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This young dark-faced 8-point timidly came into a rattle. Finally came out of the woods and checked out the nearby scrape. Didn't work it.
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Our beautiful world, pass it on.
I shot this back November in a black Kodak developing tank in the studio. The paperclip is floating on the water's surface whose tension permits the paperclip to rest slightly lower than the water's normal surface. This depression is made apparent due to the lines of perpendicular venetian blinds reflecting in the water's surface.
Strobist info: 500W hotlight snooted with blue gel on blinds and 250W hotlight snooted on paperclip from high camera left.
More Macro with my New Old Knight ... a Nikkor 28mm F/3.5
Taken with a Nikon 1 V2 , inverse Nikon Non AI 28mm, F/22, ISO 800, Shutter 1/60
What are You Think That is ?, Comment ...
Wells Cathedral is an Anglican cathedral in Wells, Somerset, England, dedicated to St Andrew the Apostle. It is the seat of the Bishop of Bath and Wells, whose cathedra it holds as mother church of the Diocese of Bath and Wells. Built as a Roman Catholic cathedral from around 1175 to replace an earlier church on the site since 705, it became an Anglican cathedral when King Henry VIII split from Rome. It is moderately sized for an English cathedral. Its broad west front and large central tower are dominant features. It has been called "unquestionably one of the most beautiful" and "most poetic" of English cathedrals.
Its Gothic architecture is mostly inspired from Early English style of the late 12th to early 13th centuries, lacking the Romanesque work that survives in many other cathedrals. Building began about 1175 at the east end with the choir. Historian John Harvey sees it as Europe's first truly Gothic structure, breaking the last constraints of Romanesque. The stonework of its pointed arcades and fluted piers bears pronounced mouldings and carved capitals in a foliate, "stiff-leaf" style. Its Early English front with 300 sculpted figures is seen as a "supreme triumph of the combined plastic arts in England". The east end retains much ancient stained glass. Unlike many cathedrals of monastic foundation, Wells has many surviving secular buildings linked to its chapter of secular canons, including the Bishop's Palace and the 15th-century residential Vicars' Close It is a Grade I listed building.
The earliest remains of a building on the site are of a late-Roman mausoleum, identified during excavations in 1980. An abbey church was built in Wells in 705 by Aldhelm, first bishop of the newly established Diocese of Sherborne during the reign of King Ine of Wessex. It was dedicated to St Andrew and stood at the site of the cathedral's cloisters, where some excavated remains can be seen. The font in the cathedral's south transept is from this church and is the oldest part of the present building. In 766 Cynewulf, King of Wessex, signed a charter endowing the church with eleven hides of land. In 909 the seat of the diocese was moved from Sherborne to Wells.
The first bishop of Wells was Athelm (909), who crowned King Æthelstan. Athelm and his nephew Dunstan both became Archbishops of Canterbury. During this period a choir of boys was established to sing the liturgy. Wells Cathedral School, which was established to educate these choirboys, dates its foundation to this point. There is, however, some controversy over this. Following the Norman Conquest, John de Villula moved the seat of the bishop from Wells to Bath in 1090. The church at Wells, no longer a cathedral, had a college of secular clergy.
The cathedral is thought to have been conceived and commenced in about 1175 by Reginald Fitz Jocelin, who died in 1191. Although it is clear from its size that from the outset, the church was planned to be the cathedral of the diocese, the seat of the bishop moved between Wells and the abbeys of Glastonbury and Bath, before settling at Wells. In 1197 Reginald's successor, Savaric FitzGeldewin, with the approval of Pope Celestine III, officially moved his seat to Glastonbury Abbey. The title of Bishop of Bath and Glastonbury was used until the Glastonbury claim was abandoned in 1219.
Savaric's successor, Jocelin of Wells, again moved the bishop's seat to Bath Abbey, with the title Bishop of Bath. Jocelin was a brother of Hugh (II) of Lincoln and was present at the signing of the Magna Carta. Jocelin continued the building campaign begun by Reginald and was responsible for the Bishop's Palace, the choristers' school, a grammar school, a hospital for travellers and a chapel. He also had a manor house built at Wookey, near Wells. Jocelin saw the church dedicated in 1239 but, despite much lobbying of the Pope by Jocelin's representatives in Rome, did not live to see cathedral status granted. The delay may have been a result of inaction by Pandulf Verraccio, a Roman ecclesiastical politician, papal legate to England and Bishop of Norwich, who was asked by the Pope to investigate the situation but did not respond. Jocelin died at Wells on 19 November 1242 and was buried in the choir of the cathedral; the memorial brass on his tomb is one of the earliest brasses in England. Following his death the monks of Bath unsuccessfully attempted to regain authority over Wells.
In 1245 the ongoing dispute over the title of the bishop was resolved by a ruling of Pope Innocent IV, who established the title as the "Bishop of Bath and Wells", which it has remained until this day, with Wells as the principal seat of the bishop. Since the 11th century the church has had a chapter of secular clergy, like the cathedrals of Chichester, Hereford, Lincoln and York. The chapter was endowed with 22 prebends (lands from which finance was drawn) and a provost to manage them. On acquiring cathedral status, in common with other such cathedrals, it had four chief clergy, the dean, precentor, chancellor and sacristan, who were responsible for the spiritual and material care of the cathedral.
The building programme, begun by Reginald Fitz Jocelin, Bishop in the 12th century, continued under Jocelin of Wells, who was a canon from 1200, then bishop from 1206. Adam Locke was master mason from about 1192 until 1230. It was designed in the new style with pointed arches, later known as Gothic, which was introduced at about the same time at Canterbury Cathedral. Work was halted between 1209 and 1213 when King John was excommunicated and Jocelin was in exile, but the main parts of the church were complete by the time of the dedication by Jocelin in 1239.
By the time the cathedral, including the chapter house, was finished in 1306, it was already too small for the developing liturgy, and unable to accommodate increasingly grand processions of clergy. John Droxford initiated another phase of building under master mason Thomas of Whitney, during which the central tower was heightened and an eight-sided Lady chapel was added at the east end by 1326. Ralph of Shrewsbury followed, continuing the eastward extension of the choir and retrochoir beyond. He oversaw the building of Vicars' Close and the Vicars' Hall, to give the men who were employed to sing in the choir a secure place to live and dine, away from the town and its temptations. He had an uneasy relationship with the citizens of Wells, partly because of his imposition of taxes, and he surrounded his palace with crenellated walls, a moat and a drawbridge.
John Harewell raised money for the completion of the west front by William Wynford, who was appointed as master mason in 1365. One of the foremost master masons of his time, Wynford worked for the king at Windsor, Winchester Cathedral and New College, Oxford. At Wells, he designed the western towers of which north-west was not built until the following century. In the 14th century, the central piers of the crossing were found to be sinking under the weight of the crossing tower which had been damaged by an earthquake in the previous century. Strainer arches, sometimes described as scissor arches, were inserted by master mason William Joy to brace and stabilise the piers as a unit.
By the reign of Henry VII the cathedral was complete, appearing much as it does today (though the fittings have changed). From 1508 to 1546, the eminent Italian humanist scholar Polydore Vergil was active as the chapter's representative in London. He donated a set of hangings for the choir of the cathedral. While Wells survived the Dissolution of the Monasteries better than the cathedrals of monastic foundation, the abolition of chantries in 1547 resulted in a reduction in its income. Medieval brasses were sold, and a pulpit was placed in the nave for the first time. Between 1551 and 1568, in two periods as dean, William Turner established a herb garden, which was recreated between 2003 and 2010.
Elizabeth I gave the chapter and the Vicars Choral a new charter in 1591, creating a new governing body, consisting of a dean and eight residentiary canons with control over the church estates and authority over its affairs, but no longer entitled to elect the dean (that entitlement thenceforward belonged ultimately to the Crown). The stability brought by the new charter ended with the onset of the Civil War and the execution of Charles I. Local fighting damaged the cathedral's stonework, furniture and windows. The dean, Walter Raleigh, a nephew of the explorer Walter Raleigh, was placed under house arrest after the fall of Bridgwater to the Parliamentarians in 1645, first in the rectory at Chedzoy and then in the deanery at Wells. His jailor, the shoe maker and city constable, David Barrett, caught him writing a letter to his wife. When he refused to surrender it, Barrett ran him through with a sword and he died six weeks later, on 10 October 1646. He was buried in an unmarked grave in the choir before the dean's stall. During the Commonwealth of England under Oliver Cromwell no dean was appointed and the cathedral fell into disrepair. The bishop went into retirement and some of the clerics were reduced to performing menial tasks.
In 1661, after Charles II was restored to the throne, Robert Creighton, the king's chaplain in exile, was appointed dean and was bishop for two years before his death in 1672. His brass lectern, given in thanksgiving, can be seen in the cathedral. He donated the nave's great west window at a cost of £140. Following Creighton's appointment as bishop, the post of dean went to Ralph Bathurst, who had been chaplain to the king, president of Trinity College, Oxford and fellow of the Royal Society. During Bathurst's long tenure the cathedral was restored, but in the Monmouth Rebellion of 1685, Puritan soldiers damaged the west front, tore lead from the roof to make bullets, broke the windows, smashed the organ and furnishings, and for a time stabled their horses in the nave.
Restoration began again under Thomas Ken who was appointed by the Crown in 1685 and served until 1691. He was one of seven bishops imprisoned for refusing to sign King James II's "Declaration of Indulgence", which would have enabled Catholics to resume positions of political power, but popular support led to their acquittal. Ken refused to take the oath of allegiance to William III and Mary II because James II had not abdicated and with others, known as the Nonjurors, was put out of office. His successor, Richard Kidder, was killed in the Great Storm of 1703 when two chimney stacks on the palace fell on him and his wife, while they were asleep in bed.
By the middle of the 19th century, a major restoration programme was needed. Under Dean Goodenough, the monuments were moved to the cloisters and the remaining medieval paint and whitewash removed in an operation known as "the great scrape". Anthony Salvin took charge of the extensive restoration of the choir. Wooden galleries installed in the 16th century were removed and the stalls were given stone canopies and placed further back within the line of the arcade. The medieval stone pulpitum screen was extended in the centre to support a new organ.
In 1933 the Friends of Wells Cathedral were formed to support the cathedral's chapter in the maintenance of the fabric, life and work of the cathedral. The late 20th century saw an extensive restoration programme, particularly of the west front. The stained glass is currently under restoration, with a programme underway to conserve the large 14th-century Jesse Tree window at the eastern terminal of the choir.
In January 2014, as part of the Bath film festival, the cathedral hosted a special screening of Martin Scorsese's The Last Temptation of Christ. This provoked some controversy, but the church defended its decision to allow the screening.
In 2021, a contemporary sculpture by Anthony Gormley was unveiled on a temporary plinth outside the cathedral.
Since the 13th century, Wells Cathedral has been the seat of the Bishop of Bath and Wells. Its governing body, the chapter, is made up of five clerical canons (the dean, the precentor, the canon chancellor, the canon treasurer, and the archdeacon of Wells) and four lay members: the administrator (chief executive), Keeper of the Fabric, Overseer of the Estate and the chairman of the cathedral shop and catering boards. The current bishop of Bath and Wells is Peter Hancock, who was installed in a service in the cathedral on 7 June 2014. John Davies has been Dean of Wells since 2016.
Employed staff include the organist and master of choristers, head Verger archivist, librarian and the staff of the shop, café and restaurant. The chapter is advised by specialists such as architects, archaeologists and financial analysts.
More than a thousand services are held every year. There are daily services of Matins, Holy Communion and Choral Evensong, as well as major celebrations of Christian festivals such as Christmas, Easter, Pentecost and saints' days. The cathedral is also used for the baptisms, weddings and funerals of those with close connections to it. In July 2009 the cathedral undertook the funeral of Harry Patch, the last British Army veteran of World War I, who died at the age of 111.
Three Sunday services are led by the resident choir in school terms and choral services are sung on weekdays. The cathedral hosts visiting choirs and does outreach work with local schools as part of its Chorister Outreach Project. It is also a venue for musical events such as an annual concert by the Somerset Chamber Choir.
Each year about 150,000 people attend services and another 300,000 visit as tourists. Entry is free, but visitors are encouraged to make a donation towards the annual running costs of around £1.5 million in 2015.
Construction of the cathedral began in about 1175, to the design of an unknown master-mason. Wells is the first cathedral in England to be built, from its foundation, in Gothic style. According to art historian John Harvey, it is the first truly Gothic cathedral in the world, its architects having entirely dispensed with all features that bound the contemporary east end of Canterbury Cathedral and the earlier buildings of France, such as the east end of the Abbey of Saint Denis, to the Romanesque. Unlike these churches, Wells has clustered piers rather than columns and has a gallery of identical pointed arches rather than the typically Romanesque form of paired openings. The style, with its simple lancet arches without tracery and convoluted mouldings, is known as Early English Gothic.
From about 1192 to 1230, Adam Lock, the earliest master-mason at Wells for whom a name is known, continued the transept and nave in the same manner as his predecessor. Lock was also the builder of the north porch, to his own design.
The Early English west front was commenced around 1230 by Thomas Norreys, with building and sculpture continuing for thirty years. Its south-west tower was begun 100 years later and constructed between 1365 and 1395, and the north-west tower between 1425 and 1435, both in the Perpendicular Gothic style to the design of William Wynford, who also filled many of the cathedral's early English lancet windows with delicate tracery.
The undercroft and chapter house were built by unknown architects between 1275 and 1310, the undercroft in the Early English and the chapter house in the Geometric style of Decorated Gothic architecture. In about 1310 work commenced on the Lady Chapel, to the design of Thomas Witney, who also built the central tower from 1315 to 1322 in the Decorated Gothic style. The tower was later braced internally with arches by William Joy. Concurrent with this work, in 1329–45 Joy made alterations and extensions to the choir, joining it to the Lady Chapel with the retrochoir, the latter in the Flowing Decorated style.
Later changes include the Perpendicular vault of the tower and construction of Sugar's Chapel, 1475–1490 by William Smyth. Also, Gothic Revival renovations were made to the choir and pulpitum by Benjamin Ferrey and Anthony Salvin, 1842–1857.
Wells has a total length of 415 feet (126 m). Like Canterbury, Lincoln and Salisbury cathedrals, it has the distinctly English arrangement of two transepts, with the body of the church divided into distinct parts: nave, choir, and retro-choir, beyond which extends the Lady Chapel. The façade is wide, with its towers extending beyond the transepts on either side. There is a large projecting porch on the north side of the nave forming an entry into the cathedral. To the north-east is the large octagonal chapter house, entered from the north choir aisle by a passage and staircase. To the south of the nave is a large cloister, unusual in that the northern range, that adjacent the cathedral, was never built.
In section, the cathedral has the usual arrangement of a large church: a central nave with an aisle on each side, separated by two arcades. The elevation is in three stages, arcade, triforium gallery and clerestory. The nave is 67 feet (20 m) in height, very low compared to the Gothic cathedrals of France. It has a markedly horizontal emphasis, caused by the triforium having a unique form, a series of identical narrow openings, lacking the usual definition of the bays. The triforium is separated from the arcade by a single horizontal string course that runs unbroken the length of the nave. There are no vertical lines linking the three stages, as the shafts supporting the vault rise above the triforium.
The exterior of Wells Cathedral presents a relatively tidy and harmonious appearance since the greater part of the building was executed in a single style, Early English Gothic. This is uncommon among English cathedrals where the exterior usually exhibits a plethora of styles. At Wells, later changes in the Perpendicular style were universally applied, such as filling the Early English lancet windows with simple tracery, the construction of a parapet that encircles the roof, and the addition of pinnacles framing each gable, similar to those around the chapter house and on the west front. At the eastern end there is a proliferation of tracery with repeated motifs in the Reticulated style, a stage between Geometric and Flowing Decorated tracery.
The west front is 100 feet (30 m) high and 147 feet (45 m) wide, and built of Inferior Oolite of the Middle Jurassic period, which came from the Doulting Stone Quarry, about 8 miles (13 km) to the east. According to the architectural historian Alec Clifton-Taylor, it is "one of the great sights of England".
West fronts in general take three distinct forms: those that follow the elevation of the nave and aisles, those that have paired towers at the end of each aisle, framing the nave, and those that screen the form of the building. The west front at Wells has the paired-tower form, unusual in that the towers do not indicate the location of the aisles, but extend well beyond them, screening the dimensions and profile of the building.
The west front rises in three distinct stages, each clearly defined by a horizontal course. This horizontal emphasis is counteracted by six strongly projecting buttresses defining the cross-sectional divisions of nave, aisles and towers, and are highly decorated, each having canopied niches containing the largest statues on the façade.
At the lowest level of the façade is a plain base, contrasting with and stabilising the ornate arcades that rise above it. The base is penetrated by three doors, which are in stark contrast to the often imposing portals of French Gothic cathedrals. The outer two are of domestic proportion and the central door is ornamented only by a central post, quatrefoil and the fine mouldings of the arch.
Above the basement rise two storeys, ornamented with quatrefoils and niches originally holding about four hundred statues, with three hundred surviving until the mid-20th century. Since then, some have been restored or replaced, including the ruined figure of Christ in the gable.
The third stages of the flanking towers were both built in the Perpendicular style of the late 14th century, to the design of William Wynford; that on the north-west was not begun until about 1425. The design maintains the general proportions, and continues the strong projection of the buttresses.
The finished product has been criticised for its lack of pinnacles, and it is probable that the towers were intended to carry spires which were never built. Despite its lack of spires or pinnacles, the architectural historian Banister Fletcher describes it as "the highest development in English Gothic of this type of façade."
The sculptures on the west front at Wells include standing figures, seated figures, half-length angels and narratives in high relief. Many of the figures are life-sized or larger. Together they constitute the finest display of medieval carving in England. The figures and many of the architectural details were painted in bright colours, and the colouring scheme has been deduced from flakes of paint still adhering to some surfaces. The sculptures occupy nine architectural zones stretching horizontally across the entire west front and around the sides and the eastern returns of the towers which extend beyond the aisles. The strongly projecting buttresses have tiers of niches which contain many of the largest figures. Other large figures, including that of Christ, occupy the gable. A single figure stands in one of two later niches high on the northern tower.
In 1851 the archaeologist Charles Robert Cockerell published his analysis of the iconography, numbering the nine sculptural divisions from the lowest to the highest. He defined the theme as "a calendar for unlearned men" illustrating the doctrines and history of the Christian faith, its introduction to Britain and its protection by princes and bishops. He likens the arrangement and iconography to the Te Deum.
According to Cockerell, the side of the façade that is to the south of the central door is the more sacred and the scheme is divided accordingly. The lowest range of niches each contained a standing figure, of which all but four figures on the west front, two on each side, have been destroyed. More have survived on the northern and eastern sides of the north tower. Cockerell speculates that those to the south of the portal represented prophets and patriarchs of the Old Testament while those to the north represented early missionaries to Britain, of which Augustine of Canterbury, St Birinus, and Benedict Biscop are identifiable by their attributes. In the second zone, above each pair of standing figures, is a quatrefoil containing a half-length angel in relief, some of which have survived. Between the gables of the niches are quatrefoils that contain a series of narratives from the Bible, with the Old Testament stories to the south, above the prophets and patriarchs, and those from the New Testament to the north. A horizontal course runs around the west front dividing the architectural storeys at this point.
Above the course, zones four and five, as identified by Cockerell, contain figures which represent the Christian Church in Britain, with the spiritual lords such as bishops, abbots, abbesses and saintly founders of monasteries on the south, while kings, queens and princes occupy the north. Many of the figures survive and many have been identified in the light of their various attributes. There is a hierarchy of size, with the more significant figures larger and enthroned in their niches rather than standing. Immediately beneath the upper course are a series of small niches containing dynamic sculptures of the dead coming forth from their tombs on the Day of Judgement. Although naked, some of the dead are defined as royalty by their crowns and others as bishops by their mitres. Some emerge from their graves with joy and hope, and others with despair.
The niches in the lowest zone of the gable contain nine angels, of which Cockerell identifies Michael, Gabriel, Raphael and Uriel. In the next zone are the taller figures of the twelve apostles, some, such as John, Andrew and Bartholomew, clearly identifiable by the attributes that they carry. The uppermost niches of the gable contained the figure of Christ the Judge at the centre, with the Virgin Mary on his right and John the Baptist on his left. The figures all suffered from iconoclasm. A new statue of Jesus was carved for the central niche, but the two side niches now contain cherubim. Christ and the Virgin Mary are also represented by now headless figures in a Coronation of the Virgin in a niche above the central portal. A damaged figure of the Virgin and Christ Child occupies a quatrefoil in the spandrel of the door.
The central tower appears to date from the early 13th century. It was substantially reconstructed in the early 14th century during the remodelling of the east end, necessitating the internal bracing of the piers a decade or so later. In the 14th century the tower was given a timber and lead spire which burnt down in 1439. The exterior was then reworked in the Perpendicular style and given the present parapet and pinnacles. Alec Clifton-Taylor describes it as "outstanding even in Somerset, a county famed for the splendour of its church towers".
The north porch is described by art historian Nikolaus Pevsner as "sumptuously decorated", and intended as the main entrance. Externally it is simple and rectangular with plain side walls. The entrance is a steeply arched portal framed by rich mouldings of eight shafts with stiff-leaf capitals each encircled by an annular moulding at middle height. Those on the left are figurative, containing images representing the martyrdom of St Edmund the Martyr. The walls are lined with deep niches framed by narrow shafts with capitals and annulets like those of the portal. The path to the north porch is lined by four sculptures in Purbeck stone, each by Mary Spencer Watson, representing the symbols of the Evangelists.
The cloisters were built in the late 13th century and largely rebuilt from 1430 to 1508 and have wide openings divided by mullions and transoms, and tracery in the Perpendicular Gothic style. The vault has lierne ribs that form octagons at the centre of each compartment, the joints of each rib having decorative bosses. The eastern range is of two storeys, of which the upper is the library built in the 15th century.
Because Wells Cathedral was secular rather than monastic, cloisters were not a practical necessity. They were omitted from several other secular cathedrals but were built here and at Chichester. Explanations for their construction at these two secular cathedrals range from the processional to the aesthetic. As at Chichester, there is no northern range to the cloisters. In monastic cloisters it was the north range, benefiting most from winter sunlight, that was often used as a scriptorium.
In 1969, when a large chunk of stone fell from a statue near the main door, it became apparent that there was an urgent need for restoration of the west front. Detailed studies of the stonework and of conservation practices were undertaken under the cathedral architect, Alban D. R. Caroe and a restoration committee formed. The methods selected were those devised by Eve and Robert Baker. W. A. (Bert) Wheeler, clerk of works to the cathedral 1935–1978, had previously experimented with washing and surface treatment of architectural carvings on the building and his techniques were among those tried on the statues.
The conservation was carried out between 1974 and 1986, wherever possible using non-invasive procedures such as washing with water and a solution of lime, filling gaps and damaged surfaces with soft mortar to prevent the ingress of water and stabilising statues that were fracturing through corrosion of metal dowels. The surfaces were finished by painting with a thin coat of mortar and silane to resist further erosion and attack by pollutants. The restoration of the façade revealed much paint adhering to the statues and their niches, indicating that it had once been brightly coloured.
The particular character of this Early English interior is dependent on the proportions of the simple lancet arches. It is also dependent on the refinement of the architectural details, in particular the mouldings.
The arcade, which takes the same form in the nave, choir and transepts, is distinguished by the richness of both mouldings and carvings. Each pier of the arcade has a surface enrichment of 24 slender shafts in eight groups of three, rising beyond the capitals to form the deeply undulating mouldings of the arches. The capitals themselves are remarkable for the vitality of the stylised foliage, in a style known as "stiff-leaf". The liveliness contrasts with the formality of the moulded shafts and the smooth unbroken areas of ashlar masonry in the spandrels. Each capital is different, and some contain small figures illustrating narratives.
The vault of the nave rises steeply in a simple quadripartite form, in harmony with the nave arcade. The eastern end of the choir was extended and the whole upper part elaborated in the second quarter of the 14th century by William Joy. The vault has a multiplicity of ribs in a net-like form, which is very different from that of the nave, and is perhaps a recreation in stone of a local type of compartmented wooden roof of which examples remain from the 15th century, including those at St Cuthbert's Church, Wells. The vaults of the aisles of the choir also have a unique pattern.
Until the early 14th century, the interior of the cathedral was in a unified style, but it was to undergo two significant changes, to the tower and to the eastern end. Between 1315 and 1322 the central tower was heightened and topped by a spire, which caused the piers that supported it to show signs of stress. In 1338 the mason William Joy employed an unorthodox solution by inserting low arches topped by inverted arches of similar dimensions, forming scissors-like structures. These arches brace the piers of the crossing on three sides, while the easternmost side is braced by a choir screen. The bracing arches are known as "St Andrew's Cross arches", in a reference to the patron saint of the cathedral. They have been described by Wim Swaan – rightly or wrongly – as "brutally massive" and intrusive in an otherwise restrained interior.
Wells Cathedral has a square east end to the choir, as is usual, and like several other cathedrals including Salisbury and Lichfield, has a lower Lady Chapel projecting at the eastern end, begun by Thomas Witney in about 1310, possibly before the chapter house was completed. The Lady Chapel seems to have begun as a free-standing structure in the form of an elongated octagon, but the plan changed and it was linked to the eastern end by extension of the choir and construction of a second transept or retrochoir east of the choir, probably by William Joy.
The Lady Chapel has a vault of complex and somewhat irregular pattern, as the chapel is not symmetrical about both axes. The main ribs are intersected by additional non-supporting, lierne ribs, which in this case form a star-shaped pattern at the apex of the vault. It is one of the earliest lierne vaults in England. There are five large windows, of which four are filled with fragments of medieval glass. The tracery of the windows is in the style known as Reticulated Gothic, having a pattern of a single repeated shape, in this case a trefoil, giving a "reticulate" or net-like appearance.
The retrochoir extends across the east end of the choir and into the east transepts. At its centre the vault is supported by a remarkable structure of angled piers. Two of these are placed as to complete the octagonal shape of the Lady Chapel, a solution described by Francis Bond as "an intuition of Genius". The piers have attached shafts of marble, and, with the vaults that they support, create a vista of great complexity from every angle. The windows of the retrochoir are in the Reticulated style like those of the Lady Chapel, but are fully Flowing Decorated in that the tracery mouldings form ogival curves.
The chapter house was begun in the late 13th century and built in two stages, completed about 1310. It is a two-storeyed structure with the main chamber raised on an undercroft. It is entered from a staircase which divides and turns, one branch leading through the upper storey of Chain Gate to Vicars' Close. The Decorated interior is described by Alec Clifton-Taylor as "architecturally the most beautiful in England". It is octagonal, with its ribbed vault supported on a central column. The column is surrounded by shafts of Purbeck Marble, rising to a single continuous rippling foliate capital of stylised oak leaves and acorns, quite different in character from the Early English stiff-leaf foliage. Above the moulding spring 32 ribs of strong profile, giving an effect generally likened to "a great palm tree". The windows are large with Geometric Decorated tracery that is beginning to show an elongation of form, and ogees in the lesser lights that are characteristic of Flowing Decorated tracery. The tracery lights still contain ancient glass. Beneath the windows are 51 stalls, the canopies of which are enlivened by carvings including many heads carved in a light-hearted manner.
Wells Cathedral contains one of the most substantial collections of medieval stained glass in England, despite damage by Parliamentary troops in 1642 and 1643. The oldest surviving glass dates from the late 13th century and is in two windows on the west side of the chapter-house staircase. Two windows in the south choir aisle are from 1310 to 1320.
The Lady Chapel has five windows, of which four date from 1325 to 1330 and include images of a local saint, Dunstan. The east window was restored to a semblance of its original appearance by Thomas Willement in 1845. The other windows have complete canopies, but the pictorial sections are fragmented.
The east window of the choir is a broad, seven-light window dating from 1340 to 1345. It depicts the Tree of Jesse (the genealogy of Christ) and demonstrates the use of silver staining, a new technique that allowed the artist to paint details on the glass in yellow, as well as black. The combination of yellow and green glass and the application of the bright yellow stain gives the window its popular name, the "Golden Window". It is flanked by two windows each side in the clerestory, with large figures of saints, also dated to 1340–45. In 2010 a major conservation programme was undertaken on the Jesse Tree window.
The panels in the chapel of St Katherine are attributed to Arnold of Nijmegen and date from about 1520. They were acquired from the destroyed church of Saint-Jean, Rouen, with the last panel having been purchased in 1953.
The large triple lancet to the nave west end was glazed at the expense of Dean Creighton at a cost of £140 in 1664. It was repaired in 1813, and the central light was largely replaced to a design by Archibald Keightley Nicholson between 1925 and 1931. The main north and south transept end windows by James Powell and Sons were erected in the early 20th century.
The greater part of the stone carving of Wells Cathedral comprises foliate capitals in the stiff-leaf style. They are found ornamenting the piers of the nave, choir and transepts. Stiff-leaf foliage is highly abstract. Though possibly influenced by carvings of acanthus leaves or vine leaves, it cannot be easily identified with any particular plant. Here the carving of the foliage is varied and vigorous, the springing leaves and deep undercuts casting shadows that contrast with the surface of the piers. In the transepts and towards the crossing in the nave the capitals have many small figurative carvings among the leaves. These include a man with toothache and a series of four scenes depicting the "Wages of Sin" in a narrative of fruit stealers who creep into an orchard and are then beaten by the farmer. Another well-known carving is in the north transept aisle: a foliate corbel, on which climbs a lizard, sometimes identified as a salamander, a symbol of eternal life.
Carvings in the Decorated Gothic style may be found in the eastern end of the buildings, where there are many carved bosses. In the chapter house, the carvings of the 51 stalls include numerous small heads of great variety, many of them smiling or laughing. A well-known figure is the corbel of the dragon-slaying monk in the chapter house stair. The large continuous capital that encircles the central pillar of the chapter house is markedly different in style to the stiff-leaf of the Early English period. In contrast to the bold projections and undercutting of the earlier work, it has a rippling form and is clearly identifiable as grapevine.
The 15th-century cloisters have many small bosses ornamenting the vault. Two in the west cloister, near the gift shop and café, have been called sheela na gigs, i. e. female figures displaying their genitals and variously judged to depict the sin of lust or stem from ancient fertility cults.
Wells Cathedral has one of the finest sets of misericords in Britain. Its clergy has a long tradition of singing or reciting from the Book of Psalms each day, along with the customary daily reading of the Holy Office. In medieval times the clergy assembled in the church eight times daily for the canonical hours. As the greater part of the services was recited while standing, many monastic or collegiate churches fitted stalls whose seats tipped up to provide a ledge for the monk or cleric to lean against. These were "misericords" because their installation was an act of mercy. Misericords typically have a carved figurative bracket beneath the ledge framed by two floral motifs known, in heraldic manner, as "supporters".
The misericords date from 1330 to 1340. They may have been carved under the direction of Master Carpenter John Strode, although his name is not recorded before 1341. He was assisted by Bartholomew Quarter, who is documented from 1343. They originally numbered 90, of which 65 have survived. Sixty-one are installed in the choir, three are displayed in the cathedral, and one is held by the Victoria and Albert Museum. New stalls were ordered when the eastern end of the choir was extended in the early 14th century. The canons complained that they had borne the cost of the rebuilding and ordered the prebendary clerics to pay for their own stalls. When the newly refurbished choir opened in 1339 many misericords were left unfinished, including one-fifth of the surviving 65. Many of the clerics had not paid, having been called to contribute a total sum of £200. The misericords survived better than the other sections of the stalls, which during the Protestant Reformation had their canopies chopped off and galleries inserted above them. One misericord, showing a boy pulling a thorn from his foot, dates from the 17th century. In 1848 came a complete rearrangement of the choir furniture, and 61 of the misericords were reused in the restructured stalls.
The subject matter of the carvings of the central brackets as misericords varies, but many themes recur in different churches. Typically the themes are less unified or directly related to the Bible and Christian theology than small sculptures seen elsewhere within churches, such as bosses. This applies at Wells, where none of the misericord carvings is directly based on a Bible story. The subjects, chosen either by the woodcarver, or perhaps by the one paying for the stall, have no overriding theme. The sole unifying elements are the roundels on each side of the pictorial subject, which all show elaborately carved foliage, in most cases formal and stylised in the later Decorated manner, but with several examples of naturalistic foliage, including roses and bindweed. Many of the subjects carry traditional interpretations. The image of the "Pelican in her Piety" (believed to feed her young on her own blood) is a recognised symbol for Christ's love for the Church. A cat playing with a mouse may represent the Devil snaring a human soul. Other subjects illustrate popular fables or sayings such as "When the fox preaches, look to your geese". Many depict animals, some of which may symbolise a human vice or virtue, or an aspect of faith.
Twenty-seven of the carvings depict animals: rabbits, dogs, a puppy biting a cat, a ewe feeding a lamb, monkeys, lions, bats, and the Early Christian motif of two doves drinking from a ewer. Eighteen have mythological subjects, including mermaids, dragons and wyverns. Five are clearly narrative, such as the Fox and the Geese, and the story of Alexander the Great being raised to Heaven by griffins. There are three heads: a bishop in a mitre, an angel, and a woman wearing a veil over hair arranged in coils over each ear. Eleven carvings show human figures, among which are several of remarkable design, conceived by the artist specifically for their purpose of supporting a shelf. One figure lies beneath the seat, supporting the shelf with a cheek, a hand and a foot. Another sits in a contorted manner supporting the weight on his elbow, while a further figure squats with his knees wide apart and a strained look on his face.
Some of the cathedral's fittings and monuments are hundreds of years old. The brass lectern in the Lady Chapel dates from 1661 and has a moulded stand and foliate crest. In the north transept chapel is a 17th-century oak screen with columns, formerly used in cow stalls, with artisan Ionic capitals and cornice, set forward over the chest tomb of John Godelee. There is a bound oak chest from the 14th century, which was used to store the chapter seal and key documents. The bishop's throne dates from 1340, and has a panelled, canted front and stone doorway, and a deep nodding cusped ogee canopy above it, with three-stepped statue niches and pinnacles. The throne was restored by Anthony Salvin around 1850. Opposite the throne is a 19th-century octagonal pulpit on a coved base with panelled sides, and steps up from the north aisle. The round font in the south transept is from the former Saxon cathedral and has an arcade of round-headed arches, on a round plinth. The font cover was made in 1635 and is decorated with the heads of putti. The Chapel of St Martin is a memorial to every Somerset man who fell in World War I.
The monuments and tombs include Gisa, bishop; † 1088; William of Bitton, bishop; † 1274; William of March, bishop; † 1302; John Droxford; † 1329; John Godelee; † 1333; John Middleton, died †1350; Ralph of Shrewsbury, died †; John Harewell, bishop; † 1386; William Bykonyll; † c. 1448; John Bernard; † 1459; Thomas Beckington; † died 1464; John Gunthorpe; † 1498; John Still; † 1607; Robert Creighton; † 1672; Richard Kidder, bishop; † 1703; George Hooper, bishop; † 1727 and Arthur Harvey, bishop; † 1894.
In the north transept is Wells Cathedral clock, an astronomical clock from about 1325 believed to be by Peter Lightfoot, a monk of Glastonbury. Its mechanism, dated between 1386 and 1392, was replaced in the 19th century and the original moved to the Science Museum in London, where it still operates. It is the second oldest surviving clock in England after the Salisbury Cathedral clock.
The clock has its original medieval face. Apart from the time on a 24-hour dial, it shows the motion of the Sun and Moon, the phases of the Moon, and the time since the last new Moon. The astronomical dial presents a geocentric or pre-Copernican view, with the Sun and Moon revolving round a central fixed Earth, like that of the clock at Ottery St Mary. The quarters are chimed by a quarter jack: a small automaton known as Jack Blandifers, who hits two bells with hammers and two with his heels. At the striking of the clock, jousting knights appear above the clock face.
On the outer wall of the transept, opposite Vicars' Hall, is a second clock face of the same clock, placed there just over seventy years after the interior clock and driven by the same mechanism. The second clock face has two quarter jacks (which strike on the quarter-hour) in the form of knights in armour.
In 2010 the official clock-winder retired and was replaced by an electric mechanism.
The first record of an organ at this church dates from 1310. A smaller organ, probably for the Lady Chapel, was installed in 1415. In 1620 an organ built by Thomas Dallam was installed at a cost of £398 1s 5d.
The 1620 organ was destroyed by parliamentary soldiers in 1643. An organ built in 1662 was enlarged in 1786 and again in 1855. In 1909–1910 an organ was built by Harrison & Harrison of Durham, with the best parts of the old organ retained. It has been serviced by the same company ever since.
Since November 1996 the cathedral has also had a portable chamber organ, by the Scottish makers, Lammermuir. It is used regularly to accompany performances of Tudor and baroque music.
The first recorded organist of Wells was Walter Bagele (or Vageler) in 1416. The post of organist or assistant organist has been held by more than 60 people since. Peter Stanley Lyons was Master of Choristers at Wells Cathedral, and Director of Music at Wells Cathedral School in 1954–1960. The choral conductor James William Webb-Jones, father of Lyons's wife Bridget (whom he married in the cathedral), was Headmaster of Wells Cathedral School in 1955–1960. Malcolm Archer was the appointed Organist and Master of the Choristers from 1996 to 2004. Matthew Owens was the appointed organist from 2005 to 2019.
There has been a choir of boy choristers at Wells since 909. Currently there are 18 boy choristers and a similar number of girl choristers, aged from eight to fourteen. The Vicars Choral was formed in the 12th century and the sung liturgy provided by a traditional cathedral choir of men and boys until the formation of an additional choir of girls in 1994. The boys and girls sing alternately with the Vicars Choral and are educated at Wells Cathedral School.
The Vicars Choral currently number twelve men, of whom three are choral scholars. Since 1348 the College of Vicars had its own accommodation in a quadrangle converted in the early 15th century to form Vicar's Close. The Vicars Choral generally perform with the choristers, except on Wednesdays, when they sing alone, allowing them to present a different repertoire, in particular plainsong.
In December 2010 Wells Cathedral Choir was rated by Gramophone magazine as "the highest ranking choir with children in the world". It continues to provide music for the liturgy at Sunday and weekday services. The choir has made many recordings and toured frequently, including performances in Beijing and Hong Kong in 2012. Its repertoire ranges from the choral music of the Renaissance to recently commissioned works.
The Wells Cathedral Chamber Choir is a mixed adult choir of 25 members, formed in 1986 to sing at the midnight service on Christmas Eve, and invited to sing at several other special services. It now sings for about 30 services a year, when the Cathedral Choir is in recess or on tour, and spends one week a year singing as the "choir in residence" at another cathedral. Although primarily liturgical, the choir's repertoire includes other forms of music, as well as performances at engagements such as weddings and funerals.
The cathedral is home to Wells Cathedral Oratorio Society (WCOS), founded in 1896. With around 160 voices, the society gives three concerts a year under the direction of Matthew Owens, Organist and Master of the Choristers at the cathedral. Concerts are normally in early November, December (an annual performance of Handel's Messiah) and late March. It performs with a number of specialist orchestras including: Music for Awhile, Chameleon Arts and La Folia.
The bells at Wells Cathedral are the heaviest ring of ten bells in the world, the tenor bell (the 10th and largest), known as Harewell, weighing 56.25 long hundredweight (2,858 kg). They are hung for full-circle ringing in the English style of change ringing. These bells are now hung in the south-west tower, although some were originally hung in the central tower.
The library above the eastern cloister was built between 1430 and 1508. Its collection is in three parts: early documents housed in the Muniment Room; the collection predating 1800 housed in the Chained Library; and the post-1800 collection housed in the Reading Room. The chapter's earlier collection was destroyed during the Reformation, so that the present library consists chiefly of early printed books, rather than medieval manuscripts. The earlier books in the Chained Library number 2,800 volumes and give an indication of the variety of interests of the members of the cathedral chapter from the Reformation until 1800. The focus of the collection is predominantly theology, but there are volumes on science, medicine, exploration, and languages. Books of particular interest include Pliny's Natural History printed in 1472, an Atlas of the World by Abraham Ortelius, printed in 1606, and a set of the works by Aristotle that once belonged to Erasmus. The library is open to the public at appointed times in the summer and presents a small exhibition of documents and books.
Three early registers of the Dean and Chapter edited by W. H. B. Bird for the Historical Manuscripts Commissioners – Liber Albus I (White Book; R I), Liber Albus II (R III) and Liber Ruber (Red Book; R II, section i) – were published in 1907. They contain with some repetition, a cartulary of possessions of the cathedral, with grants of land back to the 8th century, well before hereditary surnames developed in England, and acts of the Dean and Chapter and surveys of their estates, mostly in Somerset.
Adjacent to the cathedral is a large lawned area, Cathedral Green, with three ancient gateways: Brown's Gatehouse, Penniless Porch and Chain Gate. On the green is the 12th-century Old Deanery, largely rebuilt in the late 15th century by Dean Gunthorpe and remodelled by Dean Bathurst in the late 17th century. No longer the dean's residence, it is used as diocesan offices.
To the south of the cathedral is the moated Bishop's Palace, begun about 1210 by Jocelin of Wells but dating mostly from the 1230s. In the 15th century Thomas Beckington added a north wing, now the bishop's residence. It was restored and extended by Benjamin Ferrey between 1846 and 1854.
To the north of the cathedral and connected to it by the Chain Gate is Vicars' Close, a street planned in the 14th century and claimed to be the oldest purely residential street in Europe, with all but one of its original buildings intact. Buildings in the close include the Vicars Hall and gateway at the south end, and the Vicars Chapel and Library at the north end.
The Liberty of St Andrew was the historic liberty and parish that encompassed the cathedral and surrounding lands closely associated with it.
The English painter J. M. W. Turner visited Wells in 1795, making sketches of the precinct and a water colour of the west front, now in the Tate gallery. Other artists whose paintings of the cathedral are in national collections are Albert Goodwin, John Syer and Ken Howard.
The cathedral served to inspire Ken Follett's 1989 novel The Pillars of the Earth and with a modified central tower, featured as the fictional Kingsbridge Cathedral at the end of the 2010 television adaptation of that novel. The interior of the cathedral was used for a 2007 Doctor Who episode, "The Lazarus Experiment", while the exterior shots were filmed at Southwark Cathedral.
An account of the damage to the cathedral during the Monmouth Rebellion is included in Arthur Conan Doyle's 1889 historical novel Micah Clarke.
The cathedral provided scenes for the 2019–2020 television series The Spanish Princess.
The Aon Center in downtown Chicago is 83 floors and 1,136 feet tall. It was the tallest building in Chicago when it was completed in 1974 and was then known as the Standard Oil Building.
KFC 37418 on 0Z37 07:50 Derby R.T.C. Network Rail - Chester Middle Yard Route Learner, at Warrington Arpley 20/10/2025
Fun retro look inspired by this round at Collabor88!
-Photo taken @Boyhood Home of Johnny Cash-
Style Details @ Confessions of a SL Shopaholic
Rascando el cielo
I found really tall the buildings in Panama. I heard they have a 90 floor building and are plannig for one with 100. Using concrete it is a really huge thing.
The alpine marmot (Marmota marmota) is a large ground-dwelling squirrel, from the genus of marmots. It is found in high numbers in mountainous areas of central and southern Europe, at heights between 800 and 3,200 m (2,600–10,500 ft) in the Alps, Carpathians, Tatras and Northern Apennines. In 1948 they were reintroduced with success in the Pyrenees, where the alpine marmot had disappeared at end of the Pleistocene epoch.
The alpine marmot originates as an animal of Pleistocene cold steppe, exquisitely adapted to this ice-age climate. As such, alpine marmots are excellent diggers, able to penetrate soil that even a pickaxe would have difficulty with, and spend up to nine months per year in hibernation.
Since the disappearance of the Pleistocene cold steppe, the alpine marmot persists in the high altitude alpine meadow. During the colonization of Alpine habitat, the alpine marmot has lost most of its genetic diversity through a bottleneck effect. It could not rebuild its genetic diversity ever since, as its lifestyle adapted to the Ice Age climate slowed its rate of genomic evolution. The alpine marmot is indeed one of the least genetically diverse wild-living animals
An adult alpine marmot is between 43 and 73 cm (17–29 in) in head-and-body length and the tail measures from 13 to 20 cm (5–8 in). The body mass ranges from 1.9 to 8 kg (4.2–17.6 lb), with the animals being significantly lighter in the spring (just after hibernation) than in the autumn (just before hibernation). The alpine marmot is sometimes considered the heaviest squirrel species, although some other marmot species have a similar weight range, making it unclear exactly which is the largest. Its coat is a mixture of blonde, reddish and dark gray fur. While most of the alpine marmot's fingers have claws, its thumbs have nails.
As its name suggests, the alpine marmot ranges throughout the European Alps, ranging through alpine areas of France, Italy, Switzerland, Germany, Slovenia and Austria. They have also been introduced elsewhere with sub-populations in the Pyrenees, France's Massif Central, Jura, Vosges, Black Forest, Apennine Mountains, and the Romanian Carpathians. The Tatra marmot (Marmota marmota latirostris Kratochvíl, 1961) represents an endemic subspecies of Alpine marmot that originated during the Quaternary period. Tatra marmots inhabit Tatry Mountains and Nízke Tatry Mountains. Marmots are abundant in their core population; in the Romanian Carpathians, for example, the population is estimated at 1,500 individuals. Alpine marmots prefer alpine meadows and high-altitude pastures, where colonies live in deep burrow systems situated in alluvial soil or rocky areas.
Marmots may be seen "sun bathing", but actually this is often on a flat rock and it is believed they are actually cooling and possibly this is a strategy to deal with parasites. Marmots are temperature sensitive and an increase in temperature can cause habitat loss for the species as a whole.
Alpine marmots eat plants such as grasses and herbs, as well as grain, insects, spiders and worms. They prefer young and tender plants over any other kind, and hold food in their forepaws while eating. They mainly emerge from their burrows to engage in feeding during the morning and afternoon, as they are not well suited to heat, which may result in them not feeding at all on very warm days. When the weather is suitable, they will consume large amounts of food in order to create a layer of fat on their body, enabling them to survive their long hibernation period.
When creating a burrow, they use both their forepaws and hind feet to assist in the work—the forepaws scrape away the soil, which is then pushed out of the way by the hind feet. If there are any stones in the way, the alpine marmot will remove them with its teeth provided that the stones aren't too large. "Living areas" are created at the end of a burrow, and are often lined with dried hay, grass and plant stems. Any other burrow tunnels that go nowhere are used as toilet areas. Once burrows have been completed, they only host one family, but are often enlarged by the next generation, sometimes creating very complex burrows over time. Each alpine marmot will live in a group that consists of several burrows, and which has a dominant breeding pair. Alpine marmots are very defensive against intruders, and will warn them off using intimidating behavior, such as beating of the tail and chattering of the teeth, and by marking their territory with their scent. One can often see an alpine marmot "standing" while they keep a look-out for potential predators or other dangers. Warnings are given, by emitting a series of loud whistles, after which members of the colony may be seen running for cover.
An alpine marmot at the end of summer. Note the fattened belly.
The mating season for alpine marmots occurs in the spring, right after their hibernation period comes to a close, which gives their offspring the highest possible chance of storing enough fat to survive the coming winter. Alpine marmots are able to breed once they reach an age of two years. Dominant females tend to suppress reproduction of subordinates by being antagonistic towards them while they are pregnant which causes stress and kills the young. Once the female is pregnant, she will take bedding materials (such as grass) into the burrow for when she gives birth after a gestation period of 33–34 days. Each litter consists of between one and seven babies, though this number is usually three. The babies are born blind and will grow dark fur within several days. The weaning period takes a further forty days, during which time the mother will leave the young in the burrow while she searches for food. After this period, the offspring will come out of the burrow and search for solid food themselves. Their fur becomes the same color as adult alpine marmots by the end of the summer, and after two years they will have reached their full size. If kept in captivity, alpine marmots can live up to 15–18 years.
As the summer begins to end, alpine marmots will gather old stems in their burrows in order to serve as bedding for their impending hibernation, which can start as early as October. They seal the burrow with a combination of earth and their own faeces. Once winter arrives, alpine marmots will huddle next to each other and begin hibernation, a process which lowers their heart rate to five beats per minute and breathing to 1–3 breaths per minute. During hibernation their stored fat supplies are used slowly, which usually allows them to survive the winter. Their body temperature will drop to almost the same as the air around them, although their heart and breathing rates will speed up if the environment approaches freezing point. Some alpine marmots will starve to death due to their layers of fat running out; this is most likely to happen in younger individuals.
For more information, please visit en.wikipedia.org/wiki/Alpine_marmot
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Honesty time. I've struggled for four years to get employment in my field. I finished my 4 year bachelor's program for photography and painting back in 2012. Since then I've busted my butt to get into advertising and have a consistent income. I knew it was going to be competitive, but I definitely was not expecting to still be counting my pennies come time for rent after 4+ years post graduation. It's rough. And it gets me down. I've bounced from serving jobs to internships that lead nowhere to scraping by on freelance mixed with savings. This summer the only job I was able to find that would hire me was a landscaping crew position. Pay isn't great and it's back breaking work, but it's exactly what I need in my life right now.
I've been glued to a computer screen for too many years feeling stressed and frustrated and I began to lose my love of art, the whole reason why I decided to study photography to begin with. So this summer I'm reconnecting with nature, breathing fresh air and being outside everyday, and kicking my body back into shape. It's hard, hot, dirty, sweaty, exhausting and certainly not glamorous. It's truthfully very humbling.
Although these birds of a feather appear to be different they are classified as "California Gulls". A young couple was throwing scraps of food to a group of ground squirrels and the Gulls couldn't get in on the action. Frustration set in and the beak locking Gull gave his buddy the business. A little foot to the chest too...no bandages were need while taking this shot!
This Buck was crossing the path in front of me.... once he was across he found a fat tree and started to scrap his antlers, I tried to get a shot but he didn't give me a chance. All the trees in this area have all the bark scraped off.
© All Rights Reserved. Please do not use or reproduce this image on Websites, Blogs or any other media without my explicit permission.
Fish, any of approximately 34,000 species of vertebrate animals (phylum Chordata) found in the fresh and salt waters of the world. Living species range from the primitive jawless lampreys and hagfishes through the cartilaginous sharks, skates, and rays to the abundant and diverse bony fishes. Most fish species are cold-blooded; however, one species, the opah (Lampris guttatus), is warm-blooded.
The term fish is applied to a variety of vertebrates of several evolutionary lines. It describes a life-form rather than a taxonomic group. As members of the phylum Chordata, fish share certain features with other vertebrates. These features are gill slits at some point in the life cycle, a notochord, or skeletal supporting rod, a dorsal hollow nerve cord, and a tail. Living fishes represent some five classes, which are as distinct from one another as are the four classes of familiar air-breathing animals—amphibians, reptiles, birds, and mammals. For example, the jawless fishes (Agnatha) have gills in pouches and lack limb girdles. Extant agnathans are the lampreys and the hagfishes. As the name implies, the skeletons of fishes of the class Chondrichthyes (from chondr, “cartilage,” and ichthyes, “fish”) are made entirely of cartilage. Modern fish of this class lack a swim bladder, and their scales and teeth are made up of the same placoid material. Sharks, skates, and rays are examples of cartilaginous fishes. The bony fishes are by far the largest class. Examples range from the tiny seahorse to the 450-kg (1,000-pound) blue marlin, from the flattened soles and flounders to the boxy puffers and ocean sunfishes. Unlike the scales of the cartilaginous fishes, those of bony fishes, when present, grow throughout life and are made up of thin overlapping plates of bone. Bony fishes also have an operculum that covers the gill slits.
The study of fishes, the science of ichthyology, is of broad importance. Fishes are of interest to humans for many reasons, the most important being their relationship with and dependence on the environment. A more obvious reason for interest in fishes is their role as a moderate but important part of the world’s food supply. This resource, once thought unlimited, is now realized to be finite and in delicate balance with the biological, chemical, and physical factors of the aquatic environment. Overfishing, pollution, and alteration of the environment are the chief enemies of proper fisheries management, both in fresh waters and in the ocean. (For a detailed discussion of the technology and economics of fisheries, see commercial fishing.) Another practical reason for studying fishes is their use in disease control. As predators on mosquito larvae, they help curb malaria and other mosquito-borne diseases.
Fishes are valuable laboratory animals in many aspects of medical and biological research. For example, the readiness of many fishes to acclimate to captivity has allowed biologists to study behaviour, physiology, and even ecology under relatively natural conditions. Fishes have been especially important in the study of animal behaviour, where research on fishes has provided a broad base for the understanding of the more flexible behaviour of the higher vertebrates. The zebra fish is used as a model in studies of gene expression.
There are aesthetic and recreational reasons for an interest in fishes. Millions of people keep live fishes in home aquariums for the simple pleasure of observing the beauty and behaviour of animals otherwise unfamiliar to them. Aquarium fishes provide a personal challenge to many aquarists, allowing them to test their ability to keep a small section of the natural environment in their homes. Sportfishing is another way of enjoying the natural environment, also indulged in by millions of people every year. Interest in aquarium fishes and sportfishing supports multimillion-dollar industries throughout the world.
Fishes have been in existence for more than 450 million years, during which time they have evolved repeatedly to fit into almost every conceivable type of aquatic habitat. In a sense, land vertebrates are simply highly modified fishes: when fishes colonized the land habitat, they became tetrapod (four-legged) land vertebrates. The popular conception of a fish as a slippery, streamlined aquatic animal that possesses fins and breathes by gills applies to many fishes, but far more fishes deviate from that conception than conform to it. For example, the body is elongate in many forms and greatly shortened in others; the body is flattened in some (principally in bottom-dwelling fishes) and laterally compressed in many others; the fins may be elaborately extended, forming intricate shapes, or they may be reduced or even lost; and the positions of the mouth, eyes, nostrils, and gill openings vary widely. Air breathers have appeared in several evolutionary lines.
Many fishes are cryptically coloured and shaped, closely matching their respective environments; others are among the most brilliantly coloured of all organisms, with a wide range of hues, often of striking intensity, on a single individual. The brilliance of pigments may be enhanced by the surface structure of the fish, so that it almost seems to glow. A number of unrelated fishes have actual light-producing organs. Many fishes are able to alter their coloration—some for the purpose of camouflage, others for the enhancement of behavioral signals.
Fishes range in adult length from less than 10 mm (0.4 inch) to more than 20 metres (60 feet) and in weight from about 1.5 grams (less than 0.06 ounce) to many thousands of kilograms. Some live in shallow thermal springs at temperatures slightly above 42 °C (100 °F), others in cold Arctic seas a few degrees below 0 °C (32 °F) or in cold deep waters more than 4,000 metres (13,100 feet) beneath the ocean surface. The structural and, especially, the physiological adaptations for life at such extremes are relatively poorly known and provide the scientifically curious with great incentive for study.
Almost all natural bodies of water bear fish life, the exceptions being very hot thermal ponds and extremely salt-alkaline lakes, such as the Dead Sea in Asia and the Great Salt Lake in North America. The present distribution of fishes is a result of the geological history and development of Earth as well as the ability of fishes to undergo evolutionary change and to adapt to the available habitats. Fishes may be seen to be distributed according to habitat and according to geographical area. Major habitat differences are marine and freshwater. For the most part, the fishes in a marine habitat differ from those in a freshwater habitat, even in adjacent areas, but some, such as the salmon, migrate from one to the other. The freshwater habitats may be seen to be of many kinds. Fishes found in mountain torrents, Arctic lakes, tropical lakes, temperate streams, and tropical rivers will all differ from each other, both in obvious gross structure and in physiological attributes. Even in closely adjacent habitats where, for example, a tropical mountain torrent enters a lowland stream, the fish fauna will differ. The marine habitats can be divided into deep ocean floors (benthic), mid-water oceanic (bathypelagic), surface oceanic (pelagic), rocky coast, sandy coast, muddy shores, bays, estuaries, and others. Also, for example, rocky coastal shores in tropical and temperate regions will have different fish faunas, even when such habitats occur along the same coastline.
Although much is known about the present geographical distribution of fishes, far less is known about how that distribution came about. Many parts of the fish fauna of the fresh waters of North America and Eurasia are related and undoubtedly have a common origin. The faunas of Africa and South America are related, extremely old, and probably an expression of the drifting apart of the two continents. The fauna of southern Asia is related to that of Central Asia, and some of it appears to have entered Africa. The extremely large shore-fish faunas of the Indian and tropical Pacific oceans comprise a related complex, but the tropical shore fauna of the Atlantic, although containing Indo-Pacific components, is relatively limited and probably younger. The Arctic and Antarctic marine faunas are quite different from each other. The shore fauna of the North Pacific is quite distinct, and that of the North Atlantic more limited and probably younger. Pelagic oceanic fishes, especially those in deep waters, are similar the world over, showing little geographical isolation in terms of family groups. The deep oceanic habitat is very much the same throughout the world, but species differences do exist, showing geographical areas determined by oceanic currents and water masses.
All aspects of the life of a fish are closely correlated with adaptation to the total environment, physical, chemical, and biological. In studies, all the interdependent aspects of fish, such as behaviour, locomotion, reproduction, and physical and physiological characteristics, must be taken into account.
Correlated with their adaptation to an extremely wide variety of habitats is the extremely wide variety of life cycles that fishes display. The great majority hatch from relatively small eggs a few days to several weeks or more after the eggs are scattered in the water. Newly hatched young are still partially undeveloped and are called larvae until body structures such as fins, skeleton, and some organs are fully formed. Larval life is often very short, usually less than a few weeks, but it can be very long, some lampreys continuing as larvae for at least five years. Young and larval fishes, before reaching sexual maturity, must grow considerably, and their small size and other factors often dictate that they live in a habitat different than that of the adults. For example, most tropical marine shore fishes have pelagic larvae. Larval food also is different, and larval fishes often live in shallow waters, where they may be less exposed to predators.
After a fish reaches adult size, the length of its life is subject to many factors, such as innate rates of aging, predation pressure, and the nature of the local climate. The longevity of a species in the protected environment of an aquarium may have nothing to do with how long members of that species live in the wild. Many small fishes live only one to three years at the most. In some species, however, individuals may live as long as 10 or 20 or even 100 years.
Fish behaviour is a complicated and varied subject. As in almost all animals with a central nervous system, the nature of a response of an individual fish to stimuli from its environment depends upon the inherited characteristics of its nervous system, on what it has learned from past experience, and on the nature of the stimuli. Compared with the variety of human responses, however, that of a fish is stereotyped, not subject to much modification by “thought” or learning, and investigators must guard against anthropomorphic interpretations of fish behaviour.
Fishes perceive the world around them by the usual senses of sight, smell, hearing, touch, and taste and by special lateral line water-current detectors. In the few fishes that generate electric fields, a process that might best be called electrolocation aids in perception. One or another of these senses often is emphasized at the expense of others, depending upon the fish’s other adaptations. In fishes with large eyes, the sense of smell may be reduced; others, with small eyes, hunt and feed primarily by smell (such as some eels).
Specialized behaviour is primarily concerned with the three most important activities in the fish’s life: feeding, reproduction, and escape from enemies. Schooling behaviour of sardines on the high seas, for instance, is largely a protective device to avoid enemies, but it is also associated with and modified by their breeding and feeding requirements. Predatory fishes are often solitary, lying in wait to dart suddenly after their prey, a kind of locomotion impossible for beaked parrot fishes, which feed on coral, swimming in small groups from one coral head to the next. In addition, some predatory fishes that inhabit pelagic environments, such as tunas, often school.
Sleep in fishes, all of which lack true eyelids, consists of a seemingly listless state in which the fish maintains its balance but moves slowly. If attacked or disturbed, most can dart away. A few kinds of fishes lie on the bottom to sleep. Most catfishes, some loaches, and some eels and electric fishes are strictly nocturnal, being active and hunting for food during the night and retiring during the day to holes, thick vegetation, or other protective parts of the environment.
Communication between members of a species or between members of two or more species often is extremely important, especially in breeding behaviour (see below Reproduction). The mode of communication may be visual, as between the small so-called cleaner fish and a large fish of a very different species. The larger fish often allows the cleaner to enter its mouth to remove gill parasites. The cleaner is recognized by its distinctive colour and actions and therefore is not eaten, even if the larger fish is normally a predator. Communication is often chemical, signals being sent by specific chemicals called pheromones.
Many fishes have a streamlined body and swim freely in open water. Fish locomotion is closely correlated with habitat and ecological niche (the general position of the animal to its environment).
Many fishes in both marine and fresh waters swim at the surface and have mouths adapted to feed best (and sometimes only) at the surface. Often such fishes are long and slender, able to dart at surface insects or at other surface fishes and in turn to dart away from predators; needlefishes, halfbeaks, and topminnows (such as killifish and mosquito fish) are good examples. Oceanic flying fishes escape their predators by gathering speed above the water surface, with the lower lobe of the tail providing thrust in the water. They then glide hundreds of yards on enlarged, winglike pectoral and pelvic fins. South American freshwater flying fishes escape their enemies by jumping and propelling their strongly keeled bodies out of the water.
So-called mid-water swimmers, the most common type of fish, are of many kinds and live in many habitats. The powerful fusiform tunas and the trouts, for example, are adapted for strong, fast swimming, the tunas to capture prey speedily in the open ocean and the trouts to cope with the swift currents of streams and rivers. The trout body form is well adapted to many habitats. Fishes that live in relatively quiet waters such as bays or lake shores or slow rivers usually are not strong, fast swimmers but are capable of short, quick bursts of speed to escape a predator. Many of these fishes have their sides flattened, examples being the sunfish and the freshwater angelfish of aquarists. Fish associated with the bottom or substrate usually are slow swimmers. Open-water plankton-feeding fishes almost always remain fusiform and are capable of rapid, strong movement (for example, sardines and herrings of the open ocean and also many small minnows of streams and lakes).
Bottom-living fishes are of many kinds and have undergone many types of modification of their body shape and swimming habits. Rays, which evolved from strong-swimming mid-water sharks, usually stay close to the bottom and move by undulating their large pectoral fins. Flounders live in a similar habitat and move over the bottom by undulating the entire body. Many bottom fishes dart from place to place, resting on the bottom between movements, a motion common in gobies. One goby relative, the mudskipper, has taken to living at the edge of pools along the shore of muddy mangrove swamps. It escapes its enemies by flipping rapidly over the mud, out of the water. Some catfishes, synbranchid eels, the so-called climbing perch, and a few other fishes venture out over damp ground to find more promising waters than those that they left. They move by wriggling their bodies, sometimes using strong pectoral fins; most have accessory air-breathing organs. Many bottom-dwelling fishes live in mud holes or rocky crevices. Marine eels and gobies commonly are found in such habitats and for the most part venture far beyond their cavelike homes. Some bottom dwellers, such as the clingfishes (Gobiesocidae), have developed powerful adhesive disks that enable them to remain in place on the substrate in areas such as rocky coasts, where the action of the waves is great.
The methods of reproduction in fishes are varied, but most fishes lay a large number of small eggs, fertilized and scattered outside of the body. The eggs of pelagic fishes usually remain suspended in the open water. Many shore and freshwater fishes lay eggs on the bottom or among plants. Some have adhesive eggs. The mortality of the young and especially of the eggs is very high, and often only a few individuals grow to maturity out of hundreds, thousands, and in some cases millions of eggs laid.
Males produce sperm, usually as a milky white substance called milt, in two (sometimes one) testes within the body cavity. In bony fishes a sperm duct leads from each testis to a urogenital opening behind the vent or anus. In sharks and rays and in cyclostomes the duct leads to a cloaca. Sometimes the pelvic fins are modified to help transmit the milt to the eggs at the female’s vent or on the substrate where the female has placed them. Sometimes accessory organs are used to fertilize females internally—for example, the claspers of many sharks and rays.
In the females the eggs are formed in two ovaries (sometimes only one) and pass through the ovaries to the urogenital opening and to the outside. In some fishes the eggs are fertilized internally but are shed before development takes place. Members of about a dozen families each of bony fishes (teleosts) and sharks bear live young. Many skates and rays also bear live young. In some bony fishes the eggs simply develop within the female, the young emerging when the eggs hatch (ovoviviparous). Others develop within the ovary and are nourished by ovarian tissues after hatching (viviparous). There are also other methods utilized by fishes to nourish young within the female. In all live-bearers the young are born at a relatively large size and are few in number. In one family of primarily marine fishes, the surfperches from the Pacific coast of North America, Japan, and Korea, the males of at least one species are born sexually mature, although they are not fully grown.
Some fishes are hermaphroditic—an individual producing both sperm and eggs, usually at different stages of its life. Self-fertilization, however, is probably rare.
Successful reproduction and, in many cases, defense of the eggs and the young are assured by rather stereotypical but often elaborate courtship and parental behaviour, either by the male or the female or both. Some fishes prepare nests by hollowing out depressions in the sand bottom (cichlids, for example), build nests with plant materials and sticky threads excreted by the kidneys (sticklebacks), or blow a cluster of mucus-covered bubbles at the water surface (gouramis). The eggs are laid in these structures. Some varieties of cichlids and catfishes incubate eggs in their mouths.
Some fishes, such as salmon, undergo long migrations from the ocean and up large rivers to spawn in the gravel beds where they themselves hatched (anadromous fishes). Some, such as the freshwater eels (family Anguillidae), live and grow to maturity in fresh water and migrate to the sea to spawn (catadromous fishes). Other fishes undertake shorter migrations from lakes into streams, within the ocean, or enter spawning habitats that they do not ordinarily occupy in other ways.
The basic structure and function of the fish body are similar to those of all other vertebrates. The usual four types of tissues are present: surface or epithelial, connective (bone, cartilage, and fibrous tissues, as well as their derivative, blood), nerve, and muscle tissues. In addition, the fish’s organs and organ systems parallel those of other vertebrates.
The typical fish body is streamlined and spindle-shaped, with an anterior head, a gill apparatus, and a heart, the latter lying in the midline just below the gill chamber. The body cavity, containing the vital organs, is situated behind the head in the lower anterior part of the body. The anus usually marks the posterior termination of the body cavity and most often occurs just in front of the base of the anal fin. The spinal cord and vertebral column continue from the posterior part of the head to the base of the tail fin, passing dorsal to the body cavity and through the caudal (tail) region behind the body cavity. Most of the body is of muscular tissue, a high proportion of which is necessitated by swimming. In the course of evolution this basic body plan has been modified repeatedly into the many varieties of fish shapes that exist today.
The skeleton forms an integral part of the fish’s locomotion system, as well as serving to protect vital parts. The internal skeleton consists of the skull bones (except for the roofing bones of the head, which are really part of the external skeleton), the vertebral column, and the fin supports (fin rays). The fin supports are derived from the external skeleton but will be treated here because of their close functional relationship to the internal skeleton. The internal skeleton of cyclostomes, sharks, and rays is of cartilage; that of many fossil groups and some primitive living fishes is mostly of cartilage but may include some bone. In place of the vertebral column, the earliest vertebrates had a fully developed notochord, a flexible stiff rod of viscous cells surrounded by a strong fibrous sheath. During the evolution of modern fishes the rod was replaced in part by cartilage and then by ossified cartilage. Sharks and rays retain a cartilaginous vertebral column; bony fishes have spool-shaped vertebrae that in the more primitive living forms only partially replace the notochord. The skull, including the gill arches and jaws of bony fishes, is fully, or at least partially, ossified. That of sharks and rays remains cartilaginous, at times partially replaced by calcium deposits but never by true bone.
The supportive elements of the fins (basal or radial bones or both) have changed greatly during fish evolution. Some of these changes are described in the section below (Evolution and paleontology). Most fishes possess a single dorsal fin on the midline of the back. Many have two and a few have three dorsal fins. The other fins are the single tail and anal fins and paired pelvic and pectoral fins. A small fin, the adipose fin, with hairlike fin rays, occurs in many of the relatively primitive teleosts (such as trout) on the back near the base of the caudal fin.
The skin of a fish must serve many functions. It aids in maintaining the osmotic balance, provides physical protection for the body, is the site of coloration, contains sensory receptors, and, in some fishes, functions in respiration. Mucous glands, which aid in maintaining the water balance and offer protection from bacteria, are extremely numerous in fish skin, especially in cyclostomes and teleosts. Since mucous glands are present in the modern lampreys, it is reasonable to assume that they were present in primitive fishes, such as the ancient Silurian and Devonian agnathans. Protection from abrasion and predation is another function of the fish skin, and dermal (skin) bone arose early in fish evolution in response to this need. It is thought that bone first evolved in skin and only later invaded the cartilaginous areas of the fish’s body, to provide additional support and protection. There is some argument as to which came first, cartilage or bone, and fossil evidence does not settle the question. In any event, dermal bone has played an important part in fish evolution and has different characteristics in different groups of fishes. Several groups are characterized at least in part by the kind of bony scales they possess.
Scales have played an important part in the evolution of fishes. Primitive fishes usually had thick bony plates or thick scales in several layers of bone, enamel, and related substances. Modern teleost fishes have scales of bone, which, while still protective, allow much more freedom of motion in the body. A few modern teleosts (some catfishes, sticklebacks, and others) have secondarily acquired bony plates in the skin. Modern and early sharks possessed placoid scales, a relatively primitive type of scale with a toothlike structure, consisting of an outside layer of enamel-like substance (vitrodentine), an inner layer of dentine, and a pulp cavity containing nerves and blood vessels. Primitive bony fishes had thick scales of either the ganoid or the cosmoid type. Cosmoid scales have a hard, enamel-like outer layer, an inner layer of cosmine (a form of dentine), and then a layer of vascular bone (isopedine). In ganoid scales the hard outer layer is different chemically and is called ganoin. Under this is a cosminelike layer and then a vascular bony layer. The thin, translucent bony scales of modern fishes, called cycloid and ctenoid (the latter distinguished by serrations at the edges), lack enameloid and dentine layers.
Skin has several other functions in fishes. It is well supplied with nerve endings and presumably receives tactile, thermal, and pain stimuli. Skin is also well supplied with blood vessels. Some fishes breathe in part through the skin, by the exchange of oxygen and carbon dioxide between the surrounding water and numerous small blood vessels near the skin surface.
Skin serves as protection through the control of coloration. Fishes exhibit an almost limitless range of colours. The colours often blend closely with the surroundings, effectively hiding the animal. Many fishes use bright colours for territorial advertisement or as recognition marks for other members of their own species, or sometimes for members of other species. Many fishes can change their colour to a greater or lesser degree, by movement of pigment within the pigment cells (chromatophores). Black pigment cells (melanophores), of almost universal occurrence in fishes, are often juxtaposed with other pigment cells. When placed beneath iridocytes or leucophores (bearing the silvery or white pigment guanine), melanophores produce structural colours of blue and green. These colours are often extremely intense, because they are formed by refraction of light through the needlelike crystals of guanine. The blue and green refracted colours are often relatively pure, lacking the red and yellow rays, which have been absorbed by the black pigment (melanin) of the melanophores. Yellow, orange, and red colours are produced by erythrophores, cells containing the appropriate carotenoid pigments. Other colours are produced by combinations of melanophores, erythrophores, and iridocytes.
The major portion of the body of most fishes consists of muscles. Most of the mass is trunk musculature, the fin muscles usually being relatively small. The caudal fin is usually the most powerful fin, being moved by the trunk musculature. The body musculature is usually arranged in rows of chevron-shaped segments on each side. Contractions of these segments, each attached to adjacent vertebrae and vertebral processes, bends the body on the vertebral joint, producing successive undulations of the body, passing from the head to the tail, and producing driving strokes of the tail. It is the latter that provides the strong forward movement for most fishes.
The digestive system, in a functional sense, starts at the mouth, with the teeth used to capture prey or collect plant foods. Mouth shape and tooth structure vary greatly in fishes, depending on the kind of food normally eaten. Most fishes are predacious, feeding on small invertebrates or other fishes and have simple conical teeth on the jaws, on at least some of the bones of the roof of the mouth, and on special gill arch structures just in front of the esophagus. The latter are throat teeth. Most predacious fishes swallow their prey whole, and the teeth are used for grasping and holding prey, for orienting prey to be swallowed (head first) and for working the prey toward the esophagus. There are a variety of tooth types in fishes. Some fishes, such as sharks and piranhas, have cutting teeth for biting chunks out of their victims. A shark’s tooth, although superficially like that of a piranha, appears in many respects to be a modified scale, while that of the piranha is like that of other bony fishes, consisting of dentine and enamel. Parrot fishes have beaklike mouths with short incisor-like teeth for breaking off coral and have heavy pavementlike throat teeth for crushing the coral. Some catfishes have small brushlike teeth, arranged in rows on the jaws, for scraping plant and animal growth from rocks. Many fishes (such as the Cyprinidae or minnows) have no jaw teeth at all but have very strong throat teeth.
Some fishes gather planktonic food by straining it from their gill cavities with numerous elongate stiff rods (gill rakers) anchored by one end to the gill bars. The food collected on these rods is passed to the throat, where it is swallowed. Most fishes have only short gill rakers that help keep food particles from escaping out the mouth cavity into the gill chamber.
Once reaching the throat, food enters a short, often greatly distensible esophagus, a simple tube with a muscular wall leading into a stomach. The stomach varies greatly in fishes, depending upon the diet. In most predacious fishes it is a simple straight or curved tube or pouch with a muscular wall and a glandular lining. Food is largely digested there and leaves the stomach in liquid form.
Between the stomach and the intestine, ducts enter the digestive tube from the liver and pancreas. The liver is a large, clearly defined organ. The pancreas may be embedded in it, diffused through it, or broken into small parts spread along some of the intestine. The junction between the stomach and the intestine is marked by a muscular valve. Pyloric ceca (blind sacs) occur in some fishes at this junction and have a digestive or absorptive function or both.
The intestine itself is quite variable in length, depending upon the fish’s diet. It is short in predacious forms, sometimes no longer than the body cavity, but long in herbivorous forms, being coiled and several times longer than the entire length of the fish in some species of South American catfishes. The intestine is primarily an organ for absorbing nutrients into the bloodstream. The larger its internal surface, the greater its absorptive efficiency, and a spiral valve is one method of increasing its absorption surface.
Sharks, rays, chimaeras, lungfishes, surviving chondrosteans, holosteans, and even a few of the more primitive teleosts have a spiral valve or at least traces of it in the intestine. Most modern teleosts have increased the area of the intestinal walls by having numerous folds and villi (fingerlike projections) somewhat like those in humans. Undigested substances are passed to the exterior through the anus in most teleost fishes. In lungfishes, sharks, and rays, it is first passed through the cloaca, a common cavity receiving the intestinal opening and the ducts from the urogenital system.
Oxygen and carbon dioxide dissolve in water, and most fishes exchange dissolved oxygen and carbon dioxide in water by means of the gills. The gills lie behind and to the side of the mouth cavity and consist of fleshy filaments supported by the gill arches and filled with blood vessels, which give gills a bright red colour. Water taken in continuously through the mouth passes backward between the gill bars and over the gill filaments, where the exchange of gases takes place. The gills are protected by a gill cover in teleosts and many other fishes but by flaps of skin in sharks, rays, and some of the older fossil fish groups. The blood capillaries in the gill filaments are close to the gill surface to take up oxygen from the water and to give up excess carbon dioxide to the water.
Most modern fishes have a hydrostatic (ballast) organ, called the swim bladder, that lies in the body cavity just below the kidney and above the stomach and intestine. It originated as a diverticulum of the digestive canal. In advanced teleosts, especially the acanthopterygians, the bladder has lost its connection with the digestive tract, a condition called physoclistic. The connection has been retained (physostomous) by many relatively primitive teleosts. In several unrelated lines of fishes, the bladder has become specialized as a lung or, at least, as a highly vascularized accessory breathing organ. Some fishes with such accessory organs are obligate air breathers and will drown if denied access to the surface, even in well-oxygenated water. Fishes with a hydrostatic form of swim bladder can control their depth by regulating the amount of gas in the bladder. The gas, mostly oxygen, is secreted into the bladder by special glands, rendering the fish more buoyant; the gas is absorbed into the bloodstream by another special organ, reducing the overall buoyancy and allowing the fish to sink. Some deep-sea fishes may have oils, rather than gas, in the bladder. Other deep-sea and some bottom-living forms have much-reduced swim bladders or have lost the organ entirely.
The swim bladder of fishes follows the same developmental pattern as the lungs of land vertebrates. There is no doubt that the two structures have the same historical origin in primitive fishes. More or less intermediate forms still survive among the more primitive types of fishes, such as the lungfishes Lepidosiren and Protopterus.
The circulatory, or blood vascular, system consists of the heart, the arteries, the capillaries, and the veins. It is in the capillaries that the interchange of oxygen, carbon dioxide, nutrients, and other substances such as hormones and waste products takes place. The capillaries lead to the veins, which return the venous blood with its waste products to the heart, kidneys, and gills. There are two kinds of capillary beds: those in the gills and those in the rest of the body. The heart, a folded continuous muscular tube with three or four saclike enlargements, undergoes rhythmic contractions and receives venous blood in a sinus venosus. It passes the blood to an auricle and then into a thick muscular pump, the ventricle. From the ventricle the blood goes to a bulbous structure at the base of a ventral aorta just below the gills. The blood passes to the afferent (receiving) arteries of the gill arches and then to the gill capillaries. There waste gases are given off to the environment, and oxygen is absorbed. The oxygenated blood enters efferent (exuant) arteries of the gill arches and then flows into the dorsal aorta. From there blood is distributed to the tissues and organs of the body. One-way valves prevent backflow. The circulation of fishes thus differs from that of the reptiles, birds, and mammals in that oxygenated blood is not returned to the heart prior to distribution to the other parts of the body.
The primary excretory organ in fishes, as in other vertebrates, is the kidney. In fishes some excretion also takes place in the digestive tract, skin, and especially the gills (where ammonia is given off). Compared with land vertebrates, fishes have a special problem in maintaining their internal environment at a constant concentration of water and dissolved substances, such as salts. Proper balance of the internal environment (homeostasis) of a fish is in a great part maintained by the excretory system, especially the kidney.
The kidney, gills, and skin play an important role in maintaining a fish’s internal environment and checking the effects of osmosis. Marine fishes live in an environment in which the water around them has a greater concentration of salts than they can have inside their body and still maintain life. Freshwater fishes, on the other hand, live in water with a much lower concentration of salts than they require inside their bodies. Osmosis tends to promote the loss of water from the body of a marine fish and absorption of water by that of a freshwater fish. Mucus in the skin tends to slow the process but is not a sufficient barrier to prevent the movement of fluids through the permeable skin. When solutions on two sides of a permeable membrane have different concentrations of dissolved substances, water will pass through the membrane into the more concentrated solution, while the dissolved chemicals move into the area of lower concentration (diffusion).
The kidney of freshwater fishes is often larger in relation to body weight than that of marine fishes. In both groups the kidney excretes wastes from the body, but the kidney of freshwater fishes also excretes large amounts of water, counteracting the water absorbed through the skin. Freshwater fishes tend to lose salt to the environment and must replace it. They get some salt from their food, but the gills and skin inside the mouth actively absorb salt from water passed through the mouth. This absorption is performed by special cells capable of moving salts against the diffusion gradient. Freshwater fishes drink very little water and take in little water with their food.
Marine fishes must conserve water, and therefore their kidneys excrete little water. To maintain their water balance, marine fishes drink large quantities of seawater, retaining most of the water and excreting the salt. Most nitrogenous waste in marine fishes appears to be secreted by the gills as ammonia. Marine fishes can excrete salt by clusters of special cells (chloride cells) in the gills.
There are several teleosts—for example, the salmon—that travel between fresh water and seawater and must adjust to the reversal of osmotic gradients. They adjust their physiological processes by spending time (often surprisingly little time) in the intermediate brackish environment.
Marine hagfishes, sharks, and rays have osmotic concentrations in their blood about equal to that of seawater and so do not have to drink water nor perform much physiological work to maintain their osmotic balance. In sharks and rays the osmotic concentration is kept high by retention of urea in the blood. Freshwater sharks have a lowered concentration of urea in the blood.
Endocrine glands secrete their products into the bloodstream and body tissues and, along with the central nervous system, control and regulate many kinds of body functions. Cyclostomes have a well-developed endocrine system, and presumably it was well developed in the early Agnatha, ancestral to modern fishes. Although the endocrine system in fishes is similar to that of higher vertebrates, there are numerous differences in detail. The pituitary, the thyroid, the suprarenals, the adrenals, the pancreatic islets, the sex glands (ovaries and testes), the inner wall of the intestine, and the bodies of the ultimobranchial gland make up the endocrine system in fishes. There are some others whose function is not well understood. These organs regulate sexual activity and reproduction, growth, osmotic pressure, general metabolic activities such as the storage of fat and the utilization of foodstuffs, blood pressure, and certain aspects of skin colour. Many of these activities are also controlled in part by the central nervous system, which works with the endocrine system in maintaining the life of a fish. Some parts of the endocrine system are developmentally, and undoubtedly evolutionarily, derived from the nervous system.
As in all vertebrates, the nervous system of fishes is the primary mechanism coordinating body activities, as well as integrating these activities in the appropriate manner with stimuli from the environment. The central nervous system, consisting of the brain and spinal cord, is the primary integrating mechanism. The peripheral nervous system, consisting of nerves that connect the brain and spinal cord to various body organs, carries sensory information from special receptor organs such as the eyes, internal ears, nares (sense of smell), taste glands, and others to the integrating centres of the brain and spinal cord. The peripheral nervous system also carries information via different nerve cells from the integrating centres of the brain and spinal cord. This coded information is carried to the various organs and body systems, such as the skeletal muscular system, for appropriate action in response to the original external or internal stimulus. Another branch of the nervous system, the autonomic nervous system, helps to coordinate the activities of many glands and organs and is itself closely connected to the integrating centres of the brain.
The brain of the fish is divided into several anatomical and functional parts, all closely interconnected but each serving as the primary centre of integrating particular kinds of responses and activities. Several of these centres or parts are primarily associated with one type of sensory perception, such as sight, hearing, or smell (olfaction).
The sense of smell is important in almost all fishes. Certain eels with tiny eyes depend mostly on smell for location of food. The olfactory, or nasal, organ of fishes is located on the dorsal surface of the snout. The lining of the nasal organ has special sensory cells that perceive chemicals dissolved in the water, such as substances from food material, and send sensory information to the brain by way of the first cranial nerve. Odour also serves as an alarm system. Many fishes, especially various species of freshwater minnows, react with alarm to a chemical released from the skin of an injured member of their own species.
Many fishes have a well-developed sense of taste, and tiny pitlike taste buds or organs are located not only within their mouth cavities but also over their heads and parts of their body. Catfishes, which often have poor vision, have barbels (“whiskers”) that serve as supplementary taste organs, those around the mouth being actively used to search out food on the bottom. Some species of naturally blind cave fishes are especially well supplied with taste buds, which often cover most of their body surface.
Sight is extremely important in most fishes. The eye of a fish is basically like that of all other vertebrates, but the eyes of fishes are extremely varied in structure and adaptation. In general, fishes living in dark and dim water habitats have large eyes, unless they have specialized in some compensatory way so that another sense (such as smell) is dominant, in which case the eyes will often be reduced. Fishes living in brightly lighted shallow waters often will have relatively small but efficient eyes. Cyclostomes have somewhat less elaborate eyes than other fishes, with skin stretched over the eyeball perhaps making their vision somewhat less effective. Most fishes have a spherical lens and accommodate their vision to far or near subjects by moving the lens within the eyeball. A few sharks accommodate by changing the shape of the lens, as in land vertebrates. Those fishes that are heavily dependent upon the eyes have especially strong muscles for accommodation. Most fishes see well, despite the restrictions imposed by frequent turbidity of the water and by light refraction.
Fossil evidence suggests that colour vision evolved in fishes more than 300 million years ago, but not all living fishes have retained this ability. Experimental evidence indicates that many shallow-water fishes, if not all, have colour vision and see some colours especially well, but some bottom-dwelling shore fishes live in areas where the water is sufficiently deep to filter out most if not all colours, and these fishes apparently never see colours. When tested in shallow water, they apparently are unable to respond to colour differences.
Sound perception and balance are intimately associated senses in a fish. The organs of hearing are entirely internal, located within the skull, on each side of the brain and somewhat behind the eyes. Sound waves, especially those of low frequencies, travel readily through water and impinge directly upon the bones and fluids of the head and body, to be transmitted to the hearing organs. Fishes readily respond to sound; for example, a trout conditioned to escape by the approach of fishermen will take flight upon perceiving footsteps on a stream bank even if it cannot see a fisherman. Compared with humans, however, the range of sound frequencies heard by fishes is greatly restricted. Many fishes communicate with each other by producing sounds in their swim bladders, in their throats by rasping their teeth, and in other ways.
A fish or other vertebrate seldom has to rely on a single type of sensory information to determine the nature of the environment around it. A catfish uses taste and touch when examining a food object with its oral barbels. Like most other animals, fishes have many touch receptors over their body surface. Pain and temperature receptors also are present in fishes and presumably produce the same kind of information to a fish as to humans. Fishes react in a negative fashion to stimuli that would be painful to human beings, suggesting that they feel a sensation of pain.
An important sensory system in fishes that is absent in other vertebrates (except some amphibians) is the lateral line system. This consists of a series of heavily innervated small canals located in the skin and bone around the eyes, along the lower jaw, over the head, and down the mid-side of the body, where it is associated with the scales. Intermittently along these canals are located tiny sensory organs (pit organs) that apparently detect changes in pressure. The system allows a fish to sense changes in water currents and pressure, thereby helping the fish to orient itself to the various changes that occur in the physical environment.
Although a great many fossil fishes have been found and described, they represent a tiny portion of the long and complex evolution of fishes, and knowledge of fish evolution remains relatively fragmentary. In the classification presented in this article, fishlike vertebrates are divided into seven categories, the members of each having a different basic structural organization and different physical and physiological adaptations for the problems presented by the environment. The broad basic pattern has been one of successive replacement of older groups by newer, better-adapted groups. One or a few members of a group evolved a basically more efficient means of feeding, breathing, or swimming or several better ways of living. These better-adapted groups then forced the extinction of members of the older group with which they competed for available food, breeding places, or other necessities of life. As the new fishes became well established, some of them evolved further and adapted to other habitats, where they continued to replace members of the old group already there. The process was repeated until all or almost all members of the old group in a variety of habitats had been replaced by members of the newer evolutionary line.
The earliest vertebrate fossils of certain relationships are fragments of dermal armour of jawless fishes (superclass Agnatha, order Heterostraci) from the Upper Ordovician Period in North America, about 450 million years in age. Early Ordovician toothlike fragments from the former Soviet Union are less certainly remains of agnathans. It is uncertain whether the North American jawless fishes inhabited shallow coastal marine waters, where their remains became fossilized, or were freshwater vertebrates washed into coastal deposits by stream action.
Jawless fishes probably arose from ancient, small, soft-bodied filter-feeding organisms much like and probably also ancestral to the modern sand-dwelling filter feeders, the Cephalochordata (Amphioxus and its relatives). The body in the ancestral animals was probably stiffened by a notochord. Although a vertebrate origin in fresh water is much debated by paleontologists, it is possible that mobility of the body and protection provided by dermal armour arose in response to streamflow in the freshwater environment and to the need to escape from and resist the clawed invertebrate eurypterids that lived in the same waters. Because of the marine distribution of the surviving primitive chordates, however, many paleontologists doubt that the vertebrates arose in fresh water.
Heterostracan remains are next found in what appear to be delta deposits in two North American localities of Silurian age. By the close of the Silurian, about 416 million years ago, European heterostracan remains are found in what appear to be delta or coastal deposits. In the Late Silurian of the Baltic area, lagoon or freshwater deposits yield jawless fishes of the order Osteostraci. Somewhat later in the Silurian from the same region, layers contain fragments of jawed acanthodians, the earliest group of jawed vertebrates, and of jawless fishes. These layers lie between marine beds but appear to be washed out from fresh waters of a coastal region.
It is evident, therefore, that by the end of the Silurian both jawed and jawless vertebrates were well established and already must have had a long history of development. Yet paleontologists have remains only of specialized forms that cannot have been the ancestors of the placoderms and bony fishes that appear in the next period, the Devonian. No fossils are known of the more primitive ancestors of the agnathans and acanthodians. The extensive marine beds of the Silurian and those of the Ordovician are essentially void of vertebrate history. It is believed that the ancestors of fishlike vertebrates evolved in upland fresh waters, where whatever few and relatively small fossil beds were made probably have been long since eroded away. Remains of the earliest vertebrates may never be found.
By the close of the Silurian, all known orders of jawless vertebrates had evolved, except perhaps the modern cyclostomes, which are without the hard parts that ordinarily are preserved as fossils. Cyclostomes were unknown as fossils until 1968, when a lamprey of modern body structure was reported from the Middle Pennsylvanian of Illinois, in deposits more than 300 million years old. Fossil evidence of the four orders of armoured jawless vertebrates is absent from deposits later than the Devonian. Presumably, these vertebrates became extinct at that time, being replaced by the more efficient and probably more aggressive placoderms, acanthodians, selachians (sharks and relatives), and by early bony fishes. Cyclostomes survived probably because early on they evolved from anaspid agnathans and developed a rasping tonguelike structure and a sucking mouth, enabling them to prey on other fishes. With this way of life they apparently had no competition from other fish groups. Cyclostomes, the hagfishes and lampreys, were once thought to be closely related because of the similarity in their suctorial mouths, but it is now understood that the hagfishes, order Myxiniformes, are the most primitive living chordates, and they are classified separately from the lampreys, order Petromyzontiformes.
Early jawless vertebrates probably fed on tiny organisms by filter feeding, as do the larvae of their descendants, the modern lampreys. The gill cavity of the early agnathans was large. It is thought that small organisms taken from the bottom by a nibbling action of the mouth, or more certainly by a sucking action through the mouth, were passed into the gill cavity along with water for breathing. Small organisms then were strained out by the gill apparatus and directed to the food canal. The gill apparatus thus evolved as a feeding, as well as a breathing, structure. The head and gills in the agnathans were protected by a heavy dermal armour; the tail region was free, allowing motion for swimming.
Most important for the evolution of fishes and vertebrates in general was the early appearance of bone, cartilage, and enamel-like substance. These materials became modified in later fishes, enabling them to adapt to many aquatic environments and finally even to land. Other basic organs and tissues of the vertebrates—such as the central nervous system, heart, liver, digestive tract, kidney, and circulatory system— undoubtedly were present in the ancestors of the agnathans. In many ways, bone, both external and internal, was the key to vertebrate evolution.
The next class of fishes to appear was the Acanthodii, containing the earliest known jawed vertebrates, which arose in the Late Silurian, more than 416 million years ago. The acanthodians declined after the Devonian but lasted into the Early Permian, a little less than 280 million years ago. The first complete specimens appear in Lower Devonian freshwater deposits, but later in the Devonian and Permian some members appear to have been marine. Most were small fishes, not more than 75 cm (approximately 30 inches) in length.
We know nothing of the ancestors of the acanthodians. They must have arisen from some jawless vertebrate, probably in fresh water. They appear to have been active swimmers with almost no head armour but with large eyes, indicating that they depended heavily on vision. Perhaps they preyed on invertebrates. The rows of spines and spinelike fins between the pectoral and pelvic fins give some credence to the idea that paired fins arose from “fin folds” along the body sides.
The relationships of the acanthodians to other jawed vertebrates are obscure. They possess features found in both sharks and bony fishes. They are like early bony fishes in possessing ganoidlike scales and a partially ossified internal skeleton. Certain aspects of the jaw appear to be more like those of bony fishes than sharks, but the bony fin spines and certain aspects of the gill apparatus would seem to favour relationships with early sharks. Acanthodians do not seem particularly close to the Placodermi, although, like the placoderms, they apparently possessed less efficient tooth replacement and tooth structure than the sharks and the bony fishes, possibly one reason for their subsequent extinction.