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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by J. De Prins: 3 Jan. 2020; published: 13 Feb. 2020 351
Zootaxa 4732 (3): 351–374
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4732.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:ECF50463-FE43-400B-A11D-E0E5E9870B34
Three new ghost moths of the genus Oxycanus Walker, 1856 from Australia
(Lepidoptera: Hepialidae)
ETHAN P. BEAVER1,3, MICHAEL D. MOORE1, ALEJANDRO VELASCO-CASTRILLÓN1 & MARK I. STEVENS1,2
1South Australian Museum, Adelaide, SA 5000, Australia
2 School of Pharmacy and Medical Sciences, University of South Australia, Adelaide, Australia
3Corresponding author. E-mail: ethan.beaver@live.com.au
Abstract
Three new species of ghost moth, Oxycanus ephemerous sp. nov., O. flavoplumosus sp. nov., and O. petalous sp. nov.
are described from South Australia, New South Wales, and south-west Western Australia, respectively. We illustrate these
species and compare morphological and molecular (mtDNA COI gene) characters with similar Oxycanus Walker, 1856
species from Australia. Comparative images of Oxycanus subvaria (Walker, 1856), O. byrsa (Pfitzner, 1933), and O.
determinata (Walker, 1856) are figured. The type material of the three new species are held in the Australian National
Insect Collection, Canberra, the Western Australian Museum, Perth, and in the South Australian Museum, Adelaide.
The type specimens of Oxycanus hildae Tindale, 1964 syn. n. were also examined and the taxon is here considered
synonymous with O. subvaria. Concerns are raised about the conservation status of all three new species due to few or
localised distribution records.
Key words: COI, DNA barcodes, short-range endemics, taxonomy
Introduction
In recent years there have been several important investigations into Hepialidae in Australia, such as the revision
of Fraus Walker (Nielsen & Kristensen 1989), Elhamma Walker (Simonsen 2015), and the monograph on the
Australian members of the genera Abantiades Herrich-Schäffer, Aenetus Herrich-Schäffer, Archaeoaenetus Simon-
sen, Zelotypia Scott, and Oncopera Walker (Simonsen 2018). The only Australian genera to have not undergone
a modern revision are Jeana Tindale, and Oxycanus Walker. Oxycanus is a species-rich genus of medium to very
large-sized moths distributed across mainland New Guinea and the higher rainfall regions of Australia, particularly
in the southern, eastern and south-western regions. The adults often fly late in the season, with many species emerg-
ing en-masse only during or just after autumn and early winter rainfall events (Kallies et al. 2015). Little is known
of their subterranean larval biology for most species, however some species such as Oxycanus antipoda Herrich-
Schäffer and O. oreades Edwards & Green feed on the foliage of grasses (Hardy 1974; Edwards & Green 2011)
while O. dirempta has been recorded feeding on leaf litter of Acacia species (Tindale 1935). Many species may be
commonly encountered, although some, including those described here, are less common and may appear only at a
specific time each year.
Walker (1856) described Oxycanus for O. australis (Walker, 1856). Later Tindale (1935) revised and rede-
scribed the genus, noting differences in the proportions of the labial palps as well as in wing venation when com-
pared to the similar Jeana, and Elhamma, all of which have the typical ‘oxycanine’ wing venation (Dugdale 1994)
in which the Rs1 and Rs2+Rs3 veins share a common stalk. In Oxycanus species, the second palpomere of the labial
palp is longest, while in Jeana the basal palpomere is longest. Additionally, in Jeana the forewing vein branch of
Rs1 and Rs2 is closer to the termen than in Oxycanus (Tindale 1935). Simonsen (2015) showed that Elhamma is
unique among the Hepialidae by having only two M veins in the hindwing and a distinctly cup-shaped juxta. Gre-
han (2011) noted the similarities of the Asian genus Hepialiscus Hampson to some of the Oxycanus particularly in
regard to para-anal sclerites of the male genitalia the presence of which are helpful in differentiating Oxycanus from
BEAVER ET AL.
352 · Zootaxa 4732 (3) © 2020 Magnolia Press
the similar Australian genera Jeana and Elhamma (Grehan 2011). However, Edwards & Green (2011) noted that
some Oxycanus lack this structure, suggesting that there may be distinct morphological groups within Oxycanus.
The absence of para-anal sclerites in some species groups along with a wide range of variation in antennal morphol-
ogy may suggest that the genus is not monophyletic. Oxycanus, though too large to describe at the present time, will
be the subject of a planned taxonomic revision.
Since Tindale’s works in 1935, 1955, and 1964, only two new Oxycanus species have been recently described in
the paper by Edwards & Green (2011) and are from the alpine regions of New South Wales, Australia. All Oxycanus
species found to occur in the Australian state of Victoria were recently illustrated by Kallies et al. (2015) in which
live habitus photos, field information and figures of important morphological features were included. The addition
of O. ephemerous sp. nov., O. flavoplumosus sp. nov., and O. petalous sp. nov. brings the number of Australian
Oxycanus to 47, and world species to 79. In the years of 2018–19 all Oxycanus specimens in the collection of the
South Australian Museum were databased, sorted and re-identified by the authors, leading to the discovery of O.
flavoplumosus sp. nov. and O. ephemerous sp. nov., and field work by one author (MDM) as well as examination of
the Australian National Insect Collection, Canberra identified the presence of O. petalous sp. nov. as a distinct spe-
cies. Further collecting by the authors and members of the public has also enabled molecular work (mtDNA COI)
to be undertaken for O. ephemerous sp. nov. and O. petalous sp. nov., which we also include here in comparison to
other Oxycanus species. The three species are described here ahead of a generic revision to identify and discuss the
possible paraphyly of Oxycanus, and due to the potential conservation requirements of the new species. Edwards &
Green (2011) raised concerns due to the possible side effects of climate change on Oxycanus oreades from the alpine
and sub-alpine zones of New South Wales, due in part to its limited high altitude distribution. One species described
here, O. flavoplumosus sp. nov. is similarly known only from mountain-tops in sub-alpine woodland systems, and
may also be at risk of environmental change as a side effect of climate change.
Materials and methods
The new species described in the present article are compared only with those that they most closely resemble.
Specimens examined in this study are from:
Australian Museum, Sydney (AM), Australian National Insect Collection, Canberra (ANIC), Natural History Mu-
seum, London (NHM), Queensland Museum, Brisbane (QM), Research collection of E. P. Beaver (EPBC), South
Australian Museum, Adelaide (SAMA), Tasmanian Museum and Art Gallery, Hobart (TMAG), University of
Queensland Insect Collection (UQIC), Western Australian Museum, Perth (WAM).
Other abbreviations: FW—forewing. HW—hindwing. Location data are provided as place names and with
GPS coordinates where provided on the specimen label. Australian states are abbreviated as follows: SA—South
Australia, VIC—Victoria, NSW—New South Wales, WA—Western Australia, and TAS—Tasmania.
Specimens examined for this study other than those mentioned within the systematic treatment include the fol-
lowing:
Oxycanus byrsa (Pfitzner, 1933)—in total 22 specimens. 2 ♀, 12 ♂, Ebor, NSW, May 1939 and April 1949, 1 ♂
dissected, dissection ID 31-018407-EPB (in SAMA). 1 ♀, 6 ♂, 19 May 2017, Rocky River, 3 km NW Uralla,
NSW, 30°37’S 151°28’E, 19 May 2017, J. Clark, 1 ♀ dissected, dissection ID EPB-ANIC-13 (in ANIC). 1
♂ in EPBC, same data as ANIC specimens.
Oxycanus determinata (Walker, 1856)—in total 8 specimens. 1 ♀ Paratype, Perth WA, ‘July 08’. 2 ♂, Swan River,
WA, L.J. Newman, 1 ♂ dissected, dissection ID 31-018439-EPB. 1 ♂, S Perth, WA, 4 May 1922, dissection
ID 31-018441-EPB. 2 ♂ and 2 ♀, Esperance, South Coast, WA, 16-17 April 2017, M.D and M. Moore, 1 ♂
dissected, dissection ID 31-019870-EPB, 1 female dissected, dissection ID 31-019873-EPB (in SAMA).
Oxycanus glauteri Tindale, 1955—in total 2 paratypes, 1 ♂, 1 ♀, WA, 1906, G.C. Shortbridge (in SAMA).
Oxycanus kochi Tindale, 1955—in total 14 specimens, 1 ♂ paratype ‘New Holland’. 8 ♂, 5 ♀, Drummond Cove,
WA, 4 May 1976, N. McFarland, dissected: 3 ♂, dissection IDs 31-018721-EPB, 31-018722-EPB, EPB-
ANIC-15, 1 ♀, 31-018446-EPB. (5 ♀, 7 ♂ in SAMA, 1 ♂ in ANIC).
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 353
Additional material examined: All Oxycanus specimens in the SAMA collection (1320 specimens of 58 spe-
cies) as well as the similar Jeana (47 specimens) were sorted and re-identified by the authors prior to this study.
Morphological analyses. Scanning electron images of antennae were collected using a Hitachi TM4000Plus
Tabletop Scanning Electron Microscope provided by NewSpec Pty Ltd and imaged by Ben A. Parslow. Dissections
of genitalia were performed as described in Simonsen (2018), by removing the end of the abdomen (males) or the
entire abdomen (females) and heating in 20% KOH solution for 5–8 minutes before being cleaned in distilled water
and stored in 70% ethanol. Terminology for genitalia and wing venation follows Dugdale (1994) and Simonsen
(2018). Antennae terminology: In quadripectinate species (sensu Dugdale 1994) the longest rami are here referred
to as ‘primary rami’ while the shorter posteriorly-situated pair is termed the ‘proximal bar’ and the raised, ciliated
central process connecting the two along the ventral side of the flagellomere is referred to as the ‘ridge’ for ease of
comparison and description (Figs 3A, D, L). Wing venation diagrams were traced free-hand over photographs of
specimens. Specimen data are presented with a ‘/’ denoting a second specimen label.
Molecular analyses. DNA extraction and PCR amplification was completed at the South Australian Regional
Facility for Molecular Ecology and Evolution (SARFMEE) from a single leg from each specimen. DNA was ex-
tracted using a Gentra Puregene® DNA Purification kit (Gentra Systems Inc.) according to the manufacturer’s
protocol. We targeted the commonly used 710 bp fragment of the mitochondrial (mtDNA) cytochrome oxidase c
subunit I (COI) gene which has successfully delineated between numerous insect species (e.g. Dorey et al. 2019;
Grund et al. 2019). PCR products were amplified using the primers LCO1490 (forward) and HCO2198 (reverse)
(Folmer et al. 1994) on the thermocycler (Eppendorf Mastercycler® EP gradient) under the following conditions:
94 °C for 1 min for initial denaturation, 5 cycles of 94 °C for 1 min, annealing at 47 °C for 90 sec, and extension at
72 °C for 1 min, followed by 35 cycles at 94 °C for 60 sec, 90 sec at 51°C, and 60 sec at 72 °C, with a final extension
of 72 °C for 5 min. The PCRs were sent to Macrogen Inc. (Korea) for purification and sequencing.
All new COI sequences generated here were checked visually and sequence chromatograms (forward and
reverse) were inspected to resolve ambiguous base calls and checked for potential contamination using BlastN
(NCBI). The resulting sequences were aligned with other available GenBank accessions relevant to the current
study (see Supplementary Table S1) in Geneious R11.0.5 (https://www.geneious.com) to produce a 658 bp align-
ment. Nucleotide sequence divergence was calculated using uncorrected p-distances as implemented in PAUP*
v.4.0beta10 (Swofford 2002). Modeltest v.3.7 (Posada and Crandall 1998) was used to generate likelihood scores for
the models of evolution within PAUP*. The general time reversible model with invariant sites and gamma distribu-
tion (GTR+I+Γ) (Nei & Kumar 2000) was selected as the best model by Modeltest under the Akaike Information
Criterion (AIC), and a maximum-likelihood phylogram (and 1,000 bootstraps, Felsenstein 1985) was generated
using this model in MEGA X (Kumar et al. 2018).
Taxonomy
Family HEPIALIDAE Stephens
Genus Oxycanus Walker, 1856
Oxycanus ephemerous Beaver & Moore, sp. nov.
(Figs 1A, 3K–L, 4A–C, 7A, 10A)
Type specimens: Holotype male, SAMA. Paratypes, 2 males, SAMA.
Type locality: Inman Valley, Fleurieu Peninsula, SA, Australia.
Etymology. The specific name (Greek εφήμερος) refers to the ephemeral nature of the adult insect, which is known
to fly for a brief period at a specific time of the year, in very few locations.
Type material. HOLOTYPE, male; (SAMA). ♂, 20 May 2018, Inman Valley, Fleurieu Peninsula, South Aus-
tralia, 35°29’01.0”S, 138°26’12.2”E, 180m elevation, coll. S. Eden, pres. E. P. Beaver, / Spec. No. 1862 leg re-
moved for tissue storage MD Moore. / SAMA Database No. 31-019818.
PARATYPES, 2 males; (SAMA) 2 ♂, Amer. River, K.I. 20 May 1952, F. M. Angel / SAMA database no. 31-
019191 / Dissection ID 31-019191-EPB; second male with SAMA database no. 31-019190.
BEAVER ET AL.
354 · Zootaxa 4732 (3) © 2020 Magnolia Press
Distribution. Known from Inman Valley, Fleurieu Peninsula; and from American River, Kangaroo Island,
South Australia (Figs 11, 14).
Diagnosis. This species is similar to Oxycanus subvaria (Figs 1B; 3I–J; 4D–F, 7B, 8A & D). However, it differs
primarily in the shape and proportions of the male genitalia. In particular the vinculum is narrower and more square,
and the saccus is much broader, rounded and shorter in O. ephemerous sp. nov. than in O. subvaria. In addition, the
pseudotegumen is differently shaped in the new species. In O. ephemerous sp. nov. the ventral pseudoteguminal
arms are distinctly shorter and the basal rim is less rounded and not as sclerotized as in O. subvaria. Both species
have sternite 8 fused with the saccus (Fig. 8A & D for O. subvaria Fig. 7A for O. ephemerous sp. nov.). The anten-
nal rami of O. subvaria are equal to the width of the flagellomere in the middle of the antennae, while the rami of O.
ephemerous sp. nov. are longer than the width of the flagellomere (Fig. 3K & L). The only other Oxycanus species
to occur nearby to the known localities of O. ephemerous sp. nov. is O. niphadias (Meyrick) from the Southern
Lofty Ranges. However, this species has longer antennal rami, at least 3x the width of the flagellomere, and distinct-
ly different genitalia. The pseudotegumen in Oxycanus niphadias has a single central tooth, lacking invagination,
and the S8 is pocket-like and depressed well into the vinculum. These structural differences are obvious enough to
be observed in specimens without dissection or a microscope. The distributions of the widespread O. occidentalis
Tindale, 1935 and O. ephemerous sp. nov. may overlap at the Kangaroo Island location, however, the larger (FW: 30
mm) O. occidentalis is closely allied to O. niphadias (see discussion) except for the presence of 2–5 enlarged spines
on the pseudotegumen and extensive white patterns in the medial and sub-marginal areas of the forewing.
Description. Male. (Fig. 1A) Head: labial palpus three segmented, short, less than half length of head, directed
forwards, second palpomere longer than first, third palpomere shortest, sub-orbicular, covered in dark grey scales.
Antennae (Figs 3K–L) 7 mm, 43 flagellomeres, pale yellow, quadripectinate, each flagellomere with pair of broad,
flattened rami oppositely-situated. Length of central primary rami almost twice width of flagellomere while proxi-
mal and distal primary rami are shorter and flattened. Primary rami ciliated, hairs longer distally. Ventral surface
with short ridge adjoining thickened proximal bar, also heavily ciliated. Scape cylindrical, pedicel ovoid, flattened.
Eyes larger than head capsule, widely separate. Scales on frons and vertex dense and dark brown.
Thorax: pro- and mesothorax dorsally and ventrally covered in dark brown scales and interspersed with cin-
namon brown scales on dorsal surface, with greater suffusion of cinnamon brown on ventral surface. Wings: FW
length 22–24 mm, expanse 45–48 mm, broad, triangular. Costa slightly concave centrally, convex towards apex.
Wing venation oxycanine (Fig. 10A), vein 2A absent. HW 17–18 mm, triangular at apex, narrower, tornus rounded,
vein CuP partly membranous. Dorsal FW ground colour mix of grey and buff scales, dark brown at costa, basal
area; fringe grey; dark brown scales on all veins. Three yellow-cream spots present basally, within discal cells, in
triangular configuration, indistinct in worn or faded specimens. Some specimens with four irregular cream marginal
and sub-marginal bands from Rs2 to CuA2, with brown scales between and adjacent to these bands. Ventral surface
as for dorsal, veins highlighted with short, grey scales, cells between veins sparsely covered with long grey hair-like
scales, pale cream markings absent. Darker scales at apex and margin. Lighter brown piliform scales present along
costa and basal area, scales cinnamon brown basally. HW dorsal surface brown, darker brown at apex, sparsely
scaled and semi-translucent within discal area. Costa and veins with pale cinnamon brown scales, scales on veins
less elongate. Wing margin light cinnamon brown, fringe dark brown. Basal area and fringe along dorsum with long
light cinnamon piliform scales, tending towards orange or blush red in fresh specimens. HW ventral surface as dor-
sal, except with dark brown margin, dark brown scales over wing veins. Fringe buff-grey, sparsely covered in longer
hair-like scales. Legs dark brown dorsally and ventrally, grey on tarsus ventral surface, femur and tibia with brown
piliform scales ventrally, broadly triangular, short epiphysis on fore-tibia, aerolium present.
Abdomen: narrow, tapering, tergites at proximal end with blush-red piliform scales when fresh, gradually merg-
ing with shorter brown and cinnamon brown scales towards distal end. Rounded tuft of blush-red piliform scales
on final tergite. Ventral surface dark brown. Sternum 8 broad and rectangular, posteriorly becoming membranous,
junction with saccus indistinct.
Genitalia (Figs 4A–C, 7A, diss. no. 31-019191, paratype male). Disto-posterior arm of saccus broad, and con-
cave, trapezoidal, at right angles to sternite 8. Disto-posterior arm of vinculum narrow. Intermediate plate narrow,
rectangular, free. Pseudotegumen: basal rim narrow, globular; dorso-posterior and disto-posterior margins narrow in
postero-ventral view highly sclerotized. Dorso-distal twin processes short, apex rounded. Ventro and disto-posterior
margin of pseudotegumen in lateral or 3-quarter views are a large, curved axe-head shape, with a deep invagination
present just before ventral pseudoteguminal arms, both of which are short and fused with proximal points very short
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 355
and free. Juxta broad, subrectangular, bilobate at anterior margin. Trulleum trapezoidal, semi-transparent. Valvae:
elongate and lobate, flattened laterally, rounded apically, wider distally, sacculus short and oval, internal surface
densely covered with long curved setae. Apex of valvae in line with dorso-posterior margin of pseudotegumen.
Apodemal vinculum broad and short, posterior margin rounded. Fusion of sternum 8 and saccus forming a continu-
ous structure. Para-anal sclerite absent.
Female. Unknown.
Habitat. Oxycanus ephemerous sp. nov. is known only from South Australia, from the Fleurieu Peninsula and
Kangaroo Island bioregion (Inman Valley and American River respectively). The Inman Valley site is mainly farm-
land with some low Eucalyptus woodland and a shrub/heath and sedge understory, near a seasonal creek (S. Eden
pers. obs.). Both these locations fall within an area of annual rainfall of 400–600 mm, among the wettest parts of the
state. Similar conditions and habitats exist on the central-western part of Kangaroo Island and on the Eyre Peninsula
and Yorke Peninsula, however, the species is not recorded from those locations. Oxycanus ephemerous sp. nov.
occurs in a distinctly drier habitat compared to that of the similar O. subvaria, a species which occurs primarily in
tall Eucalyptus wet forest, rainforest, montane wet forest, and sub-alpine woodland (Edwards & Green 2011; Kal-
lies et al. 2015) in eastern Victoria, central-coastal and south-eastern NSW and Tasmania in areas of rainfall above
800–1000 mm annually.
Remarks. The two species Oxycanus ephemerous sp. nov. and O. subvaria have allopatric distributions sepa-
rated by approximatelty 630 km between Inman Valley and Melbourne. Oxycanus niphadias is the only other Oxy-
canus species currently identified as a South Australian endemic. Three specimens from the Grampians, VIC, in the
Melbourne Museum listed as O. niphadias (Kallies et al. 2015) were examined by the authors from photographs
(provided by J. Schubert) along with a specimen from Halls Gap, VIC in SAMA and all are considered to be O. oc-
cidentalis Tindale, the first records for this species from Victoria. Oxycanus occidentalis also occurs in south-west
WA (Tindale 1935) and in South Australia on the Eyre Peninsula, Riverland and Kangaroo Island (M. Hura, A. Kal-
lies pers. comm. and MDM pers. obs.). All three O. ephemerous sp. nov. specimens were collected on the 20th of
May in both 1952 and 2018, following rainfall the day before (Australian Bureau of Meteorology 2019). The larval
biology of O. ephemerous sp. nov. is unknown. A concerted effort was made by the authors to locate additional
specimens of O. ephemerous sp. nov. and O. niphadias in April and May of 2018 and 2019 without success. The ap-
parently restricted distribution and localised nature of the records raises concerns in a conservation setting, as both
species, but particularly O. niphadias, may be threatened with extinction due to urbanisation and loss of habitat.
Further work is required to determine the conservation status of these species.
Oxycanus subvaria (Walker, 1856)
(Figs 1B; 3I–J; 4D–F; 7B; 8A & D)
Elhamma subvaria Walker, 1856: 1562. Nielsen et al. 1996: 26. Nielsen et al. 2000: 858.
Lectotype: male (Fig. 1B), Lectotype, ♂ ‘Lectotype / Elhamma subvaria Wkr, Van Diemans Land / Specimen photographed E.S.
Nielsen 1984’, NHM. Lectotype selected by Ebbe S. Nielsen, here designated.
Type locality: ‘Van Diemen’s Land’ (Tasmania).
Oxycanus subvarius Walker 1856: 1575. Tindale 1935: 324.
Oxycanus lamnus Tindale 1935: 325, synonymized by Nielsen et al. 1996: 26.
Holotype male, Manly, (NSW) 31 May 1907, I18881, SAMA Database no. 31-000363’, SAMA.
Paratypes: 3 males, 1 female. ♀, Manly, (NSW), 30 May 1907, SAMA Database no. 31-000364’, two ♂, ‘Manly, SAMA
Database no. 31-000366’ and ‘SAMA Database no. 31-000367’, one ♂, ‘Clifton NSW SAMA Database no. 31-000365’,
SAMA.
Type locality: Manly, New South Wales, Australia.
Oxycanus hildae Tindale, 1964: 665. Nielsen et al. 2000: 858; syn. n.
Holotype male, Jacob Creek, (VIC), 25 April 1946, C. G. L. Gooding, I19113, SAMA Database no. 31-000517’, SAMA.
Paratype: 1 female. ♀: ‘Jacob Creek, (VIC), 25 April 1946, C. G. L. Gooding, I19113, SAMA Database no. 31-015511’,
SAMA.
Type locality: Jacob Creek, Victoria, Australia.
Additional material examined: in total 77 specimens (65 ♂, 12 ♀). 1 ♂, Manly, NSW, 23 May 1909. 1 ♂, Austin-
mer, NSW, 22 April 1923, Nicholson. 8 ♂, Ridgeway, TAS, 07 April 1950, M. Tagg. 9 ♂, 8 ♀, Moe, VIC, C.G.L.
Gooding, 23 April 1929, 5 ♂, same data except 30 April 1929, 1 ♂ same data except 30 April 1930, 1 ♀ same data
BEAVER ET AL.
356 · Zootaxa 4732 (3) © 2020 Magnolia Press
except 28 April 1937, 6 ♂, 3 ♀ same data except 16 April 1938, 12 ♂ same data except 17 April 1938, 5 ♂, same
data except 30 April 1942 (all in SAMA). 1 ♂ same data except 03 May 1948. 1 ♂, Cambridge, TAS, J.R. Cun-
ningham, 10 May 1953 (both in TMAG). 2 ♂, Croydon, VIC, B.M.W. Prieby, 03 May 1908. 2 ♂, Moondarra Dam,
VIC, M&G Coulson, 16 May 1972. 1 ♂, Trafalgar, VIC, J.A. Kershaw, 01 March 1909. 1 ♂, Hazlewood, VIC, J.H.
Courtenay, 06 May 1948. 1 ♂, Beaconsfield, VIC W.E. Drake, 05 March 1910. 5 ♂, Mt Kiera, NSW, C.E. Chad-
wick, May–June 1954. 1 ♂, Narara, NSW, B.L. Middleton, May 1947. 1 ♂, Cathcart, NSW, N.B. Tindale, 11 March
1958. 1 ♂, Lorne, VIC, J. Womerley, 19 March 1952. 2 ♂, Mt Disappointment, Whittlesea, VIC, M.D. Moore &
N. Temby, 06 March 2017. 1 ♂, no data (all in SAMA). 2 ♂, Mt Disappointment, Whittlesea, VIC, 18 March 2017,
K. Green, specimen no. 18066 and 18069. 1 ♂, Burramunga, VIC, 38°37’24.3”S 143°44’10.8”E, J. Schubert, 09
March 2019 (all in EPBC).
Dissected: 4 ♂: Moe, VIC 16 April 1938, Dissection ID 31-019120-EPB. ‘Victoria’, Dissection ID 31-019149
EPB. Ridgeway, TAS 7 April 1950, Dissection ID 31-019104-EPB. Manly, NSW, 23 May 1909, Dissection ID 31-
019096-EPB (all in SAMA).
Justification of the synonymization act. Oxycanus subvaria is highly variable in both wing pattern and colou-
ration, this coupled with subtle variation in the shape of the pseudotegumen and valva in male genitalia has led to
some taxonomic confusion, with Oxycanus lamnus representing an extreme of this variation. Oxycanus lamnus was
described by Tindale (1935) as having a larger marginal process of the pseudotegumen, and “suspensorial spines”
(interpreted as the ventral arms of pseudotegumen) as produced and curved. However, these features exist in many
examples of O. subvaria and vary among sympatric and synchronic specimens. Tindale also noted that the species
differ by the position of the markings on the wings. However, examination of a large series of O. subvaria reveals
this feature to be highly variable and insufficient for identification. Oxycanus lamnus was recognised as a synonym
of O. subvaria by Nielsen et al. (1996). A similar situation has occurred with Oxycanus hildae syn. n. that Tindale
(1964) described as having a “simple arcuate latus” of the pseudotegumen, however the ventral arms of the pseudo-
tegumen, so typical of O. subvaria are present in the O. hildae holotype, the structure is however partially contracted
into and obscured by the vinculum, which may have led to the apparent confusion. The external genitalia of the
female paratype conforms to that of other O. subvaria from Victoria. The speckled wing patterning is not unique, an
O. subvaria specimen containing the same wing pattern is in the EPB collection and a similar living specimen with
this pattern is figured in Kallies et al. (2015). One Oxycanus hildae syn. n. paratype from Cathcart NSW could not
be located. Kallies et al. (2015) speculated on the validity of O. hildae syn. n. and it became necessary to deal with,
so we state here unambiguously Oxycanus hildae Tindale (1964) is a synonym of O. subvaria (Walker, 1856).
Remarks. A lectotype status is formally recognised for Elhamma subvaria concerning a male specimen in the
NHM (Fig. 1B). The specimens intended to be designated as lectotypes for several Australian Hepialidae species
were selected by Ebbe S. Nielsen however were unpublished before his death in 2001 (see Simonsen 2018, pg. 7 for
additional information). Herewith we designate the lectotype for Elhamma subvaria species-group name. The lecto-
type specimen (Fig. 1B) was chosen by Ebbe S. Nielsen from the syntype series at the NHM, London and carries the
following labels “Lectotype / Elhamma subvaria Wkr, Van Diemans Land / Specimen photographed E.S. Nielsen
1984”. The purpose of this lectotype designation is to ensure the taxonomic stability of species-group nomenclature
within the genus Oxycanus.
Oxycanus flavoplumosus Beaver & Moore, sp. nov.
(Figs 1C–E; 3C–D; 5A–C; 7D; 8E; 10B)
Type specimens: Holotype male, SAMA. Paratypes: 80 males, SAMA, TMAG, ANIC, QM, UQIC, AM.
Type locality: Ebor, New England Tableland, New South Wales, Australia.
Etymology. The name flavoplumosus (Latin, flavo = yellow, plumosus = feathery or downy) refers to the dense yel-
low vestiture present in most specimens on the abdomen, thorax and hindwings.
Type material. HOLOTYPE, male. (SAMA) ♂, Ebor, NSW, 27 March 1941 / SAMA Database No. 31-
018764.
PARATYPES: 80 ♂ in total. 8 ♂ (SAMA): All with the label data ‘Ebor, NSW’ and differ only by way of col-
lection dates: 2 ♂ 27 Mar 1941 / SAMA no. 31-018763, 31-018765. 1 ♂ 28 Mar 1941 / SAMA no. 31-018762 /
Dissection ID 31-018762-EPB. 2 ♂ 20 April 1941 / SAMA no. 31-018766, 31-018767. 1 ♂ April 1949, / SAMA no.
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 357
31-018768. 1 ♂ 15 March 1940 / SAMA no. 31-018769. 3 ♂ 14 March 1940 / SAMA no. 31-018770, 31-018850,
31-018845. 3 ♂ (TMAG): Ebor, NSW, 18 March 1943, database numbers F2224 to F2226. 37 ♂ (ANIC): 2 ♂: Bar-
rington Tops, NSW, 31.55S 151.31E, 1370m, 23 March 2009, G. Cocking / ANIC no. 31-036845 and 31-036846.
A further 2 ♂ same data except 25 March 2009, ANIC no. 31-036847 and 31-036848; Dissection ID EPB-ANIC-
16. 1 ♂ Junction Pools, Barrington Tops, NSW, 32.02S 151.27E, 1420m, 24 March 2009, G. Cocking / ANIC no.
31-036851. 2 ♂, Gloucester Tops Road, NSW, 32.04S 151.36E, 1200m, 16 May 2007, G. Cocking / ANIC no. 31-
036849 and 31-036850. 1 ♂ Dilgry River camping area, Barrington Tops 1180m elevation, 19 March 1993 / ANIC
no. 31-036843. A further ♂ same data except 08 March 1991, ANIC no. 31-036844. Following 28 ♂ all with ‘Ebor,
NSW, B.L. Middleton’, and differ only by way of collection date: 1 ♂ 22 May 1939 / ANIC no. 31-036783. 2 ♂ 15
March 1940 / ANIC no. 31-036797 and 31-036799. 2 ♂ 20 March 1941 / ANIC no. 31-036789 and 31-036769. 2 ♂
25 March 1941 / ANIC no. 31-036786 and 31-036807. 1 ♂ 27 March 1941 / ANIC no. 31-036800. 4 ♂ 28 March
1941 / ANIC no. 31-036782, 31-036787, 31-036802, 31-036808. 1 ♂ 23 March 1943 / ANIC no. 31-036801. 2 ♂ 20
March 1943 / ANIC no. 31-036781, 31-036785. 1 ♂ 22 March 1943 / ANIC no. 31-036804. 8 ♂ 24 March 1943 /
ANIC no. 31-036784, 31-036788, and 31-036790 to 31-036795. 1 ♂ 25 March 1943 / ANIC no. 31-036798. 1 ♂ 1
April 1943 / ANIC no. 31-036805. 1 ♂ 4 April 1943 / 31-036806. 1 ♂ 19 April 1949 / 31-036803. 1 ♂ (QM): Ebor,
NSW, 15 March 1940. 2 ♂ (UQIC): 1 ♂ Dorrigo, NSW, 11 April 1968, H. Burton. 1 ♂ Ebor, NSW, 28 March 1943.
20 ♂ (AM): All are from Ebor, NSW, and differ only by way of collection date: 1 ♂ 16 March 1940 / K.570188.
1 ♂ 18 March 1941 / K.570194. 1 ♂ 18 March 1943 / K.570195. 1 ♂ 20 March 1941 / K.570193. 4 ♂ 27 March
1941 / K.570178; K.570179; K.570186; K.570191. 2 ♂ 28 March 1941 / K.570192; K.570185. 1 ♂ 20 March 1943
/ K.570177. 3 ♂ 23 March 1943 / K.570187; K.570181; K.570182. 2 ♂ 30 March 1943 / K.570184; K.570183. 1 ♂
17 April 1949 / K.570189. 1 ♂ 18 April 1949 / K.570180. 2 ♂ April 1949 / K.570190; K.570176.
Distribution. All specimens are from Ebor and Dorrigo on the New England Tableland, and the Barrington
Tops area, of the Hunter Region, New South Wales, Australia (Figs 11, 13).
Diagnosis. This species has a similar pseudotegumen shape to that of the sympatric and rarely collected Oxy-
canus byrsa (Figs 1F, 3A–B, 5A–C, 7E, 8B) however O. byrsa has the posterior margin of the pseudotegumen scal-
loped rather than smooth as it is in O. flavoplumosus sp. nov. The dorso-posterior margin is membranous in the new
species (sclerotized in O. byrsa), the para-anal sclerite is recurved (blunt in O. byrsa) and the apodemal vinculum is
broad and rounded compared with O. byrsa where the anterior margin ends in a narrow point. Wing patterns are also
much less complex in O. flavoplumosus sp. nov. compared with O. byrsa (Fig. 1F), and this species is smaller (FW:
34–41 mm) than O. byrsa which has a FW length of 50–75mm. Additionally, the antennal structure is distinct (Figs
3C–D), with the proximal bar shorter in O. flavoplumosus sp. nov. compared with that in O. byrsa. The species can
superficially resemble O. spadix Tindale, 1935 or other species which show a great degree of variation in wing pat-
tern such as O. sirpus Tindale, 1935 or the allopatric O. promiscuus Tindale, 1935 from Western Australia. However,
O. spadix is generally smaller at less than 34 mm FW length. As figured by Tindale (1935) the pseudotegumen in
all other species have prominent enlarged spines or processes, in contrast to the smooth, low pseudotegumen of O.
flavoplumosus sp. nov. Antennae are also different with rami relatively much longer in O. flavoplumosus sp. nov.
and greatly reduced in those other species previously mentioned, and from O. herdus Tindale, 1935 and O. silvanus
Tindale, 1935 where the very long rami are several times longer than the width of the flagellomere (Tindale 1935;
Kallies et al. 2015) compared with O. flavoplumosus sp. nov. which has distinctly shorter rami, however the bar-
rami are more pronounced in O. flavoplumosus sp. nov., with longer cillae.
Description. Male (Figs 1C–E) Head: Antennae (Figs 3C–D), 14 mm, 45 flagellomeres, to basal area of costa,
yellow, quadripectinate with proximal bar and ridge present, primary rami short, less than width of flagellomere,
densely ciliated at distal ends of ventral surface, ridge low, proximal bar short, rounded, only marginally wider than
flagellomere width. Scape cylindrical, pedicel ovoid, flattened. Eyes large in relation to head capsule, over half
of total head width, scales on frons and vertex dense and light brown to greyish-brown. Palps short, less than half
length of head, three segmented, directed forwards, second palpomere longer than first, third shortest, globular.
Thorax: pro- and mesothorax covered in light brown to greyish brown, and occasionally yellow, densely layered
scales, darker on ventral surface. Legs pale brown dorsally, yellow ventrally, densely scaled, epiphysis short, trian-
gular. Wings: wingspan 86–90 mm, FW: 34–41 mm, narrow, triangular. Costa mostly straight, convex towards apex.
Apex weakly pointed. Wing venation oxycanine (Fig. 10B), vein 2A small, near margin and base of jugum. HW:
32–35mm, elongate, similar shape to forewing but rounder tornus. Dorsal FW ground colour variable, yellowish
brown or brownish-grey, browner towards termen with more ochreous scales towards basal area and intermediate
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shades between. Small, cream trapezoid spot in proximal cell edged with dark brown, wider proximally, occasion-
ally a smaller spot present between M1 and M2 or Rs4 and M1. Variable darker brown suffusion especially in medial
and submarginal areas, appearing sometimes as small freckles, in other specimens as broad blotches, more notice-
able on paler or yellow specimens, spots in variable barred rows. Costa brown, darker proximally and fading into
wing ground colour distally. Outer margin with small dark brown spots between each wing vein in all specimens,
particularly between Rs1 and M3. Ventral surface uniformly ochreous-yellow or light brown with yellow piliform
scales on basal and costal areas. HW dorsal surface without markings, mustard-yellow or rarely light brown. Basal
area covered with long, dense piliform scales of same ground colour. Ventral surface as dorsal, with greater extent
of piliform scales.
Abdomen: Narrow, tapering, all tergites covered with mustard-yellow scales on all surfaces, giving most speci-
mens a broad or ‘fluffy’ appearance. S8 broad and subsquare, wider than long, corners at anterior end rounded, not
fused with saccus.
Genitalia (Figs 5A–C, 7D, dis. no. 31-018762). Pseudotegumen: basal rim broad and rounded, distal end acu-
minate, dorso and disto-posterior margins narrow in postero-ventral view, highly sclerotized; dorso-distal twin pro-
cesses large, broad and with rounded apex; deep longitudinal groove visible in lateral view, ventral ridge forming a
scythe/question mark shape; dorso-posterior margin becoming membranous towards para-anal sclerite. Ventro- and
disto-posterior margins are highly sclerotized in lateral view, ventrally flattened and smooth with a steeply curving
concavity present just before ventral pseudoteguminal arms, which are moderately long, free and pointed. Valva:
short, without ridges, lobate, parallel sided, apically rounded, flattened laterally and with short, squared sacculus,
covered with numerous long curved setae; distal end shorter than disto-posterior margin of pseudotegumen when in
closed position. Apodemal vinculum narrow and short, sub-triangular, narrow anterior margin. Saccus broad and tri-
angular, disto-posterior arm broad and concave, almost at right angles to sternite 8. Juxta square, concave. Trulleum
triangular, sclerotized. Intermediate plate narrow, sub-rectangular, free. Para-anal sclerite large, elongate, strongly
falcate, apex recurved distally, positioned between disto-posterior margin and dorso-distal twin processes, arising
proximally between each pseudotegumen.
Female. Unknown.
Variation. Some male specimens with the pseudotegumen with small raised bumps in middle, para-anal scler-
ites less strongly sclerotized.
Remarks. Similarities in genitalia and antennae as outlined in the diagnosis lead to the conclusion that O. flavo-
plumosus sp. nov. and O. byrsa are closely related. The male genitalia do not appear to vary greatly within members
of either species. Nothing is known of their larval biology. Oxycanus flavoplumosus sp. nov. has been taken between
the 8th of March and 22nd of May, which is typical for most Oxycanus species that fly during autumn rainfall on
mild nights. The species has not been collected in recent years from Ebor, and a concerted effort should be made to
confirm the presence of extant populations on the New England Tableland, and assess potential conservation obliga-
tions. More recent specimens are known from Barrington Tops NP, where the species is likely still present.
Oxycanus petalous Beaver & Moore, sp. nov.
(Figs 2A–B; 3G–H; 6A–C; 7F; 8C, 9A–B; 10C)
Type specimens: Holotype male, WAM. Paratypes: 28 males and 9 females, SAMA, ANIC.
Type locality: Boxwood Hill, Chingarrup, South Coast, Western Australia, Australia.
Etymology. The name ‘petalous’ is from the Greek petalo (πέταλο) which translates to ‘horseshoe’ and is in refer-
ence to the horseshoe-shaped eighth sternite (S8) of the male.
Type material. HOLOTYPE. (In WAM) ♂. West Aust., South Coast, Chingarrup, Boxwood Hill, 34°20’46”S,
118°43’51”E, M. Moore, 1st May [20]17 / SAMA Database no. 31-020332 / spec. no. 17123 leg removed for tis-
sue storage MD Moore. PARATYPES. 37 specimens: 9 ♀, 28 ♂. 1 ♂, Boxwood Hill, 34°18’17”S, 118°43’43”E,
M. Heath, 25th April 2015 / SAMA Database no. 31-019833 / spec. no. 19024. 1 ♂, 1♀, West Aust., Boxwood
Hill, 34°18’17”S, 118 43’43”E, Mark Heath; 25th April 2015 / SAMA Database no. 31-019832 / spec. no. 19025
leg removed for tissue storage MD Moore / MH 5; male with SAMA Database no. 31-019831. 2 ♀, March 2015,
Gleneagles Rest Area, Western Aust., 32°18’3” S 116°12’3” E, P. Kay / spec. no. 128 leg removed for tissue stor-
age MD Moore / SAMA Database No. 31-019829; second specimen with SAMA Database No. 31-019830 and
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 359
spec. no. 129; (all in SAMA). 1 ♂ 22–23 April 2000, nr. Mt Barker, Western Australia, 34.41.404S, 117.47.374E,
M. Court, to light trap, Eucalyptus globulus plantation #1, 3 / Barcode of life DNA voucher specimen sample ID:
10ANIC-09486, BOLD proc. ID: ANICM489-10 / Dissection ID EPB-ANIC-1. An additional 2 ♀ and 2 ♂ with the
same data, except ANIC no 31-036828-31-036829 for the females and 31-036810, 31-036812 for the males. An ad-
ditional 16 ♂, 3 ♀, same data except the date 20–21 April 2000, females with ANIC no’s 31-036830 to 31-036832,
males with ANIC no’s 31-036809, 31-036811, 31-036814 to 31-036827. 1 additional ♀, same data except dissection
ID EPB-ANIC-2. 1 ♂, same data except 19-20 April 2000, ANIC no. 31-036813. 2 ♂, Peak Charles NP, WA, Aust.,
32°53’56.4”S 121°11’01.0”E, 16 April 2007, MV lamp and UV-fit, A. Zwick and G. Cocking / Dissection ID EPB-
ANIC-8 and EPB-ANIC-14. 1 ♂, Stirling Range, WA, 26 Mar 1968, I.F.B. Common & M.S. Upton / Dissection ID
EPB-ANIC-7, ANIC no. 31-036833. 3 ♂, Stirling Range Nat. Park, WA, 34.23S 118.07E, 17 April 1983, I.F.B. E.S.
Nielsen & E.D. Edwards, ANIC no. 31-036834-31-036836 (all in ANIC).
FIGURE 1 (A–F). Adult male Oxycanus species: A, Oxycanus ephemerous sp. nov. holotype, ©SAMA; B, Oxycanus subvaria
lectotype; C, Oxycanus flavoplumosus sp. nov. holotype, photo by T. Simonsen, ©Trustees of the NHM; D, E, O. flavoplumosus
sp. nov. paratypes, ©SAMA; F, O. byrsa, courtesy of J. Schubert. Scales 10 mm.
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FIGURE 2 (A–D). Adult Oxycanus species: A, Oxycanus petalous sp. nov. holotype male; B, O. petalous sp. nov. paratype
female; C, O. determinata male; D O. determinata female, ©SAMA.
Distribution. All specimens taken from the far south of Western Australia, from Mount Barker; Gleneagles
Rest Area (Fig. 15); Stirling Range NP; Boxwood Hill; and Peak Charles (Fig. 11).
Diagnosis. This species falls within a small ‘WA species group’, a group distinguished from other Western Aus-
tralian species by the presence of para-anal sclerites in the male genitalia. The sclerotised horseshoe shaped sternum
8 distinguishes males from the trapezoidal sternum 8 of O. determinata (Figs 2C–D, 3E–F, 6D–F, 7C, 8F, 9C–D)
and O. kochi. Sternum 8 in O. glauteri is less distinctly curved (Tindale 1955: figure 23), while the pseudotegumen
differs markedly particularly in the shape of the dorso-posterior margin of the pseudotegumen which bears a distinct
vertical spine absent in O. petalous sp. nov. The pseudotegumen of O. petalous sp. nov. has a small spine 1/3rd the
length from the proximal end, a large medial spine, and a larger broadly triangular spine basally towards the vincu-
lum (Fig. 7F). It also has relatively greater fusion at the apex of the pseudoteguminal arms. The females can only
be reliably differentiated from O. determinata by examination of genitalia characters (Figs 9A–B). In O. petalous
sp. nov. the dorsal plate is strongly arched and has posterior, elongated finger-like projections. The sub-anal sclerite
is rectangular when viewed postero-ventrally. The lateral lobe of the antevaginal lamella is narrower, with a sub-
triangular base and is setose only at the distal end whereas in O. determinata it is mostly setose. The forewing apex
is slightly rounder than in O. determinata.
Description. Male (Fig. 2A) Head: Antennae (Figs 3G–H), 9 mm, 53 flagellomeres, to basal area of costa,
yellow, weakly bipectinate giving a general filiform appearance, primary rami very short and flattened, ciliated,
ridge present, ciliated, proximal bar absent. Scape elongate, cylindrical; pedicel ovoid, flattened. Eyes larger than
twice width of head, scales on frons and vertex dense, dark brown to black coloured. Palps long, half length of
head, three segmented, directed forward, second longer than third, with first palpomere shortest; club-shaped distal
palpomere.
Thorax: pro- and mesothorax covered in dark grey, brown, black densely layered scales, darker on ventral sur-
face. Legs dark brown, densely scaled, epiphysis present, slightly longer than wide, triangular. Fore- and mid legs
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dark, hind legs covered with bright yellow scales. Wings: FW: 25–27 mm, expanse 60 mm, elongate, broadly tri-
angular. Costa slightly concave centrally, convex towards apex. Apex pointed. Wing venation classically oxycanine
(Fig. 10C), vein 2A extends to margin, short. HW: 20–21 mm, elongate, similar shape as forewing but slightly short-
er, with sharper termen, CuP partially membranous medially. Dorsal FW ground colour dark brown or brownish-
grey, darker along costa and apex, basal area buff-grey. Lighter brown scales interspersed with darker scales across
entire wing surface. Small, angled cream bar across posterior end of discal cell in some specimens. Some specimens
with small triangular white spots in sub-marginal area between Rs4 and M2. Some specimens have a diverse range
of small rounded or trapezoidal white spots. Veins with darker scales than general wing surface, latter less densely
scaled. Outer margin with fringe of longer, pale brown scales. Ventral surface as above with piliform scales on basal
and costal areas, the latter lighter brown, lacking white spots. HW dorsal surface yellow or light brown, unmarked.
Basal area semi-translucent, finely covered with long piliform scales of same ground colour. Apex dark brown, wing
veins Rs1 to Rs3 covered in dark brown scales, others with yellow scales. Fringe dark brown, becoming yellow
towards tornus. Ventral surface as above or with bottom half of wing entirely buff or yellow, with greater extent of
piliform scales, veins edged with yellow scales.
FIGURE 3 (A–L) Antennae of male Oxycanus spp. A, B, O. byrsa; C, D, O. flavoplumosus sp. nov; E, F, O. determinata; G, H,
O. petalous sp. nov.; I, J, O. subvaria; K, L, O. ephemerous sp. nov. Abbreviations: br—bar. ra—primary rami. ri—ridge. Scale
bar in A–C is 500 µm; D is 200 µm; E, F is 500µm; G is 150µm H, I is 250 µm; J is 250 µm; K is 500 µm; L is 250 µm.
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FIGURE 4 (A–F) Oxycanus spp. male genitalia. A–C O. ephemerous sp. nov. dissection ID 31-019191-EPB; D–F O. subvaria
dissection ID 31-019120-EPB. Abbreviations: brp—basal rim of pseudotegumen. dpa—disto-posterior arm of vinculum. dsa—
disto-posterior arm of saccus. ip—intermediate plate. jx—juxta. pst—pseudotegumen. sc—saccus. scc—sacculus. s8—sternite
8. tp—twin process of pseudotegumen. tr—trulleum. v—apodemal vinculum. va—valva. vpa—ventral pseudoteguminal arms,
©SAMA.
Abdomen: Narrow, tapering, first 3–4 tergites covered with mustard-yellow to cinnamon brown coloured scales
on all surfaces, brown on posterior tergites. Sternum 8 horseshoe shaped, highly sclerotized, more than all other
sternites, anterior corners rounded, posterior corners acute. Sternum 8 unfused from apodemal vinculum, setose at
posterior margin.
Genitalia (Figs 6A–C, 7F, dis. no. 31-019833; dis. no. EPB-ANIC-1). Pseudotegumen: basal rim broad and
rounded, centrally depressed with margin upturned. Dorso-distal twin processes heavily sclerotized, elongate, leaf
shaped, and with apex rounded and cupped; in lateral view a deep groove is present, otherwise smooth; dorso-pos-
terior margin membranous, disto-posterior margin finely serrated, coming to a sharply acuminate point at apex.
Ventro- and disto-posterior margins are highly sclerotized in lateral view, broad and convex, with a small spine 1/3rd
along margin, larger spine medially, and a larger triangular spine basally towards vinculum. Ventral pseudotegumi-
nal arms indistinct, lobes fused across median. Valvae: long, narrow, distally lobate with long setae, flattened later-
ally with deep, triangular sacculus. Distally shorter than disto-posterior margin of pseudotegumen when in habitus
position. Para-anal sclerite very large relative to pseudotegumen, elongate, upright, base large, sub-triangular, with
ventral-oriented projection, apex club-shaped and curving laterally, becoming membranous along proximal rim,
positioned well back between disto-posterior margin of pseudotegumen and dorso-distal twin processes which also
curve back. Apodemal vinculum narrow, generally triangular; dorso-posterior arms deep, in lateral view anterior
end broad. Saccus broad, sclerotized, generally triangular and large, bilobed posterior-central margin, dorso-pos-
terior arms narrow, sclerotised, ventrally-oriented, membranous at end of dorso-posterior arms, where membrane
puckers and folds outwards as a fringe, in lateral view anterior end bulbous. Juxta trapezoidal, concave; anterior
margin narrow, bilobate; posterior margin straight. Trulleum sub-rectangular, sclerotised, anterior corners rounded.
Intermediate plate sclerotised, generally ovoid, anterior margin rounded, posterior margin connected to basal rim
by membrane.
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FIGURE 5 (A–F) Oxycanus spp. male genitalia. A–C O. flavoplumosus sp. nov. dissection ID 31-018762-EPB; D–F O. byrsa
dissection ID 31-018407-EPB. Abbreviations not already used: pas—para-anal sclerite, ©SAMA.
FIGURE 6 (A–F) Oxycanus spp. male genitalia. A–C, O. petalous sp. nov. dissection ID EPB-ANIC-1; D–F, O. determinata
dissection ID 31-018441-EPB, ©SAMA.
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FIGURE 7 (A–F) Oxycanus spp. male genitalia, 3-quarter view. A, O. ephemerous sp. nov. dissection ID 31-019191-EPB;
B, O. subvaria dissection ID 31-019120-EPB; C, O. determinata dissection ID 31-018441-EPB; D, O. flavoplumosus sp. nov.
dissection ID 31-018762-EPB; E, O. byrsa dissection ID 31-018407-EPB; F, O. petalous sp. nov. dissection ID EPB-ANIC-1,
©SAMA.
Female. (Fig. 2B) Head: Antennae 7 mm, 44 flagellomeres, extending to basal area of costa, yellow, weakly bi-
pectinate, filiform appearance, poorly ciliated with few narrow and long setae. Eyes proportionally similar to male,
scales on frons and vertex dense and light brown to grey. Palps longer than half length of head, three segmented,
forward-projecting. Second palpomere longest, with distally club-shaped third palpomere longer than first.
Thorax: pro- and mesothorax covered in light brown-grey densely layered scales, darker on ventral surface.
Legs light brown, densely scaled, epiphysis moderately long, triangular. Hind legs lighter as in male. Wings: FW:
39 mm, expanse 81 mm, elongate, narrowly triangular. Costa slightly concave centrally, convex towards apex.
Apex rounded. Wing venation oxycanine. HW: 30 mm, elongate, of a similar shape as forewing but shorter, with
narrower termen. Dorsal FW ground colour light brown or grey, sparsely scaled giving a generally semi-translucent
appearance but not in basal area or along costa. Scales light brown-grey on wing veins, straw coloured short scales
on surface. Small, angled cream spot across posterior end of discal cell usually present. Outer margin with fringe of
longer scales brown, becoming light buff towards basal area of tornus. Ventral surface as above with greater extent
of piliform scales on basal and costal areas. HW dorsal surface light brown to grey, unmarked. Basal area finely cov-
ered with long piliform type straw-coloured scales. Ventral surface as above, with greater extent of piliform scales.
Abdomen: broad, exceeding wing margin in habitus, first 3–4 tergites dorsally covered with straw coloured
scales on all surfaces, light brown on posterior tergites.
Genitalia (Figs 9A, B; dis. no 31-019830-EPB; EPB-ANIC-2). Dorsal plate broad, subtriangular, with two
dorso-lateral digitiform posterior projections, setose; lateral lobe of antevaginal lamella reduced, sclerotised, sub-
triangular and setose; subanal sclerite prominent, sclerotised, broadly triangular and projecting dorsally between
‘fingers’ of dorsal plate; medial lobe of antevaginal lamella broadly rectangular, highly sclerotised, setose. Bursa
copulatrix not located during dissection, presumed damaged by egg mass.
Habitat. Oxycanus petalous sp. nov. has been taken from high-rainfall Eucalyptus tall forest environments, and
from a Eucalyptus globulus plantation at Mt Barker, WA.
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FIGURE 8 (A–F) Oxycanus spp. male S8. A, O. subvaria dissection ID 31-019120-EPB; B, O. byrsa dissection ID 31-018407-
EPB; C, O. petalous sp. nov. dissection ID EPB-ANIC-1; D, lateral view of S8 and saccus connection of O. subvaria dissection
ID 31-019120-EPB; E, O. flavoplumosus sp. nov. dissection ID 31-018762-EPB; F, O. determinata dissection ID 31-018441-
EPB, ©SAMA.
Remarks. Oxycanus petalous sp. nov. along with the Western Australian endemic species O. determinata, O.
kochi and O. glauteri share specialized similarities in the male genitalia such as elongate, broad para-anal sclerites,
and a long pseudotegumen fused at the ventral pseudoteguminal arms with species-diagnostic spines present on the
pseudotegumen at varying locations. All four species also share very long, forward-facing labial palpi with the third
palpomere longer than the basal palpomere. The antennae show some differences, with broader and longer rami in
O. determinata (Fig. 3E, F) compared with those of O. petalous sp. nov. (Fig. 3G–H), which also has fewer but
longer cilia. Generally, O. petalous sp. nov. is more plainly marked when compared with O. determinata but all four
species in this group present a wing pattern and colour variations that are typical for Oxycanus species. Examination
of the male genitalia has proved to be the only reliable method of separating these species. The species in this group
may be allopatric, as O. kochi is known only from Drummond Cove approx. 480 km NW of the nearest O. petalous
sp. nov. from Gleneagles Rest Area, while O. determinata is known by historic specimens from Perth and the Swan
River district, and by more recent material collected by MDM from Esperance, (Fig. 2C, D). These distributions
suggest that O. determinata is confined to coastal habitats, while O. petalous sp. nov. is from forested environments
further inland. The exact geographic provenance of O. glauteri is unknown other than ‘Western Australia’. This situ-
ation along with the few records of O. kochi and of O. determinata indicates a need for further collecting in Western
Australia to gain a more precise understanding of the biogeography of this species-group.
Molecular analyses
The Maximum-Likelihood phylogram (Fig. 12) reveals a well supported monophyletic Oxycanus clade (compared
to our outgroup taxa) that is further divided into two well supported sister clades. These well supported clades cor-
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respond to all species that we could reliably identify from our own specimens or from images (from boldsystems.
org) for the GenBank accessions included here (see also Supplementary Table S1). These clades highlight the close
intraspecific relationship for several instances where we obtained multiple individuals within species. For example,
the greatest sequence divergence was identified within O. hamatus Tindale (p-distance = 1.5%), while all others fall
within the range of intraspecific variation and varied between 0.2–0.9%, indicating variation expected within spe-
cies included here (Supplementary Table S2). These distances are close to those reported elsewhere for Lepidoptera
that indicate up to 2% within species variation using the same COI gene (e.g. Brown et al. 1999; Hausmann et al.
2011; Simonsen 2018; Grund et al. 2019; but see Simonsen et al. 2019 for some potential caveats). For two of the
new species described here (we were not able to obtain reliable sequence data from O. flavoplumosus) we only have
a single specimen for O. ephemerous, while the eight specimens for O. petalous range from 0–0.9% within sequence
divergence (Supplementary Table S2).
FIGURE 9 (A–D) Oxycanus spp. female genitalia. A & B, O. petalous sp. nov. female genitalia dissection ID EPB-ANIC-2; C
& D O. determinata female genitalia dissection ID 31-019873-EPB. Abbreviations not already used: dp—dorsal plate. la—lat-
eral lobe of antevaginal lamella. ml—medial lobe of antevaginal lamella. pp—posterior process of dorsal plate. sas—subanal
plate, ©SAMA.
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FIGURE 10 (A–C) Oxycanus spp. male wing venation, forewing and hindwing. A, Oxycanus ephemerous sp. nov.; B, Oxyca-
nus flavoplumosus sp. nov.; C, Oxycanus petalous sp. nov.
Comparisons between Oxycanus species reveal divergences of between 2.3% between O. maculosa Felder and
O. australis and up to 9.9% between O. sirpus and O. petalous (Supplementary Table S2). For our new species O.
petalous and O. ephemerous, we found levels of divergence among all other species ranged between 2.9–9.9% and
5.2–8.1%, respectively. These comparisons are consistent with all other species included here (Supplementary Table
S2). Our COI phylogram among Oxycanus species (Figure 12) indicates that these are well supported species that
correspond to levels of morphological variation observed, including the morphological variations described for O.
flavoplumosus here. Interestingly, the two highly supported Oxycanus sister clades separate into a Western Australia
clade with the Tasmanian O. sphragidias, and a clade containing all southern and eastern Australian species with a
clade containing the Western Australian species O. perditus and the broadly distributed O. occidentalis.
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368 · Zootaxa 4732 (3) © 2020 Magnolia Press
FIGURE 11. Distribution records of three new Oxycanus species: grey circle—Oxycanus petalous sp. nov.; filled circle—Oxy-
canus ephemerous sp. nov.; empty circle—Oxycanus flavoplumosus sp. nov.
Discussion
The description of the three new species has identified several aspects of the adult morphology which bring the
monophyly of Oxycanus into question, particularly with respect to antennae and genitalia morphology. Variation in
antennal structures in Australian Oxycanus can be divided into quadripectinate (sensu Dugdale 1994) and bipecti-
nate arrangemments (Fig. 3). Quadripectinate antennae also occur in the New Zealand oxycanine genera Wiseana
Viette, Dioxycanus Dumbleton, Heloxycanus Dugdale, and Cladoxycanus (Dumbleton). In oxycanines, bipectinate
rami often have a greatly reduced ridge-like ‘central process’, that is also present in Jeana and weakly in Dum-
bletonius Dugdale. Within Oxycanus, quadripectination characterises O. stellans Tindale, 1935, O. australis, O.
subvaria, and O. occidentalis species groups (sensu Kallies et al. 2015); illustrated by Kallies et al. (2015), and O.
byrsa and O. flavoplumosus (Fig. 3A–D respectively), while bipectination appears in the O. silvanus Tindale, 1935
and O. determinata species groups (Fig 3E, F for O. determinata and 3G, H for O. petalous) and in O. aedesima
(Turner). Quadripectination is rare in Hepialidae, and is more often associated with heteroneuran Lepidoptera with
large, complex antennae such as the Saturniidae. Superficially, without the use of a microscope the antennae of most
quadripectinate Oxycanus species generally appear bipectinate, as the primary rami are longest. Observation by the
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 369
authors of a large number of male Oxycanus specimens has determined that antennae in this genus are often species
and species-group specific and are likely to be of phylogenetic significance, in some species groups the proximal
bar is reduced to a singular, raised node (such as in Oxycanus sirpus Tindale) while in O. sphragidias (Meyrick) it is
distinctive and rami-like on an otherwise bipectinate flagellomere. In Jeana, Elhamma and O. silvanus the proximal
bar is absent, which may suggest that the absence of the proximal bar is plesiomorphic. In the Western Australian
O. determinata species group there is a swelling which may be analogous to that of the bar as seen in more typical
Oxycanus.
FIGURE 12. Maximum-Likelihood (using GTR+I+Γ) phylogram of COI sequences for Oxycanus, including six outgroup taxa.
The tree with the highest log likelihood (-3955.85) is shown with branch lengths measured in the number of substitutions per
site. Bootstrap values from 1,000 pseudoreplicates greater than 50% are shown at each node. Sequences generated from this
study have been combined with GenBank sequences only when the id’s could be confirmed morphologically, and a * indicates
where we have corrected a GenBank id (see Table S1).
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FIGURE 13. Barrington Tops NP, New South Wales. Montane rainforest and wet Eucalyptus forest habitat of Oxycanus flavo-
plumosus sp. n. and Oxycanus subvaria, ©Ethan P. Beaver.
There is some confusion surrounding wing venation in this genus. Tindale (1935) figured Oxycanus australis
with 3 anal veins (interpreted as CuP with 1A and 2A) in the hind wing and applied this to the genus as a whole,
contrasting it with that of Jeana with only 2 (CuP and 1A). However, the 2A vein could not be located in any of the
Oxycanus species described here, nor in O. australis or any other species examined, signifying that Oxycanus does
not differ from Jeana in terms of the number of hind wing anal veins. The two genera differ marginally in wing
venation in that the junction of R1 and R2 branch is nearer the termen in the forewing of Jeana. It should be noted,
however, that O. subvaria and O. ephemerous lack 2A in the fore wing whereas it is present in O. petalous, O. fla-
voplumosus and O. australis. The venation typical of Oxycanus is not unlike that of New Zealand genera Cladoxy-
canus, Dioxycanus, Dumbletonius, Heloxycanus and Wiseana in terms of position and vein number (Dugdale 1994).
The wing venation diagrams figured here (Fig. 10) are typical for Oxycanus, except the species O. poeticus Tindale,
O. petalous, O. subvaria, O. ephemerous and O. sphragidias, where CuP in the hindwing becomes membranous
medially. Though this would appear to be a derived feature within the Hepialidae, the feature is also present in Dum-
bletonius unimaculatus (Salmon), Dioxycanus oreas Hudson, and Wiseana cervinata (Walker) (Dugdale 1994). In
the fore wing of Oxycanus, CuP is slightly recurved, but it is also slightly recurved in Aoraia insularis Dugdale and
is greatly recurved in Cladoxycanus minos Hudson (Dugdale, 1994), however this feature needs further investiga-
tion within additional Oxycanus spp.
Although Australian Oxycanus is similar to the New Zealand oxycanines with respect to wing venation and an-
tennal structure, all the New Zealand species lack para-anal sclerites in the males (Dugdale 1994), a feature present
in many of the Australian and New Guinean Oxycanus. Para-anal sclerites were thought to distinguish the New Zea-
land oxycanines from the Australian genera (Dugdale 1994), though they were incorrectly identified by that author
as present in Elhamma Walker (and the synonym Zauxius Viette), which was later clarified by Simonsen (2015) who
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 371
showed that this structure is not present in Elhamma. Edwards & Green (2011) discussed that para-anal sclerites
were not present in either of the species they described. Variations in the morphology of para-anal sclerites are an
effective diagnostic feature for species identification in Oxycanus where they are present, as shown with the species
described here and as discussed in Edwards & Green, (2011). These structures are also present in the Asian genera
Hepialiscus Hampson, Parahepialiscus Viette, and Xhoaphryx Viette (Chu & Wang 1985; Grehan 2012; Mielke &
Grehan 2016) but not Napialus Chu & Wang (Li & Wie 2014). Xhoaphryx is very similar to the New Guinea Oxy-
canus (Paraoxycanus Viette, sensu Viette 1953) in wing shape and male genitalia characters according to Viette’s
diagram (Viette 1953: p. 33, fig. 1), particularly with respect to the short, deep-set para-anal sclerites– however
the authors are yet to examine this species directly. The absence of para-anal sclerites in several Oxycanus species
groups in Australia suggests the possibility that the genus is not monophyletic. Indeed, one species described here
(O. ephemerous, in the subvaria species group) lacks para-anal sclerites. It is of note that the study by Brown et
al. (1999) of the phylogeny of the New Zealand oxycanines based on mtDNA COI (a different gene fragment) and
COII sequence data tentatively determined that the Australian Jeana + Oxycanus (sensu stricto, O. australis) are a
sister group to the New Zealand taxa, while the Tasmanian O. sphragidias did not fall within the Australian & New
Zealand oxycanines. The results of Brown et al. may be valid but are problematic due to sample design not including
an outgroup outside of Hepialidae as identified by Grehan (2012). Likewise, the relationships revealed in our COI
phylogeny supports all our species delimitations, but also indicated some biogeographic sorting across Oxycanus
that will benefit from greater taxon sampling and inclusion of nuclear markers to further resolve these larger scale
phylogenetic relationships in Oxycanus and within the Hepialidae.
FIGURE 14. Inman Valley, Fleurieu Peninsula, South Australia. Highly degraded habitat of Oxycanus ephemerous sp. n.,
©Ethan P. Beaver.
Further research is required to investigate the monophyly of this large genus, and the potential conservation re-
quirements of some of the species that have not seen in recent years. Females for the species Oxycanus ephemerous
and O. flavoplumosus are yet to be collected, however it is possible that they can be obtained and described in the
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future. In addition, very little is known of the larval requirements of Oxycanus or Jeana, representing a large gap in
the knowledge of the larval biology of Australian Hepialidae. A taxonomic revision of the Australian members of
the genus Oxycanus and Jeana is planned by the authors which aims to investigate the evidence of paraphyly as well
as address several species synonyms and describe new species discovered during the finalisation of this paper.
FIGURE 15. Gleneagles Rest Area, Western Australia. Sclerophyll forest habitat of Oxycanus petalous sp. n., ©P. Kay.
Acknowledgements
We are thankful to many people for assistance and support while writing this paper. We would like to thank Ben A.
Parslow (SAM, Adelaide) for the SEM photography, Peter Hudson, Matthew Shaw, and the staff and volunteers of
SAM, Adelaide for access to equipment and specimens; ArtLab Australia for access to photographic equipment. In
addition, You Ning Su, Marianne Horak and Ted Edwards (ANIC, Canberra), Simon Grove, Cathy Byrne and Kir-
rily Moore (TMAG, Hobart) Susan Wright, Christine Lambkin and Karin Koch (QM, Brisbane), Derek Smith and
Russel Cox (AM, Sydney), David Lees (NHM, London) are thanked for loan or examination of specimens under
their care. Financial support was provided by an Australian Lepidoptera Research Endowment grant to MDM and
EPB, and a Field Naturalists Society of South Australia Lirabenda Grant to MDM. We would like to thank Sophie
Eden (Inman Valley, SA) for obtaining a specimen of O. ephemerous and making it available for description, as well
as Joseph Schubert (Museum Victoria, Melbourne), Kees Green (Ferntree Gully, VIC), Oliver Hill (Burnie, TAS),
Fabian Douglas (Rainbow, VIC), Paul Kay (Perth, WA), Don Gardner (Toowoomba, QLD), Nick Fischer (Tam-
borine Mountain, QLD), John Nielsen (Canberra, ACT), Paul Hutchinson (Perth, WA), Tony Hiller (Mt Glorious,
QLD), Peter Hendry (Sheldon, QLD), John Moss (Capalaba, QLD) and Cheryl Macaulay (Northcliffe, WA) for pro-
viding additional Oxycanus specimens during the course of this study. We are thankful to Axel Kallies (University
of Melbourne), Thomas Simonsen (NHMA, Denmark) Mackenzie Kwak (National University of Singapore), John
Grehan (McGuire Center, Florida, USA) and two anonymous reviewers for ongoing discourse and useful critique
NEW GHOST MOTHS OF THE GENUS OXYCANUS FROM AUSTRALIA Zootaxa 4732 (3) © 2020 Magnolia Press · 373
of the manuscript. EPB is thankful to Aden Beaver (Adelaide, SA) for assistance with editing and preparing the
figures, and Blake Gowing’s (Adelaide, SA) editing skills were invaluable during the finalisation of the manuscript.
Some specimens were collected during fieldwork conducted in WA under permit to MDM.
References
Australian Bureau of Meteorology (2019) Climate Data Online, Commonwealth of Australia Bureau of Meteorology, ABN
92637533532. Available from: http://www.bom.gov.au/climate/data/index.shtml (accessed 10 April 2019)
Brown, B., Emberson, R.M. & Paterson, A.M. (1999) Phylogeny of “Oxycanus” lineages of hepialid moths from New Zealand
inferred from sequence variation in the mtDNA COI and II gene regions. Molecular Phylogenetics and Evolution, 13 (3),
463–473.
https://doi.org/10.1006/mpev.1999.0662
Chu, H.F. & Wang, L.Y. (1985) “Insect-herb” versus hepialids with descriptions of new genera and new species of Chinese
Hepialidae. Sinozoologia, 3, 121–134.
Dorey, J.B., Schwarz, M.P. & Stevens, M.I. (2019) Review of the bee genus Homalictus Cockerell (Hymenoptera: Halictidae)
from Fiji with description of nine new species. Zootaxa, 4674 (1), 1–46.
https://doi.org/10.11646/zootaxa.4674.1.1
Dugdale, J.S. (1994) Hepialidae (Insecta: Lepidoptera). Fauna of New Zealand, Canterbury, New Zealand, 30, 1–164.
Edwards, E.D. & Green, K. (2011) Two new species of Oxycanus Walker (Lepidoptera: Hepialidae) from Kosciuszko National
Park, one with a sub-brachypterous female. Australian Journal of Entomology, 50, 78–85.
https://doi.org/10.1111/j.1440-6055.2010.00790.x
Felsenstein, J. (1985) Confidence limits on phylogenies: An approach using the bootstrap. Evolution, 39, 783–791.
https://doi.org/10.1111/j.1558-5646.1985.tb00420.x
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome
c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
Grehan, J.R. (2011) Notes on the biogeography of eastern Asian ghost moths (Lepidoptera:
Hepialidae). Bulletin of the Buffalo Society of Natural Sciences, 40, 59–74.
Grehan, J.R. (2012) Morphological evidence for phylogenetic relationships within the Hepialidae (Lepidoptera: Exoporia). Bul-
letin of the Buffalo Society of Natural Sciences, 42, 33–62.
Grund, R., Stolarski, A. & Stevens, M.I. (2019) Rediscovery of Synemon selene (Lepidoptera: Castniidae) in the mid-north of
South Australia. Transactions of the Royal Society of South Australia, 143 (1), 86–91.
https://doi.org/10.1080/03721426.2018.1532270
Hardy, R.J. (1974) The biology and pest status of Oxycanus fuscomaculatus Walker (Lepidoptera: Hepialidae) in Tasmania.
Journal of the Australian Entomological Society, 13, 317–328.
https://doi.org/10.1111/j.1440-6055.1974.tb02211.x
Hausmann, A., Haszprunar, G. & Hebert, P.D.N. (2011) DNA Barcoding the Geometrid Fauna of Bavaria (Lepidoptera): Suc-
cesses, Surprises, and Questions. PLoS ONE, 6 (2), e17134.
https://doi.org/10.1371/journal.pone.0017134
Kallies A., Marriott P. & Hewish M. (2015) Moths of Victoria. Vol. 36. Part 6. Ghost moths, Hepialidae and Allies. Entomologi-
cal Society of Victoria, Melbourne, 36 pp. + 1 CD.
Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across
computing platforms. Molecular Biology and Evolution, 35, 1547–1549.
https://doi.org/10.1093/molbev/msy096
Li, W. & Wei, H. (2014) Notes on the genus Napialus Chu & Wang (Lepidoptera: Hepialidae), with description of a new species
from China. Zootaxa, 3793 (3), 387–392.
https://doi.org/10.11646/zootaxa.3793.3.7
Mielke, C.G.C. & Grehan, J.R. (2016) A new species of Hepialiscus Hampson, [1893], Lepidoptera: Hepialidae) from Myan-
mar. The European Entomologist, 8 (3), 133–151.
Nei, M. & Kumar S. (2000) Molecular Evolution and Phylogenetics. Oxford University Press, New York, 348 pp.
Nielsen, E.S. (1996) Hepialidae s. lat. In: Nielsen, E.S., Edwards, E.D. & Rangsi, V. (Eds.), Checklist of the Lepidoptera of
Australia. Monographs on Australian Lepidoptera, CSIRO Publishing, Collingwood, Victoria, pp. 1–529.
Nielsen, E.S. & Kristensen, N.P. (1989) Primitive Ghost Moths, Morphology and taxonomy of the Australian genus Fraus
Walker (Lepidoptera: Hepialidae s. lat.). In: Monographs on Australian Lepidoptera. Vol. 1. CSIRO Publishing, East Mel-
bourne, Victoria, pp. 1–206.
https://doi.org/10.1071/9780643105096
Nielsen, E.S., Robinson, G.S. & Wagner, D.L. (2000) Ghost-moths of the world: a global inventory and bibliography of the
Exoporia (Mnesarchaeoidea and Hepialoidea) (Lepidoptera). Journal of Natural History, 34, 823–878.
https://doi.org/10.1080/002229300299282
Pfitzner, R. (1933) s.n. In: Seitz, A. (Ed.), Die Indo-Australischen Spinner und Schwärmer. Die Grossschmetterlinge der Erde,
BEAVER ET AL.
374 · Zootaxa 4732 (3) © 2020 Magnolia Press
10, pp. 909.
Posada, D. & Crandall, K.A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
https://doi.org/10.1093/bioinformatics/14.9.817
Simonsen, S.J. (2015) Elhamma Walker (Lepidoptera: Hepialidae) revisited: adult morphology, assessment of recently proposed
synonyms and descriptions of two species. Zootaxa, 3955 (3), 301–328.
https://doi.org/10.11646/zootaxa.3955.3.1
Simonsen, T.J. (2018) Splendid Ghost Moths and their Allies, A Revision of Australian Abantiades, Oncopera, Aenetus, Ar-
chaeoaenetus and Zelotypia (Hepialidae). In: Monographs on Australian Lepidoptera. Vol. 12. CSIRO Publishing, Mel-
bourne, pp. 1–300.
https://doi.org/10.1071/9781486307487
Simonsen, T.J., Moore, M.D., Dupont, S.T. & Stevens, M.I. (2019) Testing DNA barcodes against morphology for the ‘tripec-
tinate Abantiades’ (Lepidoptera: Hepialidae) reveals a complex relationship between COI sequence data and morphology.
Austral Entomology, 58 (4), 792–799. [https://onlinelibrary.wiley.com/doi/pdf/10.1111/aen.12395]
https://doi.org/10.1111/aen.12395
Swofford, D. (2002) Paup* version 4.0. Phylogenetic Analysis Using Parsimony (and Other Methods). Sinauer Associates,
Sunderland, MA. [program]
Tindale, N.B. (1935) Revision of the Australian ghost moths (Lepidoptera Homoneura, Family Hepialidae) Part III. Records of
the South Australian Museum, 5, 275–332.
Tindale, N.B. (1955) Revision of the ghost moths (Lepidoptera Homoneura, Family Hepialidae) Part VI. Records of the South
Australian Museum, 11, 307–344.
Tindale, N.B. (1964) Revision of the ghost moths (Lepidoptera Homoneura, Family Hepialidae) Part VIII. Records of the South
Australian Museum, 14, 663–668.
Viette, P.E.L. (1953) Contribution à l’etude des Hepialidae. Lambillionea, 53, 32–35.
Walker, F. (1856) Lepidoptera Heterocera. List of the Specimens of Lepidopterous Insects in the Collection of the British Mu-
seum, 7, 1508–1808.
