The Proboscidea (; , ) are a
taxonomic
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
order
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
of
afrotherian mammals containing one living
family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
(
Elephantidae) and several extinct families. First described by
J. Illiger in 1811, it encompasses the
elephant
Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae an ...
s and their close relatives. From the mid-
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
onwards, most proboscideans were very large. The
largest land mammal
The following is a list of largest mammals by family.
Tenrecs and allies (Afrosoricida)
*The largest of these insectivorous mammals is the giant otter shrew (''Potamogale velox''), native to Central Africa. This species can weigh up to and mea ...
of all time may have been a proboscidean; ''
Palaeoloxodon namadicus'' was up to at the shoulder and may have weighed up to , almost double the weight of some
sauropods like ''
Diplodocus carnegii
''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek διπλ ...
''.
The largest extant proboscidean is the
African bush elephant
The African bush elephant (''Loxodonta africana'') is one of two extant African elephant species and one of three extant elephant species. It is the largest living terrestrial animal, with bulls reaching a shoulder height of up to and a body ...
, with a record of size of at the shoulder and .
In addition to their enormous size, later proboscideans are distinguished by tusks and long, muscular trunks, which were less developed or absent in early proboscideans.
Three
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
elephant
Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae an ...
are currently recognised: the African bush elephant, the
African forest elephant, and the
Asian elephant
The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living species of the genus ''Elephas'' and is distributed throughout the Indian subcontinent and Southeast Asia, from India in the west, Nepal in the no ...
. Elephantidae is the only surviving family of the order Proboscidea; extinct members include the
mastodon
A mastodon ( 'breast' + 'tooth') is any proboscidean belonging to the extinct genus ''Mammut'' (family Mammutidae). Mastodons inhabited North and Central America during the late Miocene or late Pliocene up to their extinction at the end of th ...
s,
gomphotheres and
stegodonts
Stegodontidae is an extinct family of proboscideans from Africa and Asia (with a single occurrence in Europe) from the Miocene (15.97 mya) to the Late Pleistocene, with some studies suggesting that some survived into the Holocene in Chi ...
. The family Elephantidae also contains several extinct groups, including the
mammoth
A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks and, ...
s and
straight-tusked elephants. The distinctive features of proboscideans include a trunk, tusks, and massive legs. Large ear flaps are present in some proboscideans, including elephants. Some also have tough but sensitive skin; others, like the woolly mammoth, have a coat. The trunk is used for breathing, bringing food and water to the mouth, and grasping objects. Tusks, which are derived from the incisor teeth, serve both as weapons and as tools for moving objects and digging. The large ear flaps assist in maintaining a constant body temperature as well as in communication. The
pillar-like legs carry their great weight.
Evolution
The earliest known proboscidean is ''
Eritherium
''Eritherium'' is an extinct genus of early Proboscidea found in the Ouled Abdoun basin (early Thanetian age), Morocco. It lived about 60 million years ago. It was first named by Emmanuel Gheerbrant in 2009 and the type species is ''Eritherium ...
'', followed by ''
Phosphatherium
''Phosphatherium escuillei'' is a basal proboscidean that lived from the Late Paleocene to the early stages of the Ypresian age until the early Thanetian some 56 million years ago in North Africa. Research has suggested that ''Phosphatherium'' ex ...
'', a small animal about the size of a fox. Both date from late
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
deposits of
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
.

Proboscideans evolved in Africa, where they increased in size and diversity during the
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
and early
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
. Proboscideans have evolved greatly over time through three major forms of radiation: radiation of primitive Lophodont forms, radiation of gomphotheres and stegodons, and radiation of elephantidae. These forms of radiation have illustrated that proboscideans characteristics such as trunk, large ears, and tusks have evolved and were appearing late in the modern form. Several primitive families from these epochs have been described, including the
Numidotheriidae
Numidotheriidae is an extinct family of primitive proboscidean that lived from the late Paleocene to the early Oligocene periods of North Africa.
Fragmentary fossils (mainly teeth) of the early Eocene genera, ''Daouitherium'' and ''Phosphatheri ...
,
Moeritheriidae, and
Barytheriidae
Barytheriidae (meaning "heavy beasts") is an extinct family of primitive proboscideans that lived during the late Eocene and early Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to ...
, all found exclusively in Africa. The
Anthracobunidae from the Indian subcontinent were also believed to be a family of proboscideans, but were excluded from the Proboscidea by Shoshani and Tassy (2005)
and have more recently been assigned to the
Perissodactyla. When Africa became connected to Europe and Asia after the shrinking of the
Tethys Sea, proboscideans migrated into Eurasia, with some families eventually reaching the Americas. Proboscideans found in Eurasia as well as Africa include the
Deinotheriidae
Deinotheriidae ("terrible beasts") is a family of prehistoric elephant-like proboscideans that lived during the Cenozoic era, first appearing in Africa, then spreading across southern Asia (Indo-Pakistan) and Europe. During that time, they cha ...
, which thrived during the
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
and into the early
Quaternary
The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS). It follows the Neogene Period and spans from 2.58 million years ...
, ''
Stegolophodon'', an early genus of the disputed family
Stegodontidae; the highly diverse
Gomphotheriidae
Gomphotheres are any members of the diverse, extinct taxonomic family Gomphotheriidae. Gomphotheres were elephant-like proboscideans, but do not belong to the family Elephantidae. They were widespread across Afro-Eurasia and North America during ...
and
Amebelodontidae
Amebelodontidae is an extinct family of large herbivorous mammals that were closely related to elephants. They were formerly assigned to Gomphotheriidae, but recent authors consider them a distinct family.
Feeding habits
In the past, Amebelodont ...
; and the
Mammutidae
Mammutidae is an extinct family of proboscideans that appeared during the Oligocene epoch and survived until the start of the Holocene. The family was first described in 1922, classifying fossil specimens of the type genus ''Mammut'' (mastodons) ...
, or mastodons.
Most proboscideans are now extinct, including all species
endemic
Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to the Americas, Europe, and northern Asia. Many of these extinctions occurred during or shortly after the last
glacial period. Recently extinct species include the
gomphotheres in the Americas, the
American mastodon of family Mammutidae in North America, numerous
stegodonts in Asia, the
mammoth
A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks and, ...
s throughout the Northern Hemisphere, and several species of
dwarf elephant
Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes (around ) in comparison with their immediate ancestors. Dwarf elephants are an example ...
s found on various islands scattered around the world.
Classification
Below is an unranked taxonomy of proboscidean genera as of 2019.
*Proboscidea Illiger, 1811
**†''
Eritherium
''Eritherium'' is an extinct genus of early Proboscidea found in the Ouled Abdoun basin (early Thanetian age), Morocco. It lived about 60 million years ago. It was first named by Emmanuel Gheerbrant in 2009 and the type species is ''Eritherium ...
'' Gheerbrant, 2009
**†''
Moeritherium'' Andrews, 1901
**†''
Saloumia'' Tabuce ''et al.'', 2019
**†
Plesielephantiformes
Plesielephantiformes is an extinct clade of large herbivorous mammals and one of two suborders of the Proboscidea, a group containing elephant
Elephants are the largest existing land animals. Three living species are currently recognise ...
Shoshani ''et al.'', 2001
***†
Numidotheriidae
Numidotheriidae is an extinct family of primitive proboscidean that lived from the late Paleocene to the early Oligocene periods of North Africa.
Fragmentary fossils (mainly teeth) of the early Eocene genera, ''Daouitherium'' and ''Phosphatheri ...
Shoshani & Tassy, 1992
****†''
Phosphatherium
''Phosphatherium escuillei'' is a basal proboscidean that lived from the Late Paleocene to the early stages of the Ypresian age until the early Thanetian some 56 million years ago in North Africa. Research has suggested that ''Phosphatherium'' ex ...
'' Gheerbrant ''et al.'', 1996
****†''
Arcanotherium
''Arcanotherium'' is an extinct genus of early proboscidean belonging to the family Numidotheriidae that lived in north Africa during the late Eocene/early Oligocene interval.
Taxonomy
''Arcanotherium'' was originally described by Court (1995) ...
'' Delmer, 2009
****†''
Daouitherium'' Gheerbrant & Sudre, 2002
****†''
Numidotherium'' Mahboubi ''et al.'', 1986
***†
Barytheriidae
Barytheriidae (meaning "heavy beasts") is an extinct family of primitive proboscideans that lived during the late Eocene and early Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to ...
Andrews, 1906
****†''
Omanitherium
''Omanitherium'' (meaning ''Oman beast'') is a genus of an extinct genus of barytheriid proboscidean
The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct familie ...
'' Seiffert ''et al.'', 2012
****†''
Barytherium'' Andrews, 1901
***†
Deinotheriidae
Deinotheriidae ("terrible beasts") is a family of prehistoric elephant-like proboscideans that lived during the Cenozoic era, first appearing in Africa, then spreading across southern Asia (Indo-Pakistan) and Europe. During that time, they cha ...
Bonaparte, 1845
****†''
Chilgatherium
''Chilgatherium'' ('Chilga beast' after the locality in which it was found) is the earliest and most primitive representative of the family Deinotheriidae. It is known from late Oligocene (27- to 28-million-year-old) fossil teeth found in the ...
'' Sanders ''et al.'', 2004
****†''
Prodeinotherium'' Ehik, 1930
****†''
Deinotherium
''Deinotherium'' was a large elephant-like proboscidean that appeared in the Middle Miocene and survived until the Early Pleistocene. Although superficially resembling modern elephants, they had notably more flexible necks, limbs adapted to a mo ...
'' Kaup, 1829
**
Elephantiformes
Elephantiformes is a suborder within the order Proboscidea that contains the elephants as well as their extinct relatives. Elephant relatives such as the mammoths and the mastodons are included in this designation.
References
Mammal s ...
Tassy, 1988
***†''
Eritreum
''Eritreum melakeghebrekristosi'' is an extinct species of proboscidean mammal, which lived in Northeast Africa during the late Oligocene some 27 million years ago, and is considered to be the missing link between modern elephants and their ance ...
'' Shoshani ''et al.'', 2006
***†''
Hemimastodon
''Hemimastodon'' ("half mastodont") is an extinct genus of proboscidean from the Late Miocene deposits of the Dera Bugti Beds in Pakistan
Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in ...
'' Pilgrim, 1912
***†''
Palaeomastodon'' Andrews, 1901
***†''
Phiomia'' Andrews & Beadnell, 1902
***
Elephantimorpha
Elephantimorpha is a group that contains the elephants as well as their extinct relatives, the gomphotheres and stegodontids. The following cladogram shows the relationships among elephantimorphs, based on hyoid
The hyoid bone (lingual bone or t ...
Tassy & Shoshani, 1997
****†
Mammutidae
Mammutidae is an extinct family of proboscideans that appeared during the Oligocene epoch and survived until the start of the Holocene. The family was first described in 1922, classifying fossil specimens of the type genus ''Mammut'' (mastodons) ...
Hay, 1922
*****†''
Losodokodon
''Losodokodon'' is an extinct genus of large herbivorous mammals belonging to the family Mammutidae. It was first described in 2009 by David Tab Rasmussen and Mercedes Gutiérrez from fossils found in the Erageleit Formation of northwestern Kenya ...
'' Rasmussen & Gutierrez, 2009
*****†''
Eozygodon
''Eozygodon'' is an extinct genus of proboscidean in the family Mammutidae. It is known from the Early Miocene
The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene Epoch made up of two stages: the Aquitanian and Burdig ...
'' Tassy & Pickford, 1983
*****†''
Zygolophodon
''Zygolophodon'' is an extinct genus of African, Asian, and European mammutid that lived from the Miocene to the Late Pliocene.
Taxonomy
''Zygolophodon'' belongs in the family Mammutidae, whose best known member is the American mastodon. ''Zy ...
'' Vacek, 1877
*****†''
Sinomammut
''Sinomammut'' (meaning "Chinese mastodon") is a proboscidean found in the Miocene of China. Only one species, ''S. tobieni'', is known, named in 2016.
Discovery and naming
It was known from GIOTC 0984-9-178, a single, fragmentary, mandible fou ...
'' Mothé ''et al.'', 2016
*****†''
Mammut
A mastodon ( 'breast' + 'tooth') is any proboscidean belonging to the extinct genus ''Mammut'' (family Mammutidae). Mastodons inhabited North and Central America during the late Miocene or late Pliocene up to their extinction at the end of the ...
'' Blumenbach, 1799
****
Elephantida
Elephantida is a group that contains the elephants as well as their extinct relatives, the gomphotheres and the stegodontids. The following cladogram shows the relationships among elephantidans, based on hyoid
The hyoid bone (lingual bone or ...
Tassy & Shoshani, 1997
*****†
Choerolophodontidae
Choerolophodontidae is an extinct family of large herbivorous mammals that were closely related to elephants. Two genera are known, ''Afrochoerodon'' and ''Choerolophodon''.
Taxonomy
Although usually classified as part of Gomphotheriidae, recent ...
Gaziry, 1976
******†''
Afrochoerodon'' Pickford, 2001
******†''
Choerolophodon'' Schlesinger, 1917
*****†
Amebelodontidae
Amebelodontidae is an extinct family of large herbivorous mammals that were closely related to elephants. They were formerly assigned to Gomphotheriidae, but recent authors consider them a distinct family.
Feeding habits
In the past, Amebelodont ...
Barbour, 1927
******†''
Afromastodon'' Pickford, 2003
******†''
Progomphotherium
''Progomphotherium'' is an extinct genus of large herbivorous mammals that were closely related to elephant
Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the A ...
'' Pickford, 2003
******†''
Eurybelodon
''Eurybelodon'' is an extinct genus of proboscidean in the family Amebelodontidae.
Taxonomy
The type specimen, a partial upper tusk, was described from Black Butte in western Oregon in 1963. It was originally assigned to the genus ''Platybelodo ...
'' Lambert, 2016
******†''
Serbelodon
''Serbelodon'' is an extinct genus of proboscidean. It had tusks and a trunk. It lived in North America during the Miocene Epoch, and it was closely related to ''Amebelodon''. They had a diet that consisted of C3 plants which include fruits, tre ...
'' Frick, 1933
******†''
Archaeobelodon
''Archaeobelodon'' is an extinct genus of proboscidean of the family Amebelodontidae that lived in Europe and North Africa (Egypt) during the Miocene from 16.9 to 16.0 Ma, living for approximately .
''Archaeobelodon'' was an ancestor of ''Plat ...
'' Tassy, 1984
******†''
Protanancus
''Protanancus'' is an extinct genus of amebelodontid proboscidean from Kenya, Pakistan and Thailand. The genus consists solely of type species ''P. macinnesi''. '' Arambourg, 1945
******†''
Amebelodon
''Amebelodon'' is a genus of extinct proboscidean belonging to Amebelodontidae (the so-called shovel-tuskers), a group of proboscideans related to the modern elephants and their close relative the mammoth. The most striking attribute of this anim ...
'' Barbour, 1927
******†''
Konobelodon
''Konobelodon'' is an extinct genus of amebelodont from southern Europe, China, and North America.
Taxonomy
''Konobelodon'' was originally coined as a subgenus of ''Amebelodon'', and was subsequently elevated to full generic rank in a 2014 re- ...
'' Lambert, 1990
******†''
Torynobelodon
''Torynobelodon'' was a genus of large herbivorous mammal related to the elephant (order Proboscidea). It lived during the late Miocene Epoch in Asia and North America.
Taxonomy
Shoshani (1996) placed ''Torynobelodon'' as a synonym of ''Platy ...
'' Barbour, 1929
******†''
Aphanobelodon
''Aphanobelodon'' is an extinct genus of proboscidean in the family Amebelodontidae.
Taxonomy
The holotype is the complete cranium of an adult female, and the paratype
In zoology and botany, a paratype is a specimen of an organism that help ...
'' Wang ''et al.'', 2016
******†''
Platybelodon
''Platybelodon'' ("flat-spear tusk") is an extinct genus of large herbivorous proboscidean mammals related to modern-day elephants. Species lived during the middle Miocene Epoch in Africa, Asia and the Caucasus.
Palaeobiology
''Platybelodon'' wa ...
'' Borissiak, 1928
*****†
Gomphotheriidae
Gomphotheres are any members of the diverse, extinct taxonomic family Gomphotheriidae. Gomphotheres were elephant-like proboscideans, but do not belong to the family Elephantidae. They were widespread across Afro-Eurasia and North America during ...
Hay, 1922
******†''
Gomphotherium
''Gomphotherium'' (; "welded beast") is an extinct genus of proboscids from the Neogene and early Pleistocene of Eurasia, Africa, North America and Asia. As of 2021, two species, ''G. annectens'' and possibly ''G. subtapiroideum'', are also kno ...
'' Burmeister, 1837
******†''
Blancotherium'' May, 2019
******†''
Gnathabelodon
''Gnathabelodon'' is an extinct genus of gomphothere (a sister group to modern elephants) endemic to North America that includes species that lived during the Middle to Late Miocene.
''"Gnathabelodon" buckneri'' Sellards, 1940 has been renamed ...
'' Barbour & Sternberg, 1935
******†''
Eubelodon
''Eubelodon'' is an extinct genus of gomphothere (a family in the order Proboscidea, which also includes modern elephants) which lived in North America during the Miocene Epoch. It contains a single species: ''Eubelodon morrilli''. Like other gom ...
'' Barbour, 1914
******†''
Stegomastodon
''Stegomastodon'' ('roof breast tooth') is an extinct genus of gomphotheres, a family of proboscideans. It ranged throughout North America from the early Blancan ~4 Ma, to the early Irvingtonian (~1.2 Ma). The South American species have be ...
'' Pohlig, 1912
******†''
Sinomastodon
''Sinomastodon'' ("Chinese mastodont") is an extinct gomphothere genus (of order Proboscidea), from the Late Miocene to the Early Pleistocene deposits of Asia ( China, Japan, Thailand, Myanmar, Kashmir, and Indonesia). It is not to be confus ...
'' Tobien ''et al.'', 1986
******†''
Notiomastodon
''Notiomastodon'' is an extinct proboscidean genus of gomphotheres (a distant relative to modern elephants) endemic to South America from the Pleistocene to the beginning of the Holocene. ''Notiomastodon'' specimens reached a size similar to that ...
'' Cabrera, 1929
******†''
Rhynchotherium
''Rhynchotherium'' is an extinct genus of proboscidea endemic to North America and Central America during the Miocene through Pliocene from 13.650 to 3.6 Ma, living for approximately .
This gomphothere had two tusks and may have evolved from ...
'' Falconer, 1868
******†''
Cuvieronius
''Cuvieronius'' is an extinct New World genus of gomphothere, named after the French naturalist Georges Cuvier. Alive, specimens typically stood about tall at the shoulder, weighed about and would have superficially resembled a modern elephant ...
'' Osborn, 1923
*****
Elephantoidea Gray, 1821
******†
Anancidae Hay, 1922
*******†''
Anancus'' Aymard, 1855
*******†''
Morrillia'' Osborn, 1924
*******†''
Paratetralophodon
Paratetralophodon is an extinct genus of proboscidean from late Neogene deposits in India and China. Although traditionally classified in the family Gomphotheriidae, recent studies find it to be more closely related to modern elephants.
''Para ...
'' Tassy, 1983
*******†''
Pediolophodon'' Lambert, 2007
*******†''
Tetralophodon'' Falconer, 1857
******†
Stegodontidae Osborn, 1918
*******†''
Stegolophodon'' Schlesinger, 1917
*******†''
Stegodon'' Falconer, 1857
******
Elephantidae Gray, 1821
*******†
Stegotetrabelodontinae Aguirre, 1969
********†''
Stegodibelodon'' Coppens, 1972
********†''
Stegotetrabelodon'' Petrocchi, 1941
********†''
Selenotherium'' Mackaye, Brunet & Tassy, 2005
*******
Elephantinae Gray, 1821
********†''
Primelephas
''Primelephas'' is a genus of Elephantinae that existed during the Miocene and Pliocene epochs. The name of the genus suggests 'first elephant'. These primitive elephantids are thought to be the common ancestor of ''Mammuthus'', the mammoths, ...
'' Maglio, 1970
********''
Loxodonta'' Anonymous, 1827
********†''
Palaeoloxodon
''Palaeoloxodon'' is an extinct genus of elephant. The genus originated in Africa during the Pliocene era, and expanded into Eurasia during the Pleistocene era. The genus contains some of the largest known species of elephants, over four metres t ...
'' Matsumoto, 1924
********†''
Mammuthus'' Brookes, 1828
********''
Elephas
''Elephas'' is one of two surviving genera in the family of elephants, Elephantidae, with one surviving species, the Asian elephant, ''Elephas maximus''.
Several extinct species have been identified as belonging to the genus, extending back to ...
'' Linnaeus, 1758
References
Bibliography
*
*
{{Authority control
Mammal orders
Selandian first appearances
Taxa named by Johann Karl Wilhelm Illiger
Extant Selandian first appearances