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The Saint Bathans mammal is a currently unnamed extinct primitive
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fu ...
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 Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 Ma to 15.97 ± 0.05 Ma (million years ago). It was pr ...
(Altonian) of
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island country ...
. A member of the
Saint Bathans fauna The St Bathans fauna is found in the lower Bannockburn Formation of the Manuherikia Group of Central Otago, in the South Island of New Zealand. It comprises a suite of fossilised prehistoric animals from the late Early Miocene (Altonian) period, ...
, it is notable for being a late-surviving "archaic" mammal species, neither a
placental Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
nor a
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a p ...
. It also provides evidence that flightless fully terrestrial mammals did in fact once live in
Zealandia Zealandia (pronounced ), also known as (Māori) or Tasmantis, is an almost entirely submerged mass of continental crust that subsided after breaking away from Gondwanaland 83–79 million years ago.Gurnis, M., Hall, C.E., and Lavier, L.L., ...
. This is in contrast to modern New Zealand, where
bats Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most b ...
,
cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel the ...
ns and seals are the only non-introduced mammals in the otherwise
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
-dominated faunas.


Discovery

The Saint Bathans mammal is currently represented by three specimens in
Te Papa The Museum of New Zealand Te Papa Tongarewa is New Zealand's national museum and is located in Wellington. ''Te Papa Tongarewa'' translates literally to "container of treasures" or in full "container of treasured things and people that spring f ...
: NMNZ S.40958, NMNZ S.41866, and NMNZ S.42214, composed of two lower jaw fragments and a femur respectively. It was part of an assemblage of fossils recovered in
Saint Bathans St Bathans, formerly named Dunstan Creek, is a former gold and coal mining town in Central Otago, New Zealand. The settlement was a centre of the Otago Gold Rush, but mining has since long ceased. It is now largely a holiday retreat due to the p ...
in 1978, in what would later be understood to be the Bannockburn Formation ( Manuherikia Group), and first described in 2006.


Description

Like most small mammal fossils, the Saint Bathans mammal material is rather incomplete, with only a lower jaw fragment and femur being known. The lower jaws are toothless, though the presence of deep tooth sockets suggests that they were toothed in life and that the teeth were lost post-mortem. They bear a long fused
mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral hal ...
, an evidently procumbent lower
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wh ...
, and five additional sockets that imply a
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolog ...
of one incisor, one canine and two double-rooted
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mout ...
s. The femur possesses a round
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may not ...
and poorly defined
neck The neck is the part of the body on many vertebrates that connects the head with the torso. The neck supports the weight of the head and protects the nerves that carry sensory and motor information from the brain down to the rest of the body. I ...
, oriented slightly dorsomedially with respect to the long axis of the shaft, and separated from the
greater trochanter The greater trochanter of the femur is a large, irregular, quadrilateral eminence and a part of the skeletal system. It is directed lateral and medially and slightly posterior. In the adult it is about 2–4 cm lower than the femoral head.Sta ...
by a marked trough. The alignment of the femur in life is hard to ascertain, but it is thought that the animal had a semi-sprawling stance, more abducted than in
theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
n mammals but nowhere near as much as in
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brains ...
s.


Phylogeny

Because of the incomplete material, it is very hard to understand the position of this taxon within
Mammaliaformes Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent ...
as a whole. Worthy et al. 2006 tentatively deemed the Saint Bathans mammal as a theriiform, being more derived than
morganucodonta Morganucodonta ("Glamorgan teeth") is an extinct order of basal Mammaliaformes, a group including crown-group mammals (Mammalia) and their close relatives. Their remains have been found in Southern Africa, Western Europe, North America, India a ...
ns,
eutriconodont Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 as a replacement name for the ...
s and
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brains ...
s but not as much as
multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, ...
s, on the basis of its femoral anatomy. As the phylogeny of non-
theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
n mammals has undergone multiple shifts since its description, new studies might be necessary.


Ecology

The Bannockburn Formation depicts a warm temperate or subtropical lakeside environment, surrounded by herbaceous
peat Peat (), also known as turf (), is an accumulation of partially decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, moors, or muskegs. The peatland ecosystem covers and is the most efficie ...
swamps A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
. Casuarinas, araucarias, podocarps,
eucalypts Eucalypt is a descriptive name for woody plants with capsule fruiting bodies belonging to seven closely related genera (of the tribe Eucalypteae) found across Australasia: ''Eucalyptus'', ''Corymbia'', '' Angophora'', '' Stockwellia'', '' Allos ...
,
palm tree The Arecaceae is a family of perennial flowering plants in the monocot order Arecales. Their growth form can be climbers, shrubs, tree-like and stemless plants, all commonly known as palms. Those having a tree-like form are called palm ...
s and
southern beech ''Nothofagus'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...
es are among the various plant species known to have grown here. As today, the local vertebrate fauna was dominated by birds: early
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refer ...
s and
adzebill The adzebills, genus ''Aptornis'', were two closely related bird species, the North Island adzebill, (''Aptornis otidiformis''), and the South Island adzebill, (''Aptornis defossor''), of the extinct family Aptornithidae. The family was endemi ...
s are represented by unnamed species, as are various representatives of groups such as
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which in ...
,
flamingos Flamingos or flamingoes are a type of wading bird in the family Phoenicopteridae, which is the only extant family in the order Phoenicopteriformes. There are four flamingo species distributed throughout the Americas (including the Caribbean ...
,
rails Rail or rails may refer to: Rail transport * Rail transport and related matters *Rail (rail transport) or railway lines, the running surface of a railway Arts and media Film * ''Rails'' (film), a 1929 Italian film by Mario Camerini * ''Rail'' ...
,
herons The herons are long-legged, long-necked, freshwater and coastal birds in the family Ardeidae, with 72 recognised species, some of which are referred to as egrets or bitterns rather than herons. Members of the genera ''Botaurus'' and ''Ixobrychus ...
, strigopoidean
parrot Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genera comprising the order Psittaciformes (), found mostly in tropical and subtropical regions. The order is subdivided into three superfamilies: the Psittac ...
s and even an early
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also ref ...
, '' Proapteryx''. However, unlike modern New Zealand it also had a varied herpetofauna: besides an early
tuatara Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language an ...
, the Saint Bathans fauna also includes meiolaniid and
pleurodire The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differen ...
turtles and possibly
mekosuchine Mekosuchinae is an extinct clade of crocodilians from the Cenozoic of Australasia. They first appear in the fossil record in the Eocene in Australia, and survived until the arrival of humans: in the Pleistocene in Australia and within the Holoce ...
crocodylids and
snakes Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more jo ...
.Scofield, R. Paul; Worthy, Trevor H. & Tennyson, Alan J.D. (2010). "A heron (Aves: Ardeidae) from the Early Miocene St Bathans Fauna of southern New Zealand.". In W.E. Boles & T.H. Worthy. Proceedings of the VII International Meeting of the Society of Avian Paleontology and Evolution (PDF). Records of the Australian Museum 62. pp. 89–104. Besides the Saint Bathans mammal, this fauna also includes mystacine bats, a group still present in modern New Zealand. Like the modern species, these were probably terrestrial foragers, the same general ecological niche proposed for the Saint Bathans mammal. Other bats, including a
vesper bat Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most diverse and widely distributed of bat famili ...
and several currently unclassified species, also existed.


References

{{Mammalia Mammals of New Zealand Extinct animals of New Zealand Miocene extinctions Miocene first appearances Undescribed vertebrate species Fossils of New Zealand