59-592: The Raphina are a clade of extinct flightless birds formerly called didines or didine birds. They inhabited the Mascarene Islands of Mauritius and Rodrigues , but became extinct through hunting by humans and predation by introduced non-native mammals following human colonisation in the 17th century. Historically, many different groups have been named for both the dodo and the Rodrigues solitaire, not all grouping them together. Most recently, it
118-534: A clade (from Ancient Greek κλάδος (kládos) 'branch'), also known as a monophyletic group or natural group , is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree . In the taxonomical literature, sometimes the Latin form cladus (plural cladi ) is used rather than the English form. Clades are
177-479: A "ladder", with supposedly more "advanced" organisms at the top. Taxonomists have increasingly worked to make the taxonomic system reflect evolution. When it comes to naming , this principle is not always compatible with the traditional rank-based nomenclature (in which only taxa associated with a rank can be named) because not enough ranks exist to name a long series of nested clades. For these and other reasons, phylogenetic nomenclature has been developed; it
236-454: A 95% confidence interval of 1688–1715. The authors also pointed out that because the last sighting before 1662 was in 1638, the dodo was probably already quite rare by the 1660s, and thus a disputed report from 1674 by an escaped slave cannot be dismissed out of hand. Although the IUCN lists the Rodrigues solitaire as going extinct by 1778, the species probably became extinct some time between
295-623: A clade can be described based on two different reference points, crown age and stem age. The crown age of a clade refers to the age of the most recent common ancestor of all of the species in the clade. The stem age of a clade refers to the time that the ancestral lineage of the clade diverged from its sister clade. A clade's stem age is either the same as or older than its crown age. Ages of clades cannot be directly observed. They are inferred, either from stratigraphy of fossils , or from molecular clock estimates. Viruses , and particularly RNA viruses form clades. These are useful in tracking
354-412: A family— or whether a described family should be acknowledged— is established and decided upon by active taxonomists . There are not strict regulations for outlining or acknowledging a family, yet in the realm of plants, these classifications often rely on both the vegetative and reproductive characteristics of plant species. Taxonomists frequently hold varying perspectives on these descriptions, leading to
413-554: A lack of widespread consensus within the scientific community for extended periods. The continual publication of new data and diverse opinions plays a crucial role in facilitating adjustments and ultimately reaching a consensus over time. The naming of families is codified by various international bodies using the following suffixes: The taxonomic term familia was first used by French botanist Pierre Magnol in his Prodromus historiae generalis plantarum, in quo familiae plantarum per tabulas disponuntur (1689) where he called
472-548: A more stable climate than Rodrigues, there was probably less need for male dodos to fight over territory. The solitaire was therefore probably the more aggressive of the two. Observations of the Rodrigues solitaire in life indicate that they were highly territorial . They presumably settled disputes by striking each other with their wings; to aid this purpose, they used the knobs on their wrists. Fractures in their wing bones also indicate that they were used in combat. It has also been suggested that these fractures may instead have been
531-440: A problem for the dodo, as they would have been used to dealing with local land crabs . The latest definite sighting of dodos, on Amber Island in 1662, may not necessarily have been the last members of the species. The last claimed sighting of a dodo was reported in the hunting records of Isaac Johannes Lamotius in 1688. Statistical analysis of these records by Roberts and Solow gives a new estimated extinction date of 1693, with
590-496: A result of isolation on islands free of predators , in accordance with Foster's rule . Historically, the dodo was assigned the genus Didus , now a junior synonym of Raphus . In 1848, a new species within the now defunct genus Didus , D. nazarenus , was named by Hugh Edwin Strickland and Alexander Gordon Melville . To house their new species, as well as the other species known at the time, Strickland and Melville named
649-422: A revised taxonomy based on a concept strongly resembling clades, although the term clade itself would not be coined until 1957 by his grandson, Julian Huxley . German biologist Emil Hans Willi Hennig (1913–1976) is considered to be the founder of cladistics . He proposed a classification system that represented repeated branchings of the family tree, as opposed to the previous systems, which put organisms on
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#1732783608391708-429: A suffix added should be e.g. "dracohortian". A clade is by definition monophyletic , meaning that it contains one ancestor which can be an organism, a population, or a species and all its descendants. The ancestor can be known or unknown; any and all members of a clade can be extant or extinct. The science that tries to reconstruct phylogenetic trees and thus discover clades is called phylogenetics or cladistics ,
767-416: A synonym of Raphus cucullatus . Illiger concluded that the dodo was related to ostriches and rheas, and so placed Inepti in the order Rasores , as the sister family to Gallinacei , Epollicati (a defunct group including Turnix and Syrrhaptes ), Columbini , and Crypturi . In 1842, Johannes Theodor Reinhardt proposed they were ground doves , based on studies of a dodo skull he had rediscovered in
826-499: Is also used with a similar meaning in other fields besides biology, such as historical linguistics ; see Cladistics § In disciplines other than biology . The term "clade" was coined in 1957 by the biologist Julian Huxley to refer to the result of cladogenesis , the evolutionary splitting of a parent species into two distinct species, a concept Huxley borrowed from Bernhard Rensch . Many commonly named groups – rodents and insects , for example – are clades because, in each case,
885-431: Is classified between order and genus . A family may be divided into subfamilies , which are intermediate ranks between the ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to the family Juglandaceae , but that family is commonly referred to as the "walnut family". The delineation of what constitutes
944-538: Is considered that the two birds can be classified in Columbidae , often under the subfamily Raphinae. The first person to suggest a close affinity to the doves was Johannes Theodor Reinhardt , whose opinions were then supported by Hugh Edwin Strickland and Alexander Gordon Melville . Recent extractions of DNA from the dodo and Rodrigues solitaire, as well as 37 species of doves, has found where in Columbidae
1003-426: Is hard to determine. Illustrations and written accounts of encounters with the dodo between its discovery and its extinction (1598–1662) are the primary evidence for its external appearance. According to most representations, the dodo had greyish or brownish plumage , with lighter primary feathers and a tuft of curly light feathers high on its rear end. The head was grey and naked, the beak green, black and yellow, and
1062-476: Is in turn included in the mammal, vertebrate and animal clades. The idea of a clade did not exist in pre- Darwinian Linnaean taxonomy , which was based by necessity only on internal or external morphological similarities between organisms. Many of the better known animal groups in Linnaeus's original Systema Naturae (mostly vertebrate groups) do represent clades. The phenomenon of convergent evolution
1121-434: Is only 60% of the weight of a larger individual. Their weight may have varied substantially due to fat cycles, meaning that individuals were fat during cool seasons, but slim during hot seasons, and may have been as low as 21 kg (46 lb) in the larger gender and 13 kg (29 lb) in the smaller. Though male pigeons are usually larger than females, there is no direct evidence for the largest specimens actually being
1180-515: Is responsible for many cases of misleading similarities in the morphology of groups that evolved from different lineages. With the increasing realization in the first half of the 19th century that species had changed and split through the ages, classification increasingly came to be seen as branches on the evolutionary tree of life . The publication of Darwin's theory of evolution in 1859 gave this view increasing weight. In 1876 Thomas Henry Huxley , an early advocate of evolutionary theory, proposed
1239-489: Is still controversial. As an example, see the full current classification of Anas platyrhynchos (the mallard duck) with 40 clades from Eukaryota down by following this Wikispecies link and clicking on "Expand". The name of a clade is conventionally a plural, where the singular refers to each member individually. A unique exception is the reptile clade Dracohors , which was made by haplology from Latin "draco" and "cohors", i.e. "the dragon cohort "; its form with
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#17327836083911298-619: Is still some controversy. It has been suggested that the weight depended on the season, and that individuals were fat during cool seasons, but less so during hot. The bird was sexually dimorphic : males were larger and had proportionally longer beaks. The beak was up to 23 centimetres (9.1 in) in length and had a hooked point. A study of the few remaining feathers on the Oxford specimen head showed that they were pennaceous (vaned feathers with barbs and quills) rather than plumaceous (downy) and most similar to those of other pigeons. The beak of
1357-459: Is thought to have been related to the crowned pigeons. It has been estimated that the group containing the dodo and the Rodrigues solitaire diverged from genera like Goura about 1.5 million years ago. However, that estimate appears highly unlikely. It was estimated that the relatives of the two species moved to the island about 35 million years ago, when a land bridge between Nazareth (Rodrigues) or St. Brandon banks and Mauritius formed. Many of
1416-439: The cantilever strength of its leg bones indicate that it could run quite fast. Unlike the Rodrigues solitaire, there is no evidence that the dodo used its wings in intraspecific combat. Though some dodo bones have been found with healed fractures, it had weak pectoral muscles and more reduced wings in comparison. The dodo may instead have used its large, hooked beak in territorial disputes. Since Mauritius received more rain and had
1475-406: The 1730s and 1760s; the exact date is unknown. Its disappearance coincided with the tortoise trade between 1730 and 1750, when traders burnt off vegetation, hunted solitaires, and released cats and pigs that preyed on eggs and chicks. In 1755, Joseph-François Charpentier de Cossigny attempted to obtain a live specimen, as he had been assured the Rodrigues solitaire still survived in remote areas of
1534-491: The 1750s or 1760s. Both birds became extinct as a consequence of human hunting and the introduction of mammals that ate the birds and their eggs. This clade is part of the order Columbiformes and contains the monotypic genera Pezophaps and Raphus . The former contains the species Pezophaps solitaria (the Rodrigues solitaire ), the latter the dodo , Raphus cucullatus . These birds reached an impressive size as
1593-437: The 17th century, but they introduced other animals, including dogs, pigs, cats, rats, and crab-eating macaques , which plundered dodo nests and competed for the limited food resources. At the same time, humans destroyed the dodo's forest habitat . The impact of these introduced animals, especially the pigs and macaques, on the dodo population is currently considered more severe than that of hunting. Rats would not have caused such
1652-460: The Columbidae is more appropriate. Many different affinities have historically been suggested for the dodo, including that it was a small ostrich , a rail , an albatross , or a vulture . Comparison of mitochondrial cytochrome b and 12S rRNA sequences isolated from a dodo tarsal and a Rodrigues solitaire femur supported their close relationship and their placement within the family of pigeons and doves Columbidae . The genetic evidence
1711-507: The Rodrigues solitaire was described as grey and brown. Females were paler than males and had light-coloured elevations on the lower neck. Sexual size dimorphism in the solitaire is perhaps the greatest in any neognath . One group, probably the males, were considerably larger than the other, measuring 90 cm (35 in) in length and weighing up to 28 kg (62 lb), whereas the smaller group, probably females, were only 70 cm (28 in) and weighed 17 kg (37 lb). This
1770-506: The Rodrigues solitaire, such as features of the skull, pelvis, and sternum, as well as their large size. It differed in other aspects, such as being more robust and shorter than the solitaire, having a larger skull and beak, a rounded skull roof , and smaller orbits . The dodo's neck and legs were proportionally shorter, and it did not possess an equivalent to the knob present on the solitaire's wrists. As no complete dodo specimens exist, its external appearance, such as plumage and colouration,
1829-401: The ancestors of both birds probably remained capable of flight for a considerable time after the separation of their lineage . The lack of mammalian herbivores competing for resources on these islands allowed the solitaire and the dodo to attain very large sizes . The DNA obtained from the Oxford specimen is degraded, and no usable DNA has been extracted from subfossil remains, so the age of
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1888-445: The availability of resources on the island, the solitaire would have more reason to evolve aggressive territorial behaviour. Several accounts state that they also defended themselves with a powerful bite. The last surviving raphine species, the Rodrigues solitaire, probably became extinct before 1778. The dodo survived until 1662 or 1690. The Rodrigues solitaire became extinct because of the introduction of feral cats and heavy hunting by
1947-635: The dissolution of the family Raphidae, and the dodo and solitaire are now placed in their own subfamily, Raphinae, in the family Columbidae. The " Réunion solitaire ", long considered a third extinct didine bird, has turned out to be an ibis ; it is now known as Threskiornis solitarius . The 2002 study indicated that the ancestors of the Rodrigues solitaire and the dodo diverged around the Paleogene - Neogene boundary. The Mascarene Islands ( Mauritius , Réunion , and Rodrigues ), are of volcanic origin and are less than 10 million years old. Therefore,
2006-572: The dodo and Rodrigues solitaire should be placed in the Gourinae subfamily along with the Goura pigeons and others, in agreement with the genetic evidence. In 2014, DNA of the only known specimen of the recently extinct spotted green pigeon ( Caloenas maculata ) was analysed, and it was found to be a close relative of the Nicobar pigeon, and thus also the dodo and Rodrigues solitaire. For many years
2065-457: The dodo and the Rodrigues solitaire were placed in a family of their own, the Raphidae (formerly Dididae), because their exact relationships with other pigeons were unresolved. Each was placed in its own monotypic family (Raphidae and Pezophapidae, respectively), as it was thought that they had evolved their similar features independently. Osteological and molecular data has since led to
2124-405: The extinction of the dodo is 1662, but some possible sightings had been made as late as 1688. The last sighting with a description was in 1662, but a statistical analysis by Roberts and Solow found that the extinction of the dodo was in 1693. The Rodrigues solitaire was killed off later than the dodo. The IUCN uses an extinction date of 1778 for the solitaire, although a more probable date would be in
2183-540: The family as a rank intermediate between order and genus was introduced by Pierre André Latreille in his Précis des caractères génériques des insectes, disposés dans un ordre naturel (1796). He used families (some of them were not named) in some but not in all his orders of "insects" (which then included all arthropods ). In nineteenth-century works such as the Prodromus of Augustin Pyramus de Candolle and
2242-451: The fundamental unit of cladistics , a modern approach to taxonomy adopted by most biological fields. The common ancestor may be an individual, a population , or a species ( extinct or extant ). Clades are nested, one in another, as each branch in turn splits into smaller branches. These splits reflect evolutionary history as populations diverged and evolved independently. Clades are termed monophyletic (Greek: "one clan") groups. Over
2301-546: The group consists of a common ancestor with all its descendant branches. Rodents, for example, are a branch of mammals that split off after the end of the period when the clade Dinosauria stopped being the dominant terrestrial vertebrates 66 million years ago. The original population and all its descendants are a clade. The rodent clade corresponds to the order Rodentia, and insects to the class Insecta. These clades include smaller clades, such as chipmunk or ant , each of which consists of even smaller clades. The clade "rodent"
2360-458: The groups divergence from other pigeons still needs to be independently verified. The dodo lost the ability to fly owing to the lack of mammalian predators on Mauritius. Another large, flightless pigeon, the Viti Levu giant pigeon ( Natunaornis gigoura ), was described in 2001 from subfossil material from Fiji . It was only slightly smaller than the dodo and the Rodrigues solitaire, and it too
2419-463: The human population. Although the dodo became extinct earlier, the causes of extinction are related and involve the introduction of animals and hunting. Like many animals that evolved in isolation from significant predators, the dodo was entirely fearless of humans. This fearlessness and its inability to fly made the dodo easy prey for sailors. The human population on Mauritius (an area of 1,860 km or 720 sq mi) never exceeded 50 people in
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2478-525: The island. After searching for 18 months and offering large rewards, he could find none. He noted that cats were blamed for decimating the species, but suspected that it was really due to hunting by humans. When he visited Rodrigues to observe the 1761 transit of Venus , Alexandre Guy Pingré encountered no solitaires, although he had been assured that they survived. Explanatory notes Citations [REDACTED] Media related to Raphinae at Wikimedia Commons Clade In biological phylogenetics ,
2537-590: The last few decades, the cladistic approach has revolutionized biological classification and revealed surprising evolutionary relationships among organisms. Increasingly, taxonomists try to avoid naming taxa that are not clades; that is, taxa that are not monophyletic . Some of the relationships between organisms that the molecular biology arm of cladistics has revealed include that fungi are closer relatives to animals than they are to plants, archaea are now considered different from bacteria , and multicellular organisms may have evolved from archaea. The term "clade"
2596-518: The latter term coined by Ernst Mayr (1965), derived from "clade". The results of phylogenetic/cladistic analyses are tree-shaped diagrams called cladograms ; they, and all their branches, are phylogenetic hypotheses. Three methods of defining clades are featured in phylogenetic nomenclature : node-, stem-, and apomorphy-based (see Phylogenetic nomenclature§Phylogenetic definitions of clade names for detailed definitions). The relationship between clades can be described in several ways: The age of
2655-474: The leg bones, features which were otherwise known only in pigeons. The raphines are sometimes separated as a distinct family Raphidae, and their affinities were for long uncertain. They were initially placed in the ratites due to their peculiar, flightlessness-related apomorphies , and a relationship to the Rallidae has also been suggested. Osteological and molecular data, however, agrees that placement in
2714-535: The legs were stout and yellowish, with black claws. Subfossil remains and remnants of the birds that were brought to Europe in the 17th century show that they were very large birds, 1 metre (3.3 feet) tall, and possibly weighing up to 23 kilograms (51 lb). The higher weights have been attributed to birds in captivity; weights in the wild were estimated to have been in the range 10.6–21.1 kg (23–47 lb). A later estimate gives an average weight as low as 10.2 kg (22 lb). This has been questioned, and there
2773-423: The males of the species, and this has only been assumed based on early works. Though the male was probably largest, this can only be confirmed by molecular sexing techniques, and not skeletal morphology alone. Little is known of the behaviour of the dodo, as most contemporary descriptions are very brief. Based on weight estimates, it has been suggested the male could reach the age of 21, and the female 17. Studies of
2832-487: The position of the dodo and solitaire within the pigeon and dove family. Goura victoria Caloenas nicobarica † Pezophaps solitaria † Raphus cucullatus Didunculus strigirostris A similar cladogram was published in 2007, inverting the placement of Goura and Didunculus and including the pheasant pigeon and the thick-billed ground pigeon at the base of the clade. Based on behavioural and morphological evidence, Jolyon C. Parish proposed that
2891-486: The raphines should be placed. Raphines are not the most primitive columbid, instead they are grouped with the Nicobar pigeon as their closest relative, with other closely related birds being the crowned pigeons and tooth-billed pigeon . A third raphine, Raphus solitarius , is now considered to be an ibis in the genus Threskiornis . Both the Rodrigues solitaire and the dodo are now extinct. A common threshold of
2950-451: The result of a hereditary bone disease rather than battle-injuries. But in all extant birds where carpal spurs and knobs are present, these are used as weapons without exceptions. Though some dodo bones have been found with healed fractures, it had weak pectoral muscles and more reduced wings in comparison with the Rodrigues solitaire. Since Rodrigues receives less rainfall and has more seasonal variation than Mauritius, which would have affected
3009-432: The royal Danish collection of Copenhagen . This view was met with ridicule, but later supported by Strickland and Melville, who suggested the common descent of the Rodrigues solitaire and the dodo in 1848, after dissecting the only known dodo specimen with soft tissue and comparing it with the few solitaire remains then available. Strickland stated that although not identical, these birds shared many distinguishing features in
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#17327836083913068-575: The seventy-six groups of plants he recognised in his tables families ( familiae ). The concept of rank at that time was not yet settled, and in the preface to the Prodromus Magnol spoke of uniting his families into larger genera , which is far from how the term is used today. In his work Philosophia Botanica published in 1751, Carl Linnaeus employed the term familia to categorize significant plant groups such as trees , herbs , ferns , palms , and so on. Notably, he restricted
3127-535: The skeletal features that distinguish the dodo and the Rodrigues solitaire, its closest relative, from pigeons have been attributed to their flightlessness. The pelvic elements were thicker than those of flighted pigeons to support the higher weight, and the pectoral region and the small wings were paedomorphic , meaning that they were underdeveloped and retained juvenile features. The skull, trunk and pelvic limbs were peramorphic , meaning that they changed considerably with age. The dodo shared several other traits with
3186-419: The solitaire was slightly hooked, and its neck and legs were long. One observer described it as the size of a swan. The skull was 170 millimetres (6.7 in) long, flattened at the top with the fore and hind parts elevated into two bony ridges structured with cancellous . A black band (a contemporary description described it as a "frontlet") appeared on its head just behind the base of the beak. The plumage of
3245-591: The spread of viral infections . HIV , for example, has clades called subtypes, which vary in geographical prevalence. HIV subtype (clade) B, for example is predominant in Europe, the Americas and Japan, whereas subtype A is more common in east Africa. Family (biology) Family ( Latin : familia , pl. : familiae ) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy . It
3304-487: The subfamily Didinae. In 1893 three species were assigned to the group Pezophaps solitarius , Didus ineptus , and the possible species Didus borbonicus . Today, only two raphine species are known, with Didus ineptus becoming a junior subjective synonym of Raphus cucullatus ; Didus? borbonicus now classified as the ibis Threskiornis solitarius ; and Didus nazarenus being identified as synonymous with Pezophaps solitarius . A suborder named in 1893 by Sharpe, Didi
3363-549: The use of this term solely within the book's morphological section, where he delved into discussions regarding the vegetative and generative aspects of plants. Subsequently, in French botanical publications, from Michel Adanson 's Familles naturelles des plantes (1763) and until the end of the 19th century, the word famille was used as a French equivalent of the Latin ordo (or ordo naturalis ). In zoology ,
3422-432: Was defined as a group including only the massive birds, that were sister to Columbidae , from the Mascarene Islands of Mauritius , Réunion , and Rodrigues . Features grouping Didi with Columbidae were the angle of the mandible and the hook at the end of the beak. In 1811, Johann Karl Wilhelm Illiger created a new family for the genus Didus . He named the family Inepti, and in it included only Didus ineptus , now
3481-556: Was interpreted as suggesting the Southeast Asian Nicobar pigeon ( Caloenas nicobarica ) to be their closest living relative among 35 analyzed species of pigeons and doves, followed by the crowned pigeons (genus Goura ) of New Guinea and the superficially dodo-like tooth-billed pigeon from Samoa . The genus of the latter is Didunculus ("little dodo"), and it was called "dodlet" by Richard Owen . The following cladogram , from Shapiro and colleagues (2002), shows
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