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60-584: See text Palaeoloxodon is an extinct genus of elephant . The genus originated in Africa during the Early Pleistocene , and expanded into Eurasia at the beginning of the Middle Pleistocene . The genus contains the largest known species of elephants, over 4 metres (13 ft) tall at the shoulders and over 13 tonnes (29,000 lb) in weight, representing among the largest land mammals ever, including

120-557: A species : see Botanical name and Specific name (zoology) . The rules for the scientific names of organisms are laid down in the nomenclature codes , which allow each species a single unique name that, for animals (including protists ), plants (also including algae and fungi ) and prokaryotes ( bacteria and archaea ), is Latin and binomial in form; this contrasts with common or vernacular names , which are non-standardized, can be non-unique, and typically also vary by country and language of usage. Except for viruses ,

180-564: A species name , in parentheses, placed between the generic name and the specific epithet : e.g. the tiger cowry of the Indo-Pacific, Cypraea ( Cypraea ) tigris Linnaeus , which belongs to the subgenus Cypraea of the genus Cypraea . However, it is not mandatory, or even customary, when giving the name of a species, to include the subgeneric name. In the International Code of Nomenclature for algae, fungi, and plants (ICNafp),

240-495: A straight-tusked elephant were collected from travertine deposits near Burgtonna in what is now Thuringia , Germany. While these remains were originally declared by the Collegium Medicum in the nearby city of Gotha to be purely mineral in nature, Wilhelm Ernst Tentzel, a polymath in the employ of the ducal court of Saxe-Gotha-Altenburg , correctly recognised that they represented the remains of an elephant. Prior to 1845,

300-643: A later homonym of a validly published name is a nomen illegitimum or nom. illeg. ; for a full list refer to the International Code of Nomenclature for algae, fungi, and plants and the work cited above by Hawksworth, 2010. In place of the "valid taxon" in zoology, the nearest equivalent in botany is " correct name " or "current name" which can, again, differ or change with alternative taxonomic treatments or new information that results in previously accepted genera being combined or split. Prokaryote and virus codes of nomenclature also exist which serve as

360-446: A lesser extent mammoths (~6%) The ancestry from L. cyclotis was more closely related to modern West African populations of the forest elephant than to other forest elephant populations, while the mammoth ancestry was basal to the split between woolly and Columbian mammoths , probably from shortly after the split between the ancestors of mammoths and Asian elephants. The hybridisation probably took place in Africa, where Palaeoloxodon

420-621: A long time and redescribed as new by a range of subsequent workers, or if a range of genera previously considered separate taxa have subsequently been consolidated into one. For example, the World Register of Marine Species presently lists 8 genus-level synonyms for the sperm whale genus Physeter Linnaeus, 1758, and 13 for the bivalve genus Pecten O.F. Müller, 1776. Within the same kingdom, one generic name can apply to one genus only. However, many names have been assigned (usually unintentionally) to two or more different genera. For example,

480-409: A reference for designating currently accepted genus names as opposed to others which may be either reduced to synonymy, or, in the case of prokaryotes, relegated to a status of "names without standing in prokaryotic nomenclature". An available (zoological) or validly published (botanical) name that has been historically applied to a genus but is not regarded as the accepted (current/valid) name for

540-630: A substantial number of lamellae (up to 19 to 21 on the third molar), though the lamellae frequency is distinctly lower than that reached by advanced mammoth species. The morphology of the teeth varies little between non-dwarf Eurasian Palaeoloxodon species, meaning that they generally cannot be distinguished based on tooth morphology alone. Species of Palaeoloxodon varied widely in size. Fully grown bulls of Palaeoloxodon recki , Palaeoloxodon antiquus , Palaeoloxodon namadicus and Chinese Palaeoloxodon grew substantially larger than living elephants, reaching or exceeding 4 metres (13 ft) tall at

600-427: A taxon; however, the names published in suppressed works are made unavailable via the relevant Opinion dealing with the work in question. In botany, similar concepts exist but with different labels. The botanical equivalent of zoology's "available name" is a validly published name . An invalidly published name is a nomen invalidum or nom. inval. ; a rejected name is a nomen rejiciendum or nom. rej. ;

660-455: A total of c. 520,000 published names (including synonyms) as at end 2019, increasing at some 2,500 published generic names per year. "Official" registers of taxon names at all ranks, including genera, exist for a few groups only such as viruses and prokaryotes, while for others there are compendia with no "official" standing such as Index Fungorum for fungi, Index Nominum Algarum and AlgaeBase for algae, Index Nominum Genericorum and

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720-462: A valid genus or a subgenus of Elephas. Cladistic analyses finding Elephas and Palaeoloxodon to not be each other's closest relatives led to the placement of Palaeoloxodon species within Elephas to be questioned by other authors. By the 2010s Palaeoloxodon was widely regarded as a valid genus separate from Elephas . Historically, Palaeoloxodon was usually considered to be closely related to

780-536: Is discouraged by both the International Code of Zoological Nomenclature and the International Code of Nomenclature for algae, fungi, and plants , there are some five thousand such names in use in more than one kingdom. For instance, A list of generic homonyms (with their authorities), including both available (validly published) and selected unavailable names, has been compiled by the Interim Register of Marine and Nonmarine Genera (IRMNG). The type genus forms

840-470: Is proportionally short and tall, with the premaxillary bones containing the tusks being flared outwards. The tusks have relatively little curvature, and are proportionally large, and somewhat twisted, with the tusk alveoli (sockets) being divergent from each other at least in Pleistocene species. These tusks could reach 4 metres (13 ft) in length, and probably over 190 kilograms (420 lb) in weight in

900-460: Is somewhat arbitrary. Although all species within a genus are supposed to be "similar", there are no objective criteria for grouping species into genera. There is much debate among zoologists about whether enormous, species-rich genera should be maintained, as it is extremely difficult to come up with identification keys or even character sets that distinguish all species. Hence, many taxonomists argue in favor of breaking down large genera. For instance,

960-497: Is suggested to have been a variable mixed feeder that consumed a considerable amount of browse . Palaeoloxodon first unambiguously appears in the fossil record in Africa during the Early Pleistocene , around 1.8 million years ago as the species Palaeoloxodon recki ileretensis (it is contested whether earlier " E. recki " subspecies are related to Palaeoloxodon ). P. recki was the dominant elephant in East Africa for most of

1020-474: Is the type species , and the generic name is permanently associated with the type specimen of its type species. Should the specimen turn out to be assignable to another genus, the generic name linked to it becomes a junior synonym and the remaining taxa in the former genus need to be reassessed. In zoological usage, taxonomic names, including those of genera, are classified as "available" or "unavailable". Available names are those published in accordance with

1080-552: Is uncertain, but claims of a Holocene survival are not substantiated for either region. The youngest dates for the Sicilian dwarf elephant P. cf . mnaidriensis are around 32-20,000 years ago, while those of Cyprus dwarf elephant P. cypriotes are around 12,000 years ago. P. tiliensis from the Greek island of Tilos was suggested to have survived as recently as 3,500 years Before Present based on preliminary radiocarbon dating done in

1140-431: Is unclear as to whether this is the result of multiple hybridisation events, or whether multiple mitochondrial lineages were introgressed in a single event. It has been found that mitochondrial genome of Chinese Palaeoloxodon specimens clustered with a P. antiquus individual from western Europe, which belonged to a separate clade than other sampled European P. antiquus specimens. The relatively low divergence between

1200-621: The International Code of Zoological Nomenclature ; the earliest such name for any taxon (for example, a genus) should then be selected as the " valid " (i.e., current or accepted) name for the taxon in question. Consequently, there will be more available names than valid names at any point in time; which names are currently in use depending on the judgement of taxonomists in either combining taxa described under multiple names, or splitting taxa which may bring available names previously treated as synonyms back into use. "Unavailable" names in zoology comprise names that either were not published according to

1260-799: The International Plant Names Index for plants in general, and ferns through angiosperms, respectively, and Nomenclator Zoologicus and the Index to Organism Names for zoological names. Totals for both "all names" and estimates for "accepted names" as held in the Interim Register of Marine and Nonmarine Genera (IRMNG) are broken down further in the publication by Rees et al., 2020 cited above. The accepted names estimates are as follows, broken down by kingdom: The cited ranges of uncertainty arise because IRMNG lists "uncertain" names (not researched therein) in addition to known "accepted" names;

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1320-749: The paraceratheres . However, this estimate is highly speculative and the author suggested that it should be "taken with a grain of salt ". In contrast, some of the island dwarf species are the smallest elephants known. The smallest species, P. cypriotes and P. falconeri , only reached 1 metre (3.3 ft) tall as fully grown adults, with fully grown adult bulls of P. falconeri having an estimated body mass of only 250 kg (550 lb). Species of Palaeoloxodon are thought to have similar social behaviour to modern elephants, with herds of adult females and juveniles, as well as solitary adult males. The African species of Palaeoloxodon , as well as P. namadicus are suggested to have been grazers, while P. antiquus

1380-404: The platypus belongs to the genus Ornithorhynchus although George Shaw named it Platypus in 1799 (these two names are thus synonyms ) . However, the name Platypus had already been given to a group of ambrosia beetles by Johann Friedrich Wilhelm Herbst in 1793. A name that means two different things is a homonym . Since beetles and platypuses are both members of the kingdom Animalia,

1440-695: The 1970s, which would make it the youngest surviving elephant in Europe, but this has not been thoroughly investigated. Evidence of interaction with Palaeoloxodon by archaic humans extends back over 1 million years ago in Africa, with a number sites with Palaeoloxodon recki in Africa showing evidence of butchery. There is extensive evidence for butchery and to a lesser extent hunting of the European straight-tusked elephant by archaic humans like Homo heidelbergensis and Neanderthals . Evidence has been found for butchery of Palaeoloxodon turkmenicus by archaic humans in

1500-587: The African Palaeoloxodon recki , the European straight-tusked elephant ( Palaeoloxodon antiquus ) and the South Asian Palaeoloxodon namadicus . P. namadicus has been suggested to be the largest known land mammal by some authors based on extrapolation from fragmentary remains, though these estimates are highly speculative. In contrast, the genus also contains many species of dwarf elephants that evolved via insular dwarfism on islands in

1560-469: The French botanist Joseph Pitton de Tournefort (1656–1708) is considered "the founder of the modern concept of genera". The scientific name (or the scientific epithet) of a genus is also called the generic name ; in modern style guides and science, it is always capitalised. It plays a fundamental role in binomial nomenclature , the system of naming organisms , where it is combined with the scientific name of

1620-769: The Indian subcontinent. Based on the association of their remains with stone artefacts, it has been suggested modern humans encountered and butchered the Japanese P. naumanni and the Cyprus dwarf elephant P. cypriotes during the Last Glacial Period . [REDACTED] Genus The composition of a genus is determined by taxonomists . The standards for genus classification are not strictly codified, so different authorities often produce different classifications for genera. There are some general practices used, however, including

1680-671: The Mediterranean, some like Palaeoloxodon falconeri less than 1 metre (3.3 ft) in shoulder height as fully grown adults, making them the smallest elephants known. The genus has a long and complex taxonomic history, and at various times, it has been considered to belong to Loxodonta or Elephas , but today is usually considered a valid and separate genus in its own right. Remains of Palaeoloxodon species have probably been noted since ancient times where their remains like those of other fossil proboscideans were interpreted as those of giants or other mythical beings. In 1695, remains of

1740-534: The Pleistocene. A population of P. recki migrated out of Africa at the beginning of the Middle Pleistocene around 800,000 years ago, diversifying into the radiation of Eurasian Palaeoloxodon species, including P. antiquus, and P. namadicus. The precise relationships of the Eurasian taxa to each other are obscure. The arrival of P. antiquus in Europe co-incides with the extinction of the temperate adapted mammoth species Mammuthus meridionalis (sometimes called

1800-442: The base for higher taxonomic ranks, such as the family name Canidae ("Canids") based on Canis . However, this does not typically ascend more than one or two levels: the order to which dogs and wolves belong is Carnivora ("Carnivores"). The numbers of either accepted, or all published genus names is not known precisely; Rees et al., 2020 estimate that approximately 310,000 accepted names (valid taxa) may exist, out of

1860-481: The distinctive parieto-occipital crest, a bone growth at the top of the skull above the nasal opening which projects forwards and overhangs the rest of the skull. The crest probably functioned to anchor muscle tissue, including the splenius as well as an additional muscle layer called the "extra splenius" (which was likely similar to the "splenius superficialis" found in Asian elephants, and which may have been an extension of

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1920-631: The end of the Middle Pleistocene. Most Eurasian species of Palaeoloxodon became extinct towards the end of the Last Glacial Period . The youngest records of P. antiquus are from the Iberian Peninsula, dating to around 44-43,000 years ago, with footprints from the southern part of the peninsula possibly extending the record to 28,000 years ago. The youngest Japanese records of P. naumanni date to around 24,000 years ago. The timing of extinction of Chinese Palaeoloxodon and Indian P. namadicus

1980-446: The form "author, year" in zoology, and "standard abbreviated author name" in botany. Thus in the examples above, the genus Canis would be cited in full as " Canis Linnaeus, 1758" (zoological usage), while Hibiscus , also first established by Linnaeus but in 1753, is simply " Hibiscus L." (botanical usage). Each genus should have a designated type , although in practice there is a backlog of older names without one. In zoology, this

2040-485: The genera Sivalika and Pilgrimia , with the former covering the Asian species and the latter covering the African and Mediterranean island dwarf species of Paleoloxodon . In 1931 Osborn named the genus Hesperoloxodon to include Palaeoloxodon antiquus . In a 1942 publication, Osborn recognised Sivalika and Pilgrimia as junior synonyms of Palaeoloxodon , while still recognising Hesperoloxodon as valid. This publication

2100-727: The generic name (or its abbreviated form) still forms the leading portion of the scientific name, for example, Canis lupus lupus for the Eurasian wolf subspecies, or as a botanical example, Hibiscus arnottianus ssp. immaculatus . Also, as visible in the above examples, the Latinised portions of the scientific names of genera and their included species (and infraspecies, where applicable) are, by convention, written in italics . The scientific names of virus species are descriptive, not binomial in form, and may or may not incorporate an indication of their containing genus; for example,

2160-484: The genus Elephas (which includes the Asian elephant ). In 1924, Matsumoto Hikoshichirō coined Palaeoloxodon , and circumscribed it as a subgenus of Loxodonta (which include the living species of African elephants). It included the " E. antiquus—namadicus group ", and he designated the Japanese " E. namadicus naumanni Mak." as its type species . Also in 1924, American paleontologist Henry Fairfield Osborn named

2220-509: The genus Elephas . In 2016, a study of the straight-tusked elephant ( P. antiquus ) mitochondrial genome and part of the nuclear genome found that the mitochondrial sequences were nested within the diversity of those of the African forest elephant , Loxodonta cyclotis, with the partial nuclear genome supporting P. antiquus as more closely related to L. cyclotis than the African bush elephant , L. africana . A later study published in 2018 by

2280-432: The idea that a newly defined genus should fulfill these three criteria to be descriptively useful: Moreover, genera should be composed of phylogenetic units of the same kind as other (analogous) genera. The term "genus" comes from Latin genus , a noun form cognate with gignere ('to bear; to give birth to'). The Swedish taxonomist Carl Linnaeus popularized its use in his 1753 Species Plantarum , but

2340-628: The largest component, with 23,236 ± 5,379 accepted genus names, of which 20,845 ± 4,494 are angiosperms (superclass Angiospermae). By comparison, the 2018 annual edition of the Catalogue of Life (estimated >90% complete, for extant species in the main) contains currently 175,363 "accepted" genus names for 1,744,204 living and 59,284 extinct species, also including genus names only (no species) for some groups. The number of species in genera varies considerably among taxonomic groups. For instance, among (non-avian) reptiles , which have about 1180 genera,

2400-429: The largest species, larger than any recorded in modern elephants. The molar teeth of Palaeoloxodon species typically show a "dot-dash-dot" wear pattern, with the enamel folds concentrated into a major central structure at the midline of the tooth, which are flanked by smaller folds on either side, and the crowns of the tooth are generally proportionally narrow. The teeth are typically very hypsodont (high crowned) with

2460-399: The lizard genus Anolis has been suggested to be broken down into 8 or so different genera which would bring its ~400 species to smaller, more manageable subsets. Subgenus In biology , a subgenus (plural: subgenera ) is a taxonomic rank directly below genus . In the International Code of Zoological Nomenclature , a subgeneric name can be used independently or included in

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2520-1076: The mitochondrial genomes of the European P. antiquus individual and the Chinese Palaeoloxodon specimens may indicate that the populations of Palaeoloxodon across Eurasia maintained gene flow with each other, but this is uncertain. Diagram of the relationships of elephant mitochondrial genomes, after Lin et al. 2023: Elephas (Asian elephant) Mammuthus (mammoths) Loxodonta africana (African bush elephant) North central African forest elephant clade Chinese Palaeoloxodon Chinese Palaeoloxodon Palaeoloxodon antiquus (Germany) West central African forest elephant clade Western African forest elephant clade Paleoloxodon antiquus (Germany) Palaeoloxodon cf. mnaidriensis (Sicily) These Mediterranean insular dwarf elephant species are almost certainly descended from P. antiquus Other indeterminate dwarf Palaeoloxodon species are known from other Greek islands, including Rhodes and Kasos . Many species of Palaeoloxodon are noted for

2580-403: The most (>300) have only 1 species, ~360 have between 2 and 4 species, 260 have 5–10 species, ~200 have 11–50 species, and only 27 genera have more than 50 species. However, some insect genera such as the bee genera Lasioglossum and Andrena have over 1000 species each. The largest flowering plant genus, Astragalus , contains over 3,000 species. Which species are assigned to a genus

2640-428: The name could not be used for both. Johann Friedrich Blumenbach published the replacement name Ornithorhynchus in 1800. However, a genus in one kingdom is allowed to bear a scientific name that is in use as a generic name (or the name of a taxon in another rank) in a kingdom that is governed by a different nomenclature code. Names with the same form but applying to different taxa are called "homonyms". Although this

2700-526: The provisions of the ICZN Code, e.g., incorrect original or subsequent spellings, names published only in a thesis, and generic names published after 1930 with no type species indicated. According to "Glossary" section of the zoological Code, suppressed names (per published "Opinions" of the International Commission of Zoological Nomenclature) remain available but cannot be used as the valid name for

2760-399: The rank of the division within the genus. Connecting terms are usually abbreviated, e.g. "subg." for "subgenus", and are not italicized. In zoological nomenclature, when a genus is split into subgenera, the originally described population is retained as the "nominotypical subgenus" or "nominate subgenus", which repeats the same name as the genus. For example, Panthera ( Panthera ) pardus ,

2820-519: The relationships of Chinese Palaeoloxodon are currently unresolved and it is unclear how many species were present in the region. The timing of the extinction of the last Paleoloxodon species in Africa, P. jolensis , is uncertain. While often suggested to have gone extinct during the Late Pleistocene , most specimens of the species are poorly dated and dating of specimens from Kenya suggests that it went extinct there around 130,000 years ago, at

2880-511: The remains of Eurasian species of Palaeoloxodon were considered to be those of woolly mammoths . The earliest species of Palaeoloxodon to be described, the European straight-tusked elephant ( Palaeoloxodon antiquus ) and the South Asian Palaeoloxodon namadicus , were named by British paleontologists Hugh Falconer and Proby Cautley in 1846-47. Prior to the description of the genus, Palaeoloxodon species were initially placed in

2940-416: The rhomboideus cervicis muscle) which wrapped around the top of the head to support it. The development of the crest is variable depending on the species, growth stage and gender, with females and juveniles having less developed or absent crests. The crest likely developed as a response to the large size of the head, which in proportional and absolute terms are the largest in size of any proboscideans. The skull

3000-495: The same authors based on the complete nuclear genome revised these results, and suggested P. antiquus resulted from reticulate evolution and had a complex hybridization history, with the majority (~60%) of its nuclear genome coming from a lineage more closely related to modern African elephants than to Asian elephants and mammoths, but which diverged before the split between the two living species, with significant introgressed ancestry from African forest elephants (~36%) and to

3060-421: The shoulder and 13 tonnes (29,000 lb) in body mass, making them some of the largest known terrestrial mammals to have ever lived. In a 2015 study, one fragmentary unlocated femur of P. namadicus described in the 19th century was estimated to have belonged to an individual 5.2 metres (17 ft) tall and 22 tonnes (49,000 lb) in weight, exceeding the estimates for the otherwise largest known land mammals,

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3120-623: The southern mammoth) and its replacement by Mammuthus trogontherii (the steppe mammoth) in Europe, with the extinction of M. meridionalis possibly in part a result of competition with P. antiquus . P. antiquus was able to disperse onto many islands in the Mediterranean , undergoing insular dwarfism and speciating into numerous distinct varieties of dwarf elephants . Palaeoloxodon fossils are abundant in China and are assigned to three species, P. namadicus, P. naumanni and P. huaihoensis. However,

3180-497: The specific name particular to the wolf. A botanical example would be Hibiscus arnottianus , a particular species of the genus Hibiscus native to Hawaii. The specific name is written in lower-case and may be followed by subspecies names in zoology or a variety of infraspecific names in botany . When the generic name is already known from context, it may be shortened to its initial letter, for example, C. lupus in place of Canis lupus . Where species are further subdivided,

3240-412: The standard format for a species name comprises the generic name, indicating the genus to which the species belongs, followed by the specific epithet, which (within that genus) is unique to the species. For example, the gray wolf 's scientific name is Canis lupus , with Canis ( Latin for 'dog') being the generic name shared by the wolf's close relatives and lupus (Latin for 'wolf') being

3300-459: The subgenus is one of the possible subdivisions of a genus. There is no limit to the number of divisions that are permitted within a genus by adding the prefix "sub-" or in other ways as long as no confusion can result. The secondary ranks of section and series are subordinate to subgenus. An example is Banksia subg. Isostylis , a subgenus of the large Australian genus Banksia . The ICNafp requires an explicit "connecting term" to indicate

3360-403: The taxon is termed a synonym ; some authors also include unavailable names in lists of synonyms as well as available names, such as misspellings, names previously published without fulfilling all of the requirements of the relevant nomenclatural code, and rejected or suppressed names. A particular genus name may have zero to many synonyms, the latter case generally if the genus has been known for

3420-566: The values quoted are the mean of "accepted" names alone (all "uncertain" names treated as unaccepted) and "accepted + uncertain" names (all "uncertain" names treated as accepted), with the associated range of uncertainty indicating these two extremes. Within Animalia, the largest phylum is Arthropoda , with 151,697 ± 33,160 accepted genus names, of which 114,387 ± 27,654 are insects (class Insecta). Within Plantae, Tracheophyta (vascular plants) make up

3480-429: The virus species " Salmonid herpesvirus 1 ", " Salmonid herpesvirus 2 " and " Salmonid herpesvirus 3 " are all within the genus Salmonivirus ; however, the genus to which the species with the formal names " Everglades virus " and " Ross River virus " are assigned is Alphavirus . As with scientific names at other ranks, in all groups other than viruses, names of genera may be cited with their authorities, typically in

3540-551: Was dominant for most of the Pliocene and Early Pleistocene, with the mammoth hybridisation suggested to have taken place earlier than the hybridisation with forest elephants. Analysis of mitochondrial genomes, including Palaeoloxodon individuals from Northern China indicates Palaeoloxodon individuals harboured multiple separate mitochondrial genome lineages derived from African forest elephants, some being more closely related to some West African forest elephant groups than to others. It

3600-435: Was the first to consider Palaeoloxodon a valid genus in its own right, an opinion followed by later authors such as Emiliano Aguirre in 1969. Later authors have considered Hesperoloxodon another synonym of Palaeoloxodon . Vincent J. Maglio in a 1973 publication controversially synonymised Palaeoloxodon with Elephas based on morphological similarities between the two genera. Later authors either considered Palaeoloxodon

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