FNZ 54 - Hierodoris (Insecta: Lepidoptera: Gelechioidea: Oecophoridae) - Introduction
Hoare, RJB 2005. Hierodoris (Insecta: Lepidoptera: Gelechoidea: Oecophoridae), and overview of Oecophoridae. Fauna of New Zealand 54, 100 pages.
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ISSN 0111-5383 (print),
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no.
54.
ISBN 0-478-09378-0 (print),
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Published 24 Dec 2005
ZooBank: http://zoobank.org/References/58787BD7-116F-42F4-B565-26BE1BC948AD
Introduction
Oecophoridae: General Remarks
Oecophoridae belong to the ‘megadiverse’ superfamily Gelechioidea, one of the largest radiations of Lepidoptera with over 16,000 described and countless undescribed species (Kaila 2004). The family Oecophoridae is worldwide in distribution and probably contains well over 4000 described and undescribed species, even in its narrowest definition (see below). It is best represented in the southern hemisphere, and particularly in Australia, where over 3000 endemic species (including undescribed taxa) are known (Common 1994; 1997; 2000). Most species with known life histories have larvae feeding on dead plant material, especially leaf-litter.
The family has a complex taxonomic history and systematists still disagree about its limits. The Stathmopodinae have usually been included in Oecophoridae as a subfamily (e.g., Common 1990; Hodges 1998) or even as a tribe of Oecophorinae (Hodges 1978). The Depressariinae and Ethmiinae, formerly treated as oecophorid subfamilies (e.g., Hodges 1978), have either been given separate family status (Common 1990; Nielsen et al. 1996) or transferred as subfamilies to an expanded Elachistidae (Minet 1990; Hodges 1998). Stenomatinae, Xyloryctinae, Deuterogoniinae and Chimabachinae, all formerly treated as oecophorid subfamilies (e.g., Hodges 1978; Palm 1989; Nielsen et al. 1996), were excluded by Hodges (1998). Thus, in the most recent classification of Gelechioidea (Hodges 1998), Oecophoridae is restricted to the two subfamilies Oecophorinae and Stathmopodinae.
In an outstanding recent contribution to the systematics of Gelechioidea, Kaila (2004) presented a cladistic analysis based on 187 morphological and 5 behavioural characters scored for 143 species representing almost the full range of recognised gelechioid lineages. Given his extensive use of characters from the immature stages as well as formerly neglected character-complexes from adults (e.g., internal thoracic structure), Kaila’s attempt to resolve relationships in the group must be regarded as the most thorough and objective to date. His results support a basal division of the superfamily into two major lineages, the ‘gelechiid lineage’ and the ‘oecophorid lineage’. The low consistency index for Kaila’s most parsimonious trees indicates the high incidence of convergent evolution amongst Gelechioidea, and whilst there is support for the monophyly of many of the lineages recognised, e.g., by Hodges (1998), a large number of groups lack unique defining apomorphies and are recovered only on the basis of homoplasious characters. Also, as pointed out by Kaila, some of the basal clades, especially in his ‘oecophorid lineage’, have relatively weak character support, and there is a lack of data for immature stages of some taxa in these groups. Because of these problems, Kaila did not recommend the use of his cladogram (Kaila 2004: fig. 1) to construct a new classification of Gelechioidea, but indicated how Hodges’ (1998) classification could be minimally rearranged to accommodate the revised hypothesis of relationships.
Kaila’s study suggests that Stathmopoda and related genera are closely allied to genera currently assigned to Batrachedridae and Coleophoridae in the ‘gelechiid lineage’. The support for this relationship is strong: four synapomorphies, two of them unique, are shared by Stathmopoda with Idioglossa, Batrachedra, Goniodoma, and Coleophora (Kaila 2004). I accept this here as sufficient evidence for the removal of Stathmopodinae from Oecophoridae.
Taxa formerly assigned to Oecophorinae in New Zealand appear to belong to two distinct groups. On the basis of Kaila’s cladogram (Kaila 2004: fig. 1), Tingena (and related genera: see below) would be referable to Oecophoridae sensu stricto, whilst Hierodoris, Izatha, and Phaeosaces (and related genera: see below) would be assigned to the ‘xyloryctid assemblage’, with Hierodoris occupying a basal position as sister to all remaining taxa in this clade. However, as Kaila emphasises, character support at the base of the ‘oecophorid lineage’ is weak, and the inclusion in the analysis of the Russian ‘oecophorine’ Martyringa ussuriella Lvovsky collapses the basal resolution in this large clade. Likewise, one of the apomorphies supporting monophyly of the sister-group of the ‘xyloryctid assemblage’ in Kaila’s cladogram (i.e. Oecophoridae s.s. + Amphisbatidae s.s. + Carcinidae + Stenomatidae + Chimabachidae + Elachistidae s.l.) is ‘mesal part of gnathos scobinate with small thorns’; this character-state also occurs in Hierodoris, but not in the species sampled by Kaila. So although there is evidence for assigning Hierodoris and its relatives to an expanded ‘Xyloryctidae’, the phylogenetic position of these genera remains potentially unstable and sensitive to taxon sampling. Therefore, for the present I have adopted a conservative approach, retaining in Oecophoridae all New Zealand taxa assigned to Oecophorinae by Dugdale (1988) except for Compsistis bifaciella Walker, which is placed in Lecithoceridae (Dugdale 1996). However, following the approach of Common (1994; 1997; 2000), I here assign the New Zealand Oecophoridae to informal genus groups and erect the ‘Hierodoris group’ (see below for definition) to accommodate the basal genera of Kaila’s ‘xyloryctid assemblage’. All New Zealand taxa not assigned to the Hierodoris group are collectively placed in ‘Oecophoridae sensu stricto’, and divided between Common’s named groups. The assembly of these groups into a stable higher classification should await further phylogenetic analysis of the basal taxa in the ‘oecophorid lineage’ (cf. Kaila 2004: 324).
Overview of New Zealand Oecophoridae
In New Zealand, Oecophoridae is the second most species-rich family of Lepidoptera (after Geometridae), with 247 known species in 28 genera. The composition of this diversity is indicated in Table 1. Numbers for the Hierodoris group are based on recent sorting of specimens held in NZAC and are probably relatively accurate; those for the Oecophoridae s.s. are based on Dugdale (1988), together with some more recent sorting of material, and are only best estimates, since this group contains many externally variable and closely similar taxa, and requires much further work. All species in the Hierodoris group are endemic, with the exception of Scieropepla typhicola Meyrick, which is shared with Australia. There are 14 species in the Oecophoridae s.s. suspected or known to be adventive (Dugdale 1988; Hoare 2001).
As revision of the Oecophoridae progresses, the species count can be expected to increase, since the number of cryptic taxa awaiting recognition in collections (especially in Tingena and related genera) probably exceeds the number of new synonymies to be made. Further new species are likely to be added as a result of field-work in poorly collected areas. The current figure for recognised genera (28) can only be regarded as a rough guide to higher level diversity, since almost no endemic genus has been satisfactorily defined to modern systematic standards. A number of New Zealand species are still placed in Australian and European genera to which they do not belong.
Although not rivalling the extraordinary Australian fauna, the New Zealand Oecophoridae still represent an impressive endemic radiation. The oecophorid fauna of Russia and adjacent countries (i.e. the former USSR) was recently catalogued by Lvovsky (2003), who listed 110 species for this entire vast area; however, at most, 86 of these belong to the Oecophoridae as here defined. In comparison, New Zealand, with less than 1/80 of the land area, probably has about three times the diversity of Oecophoridae.
A revision of the Oecophoridae of New Zealand is timely for several reasons:
- As one of the two most species-rich families of New Zealand Lepidoptera (alongside Geometridae), Oecophoridae has a high priority in the documentation of New Zealand’s biodiversity.
- The family has high conservation value, since (a) it includes a number of enigmatic or ‘lost’ species that have not been recognised for many years, e.g., Izatha rigescens Meyrick, Lathicrossa prophetica Meyrick, Chersadaula ochrogastra Meyrick (Patrick & Dugdale 2000), (b) the level of endemism at both species and genus level is nearly 100% for native taxa, and (c) the detritivorous larvae are probably ‘key players’ in recycling nutrients in forests and other ecosytems (Dugdale 1996).
- Being species-rich and numerically abundant, Oecophoridae are likely to prove useful indicators of environmental change, e.g., the drying out of forest fragments as the result of ‘edge effects’ and the predations of introduced social Hymenoptera. This especially applies to the many species with litter-feeding larvae, which are likely to be vulnerable to such environmental pressures.
- Since most adventive Lepidoptera that successfully establish in New Zealand belong to the detritivorous guild (Hoare 2001), we urgently need a basis from which to study the effects of these invaders on the native fauna. This will rely on a thorough up-to-date documentation of the taxonomy and natural history of native taxa.
- The recent revisions at the generic level of the huge Australian oecophorid fauna by Common (1994; 1997; 2000) have established a very sound context for the taxonomic reassessment of other southern hemisphere Oecophoridae.
- The present genus-level classification is confused and outdated, and a new framework is needed to understand the true diversity and relationships of the New Zealand species.
- Kaila’s (2004) phylogenetic work suggests that some New Zealand taxa, such as Hierodoris, occupy key (basal) positions in the evolutionary history of the vast gelechioid radiation; therefore New Zealand will play an important role in understanding that history, and in placing the classification of Gelechioidea on a more stable basis.
Earlier work on the New Zealand Oecophorinae
Most of the genera and species of New Zealand Oecophoridae were described by Francis Walker (especially Walker 1864) and Edward Meyrick (many publications, but especially Meyrick 1884; 1914). Walker’s generic concepts were very sketchy by modern standards: for example, he erected the genus Izatha for attactella, but placed its congeners convulsella and picarella in Gelechia and Oecophora respectively (Walker 1864). Moreover, he described I. peroneanella three times in two different genera! Meyrick, on the other hand, was a fine systematist, and much of the foundation he laid for the classification of Lepidoptera still stands. However, he based his system entirely on external characters such as scale vestiture, antennae, palpi, and especially wing venation, and dismissed the study of the genitalia as at best unnecessary, or at worst misleading (e.g., Clarke 1955: 13–14). It has now been universally recognised that the genitalia provide crucial clues to the classification of Lepidoptera at all levels, and, as a result, many of Meyrick’s generic and family concepts have had to be revised (although his species-level taxonomy remains largely sound). For example, in revising the Australian oecophorine genera, Common (1994; 1997; 2000) found it necessary to change substantially the circumscription of many genera treated by Meyrick: an extreme instance is Eulechria Meyrick, in which Meyrick (1922) included 244 species, only 37 of which are accepted by Common (1997) as correctly placed.
Alfred Philpott also described a number of New Zealand Oecophoridae, and was a pioneer in the study of the genitalia of Lepidoptera. He illustrated the genitalia of most of the oecophorid species known at the time (Philpott 1926; 1927a, b). In general, he did not attempt any redefinition of genera on the basis of his results, but continued to follow Meyrick’s systematic concepts. He was, however, able to demonstrate the existence of a number of ‘cryptic’ species, best distinguished on the basis of the genitalia (for example Gymnobathra primaria, G. levigata, and G. inaequata (Philpott 1928)).
The only complete descriptive treatment of the New Zealand Oecophoridae is that contained in G. V. Hudson’s admirable, fully illustrated account of the country’s Lepidoptera (Hudson 1928; 1939; 1950). Hudson relied on Meyrick’s classification, and accepted the latter’s arguments against a consideration of genital structures (see Hudson 1939: Introduction). His works are still very helpful (though rare and expensive), but caution is required in using them to identify species in families such as Oecophoridae, where the taxonomy of the numerous similar species is still confused, and there are a number of undescribed species Hudson did not know. Comparison of the genitalia with those of type specimens will often be required to arrive at a satisfactory diagnosis.
Until now, only two new species of New Zealand Oecophoridae have been described since 1950: Tinearupa sorenseni by Salmon & Bradley (1956) and Izatha oleariae by Dugdale (1971). Dugdale (1971) also described a new subspecies of T. sorenseni (T. s. aucklandica). All these taxa are endemic to New Zealand’s subantarctic islands.
The annotated catalogue of Dugdale (1988) summarised the current state of taxonomic knowledge, and introduced several new synonymies and the 10 species recorded in New Zealand or described since Hudson’s time. Dugdale altered the generic placement of many species in line with a more critical assessment of relationships than Meyrick’s or Hudson’s, and transferred to the family the genera Hierodoris Meyrick and Coridomorpha Meyrick, the species of which had previously been assigned to the Glyphipterigidae (either as a family or as a subfamily of the old, broad concept of Tineidae).
In an important and often overlooked contribution, Dugdale (1996) described and keyed oecophorid larvae found in beech (Nothofagus) forest leaf-litter in the Orongorongo Valley, near Wellington. This was the first work to recognise a distinction between the Hierodoris group (‘Gymnobathra group’ of Dugdale 1996) and other Oecophoridae in New Zealand, on the basis of larval characters.
Finally, Hoare (2001) added three adventive Australian Oecophoridae to the New Zealand list.
Generic groupings of Oecophoridae
Dugdale (1988) divided the New Zealand Oecophorinae into 2 major groupings: Group A, species with ocelli, containing Coridomorpha and Hierodoris, and Group B, species without ocelli, containing all remaining genera. In his revision of the Australian fauna, Common (1994, 1997, 2000) named 6 genus groups, i.e. the Wingia, Philobota, Chezala, Eulechria, Barea, and Tisobarica groups. From his study of larvae, Dugdale (1996) referred several New Zealand genera (Tingena, Trachypepla, Atomotricha, and Leptocroca s.l.) to Common’s Barea group, and distinguished a ‘Gymnobathra group’ comprising Thamnosara and litter-feeding species assigned to Gymnobathra.
Here I propose a ‘Hierodoris group’ to include the following endemic New Zealand taxa: Hierodoris (= Coridomorpha), Gymnobathra (excluding G. rufopunctella Hudson), Izatha, Lathicrossa, Phaeosaces, Thamnosara, Tinearupa, and ‘Schiffermuelleria’ orthophanes Meyrick. This represents an amalgamation and expansion of Dugdale’s (1988) Group A and his (1996) Gymnobathra group. The diagnostic features of the Hierodoris group are summarised in Table 2. Character states for Oecophoridae s.s. are based on Common (1994; 1997; 2000) and Kaila (2004) and on a study of New Zealand species of the Barea group. Those for Xyloryctinae + Blastobasinae are based largely on Kaila (2004).
The inclusion of Phaeosaces in the Hierodoris group is based on Kaila (2004), who recovered the genus as sister to Izatha on the basis of 7 shared homoplasious characters. However, I assign it to the group only tentatively, since it has some features in common with many Oecophoridae s.s. (especially the curved base of the aedeagus and the coiled bulbus and ductus ejaculatorius). Larval characters of Phaeosaces should help to resolve its relationships, but these have not yet been examined.
The following Australian genera are tentatively assigned to the Hierodoris group: Scieropepla (14 described species, including 1, S. typhicola Meyrick, shared with New Zealand), Nemotyla (1 Tasmanian alpine species, N. oribates Nielsen, McQuillan & Common) and Athrotaxivora (1 Tasmanian species, A. tasmanica McQuillan). The inclusion of Scieropepla and Nemotyla is based on their position in Kaila’s (2004) cladogram, where together with Izatha + Phaeosaces they form a monophyletic group basal to the core Xyloryctinae + Blastobasinae. Athrotaxivora was provisionally associated with Xyloryctinae by McQuillan (1998), who noted however that it lacked characters of the core Xyloryctinae. In characters illustrated by McQuillan (loc. cit.) it matches the diagnosis of the Hierodoris group given here.
Metaphrastis Meyrick from South Australia and Western Australia (redescribed by Common (2000: 401–406)) has 2 characters in common with Hierodoris, i.e. presence of ocelli and an overlay of narrow scales on the forewing, but the genitalia of both sexes differ widely from those of any genus here assigned to the Hierodoris group. In particular, the basally coiled aedeagus with its single long cornutus (Common 2000: fig. 719) would suggest a closer relationship with the Barea group of genera in Oecophoridae s.s. I agree with Common (loc. cit.) in considering Metaphrastis to have no near relationship to Hierodoris.
No claim is made here for the monophyly of the Hierodoris group, which is intended as a convenient informal association of genera that fall outside the core Oecophoridae and Xyloryctidae, whilst sharing some characters with each of these groups. Indeed, Kaila’s (2004) cladogram would indicate that the Hierodoris group is not monophyletic. However, it is possible that denser taxon sampling in this region of the phylogeny, and a better coverage of immature stages, might recover a monophyletic Hierodoris group.
Relationships of the Hierodoris group
As indicated above, the Hierodoris group appears to be related to taxa traditionally assigned to Xyloryctidae / Xyloryctinae (Kaila 2004). Probably the most convincing character supporting this relationship is the presence of paired sclerotised slits in the larval mentum: outside the Hierodoris group and Xyloryctinae, these slits are known only from a few Lecithoceridae. However, because some more derived taxa in Kaila’s ‘xyloryctid assemblage’ (Uzucha humeralis Walker and Blastobasinae) lack these slits, and because there are no data for the larvae of other taxa, their presence is not recovered as an unambiguous apomorphy supporting the monophyly of the assemblage (L. Kaila, pers. comm.).
The second character linking the Hierodoris group to the xyloryctid assemblage in Kaila’s phylogeny is the presence of a pinacular ring around the abdominal SD setae of larvae. This is again paralleled in Lecithoceridae, and in some species of Scythris and Stathmopoda; Hodges (1998) also listed it as an apomorphy of Autostichinae (Autostichidae).
Hierodoris group: overview of unrevised New Zealand genera
The largest genus in the Hierodoris group is Izatha Walker, with 26 described species (Dugdale 1988). The genus is probably monophyletic, possible apomorphies being the loss of the gnathos and uncus in the male genitalia, and the form of the aedeagus (outer wall spinose in apical half, and sclerotisation interrupted lengthwise). Many species, but not all, possess a tuft of scales on the third segment of the labial palp, and/or tufts of raised scales on the forewing, and the coloration is often beautifully cryptic. The known larvae feed in dead wood (Hudson 1928). Izatha is possibly unique amongst species-rich Lepidoptera genera in New Zealand in that there are many more species endemic to the North Island than to the South Island.
Gymnobathra Meyrick currently contains 21 species (with the removal of G. squamea, here transferred to Hierodoris). On external characters, G. rufopunctella Hudson closely resembles some members of the Barea group of genera (e.g., Euchersadaula spp.) and is almost certainly misplaced in Gymnobathra. Gymnobathra origenes Meyrick is an enigmatic taxon known from a single specimen that has unfortunately lost its abdomen; from the hindwing shape and the presence of ocelli it appears to be related to ‘Hierodoris’ insignis Philpott (Gelechiidae), but further material is needed to confirm this. The remaining species of Gymnobathra are rather heterogeneous in genitalic morphology (see Philpott 1927a) and in biology, and even without the species discussed above, the genus is unlikely to be monophyletic. Several species have larvae feeding in bark or dead wood (Hudson 1928): these include G. flavidella Walker (the type species), G. omphalota Meyrick, G. bryaula Meyrick, and G. dinocosma Meyrick. Gymnobathra calliploca Meyrick and G. levigata Philpott have leaf-litter feeding larvae that make silken nests covered with frass and leaf fragments; they construct silken runways from these shelters into the surrounding litter (Dugdale 1996). There is another litter-feeding group with case-bearing larvae; Dugdale (1996) illustrated cases of G. tholodella Meyrick and of another species, which he called G. sarcoxantha. This second species is probably in fact undescribed and represents G. coarctatella in the sense of Meyrick and Hudson (not Walker). The true G. sarcoxantha Meyrick and G. parca (Butler) probably have similar habits; this group is here considered the probable monophyletic sister-group of Hierodoris, and as such will require a new genus.
Lathicrossa currently contains 2 species, the common L. leucocentra Meyrick, and the very elusive L. prophetica Meyrick. On male genitalia, L. leucocentra appears to be closely related to Gymnobathra omphalota, and the genus may fall into synonymy with Gymnobathra when further studies are undertaken.
Thamnosara and Tinearupa each contain 1 species, although 2 subspecies of Tinearupa sorenseni Salmon & Bradley have been described (Dugdale 1971, 1988). Thamnosara sublitella (Walker) is an abundant and widespread species with larvae feeding in forest leaf-litter in a similar manner to Gymnobathra calliploca and levigata (Dugdale 1996). The male and female genitalia do not indicate any obvious close relationship to these Gymnobathra species, however, and the genus may well remain separate and monotypic. Tinearupa is confined to the subantarctic Auckland and Campbell Islands; both sexes are brachypterous. Dugdale (1971) compared T. sorenseni to Gymnobathra omphalota, but the genitalia indicate strong divergence from other described genera, e.g., in the reduced aedeagus, and the affinities of the genus are obscure.
‘Schiffermuelleria’ orthophanes is a very small species wrongly placed in the European genus Schiffermuelleria by Meyrick. Hudson (1928) considered it common, although it had formerly been very rare, and noted that most specimens were taken indoors. The species appears to have become rare again, and there are very few recent specimens. A possible relationship with Hierodoris is indicated by the presence of backward-pointing ‘teeth’ on the apex of the aedeagus, but orthophanes lacks both the invagination on S8 and the strap-like forewing scales of this genus. It also lacks ocelli. The relationships and life history of this species remain unknown, as does the reason for its apparent decline.
Other generic groups in New Zealand
The remaining species of Dugdale’s ‘Group B’ can almost all be referred to the generic groupings of Common (1994; 1997; 2000). Three of Common’s groups (the Wingia, Chezala, and Barea groups) are represented in New Zealand, probably only 2 of these by endemic species, and only 1 by endemic genera.
The Wingia group, recognised mainly by the strongly sexually dimorphic conformation of abdominal sternite 2 (apodemes elongate in females, strongly reduced in males: see Common 1994) is represented in New Zealand by 2 species. Heteroteucha dichroella (Zeller) is an Australian adventive known in New Zealand from a single specimen captured in Havelock North (HB) in 1925 (Hoare 2001) and not known to be established. The other species in this group is ‘Trachypepla’ indolescens Meyrick. This species was described from New Zealand (Meyrick 1927) and has not been recognised elsewhere. However, it is presumed adventive, as it was not collected in New Zealand prior to 1909, but has become very common in some localities in recent years, suggesting a pattern of progressive establishment. Also, the species shows relationships to the endemic northeast Australian genera Limothnes Turner, Basiplecta Common, and Idiozancla Turner, all of which it resembles in wing pattern, in the reduction of the proboscis, and in its tendency to go greasy in collections. ‘T.’ indolescens does not conform in genital morphology to any of the genera treated by Common (1994) and requires a new genus.
The Chezala group of genera as defined by Common (1997) is possibly not monophyletic: Common could find no unambiguous apomorphies linking the included genera. The group is represented in New Zealand by 4 species, only 1 of which is endemic. The Australian Tachystola acroxantha (Meyrick) (formerly placed in Parocystola) has been established here since 1886 or before (Hudson 1928). It is also established in England (Harper et al. 2002). Two abundant and now cosmopolitan pests, whose origins are obscure, are also established: these are Hofmannophila pseudospretella (Stainton) and Endrosis sarcitrella (Linnaeus). The larvae of both species are almost omnivorous, feeding on stored foodstuffs, woollen clothing and carpets, in birds’ nests, on dead insects, and other pabula. The assignment of Hofmannophila and Endrosis to this largely Australian group of genera is tentative (Common 1997). The fourth New Zealand member of the Chezala group is the endemic Prepalla austrina (Meyrick). Meyrick assigned this species at first to Saropla and later to Oxythecta. Its true relationships were worked out by Common (1997) who included it in his new genus Prepalla, along with 9 Australian species. B. Patrick has reared P. austrina, which occurs in coastal and mountainous areas of the South Island and on the central volcanic plateau of the North Island, from the small shrub Leucopogon fraseri (Epacridaceae) (Patrick 1994). No species of Prepalla has been reared in Australia (Common 1997), but as Leucopogon is a genus shared between the two countries, it seems likely that the Australian species will prove to have similar habits.
All remaining genera of New Zealand Oecophorinae can be assigned to the Barea group in the sense of Common (2000), the defining apomorphy of which is the presence in the male genitalia of a pair of fusiform appendages arising from near the basal plate of the juxta and ending near the pulvinus of the valva on each side. In contrast to most members of the Hierodoris, Wingia, and Chezala groups, species of the Barea group lack distinct lateral lobes of the juxta (however, these are also absent in Prepalla). A number of Australian species of this group have become established in New Zealand, including 5 species of Barea Walker, and 1 each of Sphyrelata Meyrick, Leptocroca Meyrick, and Atalopsis Common (see Dugdale 1988; Hoare 2001).
There is an extensive endemic radiation of the Barea group in New Zealand, encompassing species currently assigned to the genera Atomotricha Meyrick, Chersadaula Meyrick, Corocosma Meyrick, Euchersadaula Philpott, Eulechria Meyrick, Euthictis Meyrick, Leptocroca in the sense of Philpott, Locheutis Meyrick, Mermeristis Meyrick, Tingena Walker, and Trachypepla Meyrick. ‘Gymnobathra’ rufopunctella also probably belongs to this group, but dissection will be required to confirm this. This complex of closely related taxa comprises at least 140 species, and there are many problems in the taxonomy, both at the specific and generic levels. The type species of Eulechria, Euthictis, Leptocroca Meyrick (L. sanguinolenta: see above), Locheutis, and Mermeristis were all described from Australia, and with the single exception of Mermeristis, no endemic New Zealand species is correctly assigned to any of these genera (Common 2000). Known larvae are all associated with dying plant material, leaf-litter or dead wood (Patrick 1994, Dugdale 1996).