter alfalfa. In this trial, lambs provided insect control as effectively as insecticides.
Behavior
Sheep are flock animals and strongly gregarious; much sheep behavior can be understood on the basis of these tendencies. The dominance hierarchy of sheep and their natural inclination to follow a leader to new pastures were the pivotal factors in sheep being one of the first domesticated livestock species. Furthermore, in contrast to the red deer and gazelle (two other ungulates of primary importance to meat production in prehistoric times), sheep do not defend territories although they do form home ranges. All sheep have a tendency to congregate close to other members of a flock, although this behavior varies with breed, and sheep can become stressed when separated from their flock members. During flocking, sheep have a strong tendency to follow, and a leader may simply be the first individual to move. Relationships in flocks tend to be closest among related sheep: in mixed-breed flocks, subgroups of the same breed tend to form, and a ewe and her direct descendants often move as a unit within large flocks. Sheep can become hefted to one particular local pasture (heft) so they do not roam freely in unfenced landscapes. Lambs learn the heft from ewes and if whole flocks are culled it must be retaught to the replacement animals.
Flock behaviour in sheep is generally only exhibited in groups of four or more sheep; fewer sheep may not react as expected when alone or with few other sheep. Being a prey species, the primary defense mechanism of sheep is to flee from danger when their flight zone is entered. Cornered sheep may charge and butt, or threaten by hoof stamping and adopting an aggressive posture. This is particularly true for ewes with newborn lambs.
In regions where sheep have no natural predators, none of the native breeds of sheep exhibit a strong flocking behavior.
Herding
Farmers exploit flocking behavior to keep sheep together on unfenced pastures such as hill farming, and to move them more easily. For this purpose shepherds may use herding dogs in this effort, with a highly bred herding ability. Sheep are food-oriented, and association of humans with regular feeding often results in sheep soliciting people for food. Those who are moving sheep may exploit this behavior by leading sheep with buckets of feed.
Dominance hierarchy
Sheep establish a dominance hierarchy through fighting, threats and competitiveness. Dominant animals are inclined to be more aggressive with other sheep, and usually feed first at troughs. Primarily among rams, horn size is a factor in the flock hierarchy. Rams with different size horns may be less inclined to fight to establish the dominance order, while rams with similarly sized horns are more so. Merinos have an almost linear hierarchy whereas there is a less rigid structure in Border Leicesters when a competitive feeding situation arises.
In sheep, position in a moving flock is highly correlated with social dominance, but there is no definitive study to show consistent voluntary leadership by an individual sheep.
Intelligence and learning ability
Sheep are frequently thought of as unintelligent animals. Their flocking behavior and quickness to flee and panic can make shepherding a difficult endeavor for the uninitiated. Despite these perceptions, a University of Illinois monograph on sheep reported their intelligence to be just below that of pigs and on par with that of cattle. Sheep can recognize individual human and ovine faces and remember them for years; they can remember 50 other different sheep faces for over two years; they can recognize and are attracted to individual sheep and humans by their faces, as they possess similar specialized neural systems in the temporal and frontal lobes of their brains to humans and have a greater involvement of the right brain hemisphere. In addition to long-term facial recognition of individuals, sheep can also differentiate emotional states through facial characteristics.[68][69] If worked with patiently, sheep may learn their names, and many sheep are trained to be led by halter for showing and other purposes. Sheep have also responded well to clicker training. Sheep have been used as pack animals; Tibetan nomads distribute baggage equally throughout a flock as it is herded between living sites.
It has been reported that some sheep have apparently shown problem-solving abilities; a flock in West Yorkshire, England allegedly found a way to get over cattle grids by rolling on their backs, although documentation of this has relied on anecdotal accounts.
Vocalisations
Sounds made by domestic sheep include bleats, grunts, rumbles and snorts. Bleating ("baaing") is used mostly for contact communication, especially between dam and lambs, but also at times between other flock members. The bleats of individual sheep are distinctive, enabling the ewe and her lambs to recognize each other's vocalizations. Vocal communication between lambs and their dam declines to a very low level within several weeks after parturition. A variety of bleats may be heard, depending on sheep age and circumstances. Apart from contact communication, bleating may signal distress, frustration or impatience; however, sheep are usually silent when in pain. Isolation commonly prompts bleating by sheep. Pregnant ewes may grunt when in labor. Rumbling sounds are made by the ram during courting; somewhat similar rumbling sounds may be made by the ewe, especially when with her neonate lambs. A snort (explosive exhalation through the nostrils) may signal aggression or a warning, and is often elicited from startled sheep.
Lamb
In sheep breeds lacking facial wool, the visual field is wide. In 10 sheep (Cambridge, Lleyn and Welsh Mountain breeds, which lack facial wool), the visual field ranged from 298° to 325°, averaging 313.1°, with binocular overlap ranging from 44.5° to 74°, averaging 61.7°. In some breeds, unshorn facial wool can limit the visual field; in some individuals, this may be enough to cause "wool blindness". In 60 Merinos, visual fields ranged from 219.1° to 303.0°, averaging 269.9°, and the binocular field ranged from 8.9° to 77.7°, averaging 47.5°; 36% of the measurements were limited by wool, although photographs of the experiments indicate that only limited facial wool regrowth had occurred since shearing. In addition to facial wool (in some breeds), visual field limitations can include ears and (in some breeds) horns, so the visual field can be extended by tilting the head. Sheep eyes exhibit very low hyperopia and little astigmatism. Such visual characteristics are likely to produce a well-focused retinal image of objects in both the middle and long distance. Because sheep eyes have no accommodation, one might expect the image of very near objects to be blurred, but a rather clear near image could be provided by the tapetum and large retinal image of the sheep's eye, and adequate close vision may occur at muzzle length. Good depth perception, inferred from the sheep's sure-footedness, was confirmed in "visual cliff" experiments; behavioral responses indicating depth perception are seen in lambs at one day old. Sheep are thought to have colour vision, and can distinguish between a variety of colours: black, red, brown, green, yellow and white. Sight is a vital part of sheep communication, and when grazing, they maintain visual contact with each other. Each sheep lifts its head upwards to check the position of other sheep in the flock. This constant monitoring is probably what keeps the sheep in a flock as they move along grazing. Sheep become stressed when isolated; this stress is reduced if they are provided with a mirror, indicating that the sight of other sheep reduces stress.
Taste is the most important sense in sheep, establishing forage preferences, with sweet and sour plants being preferred and bitter plants being more commonly rejected. Touch and sight are also important in relation to specific plant characteristics, such as succulence and growth form.
The ram uses his vomeronasal organ (sometimes called the Jacobson's organ) to sense the pheromones of ewes and detect when they are in estrus. The ewe uses her vomeronasal organ for early recognition of her neonate lamb.
Reproduction
Sheep follow a similar reproductive strategy to other herd animals. A group of ewes is generally mated by a single ram, who has either been chosen by a breeder or (in feral populations) has established dominance through physical contest with other rams. Most sheep are seasonal breeders, although some are able to breed year-round. Ewes generally reach sexual maturity at six to eight months old, and rams generally at four to six months. However, there are exceptions. For example, Finnsheep ewe lambs may reach puberty as early as 3 to 4 months, and Merino ewes sometimes reach puberty at 18 to 20 months. Ewes have estrus cycles about every 17 days, during which they emit a scent and indicate readiness through physical displays towards rams.
In feral sheep, rams may fight during the rut to determine which individuals may mate with ewes. Rams, especially unfamiliar ones, will also fight outside the breeding period to establish dominance; rams can kill one another if allowed to mix freely. During the rut, even usually friendly rams may become aggressive towards humans due to increases in their hormone levels.
After mating, sheep have a gestation period of about five months, and normal labor takes one to three hours. Although some breeds regularly throw larger litters of lambs, most produce single or twin lambs. During or soon after labor, ewes and lambs may be confined to small lambing jugs, small pens designed to aid both careful observation of ewes and to cement the bond between them and their lambs.
A lamb's first steps
Ovine obstetrics can be problematic. By selectively breeding ewes that produce multiple offspring with higher birth weights for generations, sheep producers have inadvertently caused some domestic sheep to have difficulty lambing; balancing ease of lambing with high productivity is one of the dilemmas of sheep breeding. In the case of any such problems, those present at lambing may assist the ewe by extracting or repositioning lambs. After the birth, ewes ideally break the amniotic sac (if it is not broken during labor), and begin licking clean the lamb. Most lambs will begin standing within an hour of birth. In normal situations, lambs nurse after standing, receiving vital colostrum milk. Lambs that either fail to nurse or are rejected by the ewe require help to survive, such as bottle-feeding or fostering by another ewe.
Most lambs begin life being born outdoors. After lambs are several weeks old, lamb marking (ear tagging, docking, mulesing, and castrating) is carried out. Vaccinations are usually carried out at this point as well. Ear tags with numbers are attached, or ear marks are applied, for ease of later identification of sheep. Docking and castration are commonly done after 24 hours (to avoid interference with maternal bonding and consumption of colostrum) and are often done not later than one week after birth, to minimize pain, stress, recovery time and complications. The first course of vaccinations (commonly anti-clostridial) is commonly given at an age of about 10 to 12 weeks; i.e. when the concentration of maternal antibodies passively acquired via colostrum is expected to have fallen low enough to permit development of active immunity. Ewes are often revaccinated annually about 3 weeks before lambing, to provide high antibody concentrations in colostrum during the first several hours after lambing. Ram lambs that will either be slaughtered or separated from ewes before sexual maturity are not usually castrated. Objections to all these procedures have been raised by animal rights groups, but farmers defend them by saying they save money, and inflict only temporary pain.
Homosexuality
Sheep are the only species of mammal except for humans which exhibits exclusive homosexual behavior. About 10% of rams refuse to mate with ewes but readily mate with other rams, and thirty percent of all rams demonstrate at least some homosexual behavior. Additionally, a small number of females that were accompanied by a male fetus in utero (i.e. as fraternal twins) are freemartins (female animals that are behaviorally masculine and lack functioning ovaries).
Health
Sheep may fall victim to poisons, infectious diseases, and physical injuries. As a prey species, a sheep's system is adapted to hide the obvious signs of illness, to prevent being targeted by predators. However, some signs of ill health are obvious, with sick sheep eating little, vocalizing excessively, and being generally listless. Throughout history, much of the money and labor of sheep husbandry has aimed to prevent sheep ailments. Historically, shepherds often created remedies by experimentation on the farm. In some developed countries, including the United States, sheep lack the economic importance for drug companies to perform expensive clinical trials required to approve more than a relatively limited number of drugs for ovine use. However, extra-label drug use in sheep production is permitted in many jurisdictions, subject to certain restrictions. In the US, for example, regulations governing extra-label drug use in animals are found in 21 CFR (Code of Federal Regulations) Part 530. In the 20th and 21st centuries, a minority of sheep owners have turned to alternative treatments such as homeopathy, herbalism and even traditional Chinese medicine to treat sheep veterinary problems. Despite some favorable anecdotal evidence, the effectiveness of alternative veterinary medicine has been met with skepticism in scientific journals. The need for traditional anti-parasite drugs and antibiotics is widespread, and is the main impediment to certified organic farming with sheep.
Many breeders take a variety of preventive measures to ward off problems. The first is to ensure all sheep are healthy when purchased. Many buyers avoid outlets known to be clearing houses for animals culled from healthy flocks as either sick or simply inferior. This can also mean maintaining a closed flock, and quarantining new sheep for a month. Two fundamental preventive programs are maintaining good nutrition and reducing stress in the sheep. Restraint, isolation, loud noises, novel situations, pain, heat, extreme cold, fatigue and other stressors can lead to secretion of cortisol, a stress hormone, in amounts that may indicate welfare problems. Excessive stress can compromise the immune system. "Shipping fever" (pneumonic mannheimiosis, formerly called pasteurellosis) is a disease of particular concern, that can occur as a result of stress, notably during transport and (or) handling. Pain, fear and several other stressors can cause secretion of epinephrine (adrenaline). Considerable epinephrine secretion in the final days before slaughter can adversely affect meat quality (by causing glycogenolysis, removing the substrate for normal post-slaughter acidification of meat) and result in meat becoming more susceptible to colonization by spoilage bacteria. Because of such issues, low-stress handling is essential in sheep management. Avoiding poisoning is also important; common poisons are pesticide sprays, inorganic fertilizer, motor oil, as well as radiator coolant containing ethylene glycol.
Common forms of preventive medication for sheep are vaccinations and treatments for parasites. Both external and internal parasites are the most prevalent malady in sheep, and are either fatal, or reduce the productivity of flocks. Worms are the most common internal parasites. They are ingested during grazing, incubate within the sheep, and are expelled through the digestive system (beginning the cycle again). Oral anti-parasitic medicines, known as drenches, are given to a flock to treat worms, sometimes after worm eggs in the feces has been counted to assess infestation levels. Afterwards, sheep may be moved to a new pasture to avoid ingesting the same parasites. External sheep parasites include: lice (for different parts of the body), sheep keds, nose bots, sheep itch mites, and maggots. Keds are blood-sucking parasites that cause general malnutrition and decreased productivity, but are not fatal. Maggots are those of the bot fly and the blow-fly, commonly Lucilia sericata or its relative L. cuprina. Fly maggots cause the extremely destructive condition of flystrike. Flies lay their eggs in wounds or wet, manure-soiled wool; when the maggots hatch they burrow into a sheep's flesh, eventually causing death if untreated. In addition to other treatments, crutching (shearing wool from a sheep's rump) is a common preventive method. Some countries allow mulesing, a practice that involves stripping away the skin on the rump to prevent fly-strike, normally performed when the sheep is a lamb. Nose bots are fly larvae that inhabit a sheep's sinuses, causing breathing difficulties and discomfort. Common signs are a discharge from the nasal passage, sneezing, and frantic movement such as head shaking. External parasites may be controlled through the use of backliners, sprays or immersive sheep dips.
A wide array of bacterial and viral diseases affect sheep. Diseases of the hoof, such as foot rot and foot scald may occur, and are treated with footbaths and other remedies. Foot rot is present in over 97% of flocks in the UK. These painful conditions cause lameness and hinder feeding. Ovine Johne's disease is a wasting disease that affects young sheep. Bluetongue disease is an insect-borne illness causing fever and inflammation of the mucous membranes. Ovine rinderpest (or peste des petits ruminants) is a highly contagious and often fatal viral disease affecting sheep and goats. Sheep may also be affected by primary or secondary photosensitization. Tetanus can also afflict sheep through wounds from shearing, docking, castration, or vaccination. The organism also can be introduced into the reproductive tract by unsanitary humans who assist ewes during lambing.
A few sheep conditions are transmissible to humans. Orf (also known as scabby mouth, contagious ecthyma or soremouth) is a skin disease leaving lesions that is transmitted through skin-to-skin contact. Cutaneous anthrax is also called woolsorter's disease, as the spores can be transmitted in unwashed wool. More seriously, the organisms that can cause spontaneous enzootic abortion in sheep are easily transmitted to pregnant women. Also of concern are the prion disease scrapie and the virus that causes foot-and-mouth disease (FMD), as both can devastate flocks. The latter poses a slight risk to humans. During the 2001 FMD pandemic in the UK, hundreds of sheep were culled and some rare British breeds were at risk of extinction due to this.
Of the 600,300 sheep lost to the US economy in 2004, 37.3% were lost to predators, while 26.5% were lost to some form of disease. Poisoning accounted for 1.7% of non-productive deaths.
Predators
A lamb being attacked by coyotes with a bite to the throat
Other than parasites and disease, predation is a threat to sheep and the profitability of sheep raising. Sheep have little ability to defend themselves, compared with other species kept as livestock. Even if sheep survive an attack, they may die from their injuries or simply from panic. However, the impact of predation varies dramatically with region. In Africa, Australia, the Americas, and parts of Europe and Asia predators are a serious problem. In the United States, for instance, over one third of sheep deaths in 2004 were caused by predation. In contrast, other nations are virtually devoid of sheep predators, particularly islands known for extensive sheep husbandry. Worldwide, canids—including the domestic dog—are responsible for most sheep deaths. Other animals that occasionally prey on sheep include: felines, bears, birds of prey, ravens and feral hogs.
Sheep producers have used a wide variety of measures to combat predation. Pre-modern shepherds used their own presence, livestock guardian dogs, and protective structures such as barns and fencing. Fencing (both regular and electric), penning sheep at night and lambing indoors all continue to be widely used. More modern shepherds used guns, traps, and poisons to kill predators, causing significant decreases in predator populations. In the wake of the environmental and conservation movements, the use of these methods now usually falls under the purview of specially designated government agencies in most developed countries.
The 1970s saw a resurgence in the use of livestock guardian dogs and the development of new methods of predator control by sheep producers, many of them non-lethal. Donkeys and guard llamas have been used since the 1980s in sheep operations, using the same basic principle as livestock guardian dogs. Interspecific pasturing, usually with larger livestock such as cattle or horses, may help to deter predators, even if such species do not actively guard sheep. In addition to animal guardians, contemporary sheep operations may use non-lethal predator deterrents such as motion-activated lights and noisy alarms.
Economic importance
Main article: Agricultural economics
Global sheep stock
in 2019
Number in millions
1. China163.5 (13.19%)
2. India74.3 (5.99%)
3. Australia65.8 (5.31%)
4. Nigeria46.9 (3.78%)
5. Iran41.3 (3.33%)
6. Sudan40.9 (3.3%)
7. Chad35.9 (2.9%)
8. Turkey35.2 (2.84%)
9. United Kingdom33.6 (2.71%)
10. Mongolia32.3 (2.61%)
World total1,239.8
Source: UN Food and Agriculture Organization
Sheep are an important part of the global agricultural economy. However, their once vital status has been largely replaced by other livestock species, especially the pig, chicken, and cow. China, Australia, India, and Iran have the largest modern flocks, and serve both local and exportation needs for wool and mutton. Other countries such as New Zealand have smaller flocks but retain a large international economic impact due to their export of sheep products. Sheep also play a major role in many local economies, which may be niche markets focused on organic or sustainable agriculture and local food customers. Especially in developing countries, such flocks may be a part of subsistence agriculture rather than a system of trade. Sheep themselves may be a medium of trade in barter economies.
Domestic sheep provide a wide array of raw materials. Wool was one of the first textiles, although in the late 20th century wool prices began to fall dramatically as the result of the popularity and cheap prices for synthetic fabrics. For many sheep owners, the cost of shearing is greater than the possible profit from the fleece, making subsisting on wool production alone practically impossible without farm subsidies. Fleeces are used as material in making alternative products such as wool insulation. In the 21st century, the sale of meat is the most profitable enterprise in the sheep industry, even though far less sheep meat is consumed than chicken, pork or beef.
Sheepskin is likewise used for making clothes, footwear, rugs, and other products. Byproducts from the slaughter of sheep are also of value: sheep tallow can be used in candle and soap making, sheep bone and cartilage has been used to furnish carved items such as dice and buttons as well as rendered glue and gelatin. Sheep intestine can be formed into sausage casings, and lamb intestine has been formed into surgical sutures, as well as strings for musical instruments and tennis rackets. Sheep droppings, which are high in cellulose, have even been sterilized and mixed with traditional pulp materials to make paper. Of all sheep byproducts, perhaps the most valuable is lanolin: the waterproof, fatty substance found naturally in sheep's wool and used as a base for innumerable cosmetics and other products.
Some farmers who keep sheep also make a profit from live sheep. Providing lambs for youth programs such as 4-H and competition at agricultural shows is often a dependable avenue for the sale of sheep. Farmers may also choose to focus on a particular breed of sheep in order to sell registered purebred animals, as well as provide a ram rental service for breeding. A new option for deriving profit from live sheep is the rental of flocks for grazing; these "mowing services" are hired in order to keep unwanted vegetation down in public spaces and to lessen fire hazard.
Despite the falling demand and price for sheep products in many markets, sheep have distinct economic advantages when compared with other livestock. They do not require expensive housing, such as that used in the intensive farming of chickens or pigs. They are an efficient use of land; roughly six sheep can be kept on the amount that would suffice for a single cow or horse. Sheep can also consume plants, such as noxious weeds, that most other animals will not touch, and produce more young at a faster rate. Also, in contrast to most livestock species, the cost of raising sheep is not necessarily tied to the price of feed crops such as grain, soybeans and corn. Combined with the lower cost of quality sheep, all these factors combine to equal a lower overhead for sheep producers, thus entailing a higher profitability potential for the small farmer. Sheep are especially beneficial for independent producers, including family farms with limited resources, as the sheep industry is one of the few types of animal agriculture that has not been vertically integrated by agribusiness. However, small flocks, from 10 to 50 ewes, often are not profitable because they tend to be poorly managed. The primary reason is that mechanization is not feasible, so return per hour of labor is not maximized. Small farm flocks generally are used simply to control weeds on irrigation ditches or maintained as a hobby.
Shoulder of lamb
Sheep meat and milk were one of the earliest staple proteins consumed by human civilization after the transition from hunting and gathering to agriculture. Sheep meat prepared for food is known as either mutton or lamb, and approximately 540 million sheep are slaughtered each year for meat worldwide. "Mutton" is derived from the Old French moton, which was the word for sheep used by the Anglo-Norman rulers of much of the British Isles in the Middle Ages. This became the name for sheep meat in English, while the Old English word sceap was kept for the live animal. Throughout modern history, "mutton" has been limited to the meat of mature sheep usually at least two years of age; "lamb" is used for that of immature sheep less than a year.
In the 21st century, the nations with the highest consumption of sheep meat are the Arab states of the Persian Gulf, New Zealand, Australia, Greece, Uruguay, the United Kingdom and Ireland. These countries eat 14–40 lbs (3–18 kg) of sheep meat per capita, per annum. Sheep meat is also popular in France, Africa (especially the Arab world), the Caribbean, the rest of the Middle East, India, and parts of China. This often reflects a history of sheep production. In these countries in particular, dishes comprising alternative cuts and offal may be popular or traditional. Sheep testicles—called animelles or lamb fries—are considered a delicacy in many parts of the world. Perhaps the most unusual dish of sheep meat is the Scottish haggis, composed of various sheep innards cooked along with oatmeal and chopped onions inside its stomach. In comparison, countries such as the U.S. consume only a pound or less (under 0.5 kg), with Americans eating 50 pounds (22 kg) of pork and 65 pounds (29 kg) of beef. In addition, such countries rarely eat mutton, and may favor the more expensive cuts of lamb: mostly lamb chops and leg of lamb.
Though sheep's milk may be drunk rarely in fresh form, today it is used predominantly in cheese and yogurt making. Sheep have only two teats, and produce a far smaller volume of milk than cows. However, as sheep's milk contains far more fat, solids, and minerals than cow's milk, it is ideal for the cheese-making process. It also resists contamination during cooling better because of its much higher calcium content. Well-known cheeses made from sheep milk include the feta of Bulgaria and Greece, Roquefort of France, Manchego from Spain, the pecorino romano (the Italian word for "sheep" is pecore) and ricotta of Italy. Yogurts, especially some forms of strained yogurt, may also be made from sheep milk. Many of these products are now often made with cow's milk, especially when produced outside their country of origin. Sheep milk contains 4.8% lactose, which may affect those who are intolerant.
As with other domestic animals, the meat of uncastrated males is inferior in quality, especially as they grow. A "bucky" lamb is a lamb which was not castrated early enough, or which was castrated improperly (resulting in one testicle being retained). These lambs are worth less at market.
In science
Sheep are generally too large and reproduce too slowly to make ideal research subjects, and thus are not a common model organism. They have, however, played an influential role in some fields of science. In particular, the Roslin Institute of Edinburgh, Scotland used sheep for genetics research that produced groundbreaking results. In 1995, two ewes named Megan and Morag were the first mammals cloned from differentiated cells, also referred to as gynomerogony. A year later, a Finnish Dorset sheep named Dolly, dubbed "the world's most famous sheep" in Scientific American, was the first mammal to be cloned from an adult somatic cell. Following this, Polly and Molly were the first mammals to be simultaneously cloned and transgenic.
As of 2008, the sheep genome has not been fully sequenced, although a detailed genetic map has been published, and a draft version of the complete genome produced by assembling sheep DNA sequences using information given by the genomes of other mammals. In 2012, a transgenic sheep named "Peng Peng" was cloned by Chinese scientists, who spliced his genes with that of a roundworm (C. elegans) in order to increase production of fats healthier for human consumption.
In the study of natural selection, the population of Soay sheep that remain on the island of Hirta have been used to explore the relation of body size and coloration to reproductive success. Soay sheep come in several colors, and researchers investigated why the larger, darker sheep were in decline; this occurrence contradicted the rule of thumb that larger members of a population tend to be more successful reproductively. The feral Soays on Hirta are especially useful subjects because they are isolated.
Domestic sheep are sometimes used in medical research, particularly for researching cardiovascular physiology, in areas such as hypertension and heart failure. Pregnant sheep are also a useful model for human pregnancy, and have been used to investigate the effects on fetal development of malnutrition and hypoxia. In behavioral sciences, sheep have been used in isolated cases for the study of facial recognition, as their mental process of recognition is qualitatively similar to humans.
Cultural impact
Sheep have had a strong presence in many cultures, especially in areas where they form the most common type of livestock. In the English language, to call someone a sheep or ovine may allude that they are timid and easily led. In contradiction to this image, male sheep are often used as symbols of virility and power; the logos of the Los Angeles Rams football team and the Dodge Ram pickup truck allude to males of the bighorn sheep, Ovis canadensis.
Counting sheep is popularly said to be an aid to sleep, and some ancient systems of counting sheep persist today. Sheep also enter in colloquial sayings and idiom frequently with such phrases as "black sheep". To call an individual a black sheep implies that they are an odd or disreputable member of a group. This usage derives from the recessive trait that causes an occasional black lamb to be born into an entirely white flock. These black sheep were considered undesirable by shepherds, as black wool is not as commercially viable as white wool. Citizens who accept overbearing governments have been referred to by the Portmanteau neologism of sheeple. Somewhat differently, the adjective "sheepish" is also used to describe embarrassment.
In heraldry
In British heraldry, sheep appear in the form of rams, sheep proper and lambs. These are distinguished by the ram being depicted with horns and a tail, the sheep with neither and the lamb with its tail only. A further variant of the lamb, termed the Paschal lamb, is depicted as carrying a Christian cross and with a halo over its head. Rams' heads, portrayed without a neck and facing the viewer, are also found in British armories. The fleece, depicted as an entire sheepskin carried by a ring around its midsection, originally became known through its use in the arms of the Order of the Golden Fleece and was later adopted by towns and individuals with connections to the wool industry. A sheep on a blue field is depicted on the greater/royal arms of the king of Denmark to represent the Faroe Islands. In 2004 a modernized arms has been adopted by the Faroe Islands, which based on a 15th century coat of arms.
Religion and folklore
In antiquity, symbolism involving sheep cropped up in religions in the ancient Near East, the Mideast, and the Mediterranean area: Çatalhöyük, ancient Egyptian religion, the Cana'anite and Phoenician tradition, Judaism, Greek religion, and others. Religious symbolism and ritual involving sheep began with some of the first known faiths: Skulls of rams (along with bulls) occupied central placement in shrines at the Çatalhöyük settlement in 8,000 BCE. In Ancient Egyptian religion, the ram was the symbol of several gods: Khnum, Heryshaf and Amun (in his incarnation as a god of fertility). Other deities occasionally shown with ram features include the goddess Ishtar, the Phoenician god Baal-Hamon, and the Babylonian god Ea-Oannes. In Madagascar, sheep were not eaten as they were believed to be incarnations of the souls of ancestors.
There are many ancient Greek references to sheep: that of Chrysomallos, the golden-fleeced ram, continuing to be told through into the modern era. Astrologically, Aries, the ram, is the first sign of the classical Greek zodiac, and the sheep is the eighth of the twelve animals associated with the 12-year cycle of in the Chinese zodiac, related to the Chinese calendar. It is said in Chinese traditions that Hou ji sacrificed sheep. Mongolia, shagai are an ancient form of dice made from the cuboid bones of sheep that are often used for fortunetelling purposes.
Sheep play an important role in all the Abrahamic faiths; Abraham, Isaac, Jacob, Moses, and King David were all shepherds. According to the Biblical story of the Binding of Isaac, a ram is sacrificed as a substitute for Isaac after an angel stays Abraham's hand (in the Islamic tradition, Abraham was about to sacrifice Ishmael). Eid al-Adha is a major annual festival in Islam in which sheep (or other animals) are sacrificed in remembrance of this act. Sheep are occasionally sacrificed to commemorate important secular events in Islamic cultures. Greeks and Romans sacrificed sheep regularly in religious practice, and Judaism once sacrificed sheep as a Korban (sacrifice), such as the Passover lamb. Ovine symbols—such as the ceremonial blowing of a shofar—still find a presence in modern Judaic traditions.
Collectively, followers of Christianity are often referred to as a flock, with Christ as the Good Shepherd, and sheep are an element in the Christian iconography of the birth of Jesus. Some Christian saints are considered patrons of shepherds, and even of sheep themselves. Christ is also portrayed as the Sacrificial lamb of God (Agnus Dei) and Easter celebrations in Greece and Romania traditionally feature a meal of Paschal lamb. A church leader is often called the pastor, which is derived from the Latin word for shepherd. In many western Christian traditions bishops carry a staff, which also serves as a symbol of the episcopal office, known as a crosier, which is modeled on the shepherd's crook.
Sheep are key symbols in fables and nursery rhymes like The Wolf in Sheep's Clothing, Little Bo Peep, Baa, Baa, Black Sheep, and Mary Had a Little Lamb; novels such as George Orwell's Animal Farm and Haruki Murakami's A Wild Sheep Chase; songs such as Bach's Sheep may safely graze (Schafe können sicher weiden) and Pink Floyd's "Sheep", and poems like William Blake's "The Lamb".
The coastline of Norfolk, which is skull-shaped, protrudes into the North Sea - where it is exposed to the full force of wind and wave. Running for nearly 100 miles, it consists of a wide variety of terrains including: cliffs, beaches, sand dunes, salt-marshes, mudflats, shingle banks and spits. Over the centuries it has inspired countless writers - with its constantly changing light, its unpredictable tides and its bleak beauty. Even today, despite an influx of tourists, it still contains a wild and unspoilt quality.
On its western edge, it begins on the remote mudflats of the Wash - before heading eastwards towards the town of Kings Lynn. Steeped in maritime history, Lynn is one of Norfolk's best kept secrets. Once part of the Hanseatic League it possesses a wealth of historic buildings - most of them clustered around the quayside. In his poem King's Lynn R.N. Curry invites us to: 'Explore now, as a freak wave might explore/This town that history could have made a city'.
From Lynn the coastline veers northwards along the eastern side of the Wash past the ancient town of Castle Rising and the Queen's favourite retreat at Sandringham to Snettisham - famous for its RSPB reserve. The Manx poet David Callin visited the area and wrote a moving poem called Snettisham which begins:
This is the Wash it seems -
a last exhalation
of the dying land, or something
the sea's been working on
for ages: sketching it in,
rubbing it out,
redoing and redoing it,
never satisfied.
At Hunstanton, cliffs appear for the first time - but these are no ordinary cliffs - but strikingly striped ones composed of red and white chalk and carr-stone. L.P. Hartley used them as a backdrop in his novel The Shrimp and the Anemone and in Patrick Hamilton's novel Rope the first chapter opens with George Harvey Bone walking along them. And according to the poet Andrew McDonnell, Hamilton's fictional character is still haunting the town today - see George Harvey Bone.
Rounding the north western tip of the county - the cliffs subside - and the sand dunes and beaches of Holme-next-the-Sea take over. This is a location familiar to the nature writer Patrick Barkham who used to come with his father on family holidays. Together they would search for butterflies - an activity which sparked a life-long interest and ultimately resulted in his book The Butterfly Isles (2010).
Sheltering behind Scolt Head Island lies Brancaster - which was once a Roman settlement. Lord Nelson learnt to sail in the creeks here and the stunning beach inspired both the novelist Angela Huth in Invitation to the Married Life and also the poet John Gallas in his poem Sunrise on Brancaster Beach where he notes: 'The sea looks white,/ like gnawed skin; the wrinkled sand feels cold.'
Burnham Overy Staithe, where the tiny River Burn reaches the sea, has always provided inspiration for the poet and children's writer Kevin Crossley-Holland - particularly in the 25 poems which make up his Waterslain collection where he depicts characters from the village.
Heading along the coast road, we then pass the Holkham estate - with its magnificent Paladian mansion and landscaped parkland. Another spectacular beach here was used by filmmakers in Shakespeare in Love and the wild swimmer Roger Deakin took a dip here and recorded it in his delightful book Waterlog.
Entering the narrow, flint-walled street of Stiffkey - history seems palpable. It was here in 1937 that the writer Henry Williamson tried his hand at farming and recorded the ups and downs in The Story of a Norfolk Farm. While in the county, he also collaborated with Lilias Rider Haggard, the daughter of Henry Rider Haggard, to produce the ever-green Norfolk Life.
Next comes the tiny village of Morston - famous today for its seal tours and fine food. The poet Edwin Brock (who lived at Low Tharston next door to another poet Anthony Thwaite) wrote a beautifully haunting poem called Morston Marshes - which captures the transience of the tides and the isolated beauty of the place.
Into this muddy coastline
the North Sea seeps silently
twice a day
under the kestrel's weather eye
in the growing puddles
gulls drill the marsh
for nothing we can see
or screech their territory
like fishwives
from the tops of poles
At Blakeney the famous four-mile long sand spit known as Blakeney Point curves round protecting an area of dunes and salt marshes from the ravages of the North Sea. The town itself was once a wealthy port, but declined when its harbour silted up. The poet Thomas Thornely wrote a fine lament about the loss of its trade entitled Blakeney.
The author Jack Higgins stayed at the Blakeney Hotel in the 1970s while researching and writing his WW2 novel The Eagle Has Landed about the assassination of Winston Churchill. He drew on a number of locations - including St Mary's Church at Cley - for his fictional Studley Constable.
The charming village of Cley-next-the-Sea is renowned for its nature reserve, its church and its splendid windmill. Many bird watchers come here and stay at the George Hotel. My own poem Ornithologist hopefully captures something of the spirit of the place.
At Salthouse a fragile shingle bank protects the village from flooding - but it is regularly breached - as in 1953 and 2014. The precarious existence of the villagers is captured in a bleak but beautiful poem by John Press entitled Salthouse, Norfolk. The village originally got its name from the salt it produced and exported to the continent.
A short walk along The Quag, past Kelling Hard, brings you to the steep, shingle beach at Weybourne where cliffs suddenly reappear. Composed of sand and gravel deposited by a melting glacier - they offer little resistance against the onslaught of the North Sea. In fact, when John Taylor the Water Poet landed in Cromer in 1622 as part of his Very Merry - Wherry - Ferry Voyage he wrote:
It is an ancient market town that stands
Upon a lofty cliff of mouldering sands;
The sea against the cliffs doth daily beat,
And every tide into the land doth eat.'
Cromer has influenced many other writers over the years including Jane Austen who mentions it in Emma and Oscar Wilde who stayed at the Hotel de Paris while working on A Woman of No Importance. The town has inspired a good deal of poetry including George Barker's dramatic On A Friend's Escape from Drowning Off the Norfolk Coast. However, it's greatest claim to fame is probably that Arthur Conan Doyle found inspiration here (at Cromer Hall) for Baskerville Hall in his classic Hound of the Baskervilles adventure.
George Barker was also a regular visitor to nearby Overstrand and used it as a location for his long memorial poem entitled In Memory of David Archer.
The church at Sidestrand holds a secret. Despite appearances, it is not in its original location but was moved back from the cliff edge and rebuilt by the Victorians. However, the original cliff-top graveyard inspired the London theatre critic Clement Scott to compose his The Garden of Sleep. The poem contained the term Poppyland - which has come to describe this whole section of the Norfolk Coast. Scott helped to popularise both Cromer and Sidestrand and other affluent Victorian tourists soon followed including the sea-loving poet A.C. Swinburne.
Mundesley, with its flint and brick cottages, still retains considerable charm. It was here that the troubled poet William Cowper came in 1795 to seek refuge from his depression and stayed in a house which now bears his name.
At Bacton the cliffs subside - but then reappear for one final time at Happisburgh. Stuck on an exposed section of the coast, Happisburgh has born the brunt of coastal erosion for many years. Despite groynes and rocks and other sea defence schemes large chunks of land have been disappearing. The houses on Beach Road were demolished in 2012 and even the doughty Mrs Nierop-Reading who stayed on finally had to abandon her beloved cliff-top chalet in 2013.
Beyond Happisburgh lies Cart Gap and the lost village of Eccles-on-Sea. For many years Eccles church tower stood on the beach before finally being washed away by the sea. The village is commemorated by Anthony Thwaite's fine poem Eccles.
From Eccles onwards the Norfolk coast is dominated by sand dunes. Held together by marram grass - the dunes provide a barrier of sorts against the sea but are always vulnerable to breaches. The worst of these breaches in recent history occurred in 1953 when the sea burst through at Sea Palling destroying homes, inundating farmland and killing seven people. Today the sea remains a constant threat to this part of Norfolk:
The sea will always triumph
Tomorrow or today
Despite Norwegian rocks here
Will always find a way
For the sea knows no time
Only deepness and cold
No clocks or watches
For the sea is very old
Horsey, with its iconic windmill is another location under threat from the sea. The poet John Betjeman knew Horsey well from boating holidays on the Broads and it features in his East Anglian Bathe where he recalls walking from the staithe, swimming in the sea and Horsey Mere.
Oh when the early morning at the seaside
Took us with hurrying steps from Horsey Mere
To see the whistling bent-grass on the leeside
And then the tumbled breaker-line appear,
Winterton has always had a reputation as a dangerous place for shipping. The author Daniel Defoe dedicates a long section to it in his A Tour Through the Whole Island of Great Britain and it is off Winterton Ness that Robinson Crusoe experiences his first shipwreck.
Beyond Winterton lie the seaside resorts of Hemsby, Scratby and California with their holiday chalets and caravans and amusement arcades. Then comes Caister-on-Sea - where the Paston family had their castle and where in the 15th Century the stout-hearted Margaret Paston saw off an armed raid while her husband was in London.
The last large town on the Norfolk coast is Great Yarmouth. Today, it is a holiday destination - famous for its Pleasure Beach and attractions - but not so long ago it was a thriving fishing port where herring boats packed the quaysides. And it was the smell of fish which permeated the air when David Copperfield first arrives here - in Charles Dickens' famous autobiographical novel.
'When we got into the street (which was strange enough to me), and smelt the fish, and pitch, and oakum, and tar, and saw the sailors walking about, and the carts jangling up and down over the stones, I felt I had done so busy a place an injustice; and said as much to Peggotty, who heard my expressions of delight with great complacency, and told me it was well known (I suppose to those who had the good fortune to be born Bloaters) that Yarmouth was, upon the whole, the finest place in the universe.'
On the other side of the River Yare the coastline arrives at Gorleston - which was once part of Suffolk. From here sandy beaches run next to the promenade and the links course until we reach Hopton and the start of Suffolk proper. Behind us now lies the entire curve of Norfolk - like a 'land of lost content'.
"Now I'm smoking cigarettes
and I strive for purity,
and I slip just like the stars into obscurity.
'Cause every time I hear that melody,
well, it puts me up a tree,
and the grapefruit moon,
one star shining,
is all that I can see...."
- Tom Waits
'Grapefruit Moon'
Squid are cephalopods in the superorder Decapodiformes with elongated bodies, large eyes, eight arms and two tentacles. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.
Squid diverged from other cephalopods during the Jurassic and occupy a similar role to teleost fish as open water predators of similar size and behaviour. They play an important role in the open water food web. The two long tentacles are used to grab prey and the eight arms to hold and control it. The beak then cuts the food into suitable size chunks for swallowing. Squid are rapid swimmers, moving by jet propulsion, and largely locate their prey by sight. They are among the most intelligent of invertebrates, with groups of Humboldt squid having been observed hunting cooperatively. They are preyed on by sharks, other fish, sea birds, seals and cetaceans, particularly sperm whales.
Squid can change colour for camouflage and signalling. Some species are bioluminescent, using their light for counter-illumination camouflage, while many species can eject a cloud of ink to distract predators.
Squid are used for human consumption with commercial fisheries in Japan, the Mediterranean, the southwestern Atlantic, the eastern Pacific and elsewhere. They are used in cuisines around the world, often known as "calamari". Squid have featured in literature since classical times, especially in tales of giant squid and sea monsters.
TAXONOMY AND PHYLOGENY
Squid are members of the class Cephalopoda, subclass Coleoidea. The squid orders Myopsida and Oegopsida are in the superorder Decapodiformes (from the Greek for "ten-legged"). Two other orders of decapodiform cephalopods are also called squid, although they are taxonomically distinct from squids and differ recognizably in their gross anatomical features. They are the bobtail squid of order Sepiolida and the ram's horn squid of the monotypic order Spirulida. The vampire squid, however, is more closely related to the octopuses than to any squid.
The cladogram, not fully resolved, is based on Sanchez et al, 2018. Their molecular phylogeny used mitochondrial and nuclear DNA marker sequences; they comment that a robust phylogeny "has proven very challenging to obtain". If it is accepted that Sepiidae cuttlefish are a kind of squid, then the squids, excluding the vampire squid, form a clade as illustrated. Orders are shown in boldface; all the families not included in those orders, except Sepiadariidae and Sepiidae are in the paraphyletic order "Sepiida", are in the paraphyletic order "Oegopsida".
EVOLUTION
Crown coleoids (the ancestors of octopuses and squid) diverged at the end of the Paleozoic, in the Permian. Squid diverged during the Jurassic, but many squid families appeared in or after the Cretaceous. Both the coleoids and the teleost fish were involved in much adaptive radiation at this time, and the two modern groups resemble each other in size, ecology, habitat, morphology and behaviour, however some fish moved into fresh water while the coleoids remained in marine environments.
The ancestral coleoid was probably nautiloid-like with a strait septate shell that became immersed in the mantle and was used for buoyancy control. Four lines diverged from this, Spirulida (with one living member), the cuttlefishes, the squids and the octopuses. Squid have differentiated from the ancestral mollusc such that the body plan has been condensed antero-posteriorly and extended dorso-ventrally. What may have been the foot of the ancestor is modified into a complex set of appendages around the mouth. The sense organs are highly developed and include advanced eyes similar to those of vertebrates.
The ancestral shell has been lost, with only an internal gladius, or pen, remaining. The pen, made of a chitin-like material, is a feather-shaped internal structure that supports the squid's mantle and serves as a site for muscle attachment. The cuttlebone or sepion of the Sepiidae is calcareous and appears to have evolved afresh in the Tertiary
DESCIPTION
Squid are soft-bodied molluscs whose forms evolved to adopt an active predatory lifestyle. The head and foot of the squid are at one end of a long body, and this end is functionally anterior, leading the animal as it moves through the water. A set of eight arms and two distinctive tentacles surround the mouth; each appendage takes the form of a muscular hydrostat and is flexible and prehensile, usually bearing disc-like suckers.
The suckers may lie directly on the arm or be stalked. Their rims are stiffened with chitin and may contain minute toothlike denticles. These features, as well as strong musculature, and a small ganglion beneath each sucker to allow individual control, provide a very powerful adhesion to grip prey. Hooks are present on the arms and tentacles in some species, but their function is unclear. The two tentacles are much longer than the arms and are retractile. Suckers are limited to the spatulate tip of the tentacle, known as the manus.
In the mature male, the outer half of one of the left arms is hectocotylised – and ends in a copulatory pad rather than suckers. This is used for depositing a spermatophore inside the mantle cavity of a female. A ventral part of the foot has been converted into a funnel through which water exits the mantle cavity.
The main body mass is enclosed in the mantle, which has a swimming fin along each side. These fins are not the main source of locomotion in most species. The mantle wall is heavily muscled and internal. The visceral mass, which is covered by a thin, membranous epidermis, forms a cone-shaped posterior region known as the "visceral hump". The mollusc shell is reduced to an internal, longitudinal chitinous "pen" in the functionally dorsal part of the animal; the pen acts to stiffen the squid and provides attachments for muscles.
On the functionally ventral part of the body is an opening to the mantle cavity, which contains the gills (ctenidia) and openings from the excretory, digestive and reproductive systems. An inhalant siphon behind the funnel draws water into the mantel cavity via a valve. The squid uses the funnel for locomotion via precise jet propulsion. In this form of locomotion, water is sucked into the mantle cavity and expelled out of the funnel in a fast, strong jet. The direction of travel is varied by the orientation of the funnel. Squid are strong swimmers and certain species can "fly" for short distances out of the water.
CAMOUFLAGE
Squid make use of different kinds of camouflage, namely active camouflage for background matching (in shallow water) and counter-illumination. This helps to protect them from their predators and allows them to approach their prey.
The skin is covered in controllable chromatophores of different colours, enabling the squid to match its coloration to its surroundings. The play of colours may in addition distract prey from the squid's approaching tentacles. The skin also contains light reflectors called iridophores and leucophores that, when activated, in milliseconds create changeable skin patterns of polarized light. Such skin camouflage may serve various functions, such as communication with nearby squid, prey detection, navigation, and orientation during hunting or seeking shelter. Neural control of the iridophores enabling rapid changes in skin iridescence appears to be regulated by a cholinergic process affecting reflectin proteins.
Some mesopelagic squid such as the firefly squid (Watasenia scintillans) and the midwater squid (Abralia veranyi) use counter-illumination camouflage, generating light to match the downwelling light from the ocean surface. This creates the effect of countershading, making the underside lighter than the upperside.
Counter-illumination is also used by the Hawaiian bobtail squid (Euprymna scolopes), which has symbiotic bacteria (Aliivibrio fischeri) that produce light to help the squid avoid nocturnal predators. This light shines through the squid's skin on its underside and is generated by a large and complex two-lobed light organ inside the squid's mantle cavity. From there, it escapes downwards, some of it travelling directly, some coming off a reflector at the top of the organ (dorsal side). Below there is a kind of iris, which has branches (diverticula) of its ink sac, with a lens below that; both the reflector and lens are derived from mesoderm. The squid controls light production by changing the shape of its iris or adjusting the strength of yellow filters on its underside, which presumably change the balance of wavelengths emitted. Light production shows a correlation with intensity of down-welling light, but it is about one third as bright; the squid can track repeated changes in brightness. Because the Hawaiian bobtail squid hides in sand during the day to avoid predators, it does not use counter-illumination during daylight
PREDATOR DISTRACTION WITH INK
Squid distract attacking predators by ejecting a cloud of ink, giving themselves an opportunity to escape. The ink gland and its associated ink sac empties into the rectum close to the anus, allowing the squid to rapidly discharge black ink into the mantle cavity and surrounding water. The ink is a suspension of melanin particles and quickly disperses to form a dark cloud that obscures the escape manoeuvres of the squid. Predatory fish may also be deterred by the alkaloid nature of the discharge which may interfere with their chemoreceptors.
NERVOUS SYSTEM AND SENSE ORGANS
Cephalopods have the most highly developed nervous systems among invertebrates. Squids have a complex brain in the form of a nerve ring encircling the oesophagus, enclosed in a cartilaginous cranium. Paired cerebral ganglia above the oesophagus receive sensory information from the eyes and statocysts, and further ganglia below control the muscles of the mouth, foot, mantle and viscera. Giant axons up to 1 mm in diameter convey nerve messages with great rapidity to the circular muscles of the mantle wall, allowing a synchronous, powerful contraction and maximum speed in the jet propulsion system.
The paired eyes, on either side of the head, are housed in capsules fused to the cranium. Their structure is very similar to that of a fish eye, with a globular lens that has a depth of focus from 3 cm to infinity. The image is focused by changing the position of the lens, as in a camera or telescope, rather than changing the shape of the lens, as in the human eye. Squid adjust to changes in light intensity by expanding and contracting the slit-shaped pupil. Deep sea squids in the family Histioteuthidae have eyes of two different types and orientation. The large left eye is tubular in shape and looks upwards, presumably searching for the silhouettes of animals higher in the water column. The normally-shaped right eye points forwards and downwards to detect prey.
The statocysts are involved in maintaining balance and are analogous to the inner ear of fish. They are housed in cartilaginous capsules on either side of the cranium. They provide the squid with information on its body position in relation to gravity, its orientation, acceleration and rotation, and are able to perceive incoming vibrations. Without the statocysts, the squid cannot maintain equilibrium. Squid appear to have limited hearing, but the head and arms bear lines of hair-cells that are weakly sensitive to water movements and changes in pressure, and are analogous in function to the lateral line system of fish.
REPRODUCTIVE SYSTEM
The sexes are separate in squid, there being a single gonad in the posterior part of the body with fertilisation being external, and usually taking place in the mantle cavity of the female. The male has a testis from which sperm pass into a single gonoduct where they are rolled together into a long bundle, or spermatophore. The gonoduct is elongated into a "penis" that extends into the mantle cavity and through which spermatophores are ejected. In shallow water species, the penis is short, and the spermatophore is removed from the mantle cavity by a tentacle of the male, which is specially adapted for the purpose and known as a hectocotylus, and placed inside the mantle cavity of the female during mating.The female has a large translucent ovary, situated towards the posterior of the visceral mass. From here, eggs travel along the gonocoel, where there are a pair of white nidamental glands, which lie anterior to the gills. Also present are red-spotted accessory nidamental glands containing symbiotic bacteria; both organs are associated with nutrient manufacture and forming shells for the eggs. The gonocoel enters the mantle cavity at the gonopore, and in some species, receptacles for storing spermatophores are located nearby, in the mantle wall. In shallow-water species of the continental shelf and epipelagic or mesopelagic zones, it is frequently one or both of arm pair IV of males that are modified into hectocotyli. However, most deep-sea squid lack hectocotyl arms and have longer penises; Ancistrocheiridae and Cranchiinae are exceptions. Giant squid of the genus Architeuthis are unusual in that they possess both a large penis and modified arm tips, although whether the latter are used for spermatophore transfer is uncertain. Penis elongation has been observed in the deep-water species Onykia ingens; when erect, the penis may be as long as the mantle, head, and arms combined. As such, deep-water squid have the greatest known penis length relative to body size of all mobile animals, second in the entire animal kingdom only to certain sessile barnacles.
DIGESTIVE SYSTEM
Like all cephalopods, squids are predators and have complex digestive systems. The mouth is equipped with a sharp, horny beak mainly made of chitin and cross-linked proteins, which is used to kill and tear prey into manageable pieces. The beak is very robust, but does not contain minerals, unlike the teeth and jaws of many other organisms; the cross-linked proteins are histidine- and glycine-rich and give the beak a stiffness and hardness greater than most equivalent synthetic organic materials. The stomachs of captured whales often have indigestible squid beaks inside. The mouth contains the radula, the rough tongue common to all molluscs except bivalvia, which is equipped with multiple rows of teeth.[6] In some species, toxic saliva helps to control large prey; when subdued, the food can be torn in pieces by the beak, moved to the oesophagus by the radula, and swallowed.
The food bolus is moved along the gut by waves of muscular contractions (peristalsis). The long oesophagus leads to a muscular stomach roughly in the middle of the visceral mass. The digestive gland, which is equivalent to a vertebrate liver, diverticulates here, as does the pancreas, and both of these empty into the caecum, a pouch-shaped sac where most of the absorption of nutrients takes place. Indigestible food can be passed directly from the stomach to the rectum where it joins the flow from the caecum and is voided through the anus into the mantle cavity. Cephalopods are short-lived, and in mature squid, priority is given to reproduction; the female Onychoteuthis banksii for example, sheds its feeding tentacles on reaching maturity, and becomes flaccid and weak after spawning.
CARDIOVASCULAR AND EXCRETORY SYSTEMS
The squid mantle cavity is a seawater-filled sac containing three hearts and other organs supporting circulation, respiration, and excretion. Squid have a main systemic heart that pumps blood around the body as part of the general circulatory system, and two branchial hearts. The systemic heart consists of three chambers, a lower ventricle and two upper atria, all of which can contract to propel the blood. The branchial hearts pump blood specifically to the gills for oxygenation, before returning it to the systemic heart. The blood contains the copper-rich protein hemocyanin, which is used for oxygen transport at low ocean temperatures and low oxygen concentrations, and makes the oxygenated blood a deep, blue color. As systemic blood returns via two vena cavae to the branchial hearts, excretion of urine, carbon dioxide, and waste solutes occurs through outpockets (called nephridial appendages) in the vena cavae walls that enable gas exchange and excretion via the mantle cavity seawater.
BUOYANCY
Unlike nautiloids which have gas-filled chambers inside their shells which provide buoyancy, and octopuses which live near and rest on the seabed and do not require to be buoyant, many squid have a fluid-filled receptacle, equivalent to the swim bladder of a fish, in the coelom or connective tissue. This reservoir acts as a chemical buoyancy chamber, with the heavy metallic cations typical of seawater replaced by low molecular-weight ammonium ions, a product of excretion. The small difference in density provides a small contribution to buoyancy per unit volume, so the mechanism requires a large buoyancy chamber to be effective. Since the chamber is filled with liquid, it has the advantage over a swim bladder of not changing significantly in volume with pressure. Glass squids in the family Cranchiidae for example, have an enormous transparent coelom containing ammonium ions and occupying about two-thirds the volume of the animal, allowing it to float at the required depth. About half of the 28 families of squid use this mechanism to solve their buoyancy issues.
LARGEST AND SMALLEST
The majority of squid are no more than 60 cm long, although the giant squid may reach 13 m. The smallest species are probably the benthic pygmy squids Idiosepius, which grow to a mantle length of 10 to 18 mm, and have short bodies and stubby arms.
In 1978, sharp, curved claws on the suction cups of squid tentacles cut up the rubber coating on the hull of the USS Stein. The size suggested the largest squid known at the time.
In 2003, a large specimen of an abundant but poorly understood species, Mesonychoteuthis hamiltoni (the colossal squid), was discovered. This species may grow to 10 m in length, making it the largest invertebrate. In February 2007, a New Zealand fishing vessel caught the largest squid ever documented, weighing 495 kg and measuring around 10 m off the coast of Antarctica. Dissection showed that the eyes, used to detect prey in the deep Southern Ocean, exceeded the size of footballs; these may be among the largest eyes ever to exist in the animal kingdom.
DEVELOPMENT
The eggs of squid are large for a mollusc, containing a large amount of yolk to nourish the embryo as it develops directly, without an intervening veliger larval stage. The embryo grows as a disc of cells on top of the yolk. During the gastrulation stage, the margins of the disc grow to surround the yolk, forming a yolk sac, which eventually forms part of the animal's gut. The dorsal side of the disc grows upwards and forms the embryo, with a shell gland on its dorsal surface, gills, mantle and eyes. The arms and funnel develop as part of the foot on the ventral side of the disc. The arms later migrate upwards, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo grows. Some juvenile squid live higher in the water column than do adults. Squids tend to be short-lived; Loligo for example lives from one to three years according to species, typically dying soon after spawning.
n a well-studied bioluminescent species, the Hawaiian bobtail squid, a special light organ in the squid's mantle is rapidly colonized with Aliivibrio fischeri bacteria within hours of hatching. This light-organ colonization requires this particular bacterial species for a symbiotic relationship; no colonization occurs in the absence of A. fischeri. Colonization occurs in a horizontal manner, such that the hosts acquires its bacterial partners from the environment. The symbiosis is obligate for the squid, but facultative for the bacteria. Once the bacteria enter the squid, they colonize interior epithelial cells in the light organ, living in crypts with complex microvilli protrusions. The bacteria also interact with hemocytes, macrophage-like blood cells that migrate between epithelial cells, but the mechanism and function of this process is not well understood. Bioluminescence reaches its highest levels during the early evening hours and bottoms out before dawn; this occurs because at the end of each day, the contents of the squid's crypts are expelled into the surrounding environment. About 95% of the bacteria are voided each morning before the bacterial population builds up again by nightfall.
BEHAVIOUR
LOCOMOTION
Squid can move about in several different ways. Slow movement is achieved by a gentle undulation of the muscular lateral fins on either side of the trunk which drives the animal forward. A more common means of locomotion providing sustained movement is achieved using jetting, during which contraction of the muscular wall of the mantle cavity provides jet propulsion.
Slow jetting is used for ordinary locomotion, and ventilation of the gills is achieved at the same time. The circular muscles in the mantle wall contract; this causes the inhalant valve to close, the exhalant valve to open and the mantle edge to lock tightly around the head. Water is forced out through the funnel which is pointed in the opposite direction to the required direction of travel. The inhalant phase is initiated by the relaxation of the circular muscles causes them to stretch, the connective tissue in the mantle wall recoils elastically, the mantle cavity expands causing the inhalant valve to open, the exhalant valve to close and water to flow into the cavity. This cycle of exhalation and inhalation is repeated to provide continuous locomotion.
Fast jetting is an escape response. In this form of locomotion, radial muscles in the mantle wall are involved as well as circular ones, making it possible to hyper-inflate the mantle cavity with a larger volume of water than during slow jetting. On contraction, water flows out with great force, the funnel always being pointed anteriorly, and travel is backwards. During this means of locomotion, some squid exit the water in a similar way to flying fish, gliding through the air for up to 50 m, and occasionally ending up on the decks of ships.
FEEDING
Squid are carnivores, and, with their strong arms and suckers, can overwhelm relatively large animals efficiently. Prey is identified by sight or by touch, grabbed by the tentacles which can be shot out with great rapidity, brought back to within reach of the arms, and held by the hooks and suckers on their surface. In some species, the squid's saliva contains toxins which act to subdue the prey. These are injected into its bloodstream when the prey is bitten, along with vasodilators and chemicals to stimulate the heart, and quickly circulate to all parts of its body. The deep sea squid Taningia danae has been filmed releasing blinding flashes of light from large photophores on its arms to illuminate and disorientate potential prey.
Although squid can catch large prey, the mouth is relatively small, and the food must be cut into pieces by the chitinous beak with its powerful muscles before being swallowed. The radula is located in the buccal cavity and has multiple rows of tiny teeth that draw the food backwards and grind it in pieces. The deep sea squid Mastigoteuthis has the whole length of its whip-like tentacles covered with tiny suckers; it probably catches small organisms in the same way that flypaper traps flies. The tentacles of some bathypelagic squids bear photophores which may bring food within its reach by attracting prey.
Squid are among the most intelligent invertebrates. For example, groups of Humboldt squid hunt cooperatively, spiralling up through the water at night and coordinating their vertical and horizontal movements while foraging.
REPRODUCTION
Courtship in squid takes place in the open water and involves the male selecting a female, the female responding, and the transfer by the male of spermatophores to the female. In many instances, the male may display to identify himself to the female and drive off any potential competitors.[46] Elaborate changes in body patterning take place in some species in both agonistic and courtship behaviour. The Caribbean reef squid (Sepioteuthis sepioidea), for example, employs a complex array of colour changes during courtship and social interactions and has a range of about 16 body patterns in its repertoire.
The pair adopt a head-to-head position, and "jaw locking" may take place, in a similar manner to that adopted by some cichlid fish. The heterodactylus of the male is used to transfer the spermatophore and deposit it in the female's mantle cavity in the position appropriate for the species; this may be adjacent to the gonopore or in a seminal receptacle.
The sperm may be used immediately or may be stored. As the eggs pass down the oviduct, they are wrapped in a gelatinous coating, before continuing to the mantle cavity, where they are fertilised. In Loligo, further coatings are added by the nidimental glands in the walls of the cavity and the eggs leave through a funnel formed by the arms. The female attaches them to the substrate in strings or groups, the coating layers swelling and hardening after contact with sea water. Loligo sometimes forms breeding aggregations which may create a "community pile" of egg strings. Some pelagic and deep sea squid do not attach their egg masses, which float freely.
ECOLOGY
Squid mostly have an annual life cycle, growing fast and dying soon after spawning. The diet changes as they grow but mostly consists of large zooplankton and small nekton. In Antarctica for example, krill is the main constituent of the diet, with other food items being amphipods, other small crustaceans, and large arrow worms. Fish are also eaten, and some squid are cannibalistic.
As well as occupying a key role in the food chain, squid are an important prey for predators including sharks, sea birds, seals and whales. Juvenile squid provide part of the diet for worms and small fish. When researchers studied the contents of the stomachs of elephant seals in South Georgia, they found 96% squid by weight. In a single day, a sperm whale can eat 700 to 800 squid, and a Risso's dolphin entangled in a net in the Mediterranean was found to have eaten angel clubhook squid, umbrella squid, reverse jewel squid and European flying squid, all identifiable from their indigestible beaks. Ornithoteuthis volatilis, a common squid from the tropical Indo-Pacific, is predated by yellowfin tuna, longnose lancetfish, common dolphinfish and swordfish, the tiger shark, the scalloped hammerhead shark and the smooth hammerhead shark. Sperm whales also hunt this species extensively as does the brown fur seal. In the Southern Ocean, penguins and wandering albatrosses are major predators of Gonatus antarcticus.
HUMAN USES
IN LITERATUR AND ART
Giant squid have featured as monsters of the deep since classical times. Giant squid were described by Aristotle (4th century BC) in his History of Animals and Pliny the Elder (1st century AD) in his Natural History. The Gorgon of Greek mythology may have been inspired by squid or octopus, the animal itself representing the severed head of Medusa, the beak as the protruding tongue and fangs, and its tentacles as the snakes. The six-headed sea monster of the Odyssey, Scylla, may have had a similar origin. The Nordic legend of the kraken may also have derived from sightings of large cephalopods.
In literature, H. G. Wells' short story "The Sea Raiders" featured a man-eating squid species Haploteuthis ferox.[59] The science fiction writer Jules Verne told a tale of a kraken-like monster in his 1870 novel Twenty Thousand Leagues Under the Sea.
AS FOOD
Squid form a major food resource and are used in cuisines around the world, notably in Japan where it is eaten as ika sōmen, sliced into vermicelli-like strips; as sashimi; and as tempura. Three species of Loligo are used in large quantities, L. vulgaris in the Mediterranean (known as Calamar in Spanish, Calamaro in Italian); L. forbesii in the Northeast Atlantic; and L. pealei on the American East Coast. Among the Ommastrephidae, Todarodes pacificus is the main commercial species, harvested in large quantities across the North Pacific in Canada, Japan and China.
In English-speaking countries, squid as food is often called calamari, adopted from Italian into English in the 17th century. Squid are found abundantly in certain areas, and provide large catches for fisheries. The body can be stuffed whole, cut into flat pieces, or sliced into rings. The arms, tentacles, and ink are also edible; the only parts not eaten are the beak and gladius (pen). Squid is a good food source for zinc and manganese, and high in copper, selenium, vitamin B12, and riboflavin.
COMMERCIAL FISHING
According to the FAO, the cephalopod catch for 2002 was 3,173,272 tonnes. Of this, 2,189,206 tonnes, or 75.8 percent, was squid. The following table lists squid species fishery catches that exceeded 10,000 tonnes in 2002.
IN BIOMIMICRY
Prototype chromatophores that mimic the squid's adaptive camouflage, have been made by Bristol University researchers using an electroactive dielectric elastomer, a flexible "smart" material that changes its colour and texture in response to electrical signals. The researchers state that their goal is to create an artificial skin that provides rapid active camouflage.
The squid giant axon inspired Otto Schmitt to develop a comparator circuit with hysteresis now called the Schmitt trigger, replicating the axon's propagation of nerve impulses.
WIKIPEDIA
So I have started a module on creating abstract creative words. It's very difficult to think this way. I find it quite easy with my visual work but very hard when trying to write about it. This image is actually a double exposure. I added just the faintest hint of bare winter trees to the skyline and then thought long and hard about how to write about it all. This is Selsey's beautiful shingle beach this morning.
The hush of winter drapes itself over Selsey’s beautiful shingle beach, where the sea glows silver beneath a sky of quiet frost. The pebbles, sleek with dampness, shift and sigh beneath each step, their softened edges bearing the weight of time. The tide, a glassy exhalation, moves with a measured patience, its surface barely stirred, as though dreaming beneath the pale hush of morning.
Above, the ghost of bare winter trees etches delicate veins against the sky, their skeletal branches reaching, yearning, dissolving into mist. Shadows pool in their tangled limbs, a silent memory of leaves now lost to the turning year. The wind drifts through unseen, a cold, gentle hand brushing along the shore, neither cruel nor kind, only present. It carries the scent of brine and distant storms, a whisper of forgotten voyages.
Driftwood lies abandoned, its pale bones softened by the sea’s slow touch. Seaweed glistens in dark ribbons, curling over smooth stones like ink spilled upon a silent page. A gull wheels above, spectral in the grey light, its wings cutting through the weight of stillness. Footsteps leave brief imprints, soon erased by the tide’s patient return.
Here, the world breathes in quiet rhythm—water lapping, stones shifting, the wind threading through the empty arms of winter trees. Kelsey’s beach stands timeless, a place where the sea listens, the sky remembers, and the land waits beneath the sleeping season’s hush.
The chimpanzee - Pan troglodytes, also known as simply the chimp, is a species of great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close relative the bonobo was more commonly known as the pygmy chimpanzee, this species was often called the common chimpanzee or the robust chimpanzee. The chimpanzee and the bonobo are the only species in the genus Pan. Evidence from fossils and DNA sequencing shows that Pan is a sister taxon to the human lineage and is humans' closest living relative. The chimpanzee is covered in coarse black hair, but has a bare face, fingers, toes, palms of the hands, and soles of the feet. It is larger and more robust than the bonobo, weighing 40–70 kg (88–154 lb) for males and 27–50 kg (60–110 lb) for females and standing 150 cm (4 ft 11 in).
The chimpanzee lives in groups that range in size from 15 to 150 members, although individuals travel and forage in much smaller groups during the day. The species lives in a strict male-dominated hierarchy, where disputes are generally settled without the need for violence. Nearly all chimpanzee populations have been recorded using tools, modifying sticks, rocks, grass and leaves and using them for hunting and acquiring honey, termites, ants, nuts and water. The species has also been found creating sharpened sticks to spear small mammals. Its gestation period is eight months. The infant is weaned at about three years old but usually maintains a close relationship with its mother for several years more.
The chimpanzee is listed on the IUCN Red List as an endangered species. Between 170,000 and 300,000 individuals are estimated across its range. The biggest threats to the chimpanzee are habitat loss, poaching, and disease. Chimpanzees appear in Western popular culture as stereotyped clown-figures and have featured in entertainments such as chimpanzees' tea parties, circus acts and stage shows. Although many chimpanzees have been kept as pets, their strength, aggressiveness, and unpredictability makes them dangerous in this role. Some hundreds have been kept in laboratories for research, especially in the United States. Many attempts have been made to teach languages such as American Sign Language to chimpanzees, with limited success.
Etymology
The English word chimpanzee is first recorded in 1738. It is derived from Vili ci-mpenze or Tshiluba language chimpenze, with a meaning of "ape", or "mockman". The colloquialism "chimp" was most likely coined some time in the late 1870s. The genus name Pan derives from the Greek god, while the specific name troglodytes was taken from the Troglodytae, a mythical race of cave-dwellers.
Taxonomy and genetics
The first great ape known to Western science in the 17th century was the "orang-outang" (genus Pongo), the local Malay name being recorded in Java by the Dutch physician Jacobus Bontius. In 1641, the Dutch anatomist Nicolaes Tulp applied the name to a chimpanzee or bonobo brought to the Netherlands from Angola. Another Dutch anatomist, Peter Camper, dissected specimens from Central Africa and Southeast Asia in the 1770s, noting the differences between the African and Asian apes. The German naturalist Johann Friedrich Blumenbach classified the chimpanzee as Simia troglodytes by 1775. Another German naturalist, Lorenz Oken, coined the genus Pan in 1816. The bonobo was recognised as distinct from the chimpanzee by 1933.
Evolution
Further information: Chimpanzee–human last common ancestor
Despite a large number of Homo fossil finds, Pan fossils were not described until 2005. Existing chimpanzee populations in West and Central Africa do not overlap with the major human fossil sites in East Africa, but chimpanzee fossils have now been reported from Kenya. This indicates that both humans and members of the Pan clade were present in the East African Rift Valley during the Middle Pleistocene.
According to studies published in 2017 by researchers at George Washington University, bonobos, along with chimpanzees, split from the human line about 8 million years ago; then bonobos split from the common chimpanzee line about 2 million years ago. Another 2017 genetic study suggests ancient gene flow (introgression) between 200,000 and 550,000 years ago from the bonobo into the ancestors of central and eastern chimpanzees.
Subspecies and population status
Four subspecies of the chimpanzee have been recognised, with the possibility of a fifth:
Central chimpanzee or the tschego (Pan troglodytes troglodytes), found in Cameroon, the Central African Republic, Equatorial Guinea, Gabon, the Republic of the Congo, and the Democratic Republic of the Congo, with about 140,000 individuals existing in the wild.
Western chimpanzee (P. troglodytes verus), found in Ivory Coast, Guinea, Liberia, Mali, Sierra Leone, Guinea-Bissau, Senegal, and Ghana with about 52,800 individuals still in existence.
Nigeria-Cameroon chimpanzee (P. troglodytes ellioti (also known as P. t. vellerosus)), that live within forested areas across Nigeria and Cameroon, with 6000–9000 individuals still in existence.
Eastern chimpanzee (P. troglodytes schweinfurthii), found in the Central African Republic, South Sudan, the Democratic Republic of the Congo, Uganda, Rwanda, Burundi, Tanzania, and Zambia, with approximately 180,000–256,000 individuals still existing in the wild.
Southeastern chimpanzee, P. troglodytes marungensis, in Burundi, Rwanda, Tanzania, and Uganda. Colin Groves argues that this is a subspecies, created by enough variation between the northern and southern populations of P. t. schweinfurthii, but it is not recognised by the IUCN.
Genome
Main article: Chimpanzee genome project
Genomic information
NCBI genome ID202
Ploidydiploid
Genome size3,323.27 Mb
Number of chromosomes24 pairs
A draft version of the chimpanzee genome was published in 2005 and encodes 18,759 proteins, (compared to 20,383 in the human proteome). The DNA sequences of humans and chimpanzees are very similar and the difference in protein number mostly arises from incomplete sequences in the chimp genome. Both species differ by about 35 million single-nucleotide changes, five million insertion/deletion events and various chromosomal rearrangements. Typical human and chimpanzee protein homologs differ in an average of only two amino acids. About 30% of all human proteins are identical in sequence to the corresponding chimpanzee protein. Duplications of small parts of chromosomes have been the major source of differences between human and chimpanzee genetic material; about 2.7% of the corresponding modern genomes represent differences, produced by gene duplications or deletions, since humans and chimpanzees diverged from their common evolutionary ancestor.
Characteristics
Adult chimpanzees have an average standing height of 150 cm (4 ft 11 in). Wild adult males weigh between 40 and 70 kg (88 and 154 lb) with females weighing between 27 and 50 kg (60 and 110 lb). In exceptional cases, certain individuals may considerably exceed these measurements, standing over 168 cm (5 ft 6 in) on two legs and weighing up to 136 kg (300 lb) in captivity.
The chimpanzee is more robustly built than the bonobo but less than the gorilla. The arms of a chimpanzee are longer than its legs and can reach below the knees. The hands have long fingers with short thumbs and flat fingernails. The feet are adapted for grasping, and the big toe is opposable. The pelvis is long with an extended ilium. A chimpanzee's head is rounded with a prominent and prognathous face and a pronounced brow ridge. It has forward-facing eyes, a small nose, rounded non-lobed ears and a long mobile upper lip. Additionally, adult males have sharp canine teeth. Chimpanzees lack the prominent sagittal crest and associated head and neck musculature of gorillas.
Chimpanzee bodies are covered by coarse hair, except for the face, fingers, toes, palms of the hands, and soles of the feet. Chimpanzees lose more hair as they age and develop bald spots. The hair of a chimpanzee is typically black but can be brown or ginger. As they get older, white or grey patches may appear, particularly on the chin and lower region. Chimpanzee skin that is covered with body hair is white, while exposed areas vary: white which ages into a dark muddy colour in eastern chimpanzees, freckled on white which ages to a heavily mottled muddy colour in central chimpanzees, and black with a butterfly-shaped white mask that darkens with age in western chimpanzees. Facial pigmentation increases with age and exposure to ultraviolet light. Females develop swelling pink skin when in oestrus.
Chimpanzees are adapted for both arboreal and terrestrial locomotion. Arboreal locomotion consists of vertical climbing and brachiation. On the ground, chimpanzees move both quadrupedally and bipedally. These movements appear to have similar energy costs. As with bonobos and gorillas, chimpanzees move quadrupedally by knuckle-walking, which probably evolved independently in Pan and Gorilla. Their muscles are 50% stronger per weight than those of humans due to higher content of fast twitch muscle fibres, one of the chimpanzee's adaptations for climbing and swinging. According to Japan's Asahiyama Zoo, the grip strength of an adult chimpanzee is estimated to be 200 kg (440 lb), while other sources claim figures of up to 330 kg (730 lb).
Ecology
The chimpanzee is a highly adaptable species. It lives in a variety of habitats, including dry savanna, evergreen rainforest, montane forest, swamp forest, and dry woodland-savanna mosaic. In Gombe, the chimpanzee mostly uses semideciduous and evergreen forest as well as open woodland. At Bossou, the chimpanzee inhabits multistage secondary deciduous forest, which has grown after shifting cultivation, as well as primary forest and grassland. At Taï, it is found in the last remaining tropical rain forest in Ivory Coast. The chimpanzee has an advanced cognitive map of its home range and can repeatedly find food. The chimpanzee builds a sleeping nest in a tree in a different location each night, never using the same nest more than once. Chimpanzees sleep alone in separate nests except for infants or juvenile chimpanzees, which sleep with their mothers.
Diet
The chimpanzee is an omnivorous frugivore. It prefers fruit above all other food items but also eats leaves, leaf buds, seeds, blossoms, stems, pith, bark, and resin. A study in Budongo Forest, Uganda found that 64.5% of their feeding time concentrated on fruits (84.6% of which being ripe), particularly those from two species of Ficus, Maesopsis eminii, and Celtis gomphophylla. In addition, 19% of feeding time was spent on arboreal leaves, mostly Broussonetia papyrifera and Celtis mildbraedii. While the chimpanzee is mostly herbivorous, it does eat honey, soil, insects, birds and their eggs, and small to medium-sized mammals, including other primates. Insect species consumed include the weaver ant Oecophylla longinoda, Macrotermes termites, and honey bees. The red colobus ranks at the top of preferred mammal prey. Other mammalian prey include red-tailed monkeys, infant and juvenile yellow baboons, bush babies, blue duikers, bushbucks, and common warthogs.
Despite the fact that chimpanzees are known to hunt and to collect both insects and other invertebrates, such food actually makes up a very small portion of their diet, from as little as 2% yearly to as much as 65 grams of animal flesh per day for each adult chimpanzee in peak hunting seasons. This also varies from troop to troop and year to year. However, in all cases, the majority of their diet consists of fruits, leaves, roots, and other plant matter. Female chimpanzees appear to consume much less animal flesh than males, according to several studies. Jane Goodall documented many occasions within Gombe Stream National Park of chimpanzees and western red colobus monkeys ignoring each other despite close proximity.
Chimpanzees do not appear to directly compete with gorillas in areas where they overlap. When fruit is abundant, gorilla and chimpanzee diets converge, but when fruit is scarce gorillas resort to vegetation. The two apes may also feed on different species, whether fruit or insects. Interactions between them can range from friendly and even stable social bonding, to avoidance, to aggression and predation on part of chimpanzees.
Mortality and health
The average lifespan of a chimpanzee in the wild is relatively short, usually less than 15 years, although individuals that reach 12 years may live an additional 15 years. On rare occasions, wild chimpanzees may live nearly 60 years. Captive chimpanzees tend to live longer than most wild ones, with median lifespans of 31.7 years for males and 38.7 years for females. The oldest-known male captive chimpanzee to have been documented lived to 66 years, and the oldest female, Little Mama, was over 70 years old.
Leopards prey on chimpanzees in some areas. It is possible that much of the mortality caused by leopards can be attributed to individuals that have specialised in chimp-killing.[76] Chimpanzees may react to a leopard's presence with loud vocalising, branch shaking, and throwing objects. There is at least one record of chimpanzees killing a leopard cub after mobbing it and its mother in their den. Four chimpanzees could have fallen prey to lions at Mahale Mountains National Park. Although no other instances of lion predation on chimpanzees have been recorded, lions likely do kill chimpanzees occasionally, and the larger group sizes of savanna chimpanzees may have developed as a response to threats from these big cats. Chimpanzees may react to lions by fleeing up trees, vocalising, or hiding in silence.
The chimpanzee louse Pediculus schaeffi is closely related to the human body louse P. humanus.
Chimpanzees and humans share only 50% of their parasite and microbe species. This is due to the differences in environmental and dietary adaptations; human internal parasite species overlap more with omnivorous, savanna-dwelling baboons. The chimpanzee is host to the louse species Pediculus schaeffi, a close relative of P. humanus, which infests human head and body hair. By contrast, the human pubic louse Pthirus pubis is closely related to Pthirus gorillae, which infests gorillas. A 2017 study of gastrointestinal parasites of wild chimpanzees in degraded forest in Uganda found nine species of protozoa, five nematodes, one cestode, and one trematode. The most prevalent species was the protozoan Troglodytella abrassarti.
Behaviour
Recent studies have suggested that human observers influence chimpanzee behaviour. One suggestion is that drones, camera traps, and remote microphones should be used to record and monitor chimpanzees rather than direct human observation.
Group structure
Chimpanzees live in communities that typically range from around 15 to more than 150 members but spend most of their time traveling in small, temporary groups consisting of a few individuals. These groups may consist of any combination of age and sexes. Both males and females sometimes travel alone. This fission-fusion society may include groups of four types: all-male, adult females and offspring, adults of both sexes, or one female and her offspring. These smaller groups emerge in a variety of types, for a variety of purposes. For example, an all-male troop may be organised to hunt for meat, while a group consisting of lactating females serves to act as a "nursery group" for the young.
At the core of social structures are males, which patrol the territory, protect group members, and search for food. Males remain in their natal communities, while females generally emigrate at adolescence. Males in a community are more likely to be related to one another than females are to each other. Among males, there is generally a dominance hierarchy, and males are dominant over females. However, this unusual fission-fusion social structure, "in which portions of the parent group may on a regular basis separate from and then rejoin the rest," is highly variable in terms of which particular individual chimpanzees congregate at a given time. This is caused mainly by the large measure of individual autonomy that individuals have within their fission-fusion social groups. As a result, individual chimpanzees often forage for food alone, or in smaller groups, as opposed to the much larger "parent" group, which encompasses all the chimpanzees which regularly come into contact with each other and congregate into parties in a particular area.
Male chimpanzees exist in a linear dominance hierarchy. Top-ranking males tend to be aggressive even during dominance stability. This is probably due to the chimpanzee's fission-fusion society, with male chimpanzees leaving groups and returning after extended periods of time. With this, a dominant male is unsure if any "political maneuvering" has occurred in his absence and must re-establish his dominance. Thus, a large amount of aggression occurs within five to fifteen minutes after a reunion. During these encounters, displays of aggression are generally preferred over physical attacks.
Males maintain and improve their social ranks by forming coalitions, which have been characterised as "exploitative" and based on an individual's influence in agonistic interactions. Being in a coalition allows males to dominate a third individual when they could not by themselves, as politically apt chimpanzees can exert power over aggressive interactions regardless of their rank. Coalitions can also give an individual male the confidence to challenge a dominant or larger male. The more allies a male has, the better his chance of becoming dominant. However, most changes in hierarchical rank are caused by dyadic interactions. Chimpanzee alliances can be very fickle, and one member may suddenly turn on another if it is to his advantage.
Mutual grooming, removing lice
Low-ranking males frequently switch sides in disputes between more dominant individuals. Low-ranking males benefit from an unstable hierarchy and often find increased sexual opportunities if a dispute or conflict occurs. In addition, conflicts between dominant males cause them to focus on each other rather than the lower-ranking males. Social hierarchies among adult females tend to be weaker. Nevertheless, the status of an adult female may be important for her offspring. Females in Taï have also been recorded to form alliances. While chimpanzee social structure is often referred to as patriarchal, it is not entirely unheard of for females to forge coalitions against males. There is also at least one recorded case of females securing a dominant position over males in their respective troop, albeit in a captive environment. Social grooming appears to be important in the formation and maintenance of coalitions. It is more common among adult males than either between adult females or between males and females.
Males in Mahale National Park, Tanzania
Chimpanzees have been described as highly territorial and will frequently kill other chimpanzees, although Margaret Power wrote in her 1991 book The Egalitarians that the field studies from which the aggressive data came, Gombe and Mahale, used artificial feeding systems that increased aggression in the chimpanzee populations studied. Thus, the behaviour may not reflect innate characteristics of the species as a whole. In the years following her artificial feeding conditions at Gombe, Jane Goodall described groups of male chimpanzees patrolling the borders of their territory, brutally attacking chimpanzees that had split off from the Gombe group. A study published in 2010 found that the chimpanzees wage wars over territory, not mates. Patrols from smaller groups are more likely to avoid contact with their neighbours. Patrols from large groups even take over a smaller group's territory, gaining access to more resources, food, and females. While it was traditionally accepted that only female chimpanzees immigrate and males remain in their natal troop for life, there are confirmed cases of adult males safely integrating themselves into new communities among West African chimpanzees, suggesting they are less territorial than other subspecies.
Mating and parenting
Chimpanzees mate throughout the year, although the number of females in oestrus varies seasonally in a group. Female chimpanzees are more likely to come into oestrus when food is readily available. Oestrous females exhibit sexual swellings. Chimpanzees are promiscuous: during oestrus, females mate with several males in their community, while males have large testicles for sperm competition. Other forms of mating also exist. A community's dominant males sometimes restrict reproductive access to females. A male and female can form a consortship and mate outside their community. In addition, females sometimes leave their community and mate with males from neighboring communities.
These alternative mating strategies give females more mating opportunities without losing the support of the males in their community. Infanticide has been recorded in chimpanzee communities in some areas, and the victims are often consumed. Male chimpanzees practice infanticide on unrelated young to shorten the interbirth intervals in the females. Females sometimes practice infanticide. This may be related to the dominance hierarchy in females or may simply be pathological.
Copulation is brief, lasting approximately seven seconds. The gestation period is eight months. Care for the young is provided mostly by their mothers. The survival and emotional health of the young is dependent on maternal care. Mothers provide their young with food, warmth, and protection, and teach them certain skills. In addition, a chimpanzee's future rank may be dependent on its mother's status. Male chimpanzees continue to associate with the females they impregnated and interact with and support their offspring. Newborn chimpanzees are helpless. For example, their grasping reflex is not strong enough to support them for more than a few seconds. For their first 30 days, infants cling to their mother's bellies. Infants are unable to support their own weight for their first two months and need their mothers' support.
When they reach five to six months, infants ride on their mothers' backs. They remain in continual contact for the rest of their first year. When they reach two years of age, they are able to move and sit independently and start moving beyond the arms' reach of their mothers. By four to six years, chimpanzees are weaned and infancy ends. The juvenile period for chimpanzees lasts from their sixth to ninth years. Juveniles remain close to their mothers, but interact an increasing amount with other members of their community. Adolescent females move between groups and are supported by their mothers in agonistic encounters. Adolescent males spend time with adult males in social activities like hunting and boundary patrolling. A captive study suggests males can safely immigrate to a new group if accompanied by immigrant females who have an existing relationship with this male. This gives the resident males reproductive advantages with these females, as they are more inclined to remain in the group if their male friend is also accepted.
Communication
Chimpanzees use facial expressions, postures, and sounds to communicate with each other. Chimpanzees have expressive faces that are important in close-up communications. When frightened, a "full closed grin" causes nearby individuals to be fearful, as well. Playful chimpanzees display an open-mouthed grin. Chimpanzees may also express themselves with the "pout", which is made in distress, the "sneer", which is made when threatening or fearful, and "compressed-lips face", which is a type of display. When submitting to a dominant individual, a chimpanzee crunches, bobs, and extends a hand. When in an aggressive mode, a chimpanzee swaggers bipedally, hunched over and arms waving, in an attempt to exaggerate its size. While travelling, chimpanzees keep in contact by beating their hands and feet against the trunks of large trees, an act that is known as "drumming". They also do this when encountering individuals from other communities.
Vocalisations are also important in chimpanzee communication. The most common call in adults is the "pant-hoot", which may signal social rank and bond along with keeping groups together. Pant-hoots are made of four parts, starting with soft "hoos", the introduction; that gets louder and louder, the build-up; and climax into screams and sometimes barks; these die down back to soft "hoos" during the letdown phase as the call ends. Grunting is made in situations like feeding and greeting. Submissive individuals make "pant-grunts" towards their superiors. Whimpering is made by young chimpanzees as a form of begging or when lost from the group. Chimpanzees use distance calls to draw attention to danger, food sources, or other community members. "Barks" may be made as "short barks" when hunting and "tonal barks" when sighting large snakes.
Adult male eastern chimpanzee snatches a dead bushbuck antelope from a baboon in Gombe Stream National Park.
Hunting
When hunting small monkeys such as the red colobus, chimpanzees hunt where the forest canopy is interrupted or irregular. This allows them to easily corner the monkeys when chasing them in the appropriate direction. Chimpanzees may also hunt as a coordinated team, so that they can corner their prey even in a continuous canopy. During an arboreal hunt, each chimpanzee in the hunting groups has a role. "Drivers" serve to keep the prey running in a certain direction and follow them without attempting to make a catch. "Blockers" are stationed at the bottom of the trees and climb up to block prey that takes off in a different direction. "Chasers" move quickly and try to make a catch. Finally, "ambushers" hide and rush out when a monkey nears. While both adults and infants are taken, adult male colobus monkeys will attack the hunting chimps. Male chimpanzees hunt more than females. When caught and killed, the meal is distributed to all hunting party members and even bystanders.
Intelligence and cognition
Further information: Primate cognition
Drawing of human and chimpanzee skull and brain
Human and chimpanzee skull and brain. Diagram by Paul Gervais from Histoire naturelle des mammifères (1854).
Chimpanzees display numerous signs of intelligence, from the ability to remember symbols to cooperation, tool use, and perhaps language. They are among species that have passed the mirror test, suggesting self-awareness. In one study, two young chimpanzees showed retention of mirror self-recognition after one year without access to mirrors. Chimpanzees have been observed to use insects to treat their own wounds and those of others. They catch them and apply them directly to the injury. Chimpanzees also display signs of culture among groups, with the learning and transmission of variations in grooming, tool use and foraging techniques leading to localized traditions.
A 30-year study at Kyoto University's Primate Research Institute has shown that chimpanzees are able to learn to recognise the numbers 1 to 9 and their values. The chimpanzees further show an aptitude for eidetic memory, demonstrated in experiments in which the jumbled digits are flashed onto a computer screen for less than a quarter of a second. One chimpanzee, Ayumu, was able to correctly and quickly point to the positions where they appeared in ascending order. Ayumu performed better than human adults who were given the same test.
In controlled experiments on cooperation, chimpanzees show a basic understanding of cooperation, and recruit the best collaborators. In a group setting with a device that delivered food rewards only to cooperating chimpanzees, cooperation first increased, then, due to competitive behaviour, decreased, before finally increasing to the highest level through punishment and other arbitrage behaviours.
Great apes show laughter-like vocalisations in response to physical contact, such as wrestling, play chasing, or tickling. This is documented in wild and captive chimpanzees. Chimpanzee laughter is not readily recognisable to humans as such, because it is generated by alternating inhalations and exhalations that sound more like breathing and panting. Instances in which nonhuman primates have expressed joy have been reported. Humans and chimpanzees share similar ticklish areas of the body, such as the armpits and belly. The enjoyment of tickling in chimpanzees does not diminish with age.
Chimpanzees have displayed different behaviours in response to a dying or dead group member. When witnessing a sudden death, the other group members act in frenzy, with vocalisations, aggressive displays, and touching of the corpse. In one case chimpanzees cared for a dying elder, then attended and cleaned the corpse. Afterward, they avoided the spot where the elder died and behaved in a more subdued manner. Mothers have been reported to carry around and groom their dead infants for several days.
Experimenters now and then witness behaviour that cannot be readily reconciled with chimpanzee intelligence or theory of mind. Wolfgang Köhler, for instance, reported insightful behaviour in chimpanzees, but he likewise often observed that they experienced "special difficulty" in solving simple problems. Researchers also reported that, when faced with a choice between two persons, chimpanzees were just as likely to beg food from a person who could see the begging gesture as from a person who could not, thereby raising the possibility that chimpanzees lack theory of mind.
Tool use
Further information: Tool use by animals
Chimpanzees using twigs to dip for ants
Nearly all chimpanzee populations have been recorded using tools. They modify sticks, rocks, grass, and leaves and use them when foraging for termites and ants, nuts, honey, algae or water. Despite the lack of complexity, forethought and skill are apparent in making these tools. Chimpanzees have used stone tools since at least 4,300 years ago.
A chimpanzee from the Kasakela chimpanzee community was the first nonhuman animal reported making a tool, by modifying a twig to use as an instrument for extracting termites from their mound. At Taï, chimpanzees simply use their hands to extract termites. When foraging for honey, chimpanzees use modified short sticks to scoop the honey out of the hive if the bees are stingless. For hives of the dangerous African honeybees, chimpanzees use longer and thinner sticks to extract the honey.
Chimpanzees also fish for ants using the same tactic. Ant dipping is difficult and some chimpanzees never master it. West African chimpanzees crack open hard nuts with stones or branches. Some forethought in this activity is apparent, as these tools are not found together or where the nuts are collected. Nut cracking is also difficult and must be learned. Chimpanzees also use leaves as sponges or spoons to drink water.
West African chimpanzees in Senegal were found to sharpen sticks with their teeth, which were then used to spear Senegal bushbabies out of small holes in trees. An eastern chimpanzee has been observed using a modified branch as a tool to capture a squirrel.
Whilst experimental studies on captive chimpanzees have found that many of their species-typical tool-use behaviours can be individually learnt by each chimpanzees, a 2021 study on their abilities to make and use stone flakes, in a similar way as hypothesised for early hominins, did not find this behaviour across two populations of chimpanzees—suggesting that this behaviour is outside the chimpanzee species-typical range.
Language
Main article: Great ape language
Scientists have attempted to teach human language to several species of great ape. One early attempt by Allen and Beatrix Gardner in the 1960s involved spending 51 months teaching American Sign Language to a chimpanzee named Washoe. The Gardners reported that Washoe learned 151 signs, and had spontaneously taught them to other chimpanzees, including her adopted son, Loulis. Over a longer period of time, Washoe was reported to have learned over 350 signs.
Debate is ongoing among scientists such as David Premack about chimpanzees' ability to learn language. Since the early reports on Washoe, numerous other studies have been conducted, with varying levels of success. One involved a chimpanzee jokingly named Nim Chimpsky (in allusion to the theorist of language Noam Chomsky), trained by Herbert Terrace of Columbia University. Although his initial reports were quite positive, in November 1979, Terrace and his team, including psycholinguist Thomas Bever, re-evaluated the videotapes of Nim with his trainers, analyzing them frame by frame for signs, as well as for exact context (what was happening both before and after Nim's signs). In the reanalysis, Terrace and Bever concluded that Nim's utterances could be explained merely as prompting on the part of the experimenters, as well as mistakes in reporting the data. "Much of the apes' behaviour is pure drill", he said. "Language still stands as an important definition of the human species." In this reversal, Terrace now argued Nim's use of ASL was not like human language acquisition. Nim never initiated conversations himself, rarely introduced new words, and mostly imitated what the humans did. More importantly, Nim's word strings varied in their ordering, suggesting that he was incapable of syntax. Nim's sentences also did not grow in length, unlike human children whose vocabulary and sentence length show a strong positive correlation.
Relations with humans
In culture
Chimpanzees are rarely represented in African culture, as people find their resemblance to humans discomforting. The Gio people of Liberia and the Hemba people of the Congo have created masks of the animals. Gio masks are crude and blocky, and worn when teaching young people how not to behave. The Hemba masks have a smile that suggests drunken anger, insanity or horror and are worn during rituals at funerals, representing the "awful reality of death". The masks may also serve to guard households and protect both human and plant fertility. Stories have been told of chimpanzees kidnapping and raping women.
In Western popular culture, chimpanzees have occasionally been stereotyped as childlike companions, sidekicks or clowns. They are especially suited for the latter role on account of their prominent facial features, long limbs and fast movements, which humans often find amusing. Accordingly, entertainment acts featuring chimpanzees dressed up as humans with lip-synchronised human voices have been traditional staples of circuses, stage shows and TV shows like Lancelot Link, Secret Chimp (1970-1972) and The Chimp Channel (1999). From 1926 until 1972, London Zoo, followed by several other zoos around the world, held a chimpanzees' tea party daily, inspiring a long-running series of advertisements for PG Tips tea featuring such a party. Animal rights groups have urged a stop to such acts, considering them abusive.
Poster for the 1931 film Aping Hollywood. Media like this relied on the novelty of performing apes to carry their gags.
Chimpanzees in media include Judy on the television series Daktari in the 1960s and Darwin on The Wild Thornberrys in the 1990s. In contrast to the fictional depictions of other animals, such as dogs (as in Lassie), dolphins (Flipper), horses (The Black Stallion) or even other great apes (King Kong), chimpanzee characters and actions are rarely relevant to the plot. Depictions of chimpanzees as individuals rather than stock characters, and as central rather than incidental to the plot can be found in science fiction. Robert A. Heinlein's 1947 short story "Jerry Was a Man" concerns a genetically enhanced chimpanzee suing for better treatment. The 1972 film Conquest of the Planet of the Apes, the third sequel of the 1968 film Planet of the Apes, portrays a futuristic revolt of enslaved apes led by the only talking chimpanzee, Caesar, against their human masters.
As pets
Chimpanzees have traditionally been kept as pets in a few African villages, especially in the Democratic Republic of Congo. In Virunga National Park in the east of the country, the park authorities regularly confiscate chimpanzees from people keeping them as pets. Outside their range, chimpanzees are popular as exotic pets despite their strength and aggression. Even in places where keeping non-human primates as pets is illegal, the exotic pet trade continues to prosper, leading to injuries from attacks.
Use in research
See also: Countries banning non-human ape experimentation and Animal testing on non-human primates § Chimpanzees in the U.S.
Hundreds of chimpanzees have been kept in laboratories for research. Most such laboratories either conduct or make the animals available for invasive research, defined as "inoculation with an infectious agent, surgery or biopsy conducted for the sake of research and not for the sake of the chimpanzee, and/or drug testing". Research chimpanzees tend to be used repeatedly over decades for up to 40 years, unlike the pattern of use of most laboratory animals. Two federally funded American laboratories use chimpanzees: the Yerkes National Primate Research Center at Emory University in Atlanta, Georgia, and the Southwest National Primate Center in San Antonio, Texas. Five hundred chimpanzees have been retired from laboratory use in the US and live in animal sanctuaries in the US or Canada.
A five-year moratorium was imposed by the US National Institutes of Health in 1996, because too many chimpanzees had been bred for HIV research, and it has been extended annually since 2001. With the publication of the chimpanzee genome, plans to increase the use of chimpanzees in America were reportedly increasing in 2006, some scientists arguing that the federal moratorium on breeding chimpanzees for research should be lifted. However, in 2007, the NIH made the moratorium permanent.
Ham, the first great ape in space, before being inserted into his Mercury-Redstone 2 capsule on 31 January 1961
Other researchers argue that chimpanzees either should not be used in research, or should be treated differently, for instance with legal status as persons. Pascal Gagneux, an evolutionary biologist and primate expert at the University of California, San Diego, argues, given chimpanzees' sense of self, tool use, and genetic similarity to human beings, studies using chimpanzees should follow the ethical guidelines used for human subjects unable to give consent. A recent study suggests chimpanzees which are retired from labs exhibit a form of post-traumatic stress disorder. Stuart Zola, director of the Yerkes laboratory, disagrees. He told National Geographic: "I don't think we should make a distinction between our obligation to treat humanely any species, whether it's a rat or a monkey or a chimpanzee. No matter how much we may wish it, chimps are not human."
Only one European laboratory, the Biomedical Primate Research Centre in Rijswijk, the Netherlands, used chimpanzees in research. It formerly held 108 chimpanzees among 1,300 non-human primates. The Dutch ministry of science decided to phase out research at the centre from 2001. Trials already under way were however allowed to run their course. Chimpanzees including the female Ai have been studied at the Primate Research Institute of Kyoto University, Japan, formerly directed by Tetsuro Matsuzawa, since 1978. Some 12 chimpanzees are currently held at the facility.
Two chimpanzees have been sent into outer space as NASA research subjects. Ham, the first great ape in space, was launched in the Mercury-Redstone 2 capsule on 31 January 1961, and survived the suborbital flight. Enos, the third primate to orbit Earth after Soviet cosmonauts Yuri Gagarin and Gherman Titov, flew on Mercury-Atlas 5 on 29 November of the same year.
Field study
Jane Goodall undertook the first long-term field study of the chimpanzee, begun in Tanzania at Gombe Stream National Park in 1960. Other long-term studies begun in the 1960s include Adriaan Kortlandt's in the eastern Democratic Republic of the Congo and Toshisada Nishida's in Mahale Mountains National Park in Tanzania. Current understanding of the species' typical behaviours and social organisation has been formed largely from Goodall's ongoing 60-year Gombe research study.
Attacks
Chimpanzees have attacked humans. In Uganda, several attacks on children have happened, some of them fatal. Some of these attacks may have been due to the chimpanzees being intoxicated (from alcohol obtained from rural brewing operations) and becoming aggressive towards humans. Human interactions with chimpanzees may be especially dangerous if the chimpanzees perceive humans as potential rivals. At least six cases of chimpanzees snatching and eating human babies are documented.
A chimpanzee's strength and sharp teeth mean that attacks, even on adult humans, can cause severe injuries. This was evident after the attack and near death of former NASCAR driver St. James Davis, who was mauled by two escaped chimpanzees (in the St. James Davis chimpanzee attack) while he and his wife were celebrating the birthday of their former pet chimpanzee. Another example of chimpanzees being aggressive toward humans occurred in 2009 in Stamford, Connecticut, when a 90-kilogram (200 lb), 13-year-old pet chimpanzee named Travis attacked his owner's friend, who lost her hands, eyes, nose, and part of her maxilla from the attack.
Human immunodeficiency virus
Two primary classes of human immunodeficiency virus (HIV) infect humans: HIV-1 and HIV-2. HIV-1 is the more virulent and easily transmitted, and is the source of the majority of HIV infections throughout the world; HIV-2 occurs mostly in west Africa. Both types originated in west and central Africa, jumping from other primates to humans. HIV-1 has evolved from a simian immunodeficiency virus (SIVcpz) found in the subspecies P. t. troglodytes of southern Cameroon. Kinshasa, in the Democratic Republic of Congo, has the greatest genetic diversity of HIV-1 so far discovered, suggesting the virus has been there longer than anywhere else. HIV-2 crossed species from a different strain of HIV, found in the sooty mangabey monkeys in Guinea-Bissau.
Status and conservation
The chimpanzee is on the IUCN Red List as an endangered species. Chimpanzees are legally protected in most of their range and are found both in and outside national parks. Between 172,700 and 299,700 individuals are thought to be living in the wild, a decrease from about a million chimpanzees in the early 1900s.[189] Chimpanzees are listed in Appendix I of the Convention on International Trade in Endangered Species (CITES), meaning that commercial international trade in wild-sourced specimens is prohibited and all other international trade (including in parts and derivatives) is regulated by the CITES permitting system.
The biggest threats to the chimpanzee are habitat destruction, poaching, and disease. Chimpanzee habitats have been limited by deforestation in both West and Central Africa. Road building has caused habitat degradation and fragmentation of chimpanzee populations and may allow poachers more access to areas that had not been seriously affected by humans. Although deforestation rates are low in western Central Africa, selective logging may take place outside national parks.
Chimpanzees are a common target for poachers. In Ivory Coast, chimpanzees make up 1–3% of bushmeat sold in urban markets. They are also taken, often illegally, for the pet trade and are hunted for medicinal purposes in some areas. Farmers sometimes kill chimpanzees that threaten their crops; others are unintentionally maimed or killed by snares meant for other animals.
Infectious diseases are a main cause of death for chimpanzees. They succumb to many diseases that afflict humans because the two species are so similar. As the human population grows, so does the risk of disease transmission between humans and chimpanzees.
At 21:47 GMT, the equinox happened, and so from then on, light is destined to win over darkness. Which meant, of course, that the day before then was the shortest "day", or amount of daylight.
This is the end of the year, the build up and excitement before Christmas, and at the same time, looking back at the year, and what has happened in the previous 50 or so weeks. So, a time of mixed emotions, good and bad, happy and sad.
But I was on vacation, or not going to work.
I am not up to date, but I did all the tasks I was supposed to do, threw a few electronic grenades over the walls, and was now happy not to think of that shit for two whole weeks.
For Jools, however, there was half a day to do, and then her employers paid for all those employed at the factory to go to a fancy place in Folkestone for lunch, drinks at the bar and a bottle of wine between four folks.
It was, in short, a time for celebration. Something I realise has not happened in my job since I left operational quality, to be happy and give thanks to those we work with. And be recognised for the good job we do.
So, I was to take Jools to work, and have the car for the day.
Jools was conscious that my plan for the day involved driving to the far west of Kent, so realised I needed an early start, and not dropping her off in Hythe at seven.
We left after coffee just after six, driving through Dover and Folkestone on the main road and motorway before turning over the downs into Hythe. I dropped her off in the town, so she could get some walking in. She always didn't walk, as waves of showers swept over the town, and me as I drove back home for breakfast and do all the chores before leaving on a mini-churchcrawl.
So, back home for breakfast, more coffee, wash up, do the bird feeders and with postcodes, set out for points in the extreme west. Now, Kent is not a big county, not say, Texas big, but it takes some time to get to some parts of the west of the county. Main roads run mainly from London to the coast, so going cross-country or cross-county would take time.
At first it was as per normal up the A20 then onto the motorway to Ashford then to Maidstone until the junction before the M26 starts. One of the reasons for going later was to avoid rush hours in and around Maidstone, Tonbridge and Tunbridge Wells.
As it was, after turning down the A road, things were fine until I got to Mereworth, but from there the road began to twist and turn until it lead me into Tonbridge. Once upon a time, this was a sleepy village or small town. The the railways came and it became a major junction. The road to Penshurt took me though the one way system, then down the wide High Street, over the river Medway and up the hill the other side.
Two more turns took me to my target, through what were once called stockbroker mansions, then down a hill, with the village laid out before me just visible through the trees.
The village was built around the outskirts of Penshurst Place, home to the Sidney family since Tudor times. Just about everything is named the Leicester something, the village having its own Leicester Square, though with no cinemas, and all timber framed houses and painfully picturesque.
The church lays behind the houses, the tower in golden sandstone topped with four spirelets.
I parked the car, and armed with two cameras, several lenses and a photographer's eye, walked to the church.
The reason for coming was I can only remember a little about my previous visit, but the Leicester name thing triggered in my head the thought the memorials and tombs might be worth a revisit.
So there I was.
Gilbert Scott was very busy here, so there is little of anything prior to the 19th century, but the memorials are there. Including one which features the heads of the children of Robert Sidney (d1702) in a cloud. Including the eldest son who died, apparently, so young he wasn't named, and is recorded as being the first born.
This is in the Sidney Chapel where the great and good are buried and remembered, it has a colourful roof, or roof beams, and heraldic shields. It has a 15th century font, which, sadly, has been brightly painted so is gaudy in the extreme.
I go around getting my shots, leave a fiver for the church. Go back to the car and program Speldhurst into the sat nav.
Its just a ten minute drive, but there is no place to park anywhere near the church. I could see from my slow drive-by the porch doors closed, and I convinced myself they were locked and not worth checking out.
I went on to Groombridge, where there is a small chapel with fabulous glass. I had been here before too, but wanted to redo my shots.
It was by now pouring with rain, and as dark as twilight, I missed the church on first pass, went to the mini-roundabout only to discover that it and the other church in the village were in Sussex. I turned round, the church looked dark and was almost certainly locked. I told myself.
I didn't stop here either, so instead of going to the final village church, I went straigh to Tunbridge Wells where there was another church to revisit.
I drove into the town, over the man road and to the car park with no waiting in traffic, how odd, I thought.
It was hard to find a parking space, but high up in the parking house there were finally spaced. I parked near the stairs down, grabbed my cameras and went down.
I guess I could have parked nearer the church, but once done it would be easier to leave the town as the road back home went past the exit.
I ambled down the hill leading to the station, over the bridge and down the narrow streets, all lined with shops. I think its fair to say that it is a richer town than Dover because on one street there were three stores offering beposke designer kitchens.
The church is across the road from the Georgian square known at The Pantiles, but it was the church I was here to visit.
I go in, and there is a service underway. I decide to sit at the back and observe.
And pray.
I did not take communion, though. The only one there who didn't.
About eight elderly parishioners did, though.
I was here to photograph the ceiling, and then the other details I failed to record when we were last here over a decade ago.
I was quizzed strongly by a warden as to why I was doing this. I had no answer other than I enjoyed it, and for me that is enough.
After getting my shots, I leave and begin the slog back up to the car, but on the way keeping my promise to a young man selling the Big Issue that I would come back and buy a copy. I did better than that in that I gave him a fiver and didn't take a copy.
He nearly burst into tears. I said, there is kindness in the world, and some of us do keep our promises.
By the time I got to the car park, it was raining hard again. I had two and a half hours to get to Folkestone to pick up Jools after her meal.
Traffic into Tunbridge Wells from this was was crazy, miles and miles of queues, so I was more than happy going the other way.
I get back to the M20, cruise down to Ashford, stopping at Stop 24 services for a coffee and something to eat. I had 90 minutes to kill, so eat, drink and scroll Twitter as I had posted yet more stuff that morning. In other news: nothing changed, sadly.
At quarter past four I went to pick up Jools, stopping outside the restaurant. When she got in she declared she had been drinking piña coladas. Just two, but she was bubby and jabbering away all the way home.
With Jools having eaten out, and with snacks I had, no dinner was needed, so when suppertime came round, we dined on cheese and crackers, followed by a large slice of Christmas cake.
She was now done for Christmas too.
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The red brick church stands on a busy junction at the end of the Pantiles whose patrons it was built to serve in 1678. Within thirty years it had been extended on two occasions to more or less reach its present size. The ceiling bears the date 1678 and is rather domestic in character, based on deep circular domes with putti, palms and swags. The stained glass in the east window is based on a picture by Alex Ender and was designed by Heaton, Butler and Bayne in 1901. There is an excellent window under the north gallery designed by Lawrence Lee in 1969. The church was sympathetically restored by Ewan Christian in 1882, when the shallow chancel was added. The woodwork it contains was brought from one of Wren's City of London churches. Outside the west wall of the church, set into the footpath, is a boundary marker to show the former parish boundaries of Tonbridge and Speldhurst.
www.kentchurches.info/church.asp?p=Tunbridge+Wells+1
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The large and populous hamlet or village of TUNBRIDGE-WELLS is situated at the south-east boundary of this parish; part of it only is in Speldhurst, another part in the parish of Tunbridge, and the remainder in that of Fant, in the county of Suffex. It consists of four smaller districts, named from the hills on which they stand, Mount Ephraim, Mount Pleasant, and Mount Sion; the other is called The Wells, from their being within it, which altogether form a considerable town; but the last is the centre of business and pleasure, for there, besides the Wells themselves, are the market, public parades, assembly rooms, taverns, shops, &c. Near the Wells is the chapel, which stands remarkably in the three parishes above mentioned—the pulpit in Speldhurst, the altar in Tunbridge, and the vestry in Fant, and the stream, which parted the two counties of Kent and Suffex, formerly ran underneath it, but is now turned to a further distance from it. The right of patronage is claimed by the rector of Speldhurst, though he has never yet possessed the chapel or presented to it; the value of it is about two hundred pounds per annum, which sum is raised by voluntary subscription; divine service is performed in it every day in summer, and three times a week in winter. Adjoining to it is a charity school, for upwards of fifty poor boys and girls, which is supported by a contribution, collected at the chapel doors, two or three times a year.
The trade of Tunbridge-wells is similar to that of Spa, in Germany, and consists chiefly in a variety of toys, made of wood, commonly called Tunbridge ware, which employs a great number of hands. The wood principally used for this purpose is beech and sycamore, with yew and holly inlaid, and beautifully polished. To the market of this place is brought, in great plenty, from the South downs, in Sussex, the little bird, called the wheatear, which, from its delicacy, is usually called the English ortolan. It is not bigger in size than a lark; it is almost a lump of fat, and of a very delicious taste; it is in season only in the midst of summer, when the heat of the weather, and the fatness of it, prevents its being sent to London, which otherwise would, in all likelihood, monopolize every one of them. On the other or Suffex side of the Medway, above a mile from the Wells, are the rocks, which consist of a great number of rude eminences, adjoining to each other, several of which are seventy feet in height; in several places there are cliffs and chasms which lead quite through the midst of them, by narrow gloomy passages, which strike the beholder with astonishment.
THESE MEDICINAL WATERS, commonly called TUNBRIDGE-WELLS, lie so near to the county of Suffex that part of them are within it, for which reason they were for some time called Fant-wells, as being within that parish. (fn. 1) Their efficacy is reported to have been accidentally found out by Dudley lord North, in the beginning of the reign of king James I. Whilst he resided at Eridge-house for his health, lord Abergavenny's seat, in this neighbourhood, and that he was entirely cured of the lingering consumptive disorder he laboured under by the use of them.
The springs, which were then discovered, seem to have been seven in number, two of the principal of which were some time afterwards, by lord Abergavenny's care, inclosed, and were afterwards much resorted to by many of the middling and lower sort, whose ill health had real occasion for the use of them. In which state they continued till queen Henrietta Maria, wife of king Charles I. having been sent hither by her physicians, in the year 1630, for the reestablishment of her health, soon brought these waters into fashion, and occasioned a great resort to them from that time. In compliment to her doctor, Lewis Rowzee, in his treatise on them, calls these springs the Queen's-wells; but this name lasted but a small time, and they were soon afterwards universally known by that of Tunbridge-wells, which names they acquired from the company usually residing at Tunbridge town, when they came into these parts for the benefit of drinking the waters.
The town of Tunbridge being five miles distant from the wells, occasioned some few houses to be built in the hamlets of Southborough and Rusthall, for the accommodation of the company resorting hither, and this place now becoming fashionable, was visited by numbers for the sake of pleasure and dissipation, as well as for the cure of their infirmities; and soon after the Restoration every kind of building, for public amusements, was erected at the two hamlets above mentioned, lodgings and other buildings were built at and near the wells, the springs themselves were secured, and other conveniencies added to them. In 1664, the queen came here by the advice of her physicians, in hopes of reinstating her health, which was greatly impaired by a dangerous fever, and her success, in being perfectly cured by these waters, greatly raised the reputation of them, and the company increasing yearly, it induced the inhabitants to make every accommodation for them adjoining to the Wells, so that both Rusthall and Southborough became ruinous and deserted by all but their native inhabitants. The duke of York, with his duchess, and the two princesses their daughters, visited Tunbridge-wells in the year 1670, which brought much more company than usual to them, and raised their reputation still higher; and the annual increase continuing, it induced the lord of the manor to think of improving this humour of visiting the wells to his own profit as well as the better accommodation of the company. To effect which, he entered into an agreement with his tenants, and hired of them the herbage of the waste of the manor for the term of fifty years, at the yearly rent of ten shillings to each tenant, and then erected shops and houses on and near the walks and springs, in every convenient spot for that purpose; by which means Tunbridge wells became a populous and flourishing village, well inhabited, for whose convenience, and the company resorting thither, a chapel was likewise built, in 1684, by subscription, on some ground given by the lady viscountess Purbeck, which was, about twelve years afterwards, enlarged by an additional subscription, amounting together to near twenty-three hundred pounds.
About the year 1726, the building lease, which had been granted by the lord of the manor of Rusthall, in which this hamlet is situated, expiring, the tenants of the manor claimed a share in the buildings, as a compensation for the loss of the herbage, which was covered by his houses. This occasioned a long and very expensive law suit between them, which was at last determined in favour of the tenants, who were adjudged to have a right to a third part of the buildings then erected on the estate, in lieu of their right to the herbage; upon which all the shops and houses, which had been built on the manor waste, were divided into three lots, of which the tenants were to draw one, and the other two were to remain to the lord of the manor; the lot which the tenants drew was the middle one, which included the assembly room on the public walk, which has since turned out much the most advantageous of the three. After which long articles of agreement, in 1739, were entered into between Maurice Conyers, esq. then lord of the manor of Rusthall, and the above mentioned tenants of it, in which, among many other matters, he agreed to permit the public walks and wells, and divers other premises there, to be made use of for the public benefit of the nobility and gentry resorting thereto, and several regulations were made in them concerning the walks, wells, and wastes of the manor, and for the restraining buildings on the waste, between the lord and his tenants, according to a plan therein specified; all which were confirmed and established by an act of parliament, passed in 1740. Since which several of the royal family have honoured these wells with their presence, and numbers of the nobility and persons of rank and fashion yearly resortto them, so that this place is now in a most flourishing state, having great numbers of good houses built for lodgings, and every other necessary accommodation for the company. Its customs are settled; the employment of the dippers regulated; (fn. 2) its pleasures regulated; its markets well and plentifully supplied, at a reasonable rate, with sowl, fish, meat, every other kind of food, and every convenience added that can contribute to give health and pleasure.
¶The whole neighbourhood of Tunbridge-wells abounds with springs of mineral water, but as the properties of all are nearly the same, only those two, which at the first discovery of them were adjudged the best, are held in any particular estimation. These two wells are enclosed with a handsome triangular stone wall; over the springs are placed two convenient basons of Portland stone, with perforations at the bottom; one of them being given by queen Anne, and the other by the lord of the manor; through which they receive the water, which at the spring is extremely clear and bright. Its taste is steely, but not disagreeable; it has hardly any smell, though sometimes, in a dense air, its ferruginous exhalations are very distinguishable. In point of heat it is invariably temperate, the spring lying so deep in the earth, that neither the heat of summer, nor the cold of winter, affects it. When this water is first taken up in a large glass, its particles continue at rest till it is warmed to nearly the heat of the atmosphere, then a few airy globules begin to separate themselves, and adhere to the sides of the glass, and in a few hours a light copper coloured scum begins to float on the surface, after which an ochreous sediment settles at the bottom. Long continued rains sometimes give the water a milky appearance, but do not otherwise sensibly affect it. From the experiments of different physicians, it appears that the component parts of this water are, steely particles, marine salts, an oily matter, an ochreous substance, simple water, and a volatile vitriolic spirit, too subtile for any chemical analysis. In weight it is, in seven ounces and a quarter, four grains lighter than the German Spa (to which it is preferable on that account) and ten grains lighter than common water; with syrup of violets this water gives a deep green, as vitriols do. (fn. 3) It requires five drops of oleum sulphuris, or elixir of vitriol, to a quart of water, to preserve its virtues to a distance from the spring.
This water is said to be an impregnation of rain in some of the neighbouring eminences, which abound in iron mineral, where it is further enriched with the marine salts and all the valuable ingredients, which constitute it a light and pure chalybeate, which instantly searches the most remote recesses of the human frame, warms and invigorates the relaxed constitution, restores the weakened fibres to their due tone and elasticity, removes those obstructions to which the minuter vessels of the body are liable, and is consequently adapted to most cold chronical disorders, lowness of spirits, weak digestions, and nervous complaints. Dr. Lodowick Rowzee, of Ashford, in this county, wrote a Treatise of the Nature and Virtues of these Waters, printed in 12mo. 1671; and Dr. Patrick Madan wrote a Philosophical and Medical Essay on them, in 1687, in quarto.
The koala (Phascolarctos cinereus), sometimes called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only extant representative of the family Phascolarctidae and its closest living relatives are the wombats. The koala is found in coastal areas of the mainland's eastern and southern regions, inhabiting Queensland, New South Wales, Victoria, and South Australia. It is easily recognisable by its stout, tailless body and large head with round, fluffy ears and large, dark nose. The koala has a body length of 60–85 cm (24–33 in) and weighs 4–15 kg (9–33 lb). Fur colour ranges from silver grey to chocolate brown. Koalas from the northern populations are typically smaller and lighter in colour than their counterparts further south. These populations possibly are separate subspecies, but this is disputed.
Koalas typically inhabit open Eucalyptus woodland, as the leaves of these trees make up most of their diet. This eucalypt diet has low nutritional and caloric content and contains toxic compounds that deter most other mammals from feeding on it. Koalas are largely sedentary and sleep up to twenty hours a day. They are asocial animals, and bonding exists only between mothers and dependent offspring. Adult males communicate with loud bellows that intimidate rivals and attract mates. Males mark their presence with secretions from scent glands located on their chests. Being marsupials, koalas give birth to underdeveloped young that crawl into their mothers' pouches, where they stay for the first six to seven months of their lives. These young koalas, known as joeys, are fully weaned around a year old. Koalas have few natural predators and parasites, but are threatened by various pathogens, such as Chlamydiaceae bacteria and koala retrovirus.
Because of their distinctive appearance, koalas, along with kangaroos and emus, are recognised worldwide as symbols of Australia. They were hunted by Indigenous Australians and depicted in myths and cave art for millennia. The first recorded encounter between a European and a koala was in 1798, and an image of the animal was published in 1810 by naturalist George Perry. Botanist Robert Brown wrote the first detailed scientific description of the koala in 1814, although his work remained unpublished for 180 years. Popular artist John Gould illustrated and described the koala, introducing the species to the general British public. Further details about the animal's biology were revealed in the 19th century by several English scientists. Koalas are listed as a vulnerable species by the International Union for Conservation of Nature. Among the many threats to their existence are habitat destruction caused by agriculture, urbanisation, droughts, and associated bushfires, some related to climate change. In February 2022, the koala was officially listed as endangered in the Australian Capital Territory, New South Wales, and Queensland.
Etymology
The word "koala" comes from the Dharug gula, meaning 'no water'. Although the vowel "u" was originally written in the English orthography as "oo" (in spellings such as coola or koolah — two syllables), the spelling later became "oa" and the word is now pronounced in three syllables, possibly in error.
Adopted by white settlers, "koala" became one of several hundred Aboriginal loan words in Australian English, where it was also commonly referred to as "native bear", later "koala bear", for its supposed resemblance to a bear. It is also one of several Aboriginal words that made it into International English alongside words like "didgeridoo" and "kangaroo". The generic name, Phascolarctos, is derived from the Greek words φάσκωλος (phaskolos) 'pouch' and ἄρκτος (arktos) 'bear'. The specific name, cinereus, is Latin for 'ash coloured'.
Taxonomy
The koala was given its generic name Phascolarctos in 1816 by French zoologist Henri Marie Ducrotay de Blainville, who would not give it a specific name until further review. In 1819, German zoologist Georg August Goldfuss gave it the binomial Lipurus cinereus. Because Phascolarctos was published first, according to the International Code of Zoological Nomenclature, it has priority as the official name of the genus. French naturalist Anselme Gaëtan Desmarest coined the name Phascolarctos fuscus in 1820, suggesting that the brown-coloured versions were a different species than the grey ones. Other names suggested by European authors included Marodactylus cinereus by Goldfuss in 1820, P. flindersii by René Primevère Lesson in 1827, and P. koala by John Edward Gray in 1827.
Evolution
The koala is classified with wombats (family Vombatidae) and several extinct families (including marsupial tapirs, marsupial lions and giant wombats) in the suborder Vombatiformes within the order Diprotodontia. The Vombatiformes are a sister group to a clade that includes macropods (kangaroos and wallabies) and possums. The koala's lineage possibly branched off around 40 million years ago during the Eocene.
The modern koala is the only extant member of Phascolarctidae, a family that includes several extinct genera and species. During the Oligocene and Miocene, koalas lived in rainforests and had more generalised diets. Some species, such as the Riversleigh rainforest koala (Nimiokoala greystanesi) and some species of Perikoala, were around the same size as the modern koala, while others, such as species of Litokoala, were one-half to two-thirds its size Like the modern species, prehistoric koalas had well developed ear structures which suggests that they also made long-distance vocalisations and had a relatively inactive lifestyle. During the Miocene, the Australian continent began drying out, leading to the decline of rainforests and the spread of open Eucalyptus woodlands. The genus Phascolarctos split from Litokoala in the late Miocene, and had several adaptations that allowed it to live on a specialised eucalyptus diet: a shifting of the palate towards the front of the skull; upper teeth lined by thicker bone, molars located relatively low compared the jaw joint and with more chewing surface; smaller pterygoid fossa; and a larger gap separating the incisor teeth and the molars.
P. cinereus may have emerged as a dwarf form of the giant koala (P. stirtoni), following the disappearance of several giant animals in the late Pleistocene. A 2008 study questions this hypothesis, noting that P. cinereus and P. stirtoni were sympatric during the middle to late Pleistocene, and the major difference in the morphology of their teeth. The fossil record of the modern koala extends back at least to the middle Pleistocene.
Molecular relationship between living Diprotodontia families based on Phillips and collages (2023)
Vombatidae (wombats)
Phascolarctidae (koalas)
Acrobatidae
Tarsipedidae (honey possum)
Petauridae (wrist-winged gliders and allies)
Pseudocheiridae (ringtail possums and allies)
Macropodidae (kangaroos, wallabies and allies)
Phalangeridae (brushtail possums and cuscuses)
Burramyidae (pygmy possums)
Morphology tree of Phascolarctidae based on Beck and collages (2020)
Thylacoleonidae (extinct marsupial lion and allies)
Vombatomorphia (wombats and fossil relatives)
Phascolarctidae
Priscakoala lucyturnbullae
Madakoala spp.
Perikoala robustus
Nimiokoala greystanesi
Litokoala dicksmithi
Litokoala kutjamarpensis
Phascolarctos cinereus
Genetics and variations
Three subspecies are recognised: the Queensland koala (Phascolarctos cinereus adustus, Thomas 1923), the New South Wales koala (Phascolarctos cinereus cinereus, Goldfuss 1817), and the Victorian koala (Phascolarctos cinereus victor, Troughton 1935). These forms are distinguished by pelage colour and thickness, body size, and skull shape. The Queensland koala is the smallest of the three, with silver or grey short hairs and a shorter skull. The Victorian koala is the largest, with shaggier, brown fur and a wider skull. The geographic limits of these variations are based on state borders, and their status as subspecies is disputed. A 1999 genetic study suggests koalas exist as a cline within a single evolutionarily significant unit with limited gene flow between local populations.
Other studies have found that koala populations have high levels of inbreeding and low genetic variation. Such low genetic diversity may have been caused by declines in the population during the late Pleistocene. Rivers and roads have been shown to limit gene flow and contribute to the isolation of southeast Queensland populations. In April 2013, scientists from the Australian Museum and Queensland University of Technology announced they had fully sequenced the koala genome.
Characteristics
The koala is a robust animal with a large head and vestigial or non-existent tail. It has a body length of 60–85 cm (24–33 in) and a weight of 4–15 kg (9–33 lb), making it among the largest arboreal marsupials. Koalas from Victoria are twice as heavy as those from Queensland. The species is sexually dimorphic, with males 50% larger than females. Males are further distinguished from females by their more curved noses and the presence of chest glands, which are visible as bald patches. The female's pouch opening is secured by a sphincter which holds the young in.
The pelage of the koala is denser on the back. The back fur colour varies from light grey to chocolate brown. The belly fur is whitish; on the rump it is mottled whitish and dark. The koala has the most effective insulating back fur of any marsupial and is highly resilient to wind and rain, while the belly fur can reflect solar radiation. The koala's curved, sharp claws are well adapted for climbing trees. The large forepaws have two opposable digits (the first and second, which are opposable to the other three) that allow them to grip small branches. On the hind paws, the second and third digits are fused, a typical condition for members of the Diprotodontia, and the attached claws (which are still separate) function like a comb. The animal has a robust skeleton and a short, muscular upper body with relatively long upper limbs that contribute to its ability to scale trees. In addition, the thigh muscles are anchored further down the shinbone, increasing its climbing power.
For a mammal, the koala has a proportionally small brain, being 60% smaller than that of a typical diprotodont, weighing only 19.2 g (0.68 oz) on average. The brain's surface is fairly smooth and "primitive". It does not entirely fill up the cranial cavity, unlike in most mammals, and is lightened by large amounts of cerebrospinal fluid. It is possible that the fluid protects the brain when animal falls from a tree. The koala's small brain size may be an adaptation to the energy restrictions imposed by its diet, which is insufficient to sustain a larger brain. Because of its small brain, the koala has a limited ability to perform complex, unusual behaviours. For example, it will not eat plucked leaves on a flat surface, which conflicts with its normal feeding routine.
The koala has a broad, dark nose with a good sense of smell, and it is known to sniff the oils of individual branchlets to assess their edibility. Its relatively small eyes are unusual among marsupials in that the pupils have vertical slits, an adaptation to living on a more vertical plane. Its round ears provide it with good hearing, and it has a well-developed middle ear. The koala larynx is located relatively low in the vocal tract and can be pulled down even further. They also possess unique folds in the velum (soft palate), known as velar vocal folds, in addition to the typical vocal folds of the larynx. These features allow the koala to produce deeper sounds than would otherwise be possible for their size.
The koala has several adaptations for its poor, toxic and fibrous diet. The animal's dentition consists of the incisors and cheek teeth (a single premolar and four molars on each jaw), which are separated by a large gap (a characteristic feature of herbivorous mammals). The koala bites a leaf with the incisors and clips it with the premolars at the petiole, before chewing it to pieces with the cusped molars. Koalas may also store food in their cheek pouches before it is ready to be chewed. The partially worn molars of koalas in their prime are optimal for breaking the leaves into small particles, resulting in more efficient stomach digestion and nutrient absorption in the small intestine, which digests the eucalyptus leaves to provide most of the animal's energy. A koala sometimes regurgitates the food into the mouth to be chewed a second time.
Koalas are hindgut fermenters, and their digestive retention can last for up to 100 hours in the wild or up to 200 hours in captivity. This is made possible by their caecum—200 cm (80 in) long and 10 cm (4 in) in diameter—possibly the largest for an animal when accounting for its size. Koalas can hold food particles for longer fermentation if needed. They are more likely keep smaller particles as larger ones take longer to digest. While the hindgut is relatively large, only 10% of the animal's energy is obtained from digestion in this chamber. The koala's metabolic rate is only 50% of the typical mammalian rate, owing to its low energy intake, although this can vary between seasons and sexes. They can digest the toxic plant secondary metabolites, phenolic compounds and terpenes present in eucalyptus leaves due to their production of cytochrome P450, which breaks down these poisons in the liver. The koala replaces lost water at a lower rate than some other species like some possums. It maintains water by absorbing it in the caecum, resulting in drier faecal pellets packed with undigested fibre.
Distribution and habitat
The koala's geographic range covers roughly 1,000,000 km2 (390,000 sq mi), and 30 ecoregions. It ranges throughout mainland eastern and southeastern Australia, including the states of Queensland, New South Wales, Victoria, and South Australia. The koala was also introduced to several nearby islands. The population on Magnetic Island represents the northern limit of its range.
Fossil evidence shows that the koala's range stretched as far west as southwestern Western Australia during the late Pleistocene. They were likely driven to extinction in these areas by environmental changes and hunting by Indigenous Australians. Koalas were introduced to Western Australia at Yanchep in 2022. Koalas can be found in both tropical and temperate habitats ranging from dense woodlands to more spaced-out forests. In semi-arid climates, they prefer riparian habitats, where nearby streams and creeks provide refuge during times of drought and extreme heat.
Behaviour and ecology
Koalas are herbivorous, and while most of their diet consists of eucalypt leaves, they can be found in trees of other genera, such as Acacia, Allocasuarina, Callitris, Leptospermum, and Melaleuca. Though the foliage of over 600 species of Eucalyptus is available, the koala shows a strong preference for around 30. They prefer plant matter with higher protein over fibre and lignin. The most favoured species are Eucalyptus microcorys, E. tereticornis, and E. camaldulensis, which, on average, make up more than 20% of their diet. Despite its reputation as a picky eater, the koala is more generalist than some other marsupial species, such as the greater glider. The koala does not need to drink often as it can get enough water in the eucalypt leaves, though larger males may additionally drink water found on the ground or in tree hollows. When feeding, a koala reaches out to grab leaves with one forepaw while the other paws hang on to the branch. Depending on the size of the individual, a koala can walk to the end of a branch or must stay near the base. Each day, koalas eat up to 400 grams (14 oz) of leaves, spread over four to six feeding periods. Despite their adaptations to a low-energy lifestyle, they have meagre fat reserves and need to feed often.
Due to their low-energy diet, koalas limit their activity and sleep 20 hours a day. They are predominantly active at night and spend most of their waking hours foraging. They typically eat and sleep in the same tree, possibly for as long as a day. On warm days, a koala may rest with its back against a branch or lie down with its limbs dangling. When it gets very hot, the koala rests lower in the canopy and near the trunk, where the surface is cooler than the surrounding air. It curls up when it gets cold and wet. A koala will find a lower, thicker branch on which to rest when it gets windy. While it spends most of the time in the tree, the animal descends to the ground to move to another tree, leaping along. The koala usually grooms itself with its hind paws, with their double claws, but sometimes uses its forepaws or mouth.
Social life
Koalas are asocial animals and spend just 15 minutes a day on social behaviours. Where there are more koalas and fewer trees, home ranges are smaller and more clumped while the reverse is true for areas with fewer animals and more trees. Koala society appears to consist of "residents" and "transients", the former being mostly adult females and the latter males. Resident males appear to be territorial and dominant. The territories of dominant males are found near breeding females, while younger males must wait until they reach full size to challenge for breeding rights. Adult males occasionally venture outside their home ranges; when they do so, dominant ones retain their status. As a male climbs a new tree, he rubs his chest against it and sometimes dribbles urine. This scent-marking behaviour probably serves as communication, and individuals are known to sniff the bottom of a newly found tree. Chest gland secretions are complex chemical mixtures — about 40 compounds were identified in one analysis — that vary in composition and concentration with the season and the age of the individual.
Adult males communicate with loud bellows — "a long series of deep, snoring inhalations and belching exhalations". Because of their low frequency, these bellows can travel far through the forest. Koalas may bellow at any time of the year, particularly during the breeding season, when it serves to attract females and possibly intimidate other males. They also bellow to advertise their presence to their neighbours when they climb a different tree. These sounds signal the male's actual body size, as well as exaggerate it; females pay more attention to bellows that originate from larger males. Female koalas bellow, though more softly, in addition to making snarls, wails, and screams. These calls are produced when in distress and when making defensive threats. Squeaking and sqawking are produced when distraught; the former is made by younger animals and the latter by older ones. When another individual climbs over it, a koala makes a low closed-mouth grunt. Koalas also communicate with facial expressions. When snarling, wailing, or squawking, the animal curls the upper lip and points its ears forward. Screaming koalas pull their lips and ears back. Females form an oval shape with their lips when annoyed.
Agonistic behaviour typically consists of quarrels between individuals that are trying to pass each other in the tree. This occasionally involves biting. Strangers may wrestle, chase, and bite each other. In extreme situations, a male may try to displace a smaller rival from a tree, chasing, cornering and biting it. Once the individual is driven away, the victor bellows and marks the tree. Pregnant and lactating females are particularly aggressive and attack individuals that come too close. In general, however, koalas tend to avoid fighting due to energy costs.
Reproduction and development
A young joey, preserved at Port Macquarie Koala Hospital
Koalas are seasonal breeders, and give birth from October to May. Females in oestrus lean their heads back and shake their bodies. Despite these obvious signals, males will try to copulate with any female during this period, mounting them from behind. Because of his much larger size, a male can overpower a female. A female may scream and vigorously fight off her suitors but will accede to one that is dominant or familiar. The commotion can attract other males to the scene, obliging the incumbent to delay mating and fight off the intruders. A female may learn who is more dominant during these fights. Older males usually have accumulated scratches, scars, and cuts on the exposed parts of their noses and their eyelids.
Koalas are induced ovulators. The gestation period lasts 33–35 days, and a female gives birth to one joey (although twins do occur). As marsupials, the young are born tiny and barely formed, weighing no more than 0.5 g (0.02 oz). However, their lips, forelimbs, and shoulders are relatively advanced, and they can breathe, defecate and urinate. The joey crawls into its mother's pouch to continue the rest of its development. Female koalas do not clean their pouches, an unusual trait among marsupials.
The joey latches on to one of the female's two teats and suckles it. The female lactates for as long as a year to make up for her low energy production. Unlike in other marsupials, koala milk becomes less fatty as the joey grows in the pouch. After seven weeks, the joey has a proportionally large head, clear edges around its face, more colouration, and a visible pouch (if female) or scrotum (male). At 13 weeks, the joey weighs around 50 g (1.8 oz) and its head is twice as big as before. The eyes begin to open and hair begins to appear. At 26 weeks, the fully furred animal resembles an adult and can look outside the pouch.
Mother with joey on back
At six or seven months of age, the joey weighs 300–500 g (11–18 oz) and fully emerges from the pouch for the first time. It explores its new surroundings cautiously, clutching its mother for support. Around this time, the mother prepares it for a eucalyptus diet by producing a faecal pap that the joey eats from her cloaca. This pap comes from the cecum, is more liquid than regular faeces, and is filled with bacteria. A nine month old joey has its adult coat colour and weighs 1 kg (2.2 lb). Having permanently left the pouch, it rides on its mother's back for transportation, learning to climb by grasping branches. Gradually, it becomes more independent from its mother, who becomes pregnant again after a year, and the young is now around 2.5 kg (5.5 lb). Her bond with her previous offspring is permanently severed and she no longer allows it to suckle, but it will stay nearby until it is one-and-a-half to two years old.
Females become sexually mature at about three years of age and can then become pregnant; in comparison, males reach sexual maturity when they are about four years old, although they can experience spermatogenesis as early as two years. Males do not start marking their scent until they reach sexual maturity, though their chest glands become functional much earlier. Koalas can breed every year if environmental conditions are good, though the long dependence of the young usually leads to year-long gaps in births.
Health and mortality
Koalas may live from 13 to 18 years in the wild. While female koalas usually live this long, males may die sooner because of their more risky lives. Koalas usually survive falls from trees and can climb back up, but they can get hurt and even die, particularly inexperienced young and fighting males. Around six years of age, the koala's chewing teeth begin to wear down and their chewing efficiency decreases. Eventually, the cusps disappear completely and the animal will die of starvation. Koalas have few predators. Dingos and large pythons and some birds of prey may take them. Koalas are generally not subject to external parasites, other than ticks around the coast. The mite Sarcoptes scabiei gives koalas mange, while the bacterium Mycobacterium ulcerans skin ulcers, but even these are uncommon. Internal parasites are few and have little effect. These include the tapeworm Bertiella obesa, commonly found in the intestine, and the nematodes Marsupostrongylus longilarvatus and Durikainema phascolarcti, which are infrequently found in the lungs.[59] In a three-year study of almost 600 koalas taken to the Australia Zoo Wildlife Hospital in Queensland, 73.8% of the animals were infected with parasitic protozoal genus Trypanosoma, the most frequent of which was T. irwini.
Koalas can be subject to pathogens such as Chlamydiaceae bacteria, which can cause keratoconjunctivitis, urinary tract infection, and reproductive tract infection. Such infections are common on the mainland, but absent in some island populations. The koala retrovirus (KoRV) may cause koala immune deficiency syndrome (KIDS) which is similar to AIDS in humans. Prevalence of KoRV in koala populations suggests a trend spreading from north to south, where populations go from being completely infected to being partially uninfected.
The animals are vulnerable to bushfires due to their slow speed and the flammability of eucalypt trees. The koala instinctively seeks refuge in the higher branches, where it is vulnerable to intense heat and flames. Bushfires also break up the animal's habitat, which isolates them, decreases their numbers and creates genetic bottlenecks. Dehydration and overheating can also prove fatal. Consequently, the koala is vulnerable to the effects of climate change. Models of climate change in Australia predict warmer and drier climates, suggesting that the koala's range will shrink in the east and south to more mesic habitats.
Human relations
The first written reference to the koala was recorded by John Price, servant of John Hunter, the Governor of New South Wales. Price encountered the "cullawine" on 26 January 1798, during an expedition to the Blue Mountains, but his remarks would first be published in Historical Records of Australia, nearly a century later. In 1802, French-born explorer Francis Louis Barrallier encountered the animal when his two Aboriginal guides, returning from a hunt, brought back two koala feet they were intending to eat. Barrallier preserved the appendages and sent them and his notes to Hunter's successor, Philip Gidley King, who forwarded them to Joseph Banks. Similar to Price, Barrallier's notes were not published until 1897. Reports of the "Koolah" appeared in the Sydney Gazette in late 1803, and helped provide the impetus for King to send the artist John Lewin to paint watercolours of the animal. Lewin painted three pictures, one of which was used as a print in Georges Cuvier's Le Règne Animal (The Animal Kingdom) (1827).
Botanist Robert Brown was the first to write a formal scientific description of the koala in 1803, based on a female specimen captured near what is now Mount Kembla in the Illawarra region of New South Wales. Austrian botanical illustrator Ferdinand Bauer drew the animal's skull, throat, feet, and paws. Brown's work remained unpublished and largely unnoticed, however, as his field books and notes remained in his possession until his death, when they were bequeathed to the British Museum (Natural History) in London. They were not identified until 1994, while Bauer's koala watercolours were not published until 1989. William Paterson, who had befriended Brown and Bauer during their stay in New South Wales, wrote an eyewitness report of his encounters with the animals and this would be the basis for British surgeon Everard Home's anatomical writings on them. Home, who in 1808 published his report in the journal Philosophical Transactions of the Royal Society, coined the scientific name Didelphis coola.
George Perry would officially publish the first image of the koala in his 1810 natural history work Arcana. Perry called it the "New Holland Sloth", and his dislike for the koala, evident in his description of the animal, was reflected in the contemporary British attitudes towards Australian animals as strange and primitive the eye is placed like that of the Sloth, very close to the mouth and nose, which gives it a clumsy awkward appearance, and void of elegance in the combination they have little either in their character or appearance to interest the Naturalist or Philosopher. As Nature however provides nothing in vain, we may suppose that even these torpid, senseless creatures are wisely intended to fill up one of the great links of the chain of animated nature.
Natural history illustrator John Gould popularised the koala with his 1863 work The Mammals of Australia.
Naturalist and popular artist John Gould illustrated and described the koala in his three-volume work The Mammals of Australia (1845–1863) and introduced the species, as well as other members of Australia's little-known faunal community, to the public. Comparative anatomist Richard Owen, in a series of publications on the physiology and anatomy of Australian mammals, presented a paper on the anatomy of the koala to the Zoological Society of London. In this widely cited publication, he provided an early description of its internal anatomy, and noted its general structural similarity to the wombat. English naturalist George Robert Waterhouse, curator of the Zoological Society of London, was the first to correctly classify the koala as a marsupial in the 1840s, and compared it to fossil species Diprotodon and Nototherium, which had been discovered just recently. Similarly, Gerard Krefft, curator of the Australian Museum in Sydney, noted evolutionary mechanisms at work when comparing the koala to fossil marsupials in his 1871 The Mammals of Australia.
Britain finally received a living koala in 1881, which was obtained by the Zoological Society of London. As related by prosecutor to the society, William Alexander Forbes, the animal suffered an accidental demise when the heavy lid of a washstand fell on it and it was unable to free itself. Forbes dissected the fresh specimen and wrote about the female reproductive system, the brain, and the liver — parts not previously described by Owen, who had access only to preserved specimens. Scottish embryologist William Caldwell — well known in scientific circles for determining the reproductive mechanism of the platypus — described the uterine development of the koala in 1884, and used this new information to convincingly map out the evolutionary timeline of the koala and the monotremes.
Main article: Koala emblems and popular culture
The koala is well known worldwide and is a major draw for Australian zoos and wildlife parks. It has been featured in popular culture and as soft toys. It benefited the Australian tourism industry by over $1 billion in 1998, and this has subsequently grown. Its international popularly rose after World War II, when tourism to Australia increased and the animals were exported to zoos overseas. In 1997, about 75% of European and Japanese tourists placed the koala at the top of their list of animals to see. According to biologist Stephen Jackson: "If you were to take a straw poll of the animal most closely associated with Australia, it's a fair bet that the koala would come out marginally in front of the kangaroo". Factors that contribute to the koala's enduring popularity include its teddy bear-like appearance with childlike body proportions.
The koala is featured in the Dreamtime stories and mythology of Indigenous Australians. The Tharawal people believed that the animal helped them get to the continent by rowing the boat. Another myth tells of how a tribe killed a koala and used its long intestines to create a bridge for people from other parts of the world. How the koala lost its tail has been the subject of many tales. In one, a kangaroo cuts it off to punish the koala for its uncouth behaviour. Tribes in both Queensland and Victoria regarded the koala as a wise animal which gave valuable guidance. Bidjara-speaking people credited the koala for making trees grow in their arid lands. The animal is also depicted in rock carvings, though less so than some other species.
Early European settlers in Australia considered the koala to be a creeping sloth-like animal with a "fierce and menacing look". At the turn of the 20th century, the koala's reputation took a more positive turn. It appears in Ethel Pedley's 1899 book Dot and the Kangaroo, as the "funny native bear". Artist Norman Lindsay depicted a more anthropomorphic koala in The Bulletin cartoons, starting in 1904. This character also appeared as Bunyip Bluegum in Lindsay's 1918 book The Magic Pudding. The most well known fictional koala is Blinky Bill. Created by Dorothy Wall in 1933, the character appeared in several books and has been the subject of films, TV series, merchandise, and a 1986 environmental song by John Williamson. The koala first appeared on an Australian stamp in 1930.
The song "Ode to a Koala Bear" appears on the B-side of the 1983 Paul McCartney/Michael Jackson duet single Say Say Say. A koala is the main character in Hanna-Barbera's The Kwicky Koala Show and Nippon Animation's Noozles, both of which were animated cartoons of the early 1980s. Food products shaped like the koala include the Caramello Koala chocolate bar and the bite-sized cookie snack Koala's March. Dadswells Bridge in Victoria features a tourist complex shaped like a giant koala and the Queensland Reds rugby team has a koala as its icon.
Koala diplomacy
Several political leaders and members of royal families had their pictures taken with koalas, including Queen Elizabeth II, Prince Harry, Crown Prince Naruhito, Crown Princess Masako, Pope John Paul II, US President Bill Clinton, Soviet premier Mikhail Gorbachev and South African President Nelson Mandela At the 2014 G20 Brisbane summit, hosted by Prime Minister Tony Abbott, many world leaders, including Russian President Vladimir Putin and US President Barack Obama, were photographed holding koalas. The event gave rise to the term "koala diplomacy", which then became the Oxford Word of the Month for December 2016. The term also includes the loan of koalas by the Australian government to overseas zoos in countries such as Singapore and Japan, as a form of "soft power diplomacy", like the "panda diplomacy" practised by China.
Main article: Koala conservation
The koala was originally classified as Least Concern on the Red List, and reassessed as Vulnerable in 2014. In the Australian Capital Territory, New South Wales and Queensland, the species was listed under the EPBC Act in February 2022 as endangered by extinction. The described population was determined in 2012 to be "a species for the purposes of the EPBC Act 1999" in Federal legislation.
Australian policymakers had declined a 2009 proposal to include the koala in the Environment Protection and Biodiversity Conservation Act 1999. A 2017 WWF report found a 53% decline per generation in Queensland, and a 26% decline in New South Wales. The koala population in South Australia and Victoria appear to be abundant; however, the Australian Koala Foundation (AKF) argued that the exclusion of Victorian populations from protective measures was based on a misconception that the total koala population was 200,000, whereas they believed in 2012 that it was probably less than 100,000. AKF estimated in 2022 that there could be only 43,000–100,000.[80] This is compared with 8 to 10 million at the start of the 20th century. The Australian Government's Threatened Species Scientific Committee estimated that the 2021 koala population was 92,000, down from 185,000 two decades prior.
The koala was heavily hunted by European settlers in the early 20th century, largely for its fur. Australia exported as many as two million pelts by 1924. Koala furs were used to make rugs, coat linings, muffs, and on women's garment trimmings. The first successful efforts at conserving the species were initiated by the establishment of Brisbane's Lone Pine Koala Sanctuary and Sydney's Koala Park Sanctuary in the 1920s and 1930s. The owner of the latter park, Noel Burnet, created the first successful breeding program and earned a reputation as a top expert on the species.
One of the biggest anthropogenic threats to the koala is habitat destruction and fragmentation. Near the coast, the main cause of this is urbanisation, while in rural areas, habitat is cleared for agriculture. Its favoured trees are also taken down to be made into wood products. In 2000, Australia had the fifth highest rate of land clearance globally, having removed 564,800 hectares (1,396,000 acres) of native plants. The distribution of the koala has shrunk by more than 50% since European arrival, largely due to fragmentation of habitat in Queensland. Nevertheless, koalas live in many protected areas.
While urbanisation can pose a threat to koala populations, the animals can survive in urban areas provided enough trees are present. Urban populations have distinct vulnerabilities: collisions with vehicles and attacks by domestic dogs. Cars and dogs kill about 4,000 animals every year. To reduce road deaths, government agencies have been exploring various wildlife crossing options, such as the use of fencing to channel animals toward an underpass, in some cases adding a ledge as a walkway to an existing culvert. Injured koalas are often taken to wildlife hospitals and rehabilitation centres. In a 30-year retrospective study performed at a New South Wales koala rehabilitation centre, trauma was found to be the most frequent cause of admission, followed by symptoms of Chlamydia infection
Khajuraho, a UNESCO World Heritage Site, is a famous tourist and archaeological site known for its sculptured temples dedicated to Shiva, Vishnu, Jain patriarchs, and others. Khajuraho was one of the capitals of the Chandela kings, who from the 9th to the 12th century CE developed a large realm, which at its height included almost all of what is now Madhya Pradesh state. Khajuraho extended over 21 sq. km and contained about 85 temples, built by multiple rulers mostly between 950 to 1050. In the 12th century the Chandelas, in a period of chaos and decline, abandoned Khajuraho and moved to hill forts elsewhere. Khajuraho puttered along until the 14th century (Ibn Batuta was impressed by its temples) but was afterwards largely forgotten and overtaken by jungles, which probably saved it from the desecration that Muslim conquerors sometimes inflicted on Hindu monuments. In 1838 a British army captain, TS Burt, employed by the Asiatic Society in Calcutta, came upon information that led him to the rediscovery of the complex of temples in the jungle in Khajuraho.
Of the 85 original temples—most constructed of sandstone—about 20 are still reasonably well preserved. Both internally and externally the temples are richly carved with excellent sculptures that are frequently sensual and often sexually explicit. The temples are divided into three complexes—the western is the largest and best known, containing the magnificent Shaivite temple Kandariya Mahadev, a 31m high agglomeration of porches and turrets culminating in a spire. Khajuraho's name derives from the prevalence of khajur, or date palms, in the area.
There are many theories explaining the real reason for incorporating erotic art with the temples. One of the most popular theories is that they were meant to provide education about earthly desires. During the medieval era, young boys were sent to hermitage and practice bramhacharya till reaching maturity. The sculptures were meant to educate them about grahasthahram.
Yet another theory states that they were meant to represent kama (desire) as the third purushartha (aim of life). The sculptures are also accredited to the rise of the Tantric cults during that time. However, several experts reject these theories.
However, the most credible explanations for the erotic sculptures at Khajuraho are as follows:
1. Sign of happiness, prosperity and auspiciousness: During the medieval era there was a common belief that having erotic sculptures were considered alankaras or decorative motifs, protective and auspicious. This hypothesis is based on the authoritative religious texts like the Shilpashastras and the Brihat Samhita. According to the Brihat Samhita, mithunas (couples), goblins, creepers and erotic sculptures were meant to be carved on the temple door to bring in good luck and as a sign of auspiciousness.
2. Mock the Ascetics: Yet another theory explaining the old erotic art at Khajuraho states that the coital couples represent ascetics as well as people from the royal class. It is also believed that the sculptures of couples participating in the orgies are scenes imagined by the artists. Also, it is also said that the artists used ascetics in sensuous and passionate moods as a way to mock the extreme Tantric sects that rose during that era.
3. Code Language: According to experts, the erotic figures were used as a code language to convey Tantric doctrines and non-communicable experiences. For e.g. A sculpture wherein a washerwoman clings to an ascetic may look erotic and sensuous for the layman. However, in the Tantric language, the washerwoman represents the Kundalini energy that has ascended up to the chakras i.e. the neck of the ascetic. Thus, the erotic sculptures have a deeper meaning related to the Tantric cult.
4. Conceal the Magico-Propitiory Yantra: According to the architectural text of the Shilpa Prakasha, every temple must have a Kamakala Yantra strategically placed to protect it from evil spirits and natural calamities. However, the Yantra, which is basically a set of lines drawn symmetrically, must not be visible to the lay man. Thus, a few erotic sculptures with the head down posture were made corresponding to the lines of the Yantra and superimposed on it.
5. Non-duality: It is possible that the erotic sculptures symbolically represent the union of two opposing forces or energies like, inhalation and exhalation, in a timeless state of non-duality.
The erotic art at Khajuraho is considered to be the pinnacle of love and passion. However, during the period between 900 – 1300 AD most Hindu, Jain and Buddhist temples across western and southern India comprised of erotic art.
However, at other temples, these sculptures were carved at the plinth level, below the eye level and thus did not get noticed. It is only at Khajuraho that these sculptures were so prominently displayed on the main wall of the temples
Squid are cephalopods in the superorder Decapodiformes with elongated bodies, large eyes, eight arms and two tentacles. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.
Squid diverged from other cephalopods during the Jurassic and occupy a similar role to teleost fish as open water predators of similar size and behaviour. They play an important role in the open water food web. The two long tentacles are used to grab prey and the eight arms to hold and control it. The beak then cuts the food into suitable size chunks for swallowing. Squid are rapid swimmers, moving by jet propulsion, and largely locate their prey by sight. They are among the most intelligent of invertebrates, with groups of Humboldt squid having been observed hunting cooperatively. They are preyed on by sharks, other fish, sea birds, seals and cetaceans, particularly sperm whales.
Squid can change colour for camouflage and signalling. Some species are bioluminescent, using their light for counter-illumination camouflage, while many species can eject a cloud of ink to distract predators.
Squid are used for human consumption with commercial fisheries in Japan, the Mediterranean, the southwestern Atlantic, the eastern Pacific and elsewhere. They are used in cuisines around the world, often known as "calamari". Squid have featured in literature since classical times, especially in tales of giant squid and sea monsters.
TAXONOMY AND PHYLOGENY
Squid are members of the class Cephalopoda, subclass Coleoidea. The squid orders Myopsida and Oegopsida are in the superorder Decapodiformes (from the Greek for "ten-legged"). Two other orders of decapodiform cephalopods are also called squid, although they are taxonomically distinct from squids and differ recognizably in their gross anatomical features. They are the bobtail squid of order Sepiolida and the ram's horn squid of the monotypic order Spirulida. The vampire squid, however, is more closely related to the octopuses than to any squid.
The cladogram, not fully resolved, is based on Sanchez et al, 2018. Their molecular phylogeny used mitochondrial and nuclear DNA marker sequences; they comment that a robust phylogeny "has proven very challenging to obtain". If it is accepted that Sepiidae cuttlefish are a kind of squid, then the squids, excluding the vampire squid, form a clade as illustrated. Orders are shown in boldface; all the families not included in those orders, except Sepiadariidae and Sepiidae are in the paraphyletic order "Sepiida", are in the paraphyletic order "Oegopsida".
EVOLUTION
Crown coleoids (the ancestors of octopuses and squid) diverged at the end of the Paleozoic, in the Permian. Squid diverged during the Jurassic, but many squid families appeared in or after the Cretaceous. Both the coleoids and the teleost fish were involved in much adaptive radiation at this time, and the two modern groups resemble each other in size, ecology, habitat, morphology and behaviour, however some fish moved into fresh water while the coleoids remained in marine environments.
The ancestral coleoid was probably nautiloid-like with a strait septate shell that became immersed in the mantle and was used for buoyancy control. Four lines diverged from this, Spirulida (with one living member), the cuttlefishes, the squids and the octopuses. Squid have differentiated from the ancestral mollusc such that the body plan has been condensed antero-posteriorly and extended dorso-ventrally. What may have been the foot of the ancestor is modified into a complex set of appendages around the mouth. The sense organs are highly developed and include advanced eyes similar to those of vertebrates.
The ancestral shell has been lost, with only an internal gladius, or pen, remaining. The pen, made of a chitin-like material, is a feather-shaped internal structure that supports the squid's mantle and serves as a site for muscle attachment. The cuttlebone or sepion of the Sepiidae is calcareous and appears to have evolved afresh in the Tertiary
DESCIPTION
Squid are soft-bodied molluscs whose forms evolved to adopt an active predatory lifestyle. The head and foot of the squid are at one end of a long body, and this end is functionally anterior, leading the animal as it moves through the water. A set of eight arms and two distinctive tentacles surround the mouth; each appendage takes the form of a muscular hydrostat and is flexible and prehensile, usually bearing disc-like suckers.
The suckers may lie directly on the arm or be stalked. Their rims are stiffened with chitin and may contain minute toothlike denticles. These features, as well as strong musculature, and a small ganglion beneath each sucker to allow individual control, provide a very powerful adhesion to grip prey. Hooks are present on the arms and tentacles in some species, but their function is unclear. The two tentacles are much longer than the arms and are retractile. Suckers are limited to the spatulate tip of the tentacle, known as the manus.
In the mature male, the outer half of one of the left arms is hectocotylised – and ends in a copulatory pad rather than suckers. This is used for depositing a spermatophore inside the mantle cavity of a female. A ventral part of the foot has been converted into a funnel through which water exits the mantle cavity.
The main body mass is enclosed in the mantle, which has a swimming fin along each side. These fins are not the main source of locomotion in most species. The mantle wall is heavily muscled and internal. The visceral mass, which is covered by a thin, membranous epidermis, forms a cone-shaped posterior region known as the "visceral hump". The mollusc shell is reduced to an internal, longitudinal chitinous "pen" in the functionally dorsal part of the animal; the pen acts to stiffen the squid and provides attachments for muscles.
On the functionally ventral part of the body is an opening to the mantle cavity, which contains the gills (ctenidia) and openings from the excretory, digestive and reproductive systems. An inhalant siphon behind the funnel draws water into the mantel cavity via a valve. The squid uses the funnel for locomotion via precise jet propulsion. In this form of locomotion, water is sucked into the mantle cavity and expelled out of the funnel in a fast, strong jet. The direction of travel is varied by the orientation of the funnel. Squid are strong swimmers and certain species can "fly" for short distances out of the water.
CAMOUFLAGE
Squid make use of different kinds of camouflage, namely active camouflage for background matching (in shallow water) and counter-illumination. This helps to protect them from their predators and allows them to approach their prey.
The skin is covered in controllable chromatophores of different colours, enabling the squid to match its coloration to its surroundings. The play of colours may in addition distract prey from the squid's approaching tentacles. The skin also contains light reflectors called iridophores and leucophores that, when activated, in milliseconds create changeable skin patterns of polarized light. Such skin camouflage may serve various functions, such as communication with nearby squid, prey detection, navigation, and orientation during hunting or seeking shelter. Neural control of the iridophores enabling rapid changes in skin iridescence appears to be regulated by a cholinergic process affecting reflectin proteins.
Some mesopelagic squid such as the firefly squid (Watasenia scintillans) and the midwater squid (Abralia veranyi) use counter-illumination camouflage, generating light to match the downwelling light from the ocean surface. This creates the effect of countershading, making the underside lighter than the upperside.
Counter-illumination is also used by the Hawaiian bobtail squid (Euprymna scolopes), which has symbiotic bacteria (Aliivibrio fischeri) that produce light to help the squid avoid nocturnal predators. This light shines through the squid's skin on its underside and is generated by a large and complex two-lobed light organ inside the squid's mantle cavity. From there, it escapes downwards, some of it travelling directly, some coming off a reflector at the top of the organ (dorsal side). Below there is a kind of iris, which has branches (diverticula) of its ink sac, with a lens below that; both the reflector and lens are derived from mesoderm. The squid controls light production by changing the shape of its iris or adjusting the strength of yellow filters on its underside, which presumably change the balance of wavelengths emitted. Light production shows a correlation with intensity of down-welling light, but it is about one third as bright; the squid can track repeated changes in brightness. Because the Hawaiian bobtail squid hides in sand during the day to avoid predators, it does not use counter-illumination during daylight
PREDATOR DISTRACTION WITH INK
Squid distract attacking predators by ejecting a cloud of ink, giving themselves an opportunity to escape. The ink gland and its associated ink sac empties into the rectum close to the anus, allowing the squid to rapidly discharge black ink into the mantle cavity and surrounding water. The ink is a suspension of melanin particles and quickly disperses to form a dark cloud that obscures the escape manoeuvres of the squid. Predatory fish may also be deterred by the alkaloid nature of the discharge which may interfere with their chemoreceptors.
NERVOUS SYSTEM AND SENSE ORGANS
Cephalopods have the most highly developed nervous systems among invertebrates. Squids have a complex brain in the form of a nerve ring encircling the oesophagus, enclosed in a cartilaginous cranium. Paired cerebral ganglia above the oesophagus receive sensory information from the eyes and statocysts, and further ganglia below control the muscles of the mouth, foot, mantle and viscera. Giant axons up to 1 mm in diameter convey nerve messages with great rapidity to the circular muscles of the mantle wall, allowing a synchronous, powerful contraction and maximum speed in the jet propulsion system.
The paired eyes, on either side of the head, are housed in capsules fused to the cranium. Their structure is very similar to that of a fish eye, with a globular lens that has a depth of focus from 3 cm to infinity. The image is focused by changing the position of the lens, as in a camera or telescope, rather than changing the shape of the lens, as in the human eye. Squid adjust to changes in light intensity by expanding and contracting the slit-shaped pupil. Deep sea squids in the family Histioteuthidae have eyes of two different types and orientation. The large left eye is tubular in shape and looks upwards, presumably searching for the silhouettes of animals higher in the water column. The normally-shaped right eye points forwards and downwards to detect prey.
The statocysts are involved in maintaining balance and are analogous to the inner ear of fish. They are housed in cartilaginous capsules on either side of the cranium. They provide the squid with information on its body position in relation to gravity, its orientation, acceleration and rotation, and are able to perceive incoming vibrations. Without the statocysts, the squid cannot maintain equilibrium. Squid appear to have limited hearing, but the head and arms bear lines of hair-cells that are weakly sensitive to water movements and changes in pressure, and are analogous in function to the lateral line system of fish.
REPRODUCTIVE SYSTEM
The sexes are separate in squid, there being a single gonad in the posterior part of the body with fertilisation being external, and usually taking place in the mantle cavity of the female. The male has a testis from which sperm pass into a single gonoduct where they are rolled together into a long bundle, or spermatophore. The gonoduct is elongated into a "penis" that extends into the mantle cavity and through which spermatophores are ejected. In shallow water species, the penis is short, and the spermatophore is removed from the mantle cavity by a tentacle of the male, which is specially adapted for the purpose and known as a hectocotylus, and placed inside the mantle cavity of the female during mating.The female has a large translucent ovary, situated towards the posterior of the visceral mass. From here, eggs travel along the gonocoel, where there are a pair of white nidamental glands, which lie anterior to the gills. Also present are red-spotted accessory nidamental glands containing symbiotic bacteria; both organs are associated with nutrient manufacture and forming shells for the eggs. The gonocoel enters the mantle cavity at the gonopore, and in some species, receptacles for storing spermatophores are located nearby, in the mantle wall. In shallow-water species of the continental shelf and epipelagic or mesopelagic zones, it is frequently one or both of arm pair IV of males that are modified into hectocotyli. However, most deep-sea squid lack hectocotyl arms and have longer penises; Ancistrocheiridae and Cranchiinae are exceptions. Giant squid of the genus Architeuthis are unusual in that they possess both a large penis and modified arm tips, although whether the latter are used for spermatophore transfer is uncertain. Penis elongation has been observed in the deep-water species Onykia ingens; when erect, the penis may be as long as the mantle, head, and arms combined. As such, deep-water squid have the greatest known penis length relative to body size of all mobile animals, second in the entire animal kingdom only to certain sessile barnacles.
DIGESTIVE SYSTEM
Like all cephalopods, squids are predators and have complex digestive systems. The mouth is equipped with a sharp, horny beak mainly made of chitin and cross-linked proteins, which is used to kill and tear prey into manageable pieces. The beak is very robust, but does not contain minerals, unlike the teeth and jaws of many other organisms; the cross-linked proteins are histidine- and glycine-rich and give the beak a stiffness and hardness greater than most equivalent synthetic organic materials. The stomachs of captured whales often have indigestible squid beaks inside. The mouth contains the radula, the rough tongue common to all molluscs except bivalvia, which is equipped with multiple rows of teeth.[6] In some species, toxic saliva helps to control large prey; when subdued, the food can be torn in pieces by the beak, moved to the oesophagus by the radula, and swallowed.
The food bolus is moved along the gut by waves of muscular contractions (peristalsis). The long oesophagus leads to a muscular stomach roughly in the middle of the visceral mass. The digestive gland, which is equivalent to a vertebrate liver, diverticulates here, as does the pancreas, and both of these empty into the caecum, a pouch-shaped sac where most of the absorption of nutrients takes place. Indigestible food can be passed directly from the stomach to the rectum where it joins the flow from the caecum and is voided through the anus into the mantle cavity. Cephalopods are short-lived, and in mature squid, priority is given to reproduction; the female Onychoteuthis banksii for example, sheds its feeding tentacles on reaching maturity, and becomes flaccid and weak after spawning.
CARDIOVASCULAR AND EXCRETORY SYSTEMS
The squid mantle cavity is a seawater-filled sac containing three hearts and other organs supporting circulation, respiration, and excretion. Squid have a main systemic heart that pumps blood around the body as part of the general circulatory system, and two branchial hearts. The systemic heart consists of three chambers, a lower ventricle and two upper atria, all of which can contract to propel the blood. The branchial hearts pump blood specifically to the gills for oxygenation, before returning it to the systemic heart. The blood contains the copper-rich protein hemocyanin, which is used for oxygen transport at low ocean temperatures and low oxygen concentrations, and makes the oxygenated blood a deep, blue color. As systemic blood returns via two vena cavae to the branchial hearts, excretion of urine, carbon dioxide, and waste solutes occurs through outpockets (called nephridial appendages) in the vena cavae walls that enable gas exchange and excretion via the mantle cavity seawater.
BUOYANCY
Unlike nautiloids which have gas-filled chambers inside their shells which provide buoyancy, and octopuses which live near and rest on the seabed and do not require to be buoyant, many squid have a fluid-filled receptacle, equivalent to the swim bladder of a fish, in the coelom or connective tissue. This reservoir acts as a chemical buoyancy chamber, with the heavy metallic cations typical of seawater replaced by low molecular-weight ammonium ions, a product of excretion. The small difference in density provides a small contribution to buoyancy per unit volume, so the mechanism requires a large buoyancy chamber to be effective. Since the chamber is filled with liquid, it has the advantage over a swim bladder of not changing significantly in volume with pressure. Glass squids in the family Cranchiidae for example, have an enormous transparent coelom containing ammonium ions and occupying about two-thirds the volume of the animal, allowing it to float at the required depth. About half of the 28 families of squid use this mechanism to solve their buoyancy issues.
LARGEST AND SMALLEST
The majority of squid are no more than 60 cm long, although the giant squid may reach 13 m. The smallest species are probably the benthic pygmy squids Idiosepius, which grow to a mantle length of 10 to 18 mm, and have short bodies and stubby arms.
In 1978, sharp, curved claws on the suction cups of squid tentacles cut up the rubber coating on the hull of the USS Stein. The size suggested the largest squid known at the time.
In 2003, a large specimen of an abundant but poorly understood species, Mesonychoteuthis hamiltoni (the colossal squid), was discovered. This species may grow to 10 m in length, making it the largest invertebrate. In February 2007, a New Zealand fishing vessel caught the largest squid ever documented, weighing 495 kg and measuring around 10 m off the coast of Antarctica. Dissection showed that the eyes, used to detect prey in the deep Southern Ocean, exceeded the size of footballs; these may be among the largest eyes ever to exist in the animal kingdom.
DEVELOPMENT
The eggs of squid are large for a mollusc, containing a large amount of yolk to nourish the embryo as it develops directly, without an intervening veliger larval stage. The embryo grows as a disc of cells on top of the yolk. During the gastrulation stage, the margins of the disc grow to surround the yolk, forming a yolk sac, which eventually forms part of the animal's gut. The dorsal side of the disc grows upwards and forms the embryo, with a shell gland on its dorsal surface, gills, mantle and eyes. The arms and funnel develop as part of the foot on the ventral side of the disc. The arms later migrate upwards, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo grows. Some juvenile squid live higher in the water column than do adults. Squids tend to be short-lived; Loligo for example lives from one to three years according to species, typically dying soon after spawning.
n a well-studied bioluminescent species, the Hawaiian bobtail squid, a special light organ in the squid's mantle is rapidly colonized with Aliivibrio fischeri bacteria within hours of hatching. This light-organ colonization requires this particular bacterial species for a symbiotic relationship; no colonization occurs in the absence of A. fischeri. Colonization occurs in a horizontal manner, such that the hosts acquires its bacterial partners from the environment. The symbiosis is obligate for the squid, but facultative for the bacteria. Once the bacteria enter the squid, they colonize interior epithelial cells in the light organ, living in crypts with complex microvilli protrusions. The bacteria also interact with hemocytes, macrophage-like blood cells that migrate between epithelial cells, but the mechanism and function of this process is not well understood. Bioluminescence reaches its highest levels during the early evening hours and bottoms out before dawn; this occurs because at the end of each day, the contents of the squid's crypts are expelled into the surrounding environment. About 95% of the bacteria are voided each morning before the bacterial population builds up again by nightfall.
BEHAVIOUR
LOCOMOTION
Squid can move about in several different ways. Slow movement is achieved by a gentle undulation of the muscular lateral fins on either side of the trunk which drives the animal forward. A more common means of locomotion providing sustained movement is achieved using jetting, during which contraction of the muscular wall of the mantle cavity provides jet propulsion.
Slow jetting is used for ordinary locomotion, and ventilation of the gills is achieved at the same time. The circular muscles in the mantle wall contract; this causes the inhalant valve to close, the exhalant valve to open and the mantle edge to lock tightly around the head. Water is forced out through the funnel which is pointed in the opposite direction to the required direction of travel. The inhalant phase is initiated by the relaxation of the circular muscles causes them to stretch, the connective tissue in the mantle wall recoils elastically, the mantle cavity expands causing the inhalant valve to open, the exhalant valve to close and water to flow into the cavity. This cycle of exhalation and inhalation is repeated to provide continuous locomotion.
Fast jetting is an escape response. In this form of locomotion, radial muscles in the mantle wall are involved as well as circular ones, making it possible to hyper-inflate the mantle cavity with a larger volume of water than during slow jetting. On contraction, water flows out with great force, the funnel always being pointed anteriorly, and travel is backwards. During this means of locomotion, some squid exit the water in a similar way to flying fish, gliding through the air for up to 50 m, and occasionally ending up on the decks of ships.
FEEDING
Squid are carnivores, and, with their strong arms and suckers, can overwhelm relatively large animals efficiently. Prey is identified by sight or by touch, grabbed by the tentacles which can be shot out with great rapidity, brought back to within reach of the arms, and held by the hooks and suckers on their surface. In some species, the squid's saliva contains toxins which act to subdue the prey. These are injected into its bloodstream when the prey is bitten, along with vasodilators and chemicals to stimulate the heart, and quickly circulate to all parts of its body. The deep sea squid Taningia danae has been filmed releasing blinding flashes of light from large photophores on its arms to illuminate and disorientate potential prey.
Although squid can catch large prey, the mouth is relatively small, and the food must be cut into pieces by the chitinous beak with its powerful muscles before being swallowed. The radula is located in the buccal cavity and has multiple rows of tiny teeth that draw the food backwards and grind it in pieces. The deep sea squid Mastigoteuthis has the whole length of its whip-like tentacles covered with tiny suckers; it probably catches small organisms in the same way that flypaper traps flies. The tentacles of some bathypelagic squids bear photophores which may bring food within its reach by attracting prey.
Squid are among the most intelligent invertebrates. For example, groups of Humboldt squid hunt cooperatively, spiralling up through the water at night and coordinating their vertical and horizontal movements while foraging.
REPRODUCTION
Courtship in squid takes place in the open water and involves the male selecting a female, the female responding, and the transfer by the male of spermatophores to the female. In many instances, the male may display to identify himself to the female and drive off any potential competitors.[46] Elaborate changes in body patterning take place in some species in both agonistic and courtship behaviour. The Caribbean reef squid (Sepioteuthis sepioidea), for example, employs a complex array of colour changes during courtship and social interactions and has a range of about 16 body patterns in its repertoire.
The pair adopt a head-to-head position, and "jaw locking" may take place, in a similar manner to that adopted by some cichlid fish. The heterodactylus of the male is used to transfer the spermatophore and deposit it in the female's mantle cavity in the position appropriate for the species; this may be adjacent to the gonopore or in a seminal receptacle.
The sperm may be used immediately or may be stored. As the eggs pass down the oviduct, they are wrapped in a gelatinous coating, before continuing to the mantle cavity, where they are fertilised. In Loligo, further coatings are added by the nidimental glands in the walls of the cavity and the eggs leave through a funnel formed by the arms. The female attaches them to the substrate in strings or groups, the coating layers swelling and hardening after contact with sea water. Loligo sometimes forms breeding aggregations which may create a "community pile" of egg strings. Some pelagic and deep sea squid do not attach their egg masses, which float freely.
ECOLOGY
Squid mostly have an annual life cycle, growing fast and dying soon after spawning. The diet changes as they grow but mostly consists of large zooplankton and small nekton. In Antarctica for example, krill is the main constituent of the diet, with other food items being amphipods, other small crustaceans, and large arrow worms. Fish are also eaten, and some squid are cannibalistic.
As well as occupying a key role in the food chain, squid are an important prey for predators including sharks, sea birds, seals and whales. Juvenile squid provide part of the diet for worms and small fish. When researchers studied the contents of the stomachs of elephant seals in South Georgia, they found 96% squid by weight. In a single day, a sperm whale can eat 700 to 800 squid, and a Risso's dolphin entangled in a net in the Mediterranean was found to have eaten angel clubhook squid, umbrella squid, reverse jewel squid and European flying squid, all identifiable from their indigestible beaks. Ornithoteuthis volatilis, a common squid from the tropical Indo-Pacific, is predated by yellowfin tuna, longnose lancetfish, common dolphinfish and swordfish, the tiger shark, the scalloped hammerhead shark and the smooth hammerhead shark. Sperm whales also hunt this species extensively as does the brown fur seal. In the Southern Ocean, penguins and wandering albatrosses are major predators of Gonatus antarcticus.
HUMAN USES
IN LITERATUR AND ART
Giant squid have featured as monsters of the deep since classical times. Giant squid were described by Aristotle (4th century BC) in his History of Animals and Pliny the Elder (1st century AD) in his Natural History. The Gorgon of Greek mythology may have been inspired by squid or octopus, the animal itself representing the severed head of Medusa, the beak as the protruding tongue and fangs, and its tentacles as the snakes. The six-headed sea monster of the Odyssey, Scylla, may have had a similar origin. The Nordic legend of the kraken may also have derived from sightings of large cephalopods.
In literature, H. G. Wells' short story "The Sea Raiders" featured a man-eating squid species Haploteuthis ferox.[59] The science fiction writer Jules Verne told a tale of a kraken-like monster in his 1870 novel Twenty Thousand Leagues Under the Sea.
AS FOOD
Squid form a major food resource and are used in cuisines around the world, notably in Japan where it is eaten as ika sōmen, sliced into vermicelli-like strips; as sashimi; and as tempura. Three species of Loligo are used in large quantities, L. vulgaris in the Mediterranean (known as Calamar in Spanish, Calamaro in Italian); L. forbesii in the Northeast Atlantic; and L. pealei on the American East Coast. Among the Ommastrephidae, Todarodes pacificus is the main commercial species, harvested in large quantities across the North Pacific in Canada, Japan and China.
In English-speaking countries, squid as food is often called calamari, adopted from Italian into English in the 17th century. Squid are found abundantly in certain areas, and provide large catches for fisheries. The body can be stuffed whole, cut into flat pieces, or sliced into rings. The arms, tentacles, and ink are also edible; the only parts not eaten are the beak and gladius (pen). Squid is a good food source for zinc and manganese, and high in copper, selenium, vitamin B12, and riboflavin.
COMMERCIAL FISHING
According to the FAO, the cephalopod catch for 2002 was 3,173,272 tonnes. Of this, 2,189,206 tonnes, or 75.8 percent, was squid. The following table lists squid species fishery catches that exceeded 10,000 tonnes in 2002.
IN BIOMIMICRY
Prototype chromatophores that mimic the squid's adaptive camouflage, have been made by Bristol University researchers using an electroactive dielectric elastomer, a flexible "smart" material that changes its colour and texture in response to electrical signals. The researchers state that their goal is to create an artificial skin that provides rapid active camouflage.
The squid giant axon inspired Otto Schmitt to develop a comparator circuit with hysteresis now called the Schmitt trigger, replicating the axon's propagation of nerve impulses.
WIKIPEDIA