47-529: 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, with an age range of 19–16 million years ago. The layer in which the fossils are found derives from littoral zone sediments deposited in
94-430: A crocodilian , a rhynchocephalian (a relative of tuatara ), geckos , skinks , a primitive mammal , several species of bats , and several kinds of birds , especially waterbirds . Of tree-dwelling birds, parrots outnumber pigeons thirty to one. Proapteryx , a basal form of kiwi , is known from there. The Miocene ecosystem was recovering from the ‘ Oligocene drowning ’ a few million years earlier, when up to 80% of
141-523: A natural reservoir of many pathogens , such as rabies ; and since they are highly mobile, social, and long-lived, they can readily spread disease among themselves. If humans interact with bats, these traits become potentially dangerous to humans. Some bats are also predators of mosquitoes , suppressing the transmission of mosquito-borne diseases . Depending on the culture, bats may be symbolically associated with positive traits, such as protection from certain diseases or risks, rebirth, or long life, but in
188-973: A 2005 DNA study. A 2013 phylogenomic study supported the two new proposed suborders. Yangochiroptera (as above) [REDACTED] Pteropodidae (megabats) [REDACTED] Megadermatidae (false vampire bats) [REDACTED] horseshoe bats and allies [REDACTED] The 2003 discovery of an early fossil bat from the 52-million-year-old Green River Formation , Onychonycteris finneyi , indicates that flight evolved before echolocative abilities. Onychonycteris had claws on all five of its fingers, whereas modern bats have at most two claws on two digits of each hand. It also had longer hind legs and shorter forearms, similar to climbing mammals that hang under branches, such as sloths and gibbons . This palm-sized bat had short, broad wings, suggesting that it could not fly as fast or as far as later bat species. Instead of flapping its wings continuously while flying, Onychonycteris probably alternated between flaps and glides in
235-495: A distinctly subtropical Australian climate and the surrounding vegetation was characterised by casuarinas , eucalypts and palms as well as podocarps , araucarias and southern beeches . The fossiliferous layer has been exposed at places along the Manuherikia River and at other sites in the vicinity of the historic gold mining town of St Bathans . The fauna consists of a variety of vertebrates, including fish ,
282-475: A large fruit pigeon, possibly related to the modern Hemiphaga species. The Zealandian dove is similar to the Nicobar pigeon . Several Gruiformes have been described. The St Bathans adzebill ( Aptornis proasciarostratus ) was only slightly smaller than its more recent descendants. There were two flightless rails : the common Priscaweka parvales and uncommon Litorallus livezeyi . Priscaweka parvales
329-471: A relative of the plains-wanderer, and Sansom's plover ( Neilus sansomae ), a plover-like bird of uncertain affinities but possibly related to sheathbills and the Magellanic plover. Petrels are seabirds in the order Procellariformes . This group includes albatrosses. Petrels today make up most of all species of seabird, and the order is the only order of birds to be entirely marine. One species of petrel
376-535: A shallow, freshwater lake, with an area of 5600 km from present day Central Otago to Bannockburn and the Nevis Valley in the west; to Naseby in the east; and from the Waitaki Valley in the north to Ranfurly in the south. The lake was bordered by an extensive floodplain containing herbaceous and grassy wetland habitats with peat -forming swamp –woodland. At that time the climate was warm with
423-445: A shift from -k- to -t- (to Modern English bat ) influenced by Latin blatta , ' moth, nocturnal insect ' . The word bat was probably first used in the early 1570s. The name Chiroptera derives from Ancient Greek : χείρ – cheir , ' hand ' and πτερόν – pteron , ' wing ' . The delicate skeletons of bats do not fossilise well; it is estimated that only 12% of bat genera that lived have been found in
470-777: A sister taxon to odd-toed ungulates (Perissodactyla). Euarchontoglires (primates, treeshrews, rodents, rabbits) [REDACTED] Eulipotyphla (hedgehogs, shrews, moles, solenodons) [REDACTED] Chiroptera (bats) [REDACTED] Pholidota (pangolins) [REDACTED] Carnivora (cats, hyenas, dogs, bears, seals, weasels) [REDACTED] [REDACTED] Perissodactyla (horses, tapirs, rhinos) [REDACTED] Cetartiodactyla (camels, ruminants, whales) [REDACTED] [REDACTED] The flying primate hypothesis proposed that when adaptations to flight are removed, megabats are allied to primates by anatomical features not shared with microbats and thus flight evolved twice in mammals. Genetic studies have strongly supported
517-429: A small hawk , have been found, but await formal description. Two parrot genera are represented. Heracles is represented by its sole species, Heracles inexpectatus , the largest known parrot, weighing 7 kilograms and standing 1 meter tall. Nelepsittacus is represented by at least four species. These vary drastically in size, suggesting that they occupied a wide variety of ecological niches, having diversified in
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#1732773093306564-475: Is a basal theriiform mammal, the St Bathans mammal . Several species of mystacine bats are also known, as well as a vesper bat and several incertae sedis species. This bat fauna included Vulcanops , a giant burrowing bat three times the size of today’s relatives, and more closely related to South American bats. This suggests that small land mammals were a common component of New Zealand's fauna in
611-498: Is common. The dabbling duck Matanas enrightii remains poorly known as only a few fossils have been found. Palaelodids are ancient relatives of flamingos. The new species from St Bathans ( Palaelodus aotearoa ) is smaller than, and morphologically distinct from, the Late Oligocene-Early Miocene Palaelodus wilsoni from Australia. Two pigeon species have been described. Rupephaps is
658-668: Is grouped as the Fiddlers Member. Extensive coal swamps developed between the rivers and their flood basins. Individual flood basin lakes began to coalesce into a true lacustrine system – Lake Manuherikia . Its muddy and sandy sediments are grouped as the Bannockburn Formation. Around the lake margins, distinctive shoreline facies developed. The most widespread of these is the Kawarau Member which contains diverse plant fossils. Distinctive facies within
705-639: Is known from the St Bathans Fauna – a diving petrel in the same genus as modern diving petrels, the Miocene diving petrel ( Pelecanoides miokuaka ). At least two herons are known: Pikaihao bartlei and Matuku otagoense . The former is a bittern , while the latter is a much larger species that appears to be basal within Ardeidae (the herons). One eagle , similar in size to a wedge-tailed eagle , and another bird of prey, similar in size to
752-1147: Is not believed to originate more than 23 mya. Pteropodidae (megabats) [REDACTED] Megadermatidae (false vampire bats) [REDACTED] Craseonycteridae (Kitti's hog-nosed bat) [REDACTED] Rhinopomatidae (mouse-tailed bats) [REDACTED] Hipposideridae (Old World leaf-nosed bats) [REDACTED] Rhinolophidae (horseshoe bats) [REDACTED] Miniopteridae (long winged bat) [REDACTED] Noctilionidae (fisherman bats) [REDACTED] Mormoopidae ( Pteronotus ) [REDACTED] Mystacinidae (New Zealand short-tailed bats) [REDACTED] Thyropteridae (disc-winged bats) Furipteridae [REDACTED] Mormoopidae ( Mormoops ) [REDACTED] Phyllostomidae (New World leaf-nosed bats) [REDACTED] Molossidae (free-tailed bats) [REDACTED] Emballonuridae (sac-winged bats) [REDACTED] Myzopodidae (sucker-footed bats) Emballonuridae ( Taphozous ) [REDACTED] Natalidae (funnel-eared bats) [REDACTED] Vespertilionidae (vesper bats) [REDACTED] Genetic evidence indicates that megabats originated during
799-660: The Museum of New Zealand Te Papa Tongarewa . In 2016 Vanesa De Pietri was awarded a Royal Society of New Zealand Marsden Fast Start grant to study the shorebird fossils. This long-running (since 2000) collaborative research programme also includes scientists from the University of New South Wales in Sydney and from the University of Queensland in Brisbane. Surprisingly, given modern New Zealand's dearth of land mammals, there
846-539: The depocenter , river deltas developed and large thicknesses of coal built up on upper delta plains. These are termed the Blackstone Delta and Teviot Delta. Fully lacustrine conditions ensued with the submergence of these deltas and for much of the mid Miocene, Lake Manuherikia was without known bounds. Rising mountains in the Late Miocene - Pliocene eventually deluged Lake Manuherikia with gravel –
893-399: The echolocating microbats . But more recent evidence has supported dividing the order into Yinpterochiroptera and Yangochiroptera , with megabats as members of the former along with several species of microbats. Many bats are insectivores , and most of the rest are frugivores (fruit-eaters) or nectarivores (nectar-eaters). A few species feed on animals other than insects; for example,
940-431: The flying foxes , with the giant golden-crowned flying fox ( Acerodon jubatus ) reaching a weight of 1.6 kg (3.5 lb) and having a wingspan of 1.7 m (5 ft 7 in). The second largest order of mammals after rodents , bats comprise about 20% of all classified mammal species worldwide, with over 1,400 species. These were traditionally divided into two suborders: the largely fruit-eating megabats , and
987-542: The monophyly of bats and the single origin of mammal flight. An independent molecular analysis trying to establish the dates when bat ectoparasites ( bedbugs ) evolved came to the conclusion that bedbugs similar to those known today (all major extant lineages, all of which feed primarily on bats) had already diversified and become established over 100 mya (i.e., long before the oldest records for bats, 52 mya), suggesting that they initially all evolved on non-bat hosts and "bats were colonized several times independently, unless
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#17327730933061034-469: The vampire bats feed on blood . Most bats are nocturnal , and many roost in caves or other refuges; it is uncertain whether bats have these behaviours to escape predators . Bats are present throughout the world, with the exception of extremely cold regions. They are important in their ecosystems for pollinating flowers and dispersing seeds; many tropical plants depend entirely on bats for these services. Bats provide humans with some direct benefits, at
1081-677: The Kawarau Member have been termed the Cromwell Submember and the Ewing Submember. The Ewing is typically intercalated with the Lauder Member, an informal unit of the Bannockburn Formation. This contains a fossil fauna (the Saint Bathans Fauna ) of birds, fish, and a crocodilian, New Zealand's first known terrestrial mammal, as well as bats. Stromatolites are also present. At two locations around
1128-471: The Manuherikia Group is basically Miocene in age. The plant macrofossils , common in the lower Manuherikia Group, are mostly Early Miocene , perhaps with some in the earliest Middle Miocene . Following Douglas, the primary subdivision of the Manuherikia Group is into a lower, fluvial (commonly with coal) Dunstan Formation and an upper, lacustrine Bannockburn Formation . The oldest unit of
1175-580: The Manuherikia Group is the Saint Bathans Member of the Dunstan Formation, consisting of the braided-river fill of valleys incised into the basement rocks. Traces of three St Bathans paleovalleys are known – one at Blue Lake , adjacent to St Bathans , is the best exposed. The St Bathans paleovalleys eventually filled, sedimentation spread out over a broader area, and the fluvial character became dominantly meandering . This material
1222-528: The Maori Bottom, or Maniototo Conglomerate . An extremely diverse macroflora flora, based on well-preserved cuticle. There are two cycad-like taxa: The conifers total 16 species in 12 genera, including: Other gymnosperms: Monocots include: The dicots include: Bats (traditional): (present): Bats are flying mammals of the order Chiroptera ( / k aɪ ˈ r ɒ p t ər ə / ). With their forelimbs adapted as wings , they are
1269-580: The Miocene, with even bats being significantly more diverse than today. New Zealand's two modern palaeognath clades, the kiwi and moa , have early representatives in the fauna. The former is represented by the diminutive, possibly volant Proapteryx . The latter is represented by several bones and egg shells of currently unnamed species, but already identifiable as true moa, being large sized and flightless. The fact that moa are already recognisably modern in anatomy, and possibly ecology, while kiwis are fairly unspecialised and probably still flighted, confirms
1316-492: The West, bats are popularly associated with darkness, malevolence, witchcraft, vampires , and death. An older English name for bats is flittermouse , which matches their name in other Germanic languages (for example German Fledermaus and Swedish fladdermus ), related to the fluttering of wings. Middle English had bakke , most likely cognate with Old Swedish natbakka ( ' night-bat ' ), which may have undergone
1363-614: The air. This suggests that this bat did not fly as much as modern bats, but flew from tree to tree and spent most of its time climbing or hanging on branches. The distinctive features of the Onychonycteris fossil also support the hypothesis that mammalian flight most likely evolved in arboreal locomotors, rather than terrestrial runners. This model of flight development, commonly known as the "trees-down" theory, holds that bats first flew by taking advantage of height and gravity to drop down on to prey, rather than running fast enough for
1410-463: The ancestors of extinct Pleistocene-Holocene Cnemiornis goose, and those of a second possible goose species have been found. In both instances, there is not enough material currently to erect species. Stiff-tailed ducks dominate the fauna with Manuherikia lacustrina , M. minuta , M. douglasi , Dunstanneta johnstoneorum and a further undescribed species of Manuherikia . One species of shelduck , Miotadorna sanctibathansi , has been found and
1457-417: The ancient relatives of today's bullies , galaxiids , and the extinct New Zealand grayling . As well as fishes, shellfish, including freshwater mussels, and freshwater crayfish dominated the aquatic life in the palaeolake Manuherikia. A new species of St Bathans freshwater limpet, Latia manuherikia , was described by malacologist Bruce Marshall in 2011. This was both the first known fossil Latia and
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1504-563: The cost of some disadvantages. Bat dung has been mined as guano from caves and used as fertiliser. Bats consume insect pests, reducing the need for pesticides and other insect management measures. They are sometimes numerous enough and close enough to human settlements to serve as tourist attractions, and they are used as food across Asia and the Pacific Rim . However, fruit bats are frequently considered pests by fruit growers. Due to their physiology, bats are one type of animal that acts as
1551-574: The current land area of New Zealand was submerged. The wildlife that lived in, on, and around the palaeolake Manuherikia was uniquely New Zealand, which strongly suggesting that some emergent land remained during this near drowning event. Marked global cooling and drying during the Miocene, Pliocene and the Pleistocene Ice Ages resulted in the extinction of the 'subtropical' elements of the St Bathans fauna. Those that survived adapted to
1598-528: The dynamic geological and climatic changes, and would form part of the enigmatic fauna that characterised New Zealand when humans arrived in the late 13th century. Research on the St Bathans fauna is led by Trevor Worthy , a New Zealander based in Flinders University , Adelaide. Other key scientists involved include Jenny Worthy from Flinders University, Paul Scofield and Vanesa De Pietri from Canterbury Museum , and Alan Tennyson from
1645-483: The early Eocene , and belong within the four major lines of microbats. Two new suborders have been proposed; Yinpterochiroptera includes the Pteropodidae , or megabat family, as well as the families Rhinolophidae , Hipposideridae , Craseonycteridae , Megadermatidae , and Rhinopomatidae . Yangochiroptera includes the other families of bats (all of which use laryngeal echolocation), a conclusion supported by
1692-546: The evolutionary origin of bats has been grossly underestimated." Fleas , as a group, are quite old (most flea families formed around the end of the Cretaceous ), but no analyses have provided estimates for the age of the flea lineages associated with bats. The oldest known members of a different lineage of bat ectoparasites ( bat flies ), however, are from roughly 20 mya, well after the origin of bats. The bat-ectoparasitic earwig family Arixeniidae has no fossil record, but
1739-491: The first fossil mammals whose colouration has been discovered: both were reddish-brown. Bats were formerly grouped in the superorder Archonta , along with the treeshrews (Scandentia), colugos (Dermoptera), and primates . Modern genetic evidence now places bats in the superorder Laurasiatheria , with its sister taxon as Ferungulata , which includes carnivorans , pangolins , odd-toed ungulates , even-toed ungulates , and cetaceans . One study places Chiroptera as
1786-552: The first record of this genus from the South Island. Notable examples of absent taxa include marsupials , snakes , agamid and varanid lizards, lungfish , eels , cockatoos , and all but one lineage ( bellbirds and tūī ) of the 80 species of Australian honeyeaters . Bannockburn Formation The Manuherikia Group is a fluvial - lacustrine sedimentary fill in the Central Otago area of New Zealand, at
1833-636: The fossil record. Most of the oldest known bat fossils were already very similar to modern microbats, such as Archaeopteropus (32 million years ago). The oldest known bat fossils include Archaeonycteris praecursor and Altaynycteris aurora (55-56 million years ago), both known only from isolated teeth. The oldest complete bat skeleton is Icaronycteris gunnelli (52 million years ago), known from two skeletons discovered in Wyoming. The extinct bats Palaeochiropteryx tupaiodon and Hassianycteris kumari , both of which lived 48 million years ago, are
1880-467: The modern tuatara , geckos , and skinks . However, there are also several species not seen in modern-day New Zealand, such as a mekosuchine crocodile up to 3 metres in length and pleurodire and meiolaniid turtles . This suggests that New Zealand's herpetofauna was much richer in this epoch, probably because its climate was considerably warmer than today. The vast majority of the bones excavated from St Bathans are those of freshwater fish such as
1927-446: The only mammals capable of true and sustained flight . Bats are more agile in flight than most birds, flying with their very long spread-out digits covered with a thin membrane or patagium . The smallest bat, and arguably the smallest extant mammal , is Kitti's hog-nosed bat , which is 29–34 mm (1.1–1.3 in) in length, 150 mm (5.9 in) across the wings and 2–2.6 g (0.071–0.092 oz) in mass. The largest bats are
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1974-617: The previous suspicions that neither clade is closely related and that they arrived in New Zealand independently: moa arrived and became flightless earlier in the Cenozoic , while kiwi were then recent arrivals. Anseriforms (waterfowl) dominate the fauna. At least nine species are recognised from St Bathans, making it the richest waterfowl fauna in the world. All the waterfowl species are unique to New Zealand. Bones attributable to Cape Barren goose ( Cereopsis spp.), thought to represent
2021-406: The relative absence of other parrots. A New Zealand wren , Kuiornis indicator , is known from these deposits, possibly similar to the modern rifleman . Two or three other passerine species remain undescribed. The St Bathans fauna is rich in reptile and amphibian remains. Several groups present in modern New Zealand are represented, such as leiopelmatid frogs , a sphenodontian similar to
2068-460: The sediments were deposited in a series of small, interconnected basins between the mountain ranges, e.g. Park, later workers, like Cotton argued that the sediments had been isolated by later mountain growth. Douglas placed the Manuherikia Group sediments into a coherent genetic context. He drew attention to Manuherikia Group sediments on the top of mountain ranges, and to the observation that sequences in distinct basins were similar. His conclusion
2115-463: The site of the prehistoric Lake Manuherikia . The area consists of a valley and ridge topography , with a series of schist - greywacke mountains at roughly ninety degrees to each other. The Manuherika Group occurs in the current basins, and occasionally on the mountains themselves. One of the earliest geologists to work in the area, McKay , understood that the Manuherikia Group was probably originally continuous. Although some workers came to believe
2162-428: Was no bigger than a sparrow. Charadriiformes , including gulls, terns, noddies, snipes, dotterels, plovers, jacanas, oystercatchers, sheathbills and the plains-wanderer , are a large group of birds that are mostly found in marine or semi-marine environments. There are about 350 species, and they are mostly small to medium-sized. Two of these are known from St Bathans, the New Zealand lake-wanderer ( Hakawai melvillei ),
2209-417: Was that the Manuherikia Group is the result of sedimentation in a single (except for the earliest stages) very large basin that was later intruded by the growth of the mountain ranges. This basin ultimately grew into a single huge lake – Lake Manuherikia – that extended over some 5,600 km (2,200 sq mi). Palynological work by Couper, Mildenhall, Mildenhall and Pocknall has indicated that
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