Ornithology (from Greek: ?????, ???????, ornis, ornithos, "bird"; and ?????, logos, "knowledge") is the branch of zoology concerned with the study of birds. Several aspects of the study of ornithology differ from closely related disciplines, due partly to the high visibility and the aesthetic appeal of birds.^[1] Most marked among these is the extent of field studies undertaken by amateur volunteers working within the parameters of strict scientific methodology.

The science of ornithology has a long history and studies on birds aided the introduction and refinement of key concepts in evolution, behaviour and ecology such as those of species, speciation, instinct, learning, ecological niches, guilds, island biogeography, phylogeography and conservation.^[2] While early ornithology was principally concerned with descriptions and distributions of species, ornithologists today concern themselves with answering specific questions, often using birds as models to test hypotheses and predictions based on theory. A wide range of tools and techniques are used in ornithology and innovations are constantly made.^[3]

History

The history of ornithology reflects trends in the the history of biology. Historic trends include the move from descriptions to the identification of patterns and the search for processes leading to the emergence of these patterns.

Early studies

Belon's comparison of birds and humans in Book of Birds, 1555

Cultures around the world have rich vocabularies related to birds.^[5] Traditional bird names are often based on detailed knowledge of the behaviour, with many names being onomatopoeic, many still in use.^[6] Traditional knowledge continues to be of importance especially due to their relevance in conservation.^[7] Most of this information is passed on through oral traditions (see ethno-ornithology).^[8][9] Hunting of birds would also have required considerable knowledge of their habits. Poultry farming and falconry were practised from early times in many parts of the world. Artificial incubation of poultry was practised in China (246 BC) and Egypt (at least 400 BC).^[10] The Egyptians also showed a great deal of knowledge of birds through their use of bird symbols in hieroglyphs, many of which, though stylized, are still recognizable.

Cover of Ulisse Aldrovandi's Ornithology, 1599

Aristotle in 350 BC in his Historia Animalium^[12] noted the habit of bird migration, moulting, egg laying and life spans. He also introduced several incorrect concepts such as the idea that swallows hibernated in winter. He however noted that cranes travelled from the steppes of Scythia to the marshes at the headwaters of the Nile. The idea of swallow hibernation became so well established that, even as late as in 1878, Elliott Coues listed as many as 182 contemporary publications dealings with the hibernation of swallows and no evidence to contradict the theory.^[13][14] Similar misconceptions existed regarding the breeding of Barnacle geese. Their nests had not been seen and it was believed that they grew by transformations of goose barnacles, an idea that became prevalent from around the 11th century and noted by Bishop Giraldus Cambrensis (Gerald of Wales) in Topographia Hiberniae (1187).^[15]

The origins of falconry have been traced to Mesopotamia and the earliest record comes from the reign of Sargon II (722?705 BC). Falconry made its entry to Europe only after AD 400, brought in from the East after invasions by the Huns and Allans. Frederick II, Holy Roman Emperor of Hohenstaufen (1194 ? 1250) learnt about Arabian falconry during wars in the region and obtained an Arabic treatise on falconry by Moamyn. He had this work translated into Latin and also conducted experiments on birds in his menagerie. By sealing the eyes of vultures and placing food nearby, he concluded that they found food by sight, and not by smell. He also developed methods to keep and train falcons. The studies that he undertook over nearly 30 years, were published in 1240 as De Arte Venandi cum Avibus (The Art of Hunting with Birds), considered one of the earliest studies on bird behaviour.^[16]

Several early German and French scholars compiled old works and conducted new research on birds. These included Guillaume Rondelet who described his observations in the Mediterranean and Pierre Belon who described the fish and birds that he had seen in France and the Levant. Belon's Book of Birds (1555) is a folio volume with descriptions of some two hundred species. His comparison of the skeleton of humans and birds is considered as a landmark in comparative anatomy.^[17] Volcher Coiter (1534-1576), a Dutch anatomist made detailed studies of the internal structures of birds and produced a classification of birds, De Diferentiis Avium (around 1572), that was based on structure and habits.^[18] Konrad Gesner wrote the Vogelbuch and Icones avium omnium around 1557. Like Gesner, Ulisse Aldrovandi, an encyclopedic naturalist began a 14-volume natural history with three volumes on birds, entitled ornithologiae hoc est de avibus historiae libri XII which was published from 1599 to 1603. Aldrovandi showed great interest in plants and animals and his work included 3000 drawings of fruits, flowers, plants and animals, published in 363 volumes. His Ornithology alone covers 2000 pages and included such aspects as the chicken and poultry techniques.^[19][20]William Turner's Historia Avium ("History of Birds"), published at Cologne in 1544, was another early ornithological work. He notes that the kite in cities of England would snatch the meat out of the hands of children. In his day the Osprey was well known but disliked for it was believed to empty their fishponds; anglers used to mix their bait with its fat. Turner's work was written in a tone that reflected violent times; quite unlike Gilbert White's The Natural History and Antiquities of Selborne which was written in a more tranquil era.^[17][21] In the 17th century Francis Willughby (1635-1672) and John Ray (1627-1705) came up with the first major system of bird classification that was based on function and morphology rather than on form or behavior. Willughby's Ornithologiae libri tres (1676) completed by John Ray is sometimes considered to mark the beginning of scientific ornithology. Ray also worked on Ornithologia which was published posthumously in 1713 as Synopsis methodica avium et piscium.^[22] The earliest list of British birds, Pinax Rerum Naturalium Britannicarum was written by Christopher Merrett in 1667, however it was not considered of value by many including John Ray.^[23]

An Experiment on a Bird in the Air Pump, 1768

Scientific studies

Early bird study focused on collectibles such as eggs and nests

The bird collectors of the Victorian era observed the variations in bird forms and habits across geographic regions, noting local specialization and variation in widespread species. The collections of museums and private collectors grew with contributions from various parts of the world. The naming of species with binomials and the organization of birds into groups based on their similarities became the main work of museum specialists. The variations in widespread birds across geographical region caused the introduction of trinomial names.

The search for patterns in the variations of birds was attempted by many. Early ornithologists like William Swainson followed the Quinarian system and this was replaced by more complex "maps" of affinities in works by Hugh Edwin Strickland and Alfred Russell Wallace.^[28]

The Galapagos finches were especially influential in the development of Charles Darwin's theory of evolution. His contemporary Alfred Russel Wallace also noted these variations and the geographical separations between different forms leading to the study of biogeography. Wallace was influenced by the work of Philip Lutley Sclater on the distribution patterns of birds.^[29]

Quinarian system of bird classification by Swainson

Early ornithologists were preoccupied with matters of species identification. In 1901 Robert Ridgway wrote in the introduction to The Birds of North and Middle America that: This early idea that the study of living birds was merely recreation held sway until ecological theories became the predominant focus of ornithological studies.^[2][27] The study of birds in their habitats was particularly advanced in Germany with bird ringing stations established as early as 1903. By the 1920s the Journal für Ornithologie included many papers on the behavior, ecology, anatomy and physiology, many written by Erwin Stresemann. Ornithology in the United States continued to dominated by studies of morphological variations, species identities and geographic distributions, until it was influenced by Stresemann's student Ernst Mayr.^[31] In Britain, some of the earliest ornithological works that used the word ecology appeared in 1915.^[32] The Ibis however resisted the introduction of these new methods of study and it was not until 1943 that any paper on ecology appeared.^[27] The work of David Lack on population ecology was pioneering. Newer quantitative approaches were introduced for the study of ecology and behavior and this was not readily accepted. For instance, Claud Ticehurst wrote: David Lack's studies on population ecology sought to find the processes involved in the regulation of population based on the evolution of optimal clutch sizes. He concluded that population was regulated primarily by density-dependent controls, and also suggested that natural selection produces life-history traits that maximize the fitness of individuals. Others like Wynne-Edwards interpreted population regulation as a mechanism that aided the "species" rather than individuals. This led to widespread and sometimes bitter debate on what constituted the "unit of selection".^[30] Lack also pioneered the use of many new tools for ornithological research including the idea of using radar to study bird migration.

Birds were also widely used in studies of the niche hypothesis and Georgii Gause's competitive exclusion principle. Work on resource partitioning and the structuring of bird communities through competition were made by Robert MacArthur. Patterns of biodiversity also became a topic interest. Work on the relationship of the number of species to area and its application in the study of Island biogeography was pioneered by E. O. Wilson and Robert MacArthur.^[30] These studies led to the development of the discipline of landscape ecology.

A mounted specimen of a Red-footed Falcon.

The study of imprinting behaviour in ducks and geese by Konrad Lorenz and the studies of instinct in Herring Gulls by Nicolaas Tinbergen, led to the establishment of the field of ethology. The study of learning became particularly of interest and the study of bird song has been a model for studies in neuro-ethology. The role of hormones and physiology in the control of behaviour has also been aided by bird models. These have helped in the study of circadian and seasonal cycles. Studies on migration have attempted to answer questions on the evolution of migration, orientation and navigation.^[30]

The growth of genetics and the rise of molecular biology led to the application of the gene-centered view of evolution to explain avian phenomena. Studies on kinship and altruism, such as helpers, became of particular interest. The idea of inclusive fitness was used to interpret observations on behaviour and life-history and birds were widely used models for testing hypotheses based on theories postulated by W. D. Hamilton and others.^[30]

The new tools of molecular-biology changed the study of bird systematics. Systematics changed from being based on phenotype to the underlying genotype. The use of techniques such as DNA-DNA hybridization to study evolutionary relationships was pioneered by Charles Sibley and Jon Edward Ahlquist resulting in what is called the Sibley-Ahlquist taxonomy. These early techniques have been replaced by newer techniques based on mitochondrial DNA sequences and molecular phylogenetics approaches that make use of computational procedures for sequence alignment, construction of phylogenetic trees and calibration of molecular clocks to infer evolutionary relationships.^[36][37] Molecular techniques are also widely used in studies of avian population biology and ecology.^[38]

Rise to popularity

The use of field glasses or telescopes for bird observation began in the 1820s and 1830s with pioneers like J. Dovaston (who also a pioneered in the use of bird-feeders), but it was not until the 1880s that instruction manuals began to insist on the use of optical aids such as "a first-class telescope or field glass."^[39][40]

Page from an early field guide by Florence Augusta Merriam Bailey

The interest in birdwatching grew in popularity in many parts of the world and it was realized that there was a possibility for amateurs to contribute to the professional biology. As early as 1916, Julian Huxley wrote a two part article in the Auk, noting the tensions between amateurs and professionals and suggesting the possibility that the "vast army of bird-lovers and bird-watchers could begin providing the data scientists needed to address the fundamental problems of biology."^[42][43]

Organizations were started in many countries and these grew rapdily in membership, most notable among them being the Royal Society for the Protection of Birds (RSPB) in Britain and the Audubon Society in the US. The Audubon Society started in 1885. Both these organization were started with the primary objective of conservation. The RSPB, born in 1889, grew from a small group of women in Croydon who met regularly and called themselves the Fur, Fin and Feather Folk and who took a pledge "to refrain from wearing the feathers of any birds not killed for the purpose of food, the Ostrich only exempted." The organization did not allow men as members initially, avenging a policy of the British Ornithologists' Union to keep out women.^[25] Unlike the RSPB, which was primarily conservation oriented, the British Trust for Ornithology (BTO) was started in 1933 with the aim of advancing ornithological research. Members were often involved in collaborative ornithological projects. These projects have resulted in atlases which detail the distribution of bird species across Britain.^[44] In the United States, the Breeding Bird Surveys, conducted by the US Geological Survey have also produced atlases with information on breeding densities and changes in the density and distribution over time. Other volunteer collaborative ornithology projects were subsequently established in other parts of the world.^[45]

Techniques

The tools and techniques of ornithology are varied and new inventions and approaches are quickly incorporated. The techniques may be broadly dealt under the categories of those that are applicable to specimens and those that are used in the field, however the classification is imperfect as many of the newer non-destructive sampling and analysis techniques are applicable in both the laboratory and field.

Collections

Bird preservation techniques

Morphometric measurements of birds are important in systematics

Other methods of preservation include the storage of specimens in spirit. Such wet-specimens have special value in physiological and anatomical study, apart from providing better quality of DNA for molecular studies.^[49] Freeze drying of specimens has also been attempted in more recent times. While the technique has advantages in that it preserves stomach contents and anatomy, it may have the same problems as dry skins in that shrinkage can occur leading to errors in morphometrics.^{[50] [51]}

In the field

The study of birds in the field was helped enormously by improvements in optics. Photography made it possible to document birds in the field with great accuracy. High power spotting scopes today allow observers to detect minute morphological differences that were earlier possible only by examination of the specimen in the hand.^[52]

A bird caught in a mist net

A Californian Condor marked with wing tags

Captured birds are often marked for future recognition. Rings or bands provide long-lasting identification but require capture for the information on them to be read. Field identifiable marks such as coloured bands, wing tags or dyes enable short-term studies where individual identification is required. Mark and recapture techniques make demographic studies possible. Ringing has traditionally been used in the study of migration. In recent times satellite transmitters provide the ability to track migrating birds in near real-time.^[56]

Techniques for estimating population density include point counts, transects and territory mapping. Observations are made in the field using carefully designed protocols and the data may be analysed to estimate bird diversity, relative abundance or absolute population densities.^[57]

In the laboratory

An Emlen funnel is used to study the orientation behaviour in migratory birds

Studies in bird behaviour include the use of tamed and trained birds in captivity. Studies on bird intelligence and song learning have been largely laboratory based. Field researchers may make use of a wide range of techniques such as the use of dummy owls to elicit mobbing behaviour, dummy males or the use of call playback to elicit territorial behaviour and thereby to establish the boundaries of bird territories.^[60]

Studies of bird migration including aspects of navigation, orientation and physiology are often studied using captive birds in special cages that record their activities. The Emlen funnel for instance makes use of a cage with an inkpad at the centre and a conical floor where the ink marks can be counted to identify the diretion in which the bird attempts to fly. The funnel can have a transparent top and visible cues such as the direction of sunlight may be controlled using mirrors or the positions of the stars simulated in a planetarium.^[61]

Collaborative studies

Summer distribution and abundance of Canada Goose using data from the North American Breeding Bird Surveys 1994-2003

Applications

Wild birds impact many human activities while domesticated birds are important sources of eggs, meat, feathers and other products. Applied and economic ornithology aim to reduce the ill effects of problem birds and enhance gains from beneficial species.

Red-billed Quelea are a major agricultural pest in parts of Africa.

Large flocks of pigeons and starlings in cities are often considered as a nuisance and techniques to reduce their populations or their impacts are constantly innovated.^[74][75] Birds are also of medical importance and their role as carriers of human diseases such as Japanese Encephalitis, West Nile Virus and H5N1 have been widely recognised.^[76][77] Bird strikes and the damage they cause in aviation are of particularly great importance, due to the fatal consequences and the level of economic losses caused. It has been estimated that the airline industry incurs worldwide damages of US $ 1.2 billion each year.^[78]

Many species of birds have been driven to extinction by human activities. Bird conservation requires specialized knowledge in aspects of biology, ecology and may require the use of very location specific approaches. Ornithologists contribute to conservation biology by studying the ecology of birds in the wild and identifying the key threats and ways of enhancing the survival of species.^[79] Critically endangered species such as the California Condor have had to be captured and bred in captivity. Such ex-situ conservation measures may be followed by re-introduction of the species into the wild.^[80]

See also

• Avian ecology field methods
• Avian range expansion
• Bird observatory
• List of famous ornithologists
• List of ornithological societies
• List of ornithology journals

References

External links

• Ornithologie (1773-1792) Francois Nicholas Martinet Digital Edition Smithsonian Digital Libraries
• List of oldest ornithological organisations in the world
• History of ornithology in North America
• History of ornithology in China
• Hill ornithology collections
• History of ornithology (in Italian)

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