Journal of Insect Conservation 8: 209–220, 2004. Ó 2004 Kluwer Academic Publishers. Printed in the Netherlands.
An overview of the Tasmanian geometrid moth fauna (Lepidoptera: Geometridae) and its conservation status Peter B. McQuillan School of Geography and Environmental Studies, University of Tasmania, Private Bag 78, Hobart, Tasmania 7001 (e-mail: [email protected]) Received 16 June 2004; accepted in revised form 18 June 2004
Key words: Australia, Development pressure, Distribution, Forestry, Island, Threats
Abstract A brief review of the geometrid fauna of the large island of Tasmania and a simple analysis of its conservation status and threats are presented. The fauna comprises 310 species of which Ennominae contribute slightly less than half the total and Larentiinae one third; 23% of the geometrid fauna is endemic at species level. Mixed eucalypt-rainforest is identiﬁed as the richest wet forest habitat in geometrid species. Using distribution data at 10 km resolution, the most widespread and most restricted taxa are identiﬁed. The conservation status of Lepidoptera living above 800 m is relatively good. However, coastal species and those associated with herb-rich native grasslands are under some pressure from habitat change. Three species of geometrid moths are listed as threatened in Tasmania’s Threatened Species Protection Act 1995 but several others may qualify for listing.
Introduction Tasmania is a large (68,000 km2) continental island separated from mainland Australia by the 150 km wide Bass Strait and has been an intermittent island and mesic refuge since the Miocene (Figure 1). Situated between latitudes 40– 43°S, Tasmania is noteworthy for its very diverse range of habitats, from temperate rainforest to alpine herbﬁelds, resulting from its mountainous terrain, maritime climate and strong longitudinal rainfall gradient. It is a key stronghold of the socalled Bassian biome which occupies the moist southern temperate part of Australia. With a low human population density (7/km2) the island has generally suﬀered less severe environmental degradation than the adjacent mainland (RPDC 2003).
The family Geometridae is a large and conspicuous element in the Lepidoptera fauna of the island. Although Tasmania only comprises about 1% of the land area of Australia, it is home to 24% of the described geometrid fauna (310 of 1312 species, although this will probably decline to about 15% when the continental fauna is more fully known). This paper presents a summary of the Tasmanian geometrid fauna and its association with various habitats, and seeks to identify those taxa which are of conservation interest along with actual and emerging threats to their survival. Methods These records are taken from my database of Tasmanian Geometridae records and foodplants maintained at the University of Tasmania. Sam-
210 possible, sampling was conducted on nights suitable for moth activity where temperatures were at or above average and little or no rain was falling. Only data collected since 1975 was admitted to the analysis due to uncertainties in identiﬁcation in early literature records. The checklist draws upon the early records summarised and extended by Turner (1926a, 1926b, 1928, 1939) and supplemented by more recent surveys conducted largely by the author since 1980, plus Edwards and McQuillan (1998). Species which occurred in 20 or more squares were considered to be widespread, while those occurring in less than 3 squares were determined to be restricted and possibly worthy of future conservation assessment. Nomenclature largely follows McQuillan and Edwards 1996). Results Fauna Figure 1. Map showing location of Tasmania with Tasmanian Wilderness World Heritage Area highlighted.
pling for geometrid moths has covered every month of the year and represents collections from a total of 166 separate 10-km grid squares (ca 20% of the state). Survey methods include light trapping at night, opportunistic hand collecting by day, and rearing of larvae beaten from foodplants. Light trapping used both 12 volt, 8 watt ultraviolet lights and 240 volt, 125 watt mercury vapour lights. The collection eﬀort is uneven geographically, but 34 squares have been sampled on more than 20 separate occasions while 27 have only been sampled once. Six squares have been sampled on more than 50 days. Although weather conditions can strongly inﬂuence sampling eﬃciency, records are additive and the inﬂuence of poor nights is diluted in the multiply sampled squares. Where
The Checklist (Appendix 1) shows that the fauna totals 310 species distributed within 119 genera. Table 1 dissects the composition of the fauna by subfamily. Endemism is 23% at family level, and is highest in the subfamily Ennominae and lowest in Oenochrominae and Sterrhinae. The larentiine Chrysolarentia (tribe Xanthorhoini) is outstanding as the largest genus with 35 species, followed by the ennomine Amelora with ten. Both genera are associated with herbs as larvae. Larentiinae tend to have more species per genus than other subfamilies. The Sterrhinae are poorly represented although an endemic genus and species, Dasybela achroa, is present. Endemism is notably low in the ennomine tribe Boarmiini which has 29 species of which only one is restricted to south west Tasmania. At least 4 boarmiines are polyphagous and others feed on
Table 1. Composition of the Tasmanian Geometridae by subfamily Subfamily
Endemic spp. (%)
Largest genus (spp.)
Ennominae Larentiinae Geometrinae Oenochrominae s.l. Oenochrominae s.s. Sterrhinae Total
211 widespread hosts such as Acacia, Eucalyptus and Exocarpos.
rinae are thought to be so localised. Only 16 of the restricted species are known from a secure reserve.
Widespread and restricted species Forest species Table 2 lists the taxa which meet the deﬁnitions of widespread and restricted. The 32 widespread species all occur on mainland Australia in addition to Tasmania, whereas 14 of the 35 restricted species are endemic to the island and one, Epyaxa sodaliata, may be a vagrant. No Geomet-
Mixed forests, which are intermediate between tall eucalypt and rainforest showed the highest diversity of geometrid species (about half the state total), with rainforest less than half as rich (Table 3).
Table 2. List of the most widespread (20 or more squares) and most restricted (less than 3 squares) Tasmanian Geometridae based on 10-km grid squares (sqs) occupied. Habitats for restricted species: A = alpine, DF = dry eucalypt forest, G = grassland, H = health, M = moorland, MF = mixed forest, R = rainforest, WT = wet tall forest. Widespread species Ennominae Didymoctenia exsuperata Casbia melanops Scioglyptis lyciaria Phelotis cognata Archephanes zalosema Idiodes apicata Ectropis excursaria Oenochrominae Epidesmia hypenaria Geometrinae Hypographa phlegetonaria Chlorocoma assimilis Heliomystis electrica Sterrhinae Scopula rubraria Idaea costaria Idaea inversata Larentiinae Epyaxa subidaria Hydriomena severata Chloroclystis ﬁlata Hydriomena squamulata Microdes villosata Anachloris uncinata Microdes squamulata Poecilasthena pulchraria Chrysolarentia vicissata Chrysolarentia epicteta Epicyme rubropunctaria Chloroclystis testulata Eccymatoge morphna Hydriomena microcyma Anachloris subochraria Chrysolarentia heliacaria Chrysolarentia mecynata Xanthorhoe anthracinata
212 Table 3. Summary of geometrid genus and species level diversity in three wet forest types in Tasmania Forest type
Tall eucalypt Mixed forest Rainforest
188 151 92
10 14 6
29 37 16
22 25 14
116 168 73
Formally listed species Three geometrid moths are formally listed as threatened in Tasmania under the state’s Threatened Species Protection Act 1995:
analysis of the family. Preliminary insights from molecular data for example, suggest that the endemic Dirce-Acalyphes clade of alpine moths is better treated as ennomine rather that archiearine (C. Young, pers. comm.). The relatively high endemism in the fauna has analogues in some beetle families such as Lucanidae (>80%) and may reﬂect the diversity of Tasmania’s habitats and its relative isolation. Bass Strait has been both a bridge and a barrier to faunal interchange with the mainland and serves to reinforce the unique character of the fauna. Sister species of Lepidoptera are commonly separated by Bass Strait (e.g. Aenetus, Argyninna) or the same species may occupy diﬀerent habitats on either side of the Strait such as the butterﬂy Oreisplanus munionga.
Chrysolarentia decisaria Endangered Last collected in Tasmania near Cressy in 1904, this species was presumed extinct until a precarious population was found at Township Lagoon in the Midlands in 1996. The tiny area (10 ha) is reserved for multiple natural values but is surrounded by farmland. It appears to be rare in Victoria with few recent records.
Amelora acontistica Vulnerable This coastal saltmarsh/dune species is known from 2 grid squares in south eastern Tasmania. It also occurs in similar habitat on Kangaroo Island, South Australia.
Dasybela achroa Vulnerable A coastal saltmarsh/dune species known from 1 grid square in south eastern Tasmania where it is endemic.
Discussion High conservation value attaches to those species which are rare, restricted, endemic to Tasmania, or which are phylogenetically important. A number of species have been shown to meet these criteria but the latter value remains poorly appreciated in the absence of more comprehensive cladistic
Grassland species Grassland dependant species include some of the most widespread as well as the rarest and most restricted taxa. The large guild of xanthorhoine larentiines (50+ spp., notably Epyaxa, Chrysolarentia and an undescribed genus) in Tasmania is responsive to grassland management, an outcome driven to some extent by foodplant availability (McQuillan 1999) and plant community dynamics. In contrast to mainland Australia, grazing pressure by native mammals is high in the native grasslands of Tasmania and especially in subalpine areas (Bridle and Kirkpatrick 1999). So-called marsupial lawns are maintained in many areas by the grazing pressure of wombats, wallabies and other small macropods. Poa hiemata and other grasses are continually trimmed to low stature. Consequently, these grasslands are rich in taxa of small interstitial herbs which serve as geometrid foodplants. Another locally important agent of grass suppression is grazing by invertebrates. The hepialid moth genus Oncopera is something of a keystone taxon in Tasmanian Poa tussock grasslands, capable of reaching local densities of more than 50 late instar larvae per square metre in winter, although dense populations become self-limiting through disease or starvation. The nocturnal larvae do not readily climb but chew through stems near the base of large tussocks causing the grass to fall as detritus on the ground where it forms a loose mat. This decays fairly quickly and small herbs germinate in
213 the sunlit open patches. Within about 5 years the tussock canopy is restored to its usual size and the biomass of ground level herbs declines rapidly. In eastern and southern Tasmania the endemic O. intricata is most common at lower elevations down to sealevel, but is increasingly replaced by O. rufobrunnea above 400 m and in higher rainfall grasslands and grassy woodlands in the north and north west. Although similar biologically, they diﬀer in their phenology, with O. intricata having a pupal aestivation. These habitats are ecologically dynamic at small geographical scales and interactions can be complex, involving ﬁre, cold air drainage, frost, competition and grazing. Ecotones can be sharp or diﬀuse and cold air drainage is increasingly inﬂuential in suppressing trees and shrubs in valley bottoms above 500 m elevation. The lepidopteran fauna of these grasslands is locally rich (at least 11 families feed on Poaceae alone in Tasmania) although Castniidae are absent from the island. Endemism in the geometrids is highest in the montane grasslands and herbﬁelds where ﬁre is uncommon due to high moisture levels and cool conditions. Changes in the soil fertility of these grasslands due to ‘improvement’ can be detrimental to native Lepidoptera. The addition of phosphatic fertilisers (usually superphosphate) or farm animal excreta to the ground layer promotes the vertical shoot growth of invasive annuals, increasing their leaf area index and gradually outcompeting native plants of low stature (e.g. O’Dwyer and Attiwill 1999). The wind dispersed seeds of composites (e.g. Hypochaeris radicata) and grasses (Vulpia spp.) colonise bare ground and their establishment is favoured by increased nutrient status (Specht 1963). In the richest sites Arctotheca calendula can establish, favouring Ciampa arietaria one of the few geometrids to eat introduced herbs. Other weedy annuals such as Plantago lanceolata may promote locally high densities of the widespread Scopula rubraria and Epyaxa subidaria. Modest diversity of native moths can be maintained in moderately disturbed sites (McQuillan 1999). There are parallels to grassland butterﬂies dependent on native Poa tussock grasses as larval food. Oreixenica ptunarra is a rare Tasmanian endemic which can tolerate moderate levels of sheep grazing provided tussocks are maintained, and can use nectar from autumn-ﬂowering weeds, especially Hypochaeris, in the absence of native sources.
Eucalyptus forest species Tall eucalypt forests, with their sparse canopies reaching to 95 metres, shed large amounts of woody debris and create special microclimates which favour a suite of tall shrubs and the development of bryophytes and ferns near the ground. In the absence of wildﬁre, the geometrid fauna is gradually enriched by a trend towards increasing tree diversity as rainforest elements invade and begin to suppress the juvenile eucalypt trees. These mixed forests support a larger diversity of Geometridae than adjacent eucalypt forest or rainforest, the latter being the climax vegetation on some sites. Mixed forests may take from one to ﬁve centuries to develop. Dry eucalypt forests and woodlands typical of the east coast may have grassy understoreys and a sparse shrub layer including foodplants such as Fabaceae, Epacridaceae and Santalaceae. The understorey Acacia shrubs are an important foodplant for genera such as Thalaina and Ectropis. Common polyphagous forest taxa include Chlenias and Androchela. Forest dwelling geometrids include many species which use canopy Eucalyptus as foodplants (McFarland 1988). Paralaea and Mnesampela may reach high population densities and achieve local pest status (Elliott et al. 1980, McQuillan 1985). Several genera feeding on Eucalyptus include related species pairs which are separated by altitude into a lowland and a restricted, sometimes endemic, highland sibling; for example, Plesanemma fucata–P. altafucata (McQuillan 1984), Paralaea beggaria–P. chionoptera and Cyneoterpna wilsoni–C. alpina. In some respects, eucalypt forests in Tasmania support an attenuated mainland fauna e.g. arboreal taxa such as Mochlotona are missing, but they are also enriched by local speciation in otherwise widespread Bassian genera such as Paralaea.
Rainforest species Temperate rainforests, with Nothofagus, Atherosperma and Eucryphia as dominant tree genera, have a more limited but distinctive moth fauna. Tasmanian rainforests underwent signiﬁcant range contraction during the Pleistocene ice ages (Kirkpatrick and Fowler 1998) and this frag-
214 mentation may have caused incremental losses in the insect fauna which were possibly additive over time. Nothofagus feeders include Chlenias and Euloxia among the Geometridae as well as nepticulids, tortricids and a roeslerstammid. Archephanes zalosema is on Tasmannia (Winteraceae) (Young and McQuillan 2003) and several boarmiine taxa feed on woody plants including Atherosperma (Monimiaceae). The enigmatic larentiine genus Chaetolopha (Schmidt 2002) is associated with soft ferns in the forest understorey, while Austrocidaria erasta is an uncommon endemic associated with Coprosma (Rubiaceae). Several lithinine Idiodes are common in the vicinity of tree ferns (Dicksonia antarctica).
Saltmarsh species Saltmarshes occupy some of the driest parts of the state (450 mm annual rainfall) but have limited extent (Richardson et al. 1998). Typical geometrid species of this habitat are Amelora acontistica, Dasybela achroa, Scopula rubraria and Epyaxa hypogramma, all known or suspected to be associated with herbaceous foodplants. The ﬁrst two species are listed as vulnerable under the Threatened Species Protection Act 1995. Although Chlenias species are known to feed on Chenopodiaceae on mainland Australia, Tasmanian saltbushes do not support the genus. Some saltmarshes also support the butterﬂy Theclinesthes serpentata lavara (Couchman) which is regarded as rare under the Act.
Alpine species The highlands of the south eastern mainland have strong faunal links in Lepidoptera to Tasmania, but there is a higher diversity of alpine geometrids in the latter. The maritime climate keeps minimum temperatures suﬃciently high to allow alpine shrubs to occur at the summit of most mountains, although herbﬁelds are widespread. Annual rainfall can locally reach 5000 mm. Novelty in the Tasmanian moth fauna increases with elevation up to 1600 m. Noteworthy alpine taxa include Dirce and Acalyphes and diversiﬁcation in the geometrid tribe Xanthorhoini.
Threats Although a large proportion of the Tasmanian landmass above 800 m elevation is reserved (RPDC 2003), elsewhere a number of prevailing theats mean that the prognosis for some of the Tasmanian geometrid fauna remains uncertain, and could worsen unless the pressures are suitably abated. Land clearing There is little doubt that geometrid populations can decline as forest cover is depleted. In South Australia, the large oenochromine Gastrophora henricaria was present in the tall eucalypt forests of the Mount Lofty Ranges in the nineteenth century (Guest 1887) but now persists at a single locality following clearing and fragmentation of its habitat in the century since. Native forest felling is at unprecedented levels in Tasmania since 1997, with ﬁve million tonnes of woodchips exported annually. Several thousand hectares of native habitat are cleared annually much of it of types poorly represented in the conservation estate (Kirkpatrick 1991). Tasmania remains the only state in Australia without land clearing legislation and this is unlikely to change in the near future. The loss and degradation of ice age refugia in eastern Tasmania is likely to be serious (Kirkpatrick and Fowler 1998). Inundation of native grasslands Recent state government policy to double the gross value of agricultural production in the state within a decade is predicated on a dramatic increase in irrigation and dams along with intensiﬁcation of farming. Creation of water markets is encouraging farmers to dam the upper reaches of streams on their properties and the incremental loss of herbenriched grassy valley bottoms to innundation is a serious problem. These streams are often on shallow gradients and their damming inundates and reconﬁgures large areas of native habitat. Global warming Climate change will increase dryness in the east of Tasmania (CSIRO 2001) as well as stress alpine habitats and foodplants. Climate change is likely to be magniﬁed at higher latitudes and altitudes, and therefore the Tasmanian biota in general may be at greater risk from this pressure than species most parts of Australia.
215 Decline in the range of foodplants Some insect species are rare because their foodplants are rare (Hopkins et al. 2002). However, with the exception of conifer-feeding geometrids (Dirce, Acalyphes, Corula) in Tasmania, few are likely to qualify as rare for this reason. Grazing management changes weediness leading to the local shading out of foodplants which is identiﬁed as a threat to geometrids elsewhere (e.g. Sinclair 2002). Sheep grazing has been a major seasonal land use in Tasmania’s native grasslands since the 1830s. Sheep are run at generally low stocking rates in ‘unimproved’ native grasslands and grassy woodlands and may keep tussock crowns small. However, their selective feeding can favour unpalatable shrubs such as Hakea and Ozothamnus to form dense stands which can exclude native herbs. Coastal habitat loss Burnley and Murphy (2004) document an accelerating social trend of movement of people from urban centres to the coastal fringe in eastern and southern Australia, leading to strip development on fargile coastal habitats. In Tasmania the state coastal policy is moribund at a time of rapid alienation of the coast for private housing and tourist developments. Sinclair (2002) discussed the threats to a coastal geometrid in New Zealand and noted the need for eﬀective negotiations with landowners for sympathetic land management. Inappropriate ﬁre regimes Although ﬁre is a natural and important factor in the ecology of many lowland plant communities in Tasmania most alpine tree species are highly intolerant of ﬁre. Approximately 40% of the stands of pencil pines in Tasmania have been destroyed by ﬁre in the last century and the risk of ﬁres escaping from management burns in adjacent eucalypt forests is signiﬁcant (Kiernan 2002). The threat of ﬁre to pencil pines has been reduced by bans on open ﬁres in adjacent camping areas at Lake Ada and Mount Doris. Fire regimes in eucalypt forests have dramatically changed since European settlement. In the managed tall eucalypt forests and mixed forests (eucalypts with rainforest) of Tasmania most tree biomass is periodically removed and the remaining material windrowed, dried and hot-burned. Despite claims that this process recapitulates natural
conﬂagrations, recent research highlights the patchy nature of wildﬁres and the fact that up to half the mature trees in a landscape may survive each ﬁre passage (Lindenmayer et al. 2000; Lindenmayer and McCarthy 2002).
Predatory vespid wasps Two species of predatory social wasps are now established in Tasmania (Bashford 2001). Vespids are active in most parts of the state, including nesting at 1,200 m on Mount Wellington (pers. obs.). European wasps (Vespula germanica) have been observed taking the diurnal adults of the uncommon Chrysolarentia hedylepta and a threatened endemic butterﬂy Oreixenica ptunarra in early autumn (P. Bell, pers. comm.). These wasps establish large perennial nests in the benign Tasmanian environment and their inﬂuence on insect populations is expected to increase over time.
Disease Phytophthora root pathogens threatens the survival of some foodplants notably the pencil pine Athrotaxis cupressoides which is the foodplant of Dirce aesiodora and several other endemic geometrids. An exotic Phytophthora recently discovered on pencil pines in the Pine Lake region (1,200 m elevation) of the Central Plateau has killed many trees and an area of 92,000 ha was quarantined to avoid further spread (Driessen 1999). P. cinnamomi is spreading rapidly through Tasmania below 500 m elevation and is causing progressive depletion of Proteaceae and Epacridaceae, both signiﬁcant geometrid foodplant families (Brown et al. 2002).
Lack of knowledge Changes in land cover are proceeding quickly in some parts of Tasmania with little appreciation of the impacts on the insect fauna. Large areas of the state remain unsurveyed for geometrid communities although most habitat types have been sampled repeatedly. Nevertheless, at least 25 new species has been found in the last 24 years. The relative lack of understanding of the phylogenetic relationships within the family may obscure the conservation merits of some taxa.
216 The Tasmanian conservation estate Conserved land in Tasmania comprises a large range of tenures, from the world heritage area to state reserves and privately covenanted land. Although the conservation estate is extensive in area, it is highly skewed in terms of the vegetation types represented, with the expected emphasis on ‘worthless lands’ (Mendel and Kirkpatrick 2002) and is highly fragmented across the landscape. In the last decade extensive areas of remaining state forests (largely unallocated crown land) have been earmarked for timber harvest, notably low value woodchips. The competing interests of the forest/mining lobby, conservationists and the state and federal governments were negotiated and formalised in a Regional Forest Agreement (RFA 2002) which seeks to conserve 15% of the notional extent of each forest type present at 1750 (preEuropean settlement). These forest types are based on the identity and stature of the dominant tree species and are assumed to be adequate surrogates for the native biota in general - an untested assumption for Lepidoptera in Australia. Another shortcoming is that the Tasmanian RFA conspicuously failed to recognise the IBRA bioregions in Tasmania resulting in a loss of representation at the bioregional level. Some treeless lowland communities are not represented in any reserve. This situation is unlikely to change in the short term given that extractive industries remain very inﬂuential in determining conservation outcomes. The woodchip industry clearfells thousands of hectares of native forest annually and enjoys real or eﬀective exemption from numerous government acts along with eﬀective self-regulation. In recent years the mining industry was able to veto the inclusion of important areas of habitat in western Tasmania into the world heritage area (e.g. the Mount Reid volcanics). Some National Parks have an unfortunate history of later excision of tall forest with commercial potential and ‘salvage harvesting’ after ﬁre (Kiernan 2002). Coastal reserves are under increasing development pressure, including a site near Hobart adjacent to one of only two known locations for Amelora acontistica. In summary, the geometrid fauna is an important and conspicuous component of Tasmania’s biodiversity and some taxa will need careful attention to guarantee their survival in the short to medium term.
Appendix 1. Checklist of Tasmanian Geometridae, together with range (number of 10-km grid squares recorded from) and presence in a National Park or the Tasmanian Wilderness World Heritage Area (NP), equivalent to IUCN protected area categories I and II. e denotes endemic Species
Ennominae Acalyphes philorites Turner e Acalyphes sp. [Tarn Shelf] e Dirce aesiodora Turner e Dirce lunaris (Meyrick) e Dirce oriplancta Turner e Dirce solaris (Meyrick) e Dirce sp. [Mt Field] e ‘Dirce’ sp. [Mt Wellington] e Authaemon stenonipha Turner Amelora acontistica (Turner) Amelora acromegala McQuillan e Amelora adusta Turner Amelora arotraea (Meyrick) Amelora leucaniata (Guene´e) Amelora nebulosa McQuillan e Amelora oritropha Turner Amelora sp [Mt Nelson] e Amelora sparsularia (Guene´e) Amelora syscia (Turner) Amelora zophopasta Turner ‘Amelora’ oncerodes Turner Furcatrox australis (Rosenstock) Furcatrox crenulata Turner e Furcatrox furneauxi McQuillan e Furcatrox paracus McQuillan e Furcatrox pervaga McQuillan e Furcatrox procera McQuillan e Furcatrox serrula McQuillan e Lackrana carbo McQuillan e Lackrana durafrons McQuillan e Dolabrossa amblopa Guest Dolabrossa suﬀusa (Turner) e Androchela milvaria (Guene´e) Androchela newmannaria (Guene´e) e Androchela smithi McQuillan e Cassythaphaga macarta (Turner) Hypsitropha euschema Turner e Synzeuxis penthina Turner Aphantes melanochorda (Turner) e Neoteristis paraphanes Meyrick Neoteristis sp. [Mt Reid] e Drymoptila temenitis (Guest) Archephanes zalosema Turner e Rhynchopsota delogramma Lower Bradyctena trynchoptila Turner e Corula geometroides Walker Mictodoca callipolia (Turner) e Mictodoca toxeuta Meyrick Conosara castanea Meyrick Cycloprorodes melanoxysta (Meyrick) Ciampa arietaria (Guene´e)
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