Environment Australia, 1999
ISBN 0 642 2546 363
Taxonomy and selection of taxa for this Action Plan
Terry Reardon Evolutionary Biology Unit, South Australian Museum
The purpose of this chapter is to:
- review the current state of Australian bat taxonomy;
- describe the decision making process for the selection of taxonomic units;
- summarise the justifications for the selection of taxonomic units where these selections were based on unpublished data; and
- list the known forms of Australian Chiroptera.
In the most recent published review of the state of Australian bat taxonomy, Parnaby (1991) argued that at least two thirds of the then recognised bat species were in need of taxonomic clarification. While several new taxonomic studies have since been completed or commenced (involving species of Pteropus, Nyctimene, Miniopterus, Myotis, Hipposideros, Rhinolophus and Macroderma) most have served only to elucidate further problems. Thus, if the same four criteria used by Parnaby (1991) are applied to the currently known species, then the proportion of species requiring taxonomic clarification has not significantly altered since 1991.
The current problems facing species level systematics within Australian Chiroptera are summarised in Table 2.1. The problems identified have been compiled from both published and unpublished studies, and from the considered opinions of experienced bat biologists familiar with the taxa in question.
|Genus||species||Problem summary||Current research status|
|Pteropus||sp. (Moa Island)||Torres Strait form recognised as a new species based on morphology (Richards and Hall in prep.). This has been disputed (Coles 1997 and Tidemann 1997).||Further taxonomic work in progress (Richards and Hall – University of Queensland)|
|Pteropus||alecto||Two distinct size classes in Australia (Vardon and Tidemann 1997).||unknown|
|Pteropus||macrotis||Suggested that the subspecies epularius will be elevated to full species (Richards and Hall pers. comm.).||unknown|
|Nyctimene||robinsoni||Genetic data failed to distinguish this taxon from New Guinea albiventer (Donnellan et al. 1995). Northern NSW specimens also need to be compared with Queensland specimens (Parnaby pers. comm.).||Genetic work in progress (Irwin, Hall and Moritz – University of Queensland).|
|Nyctimene||cephalotes||Specimens from Torres Strait and a specimen from Cape York designated Nyctimene sp. (cepholotes form) appear morphologically distinct (Hall and Richards pers. comm.).||Genetic work in progress (Irwin, Hall and Moritz – University of Queensland).|
|Hipposideros||diadema||Two subspecies are recognised and are expected to be elevated to species.||Genetic study in progress (Adams – SA Museum).|
|Saccolaimus||saccolaimus||Queensland and NT populations are allopatric and separated by a large distance – possibly two species.||unknown|
|Rhinolophus||all forms||Species boundaries remain unresolved – possibly four species present.||Genetic (Reardon and Cooper – SA Museum) and morphological and echolocation studies (Coles – University of Sydney, and Clague – University of Queensland) in progress.|
|Rhinonicteris||aurantius||Pilbara form differs in morphology and echolocation call (McKenzie and Start pers. comm.)||unknown|
|Myotis||adversus subsp. indet.||The validity of this species is questioned (Reardon).||Genetic studies in progress (Reardon, Cooper, Day – SA Museum).|
|Nyctophilus||timoriensis||Sibling species proposed (Parnaby).||Genetic and morphological studies mostly completed (Parnaby – Australian Museum, Adams – SA Museum); funding required to complete.|
|Nyctophilus||bifax||Sibling species proposed (Parnaby).||as above|
|Nyctophilus||gouldi||Two disjunct populations present, requires taxonomic resolution. There is also morphological variation amongst eastern forms (Parnaby pers. comm.).||Morphological studies mostly completed (Parnaby – Australian Museum); funding required to complete.|
|Pipistrellus||murrayi||Taxonomic affinities to P. tenuis requires clarification (Lumsden pers. comm.).||unknown|
|Miniopterus||schreibersii complex||Sibling species proposed (Reinhold, Reardon, Adams and Lara, in prep – University of Queensland, SA Museum in prep.).||Further genetic and morphological studies in progress (Cardinal – University of Melbourne).|
|Murina||florium||The taxonomic relationship between the Iron Range specimen and Wet Tropics specimens is unresolved (Coles pers. comm.).||unknown|
|Scotorepens||sp. nov.||A morphologically distinct form recognised, taxonomic status unresolved (Parnaby 1992).||Genetic study in progress (Reardon – SA Museum).|
|Scotorepens||balstoni||Morphological and genetic variation may indicate cryptic species (Parnaby pers. comm., Baverstock et al. 1987).||unknown|
|Scotorepens||greyii/sanborni||Species boundaries in this group remain equivocal (Kitchener and Caputi 1985; Parnaby pers. comm.).||unknown|
|Vespadelus||regulus||Genetic discontinuity warrants further examination (Adams et al. 1987).||unknown|
|Vespadelus||vulturnus||Morphological variants suggestive of cryptic species (Parnaby, Hoye pers. comm.)|
|Chalinolobus||morio||Morphological and roosting habitat variation suggests further investigation (Hall and Richards, Parnaby, Hoye pers. comm.).||unknown|
|Mormopterus||most species||Species boundaries unresolved formally.||Genetics published (Adams et al. 1988), morphology finished but requires examination of types, funding required for completion (McKenzie – WA CALM, and Reardon – SA Museum). Funding required to complete.|
Moreover, changes in generic level systematics in the following genera can be expected from comparisons of Australian representatives with supposed extra limital congeners: Macroglossus, Nyctimene, Rhinolophus, Hipposideros, Saccolaimus, Taphozous, Chalinolobus, Kerivoula, Miniopterus, Myotis, Murina, Chaerephon and Tadarida.
It is clear that Australian bat taxonomy is currently in an extremely parlous state, despite the fact that so many of the problems outlined above have been known to bat biologists for over a decade.
If the problems are so readily identifiable, why then has Australian bat taxonomy been so neglected? The answer appears to lie in a combination of lack of commitment to bat taxonomy by professional mammalian systematists, who at least have the institutional resources to undertake the necessary work, and lack of resources (both institutional and financial) available for those who are willing to undertake the necessary work but whose main profession is not taxonomy. The latter circumstance is exemplified by the current status of two major generic revisions, namely Nyctophilus and Mormopterus. These two genera account for 22% of all taxa listed in this Action Plan and comprise nine undescribed species. Both revisions were all but completed many years ago, but their publication has stalled through lack of financial support to the respective investigators involved.
One area of bat systematics which has burgeoned in recent years is that involving molecular biology. The molecular approach to taxonomy has already made an important contribution to resolving cryptic species problems in Australian bats, and will increasingly influence all hierarchical levels of bat systematics. Perhaps the most important role for molecular biology will be in defining ‘Management Units’ and ‘Evolutionarily Significant Units’ which can be used for prioritising conservation effort (Moritz 1994). Thus far, the only Australian bat species studied at this level is the ghost bat, Macroderma gigas (Worthington-Wilmer et al. 1994).
Taxonomic research is fundamental to the conservation of Australia’s bat fauna because it is impossible to conserve a taxon until it has been described. The slow pace of resolution of taxonomic problems is recognised as a major impediment to bat conservation in Australia. As a result support for taxonomic research has been set as a high priority in this Action Plan.
The definition of taxa for the Action Plan has involved rigorous consultation with Australian bat researchers. Mr Greg Richards and Dr Les Hall initially compiled a ‘List of Australian Bat Taxa’ for the July 1996 Draft Action Plan, based primarily on Mahoney and Walton (1988), but modified to accommodate subsequently published accounts and their knowledge of unpublished data. This list formed the basis for discussion at the June 1997 workshop (refer Chapter 1, this volume). Several amendments (additions and deletions) were made to the list, again based on newly available and unpublished data. Inevitably, there was debate over some decisions and, while not all workshop participants supported all the decisions, a list was finally agreed upon. Further debate on particular taxa ensued over the following 12 months. In April 1998 the Editorial Panel convened, considered arguments for defining taxa still in dispute, and determined a final list of taxa for the Action Plan.
The IUCN Red List Categories booklet explicitly states that criteria for listing of taxa can be applied to ‘any taxonomic unit at or below the species level … including forms that are not yet formally described’ (IUCN 1994). The guiding principle adopted by Richards and Hall, the June workshop participants and the Editorial Panel was that for the Action Plan to be relevant for at least five years, it should take account of all data available, including unpublished data, when defining taxonomic units.
In deciding to use unpublished taxonomic studies (those which propose the recognition of new species) and published genetic studies (which delineate species boundaries but have no formal diagnoses or nomenclature), the Editorial Panel has been careful to avoid pre-empting unpublished scientific taxonomic names. It has also been recognised that considerable time may pass before the backlog of species-level taxonomic problems are solved, and therefore the nomenclature used here may become widely adopted. The application of names for some taxonomic units proved to be challenging.
Common names have been given to each taxon, whether it be a formally described species or subspecies, anticipated new species or conservation unit. The common names used were derived either from Richards et al. (1993) or newly applied by the Editorial Panel.
Scientific names have been dealt with in the following manner. Since most of the proposed (unpublished) new species are elevations from subspecies, the taxa are simply referred to by their subspecific names (denoted in Table 2.2 with an asterisk). It should be noted that not all subspecies listed are proposed for elevation to full species. Where anticipated species complexes have been delineated, each taxon is referred to by the parent species name but distinguished as a geographic ‘form’. For some proposed or suspected new species, the term ‘sp.’ is used with the corresponding generic name. The naming of taxa in the genus Mormopterus has been particularly difficult and it remains somewhat clumsily dealt with here. Four of the species have existing names which are considered to be correctly assigned, but the identities of the remaining five are uncertain and are therefore referred to as ‘sp.’ together with a reference to the population defined in the published genetic study of Adams et al. (1988).
Because the selection of taxa has relied so heavily upon unpublished data and opinion, and because some of the decisions to include or leave out specific taxa have been strongly debated, the rationale for the selection or omission of taxa are outlined below.
Pteropus sp. (Moa Island)
There has been considerable contention over the recognition of this taxon as being distinct from P. alecto, (both are sympatric on Moa Island, Torres Strait). This putative new species was first distinguished by G.C. Richards and L.S. Hall based on morphology (it is smaller in many characters, significantly different in cranial characters and has a different shaped baculum). A paper describing the new species was submitted to the Australian Zoologist and was ‘returned to the authors for revision’ – this paper is currently being revised (Richards and Hall in prep.). The June workshop discussed this issue at length and finally agreed (but with dissenters) to include it on the list. Subsequently, the specimens used in the Richards et al. study were re-examined and those specimens previously assigned to the new ‘species’ were judged to be either juvenile or neotenous P. alecto (R. B. Coles 1997; C.R. Tidemann 1997). Richards and Hall (pers. comm.) reject these findings. This issue remains unresolved. The Editorial Panel, having considered the evidence, agreed to include this taxon as Pteropus sp. in the Action Plan, but to make taxonomic resolution a precursor to allocation of further funding for conservation.
Nyctimene cephalotes and N. cf. vizcaccia
Specimens referrable to N. cephalotes have been collected from Silver Plains, Cape York and from Moa Island, Torres Strait. The forms from each locality are morphologically distinct (Hall and Richards pers. comm.). The Moa Island specimens were previously identified as N. vizcaccia (Richards 1995), however Bonnacorso (in press) refers to the same specimens as being N. cephalotes.
The species level taxonomy of Australo-Papuan Nyctimene is not fully resolved (Donnellan et al. 1995). The Editorial Panel has accepted the opinion of Hall and Richards that the two forms are morphologically distinct but treat both forms under the name N. cephalotes.
The species-level taxonomy of Australian rhinolophids has been problematical. Two species are recognised formally, namely R. megaphyllus and R. philippinensis. For nearly two decades a third form, intermediate in size between the two named species, has been known. This form has a distinct echolocation call (Coles 1993) and, although morphologically distinct, has nose-leaf characters similar to R. philippinensis (Churchill 1998). Recent genetic analysis (at a level usually capable of distinguishing between individuals at the population level) has failed to distinguish between the intermediate size form and R. philippinensis. Most of the workshop panel who are familiar with these forms, agree on treating the intermediate size form as a separate taxon. They have both been named here as R. philippinensis, large and small form, although it is not resolved which of the two, or in fact if either, are referable to true philippinensis.
The recognition of two forms within R. megaphyllus is based on recent convincing DNA evidence (Cooper, Reardon and Skilins 1998). Although the distribution boundaries have not been well defined, the populations are essentially separated into north and south forms. To distinguish them, the earlier discarded subspecies names have been reapplied: R. m. megaphyllus for the southern form and R. m. ignifer for the northern form.
Morphological characters of the Pilbara population reflect a functional requirement to forage efficiently in low productivity environments of the arid Pilbara region. On the basis of the differences listed below (N.L. McKenzie and A.N. Start pers. comm.), two taxa are recognised in this Action Plan.
|Region||Echolocation frequency for CF component (kHz) (range)||Wing aspect ratio (s.d.)||Wing loading (g/cm²) (s.d.)||N|
|Pilbara||125 (122–128)||6.54 (0.04)||0.066 (0.002)||6|
|Kimberley||112 (110–119)||6.29 (0.01)||0.058 (0.04)||4|
Although the species level taxonomy of Mormopterus is now mostly resolved in an informal sense, the inconsistent use of common names has served to make this genus one of the most poorly known in Australia. Independently conducted revisions based on genetics (Adams et al. 1988) and morphology (McKenzie unpub.) are in agreement on species boundaries. Based on the these taxonomic revisions, nine taxa are recognised. However, nomenclatural clarification is contingent upon examination of type material, which is held in overseas institutions.
Parnaby (1992) reported on a new form of Scotorepens from coastal eastern Australia. This form differs on skull, dental and external morphology from S. balstoni and S. orion, and differs from S. greyii on skull morphology (Parnaby unpub.). This taxon has been recognised as a distinct species, Scotorepens sp., for the purposes of the Action Plan.
The 1997 workshop accepted the opinion of Parnaby (1988), based on his revision of the genus Nyctophilus, that N. timoriensis contained three morphologically distinct forms.
Myotis adversus complex
The most recently published taxonomic revision of Myotis adversus is by Kitchener et al. (1995). Based on morphological variation, M. adversus (sensu lato) was split into three species – M. macropus, M. adversus and M. moluccarum. Whilst there has been widespread acceptance of two of the species in Australia, there remains considerable scepticism over the third (M. adversus). The main arguments are that the evidence for recognising M. adversus in Australia was based on measurements of only a single specimen (from near Lismore New South Wales), and that the Lismore specimen was the only New South Wales specimen included in the study – there was a large gap in sampling in New South Wales.
The nearest locality elsewhere for M. adversus (sensu stricto) is in Indonesia. Kitchener et al. (op cit.) were tentative about the placement of the New South Wales specimen in adversus, and inspection of the Discriminate Function Analysis (DFA) plots reveals that the placement of this specimen is speculative with respect to all three species. The reliability of the DFA analysis is further diminished because no other New South Wales specimens were included.
Analysis of mitochondrial DNA sequence data (Day, Cooper and Reardon unpub.), which used the same Lismore specimen included in the Kitchener et al. study, found:
- Indonesian adversus forms a distinct clade from the clade that includes all Australian Myotis;
- there are two clades within Australia but the genetic distance between them gives only weak support for the recognition of two species; and
- the Lismore specimen and one south-eastern Queensland specimen fall in the macropus clade.
On this evidence, M. adversus has been excluded from the list of Australian taxa.
In a taxonomic review of Australo-Papuan Miniopterus, Reinhold (1997) presented molecular and morphological evidence to support the recognition of at least two species within M. schreibersii within Australia. Although formal publication of these results will be some months away, it is expected that M. s. orianae will be elevated to full species status, leaving M. s. oceanensis as the form in eastern Australia. Reinhold did not examine the extreme southern populations of M. schreibersii (those from Naracoorte, South Australia and Warrnambool, Victoria) which have been considered as being distinct from eastern Australian forms based on morphology, genetics and breeding patterns (Hamilton-Smith 1972, Hand 1980, Maeda 1982, Wilson 1982, Cardinal 1997, Reardon unpub.). Indeed, Maeda (1982) formally recognised the southern forms as part of the M. macrodens group, which is otherwise widely distributed through Indonesia and Malaysia. However Maeda’s work has not been widely accepted. The Editorial Panel felt there was sufficient evidence to recognise three forms of M. schreibersii in Australia, the two named subspecies, orianae and oceanensis , and the geographical variant based on the Naracoorte population (southern form).
The list ultimately adopted by the Editorial Panel contains 90 taxa (Table 2.2).
|Pteropodidae (Fruit bats)|
|Pteropus poliocephalus Temminck,1825||Grey-headed Flying-fox|
|Pteropus scapulatus Peters,1862||Little Red Flying-fox|
|Pteropus alecto gouldi Peters, 1867||Black Flying-fox|
|Pteropus sp.* (Moa Island)||Torresian Flying-fox|
|Pteropus brunneus Dobson,1878||Percy Island Flying-fox|
|Pteropus conspicillatus Gould,1850||Spectacled Flying-fox|
|Pteropus macrotis epularius * Ramsay, 1878||Large-eared Flying-fox|
|Pteropus melanotus natalis Thomas, 1887||Christmas Island Flying-fox|
|Dobsonia moluccensis magna Thomas, 1904||Bare-backed Fruit Bat|
|Nyctimene robinsoni Thomas, 1904||Eastern Tube-nosed Bat|
|Nyctimene cephalotes (Pallas, 1767)||Torresian Tube-nosed Bat|
|Syconycteris australis (Peters,1867)||Eastern Blossom Bat|
|Macroglossus minimus nanus Matschie, 1899||Northern Blossom Bat|
|Megadermatidae (Ghost bats)|
|Macroderma gigas (Dobson, 1880)||Ghost Bat|
|Rhinolophidae (Horseshoe bats)|
|Rhinolophus megaphyllus ignifer* Allen, 1933||Northern Horseshoe Bat|
|Rhinolophus megaphyllus megaphyllus* Gray, 1834||Southern Horseshoe Bat|
|Rhinolophus philippinensis Waterhouse, 1843:|
|(large form)||Greater Large-eared Horseshoe Bat|
|(small form)||Lesser Large-eared Horseshoe Bat|
|Hipposideridae (Leaf-nosed bats)|
|Hipposideros ater aruensis Gray, 1858||Eastern Dusky Leaf-nosed Bat|
|Hipposideros ater gilberti Johnson,1959||Western Dusky Leaf-nosed Bat|
|Hipposideros cervinus (Gould,1854)||Fawn Leaf-nosed Bat|
|Hipposideros diadema inornatus* McKean, 1970||Arnhem Leaf-nosed Bat|
|Hipposideros diadema reginae* Troughton, 1937||Diadem Leaf-nosed Bat|
|Hipposideros semoni Matschie, 1903||Semon's Leaf-nosed Bat|
|Hipposideros stenotis Thomas, 1913||Northern Leaf-nosed Bat|
|Rhinonicteris aurantius (Gray, 1845)||Orange Leaf-nosed Bat|
|Rhinonicteris aurantius (Pilbara form)||Pilbara Leaf-nosed Bat|
|Emballonuridae (Sheathtail bats)|
|Taphozous australis Gould, 1854||Coastal Sheathtail Bat|
|Taphozous georgianus Thomas, 1915||Common Sheathtail Bat|
|Taphozous hilli Kitchener, 1980||Hill's Sheathtail Bat|
|Taphozous kapalgensis McKean and Friend, 1979||Arnhem Sheathtail Bat|
|Taphozous troughtoni Tate, 1952||Troughton's Sheathtail Bat|
|Saccolaimus flaviventris (Peters, 1867)||Yellow-bellied Sheathtail Bat|
|Saccolaimus mixtus Troughton, 1925||Papuan Sheathtail Bat|
|Saccolaimus saccolaimus nudicluniatus (De Vis, 1905)||Bare-rumped Sheathtail Bat|
|Molossidae (Freetail bats)|
|Tadarida australis (Gray, 1838)||White-striped Freetail Bat|
|Chaerephon jobensis plicatus (Thomas, 1906)||Northern Freetail Bat|
|Mormopterus beccarii Peters, 1881||Beccari's Freetail Bat|
|Mormopterus loriae cobourgiana* Johnson, 1959||Little North-western Freetail Bat|
|Mormopterus loriae ridei* Felten, 1964||Little North-eastern Freetail Bat|
|Mormopterus norfolkensis (Gray,1839)||East Coast Freetail Bat|
|Mormopterus sp.* (form sp.4, Populations P,Q and R, in Adams et al. 1988)||South-eastern Freetail Bat|
|Mormopterus sp.* (form sp.3 in Adams et al. 1988)||Inland Freetail Bat|
|Mormopterus sp.* (form sp.4, Population O, in Adams et al. 1988)||South-western Freetail Bat|
|Mormopterus sp.* (form sp.2 in Adams et al. 1988)||Eastern Freetail Bat|
|Mormopterus sp.* (form sp.6 in Adams et al. 1988)||Hairy Rostrum Freetail Bat|
|Vespertilionidae (Ordinary bats)|
|Chalinolobus dwyeri Ryan, 1966||Large-eared Pied Bat|
|Chalinolobus gouldii (Gray, 1841)||Gould's Wattled Bat|
|Chalinolobus morio (Gray, 1841)||Chocolate Wattled Bat|
|Chalinolobus nigrogriseus (Gould, 1856)||Hoary Wattled Bat|
|Chalinolobus picatus (Gould, 1852)||Little Pied Bat|
|Falsistrellus mackenziei Kitchener, Caputi and Jones, 1986||Western False Pipistrelle|
|Falsistrellus tasmaniensis (Gould, 1858)||Eastern False Pipistrelle|
|Kerivoula papuensis Dobson, 1878||Golden-tipped Bat|
|Miniopterus australis (Tomes, 1858)||Little Bent-wing Bat|
|Miniopterus schreibersii orianae* Thomas, 1922||Northern Bent-wing Bat|
|Miniopterus schreibersii oceanensis Maeda, 1982||Eastern Bent-wing Bat|
|Miniopterus schreibersii (Kuhl, 1819)|
|(Southern form)||Southern Bent-wing Bat|
|Murina florium Thomas, 1908||Tube-nosed Insectivorous Bat|
|Myotis macropus Gould, 1855||Southern Myotis|
|Myotis moluccarum Thomas, 1915||Northern Myotis|
|Nyctophilus arnhemensis Johnson, 1959||Arnhem Long-eared Bat|
|Nyctophilus bifax bifax* Thomas, 1915||Northern Long-eared Bat|
|Nyctophilus bifax daedelus* Thomas, 1915||Pallid Long-eared Bat|
|Nyctophilus geoffroyi Leach, 1821||Lesser Long-eared Bat|
|Nyctophilus gouldi Tomes, 1858||Gould's Long-eared Bat|
|Nyctophilus howensis McKean, 1975||Lord Howe Long-eared Bat|
|Nyctophilus timoriensis sherrini* Thomas, 1915||Tasmanian Long-eared Bat|
|Nyctophilus timoriensis major* Gray, 1844||Western Long-eared Bat|
|Nyctophilus timoriensis (Geoffroy, 1806):|
|(Central form)||Central Long-eared Bat|
|(South-eastern form)*||Eastern Long-eared Bat|
|Nyctophilus walkeri Thomas, 1892||Pygmy Long-eared Bat|
|Pipistrellus adamsi Kitchener, Caputi and Jones, 1986||Cape York Pipistrelle|
|Pipistrellus murrayi Andrews, 1900||Christmas Island Pipistrelle|
|Pipistrellus westralis Koopman, 1984||Northern Pipistrelle|
|Scoteanax rueppellii (Peters, 1866)||Greater Broad-nosed Bat|
|Scotorepens balstoni (Thomas, 1906)||Inland Broad-nosed Bat|
|Scotorepens greyii (Gray, 1843)||Little Broad-nosed Bat|
|Scotorepens orion (Troughton, 1937)||South-eastern Broad-nosed Bat|
|Scotorepens sanborni (Troughton, 1937)||Northern Broad-nosed Bat|
|Scotorepens sp.*||Central-eastern Broad-nosed Bat|
|Vespadelus baverstocki Kitchener, Jones and Caputi, 1987||Inland Forest Bat|
|Vespadelus caurinus (Thomas, 1914)||Northern Cave Bat|
|Vespadelus darlingtoni (Allen, 1933)||Large Forest Bat|
|Vespadelus douglasorum Kitchener, 1976||Yellow-lipped Cave Bat|
|Vespadelus finlaysoni Kitchener, Jones and Caputi, 1987||Finlayson's Cave Bat|
|Vespadelus pumilus (Gray, 1841)||Eastern Forest Bat|
|Vespadelus regulus (Thomas, 1906)||Southern Forest Bat|
|Vespadelus troughtoni Kitchener, Jones & Caputi, 1987||Eastern Cave Bat|
|Vespadelus vulturnus (Thomas, 1914)||Little Forest Bat|
Note: * taxa which are expected to be recognised as full species.
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