Four new species of Splendid Ghost Moths (Lepidoptera: Hepialidae: Aenetus) from Australia and Papua New Guinea

Abstract

Four new Aenetus Herrich-Schäffer species are described from northern Australasia; Aenetus simonseni sp. nov. from the top-end of the Northern Territory, Australia, A. maiasinus sp. nov. from the Kimberley region of Western Australia, A. trigonogrammus sp. nov. from south-eastern Queensland, Australia, and A. albadamanteum sp. nov. from eastern Papua New Guinea. Aenetus simonseni sp. nov. and A. maiasinus sp. nov. appear to belong to the tegulatus-group of species (sensu Grehan et al. 2018), A. trigonogrammus sp. nov. is part of the splendens-group of species (sensu Simonsen 2018), while A. albadamanteum sp. nov. shares morphological similarities with A. hampsoni (Joicey & Noakes, 1914), A. crameri Viette, 1956, and A. toxopeusi Viette, 1956, from New Guinea, and A. cohici Viette, 1961 from New Caledonia. The four new species are illustrated and compared with superficially similar species in morphology and, for two species, molecular (mtDNA COI gene) sequences.

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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by J. De Prins: 17 Jun. 2020; published: 8 Jul. 2020 449
Zootaxa 4809 (3):449–474
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4809.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:ZooBank:5E6909C3-5596-4CC2-AD10-B1B531FB5098
Four new species of Splendid Ghost Moths (Lepidoptera: Hepialidae: Aenetus)
from Australia and Papua New Guinea
ETHAN P. BEAVER1,2,*, MICHAEL D. MOORE1,3, JOHN R. GREHAN6,
ALEJANDRO VELASCO-CASTRILLÓN1,4 & MARK I. STEVENS1,5,7
1Biological and Earth Sciences, South Australian Museum, Adelaide, SA 5000, Australia
2
�
ethan.beaver@live.com.au; https://orcid.org/0000-0002-0613-7046
3
�
michael.moore@samuseum.sa.gov.au; https://orcid.org/0000-0002-8796-3330
4
�
a.velascocastrillon@gmail.com; https://orcid.org/0000-0002-3516-6655
5
�
mark.stevens@samuseum.sa.gov.au; https://orcid.org/0000-0003-1505-1639
6Research Associate. McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, 3215 Hull Rd, Gaines-
ville, FL 32611, USA.
�
calabar.john@gmail.com; https://orcid.org/0000-0002-3119-1140
7University of South Australia, Clinical and Health Sciences, SA 5000, Australia
*Corresponding author
Abstract
Four new Aenetus Herrich-Schäffer species are described from northern Australasia; Aenetus simonseni sp. nov. from
the top-end of the Northern Territory, Australia, A. maiasinus sp. nov. from the Kimberley region of Western Australia,
A. trigonogrammus sp. nov. from south-eastern Queensland, Australia, and A. albadamanteum sp. nov. from eastern
Papua New Guinea. Aenetus simonseni sp. nov. and A. maiasinus sp. nov. appear to belong to the tegulatus-group of
species (sensu Grehan et al. 2018), A. trigonogrammus sp. nov. is part of the splendens-group of species (sensu Simonsen
2018), while A. albadamanteum sp. nov. shares morphological similarities with A. hampsoni (Joicey & Noakes, 1914), A.
crameri Viette, 1956, and A. toxopeusi Viette, 1956, from New Guinea, and A. cohici Viette, 1961 from New Caledonia.
The four new species are illustrated and compared with superficially similar species in morphology and, for two species,
molecular (mtDNA COI gene) sequences.
Key words: Aenetus albadamanteum, Aenetus maiasinus, Aenetus simonseni, Aenetus trigonogrammus, Australasia,
biogeography, COI, DNA barcodes, taxonomy
Introduction
Aenetus Herrich-Schäffer, 1855 is a genus of medium to very large-sized moths found broadly across the higher
rainfall regions of Indonesia and Australasia, including Australia, with 22 species, New Guinea & the Moluccas,
with 10 species, and Sumatra, New Caledonia, and New Zealand, each with a single recorded species (Viette 1961;
Dugdale 1994; Simonsen 2018; Grehan et al. 2018; Beaver 2019a; Beaver 2019b). Among lepidopterists Aenetus
are well-known as the ‘Splendid Ghost Moths’ for their often brilliant green to pink colouration, and they are famous
for their secretive nature and propensity to fly primarily during rainfall and mist at night or dawn.
The larval biology has been studied extensively for the New Zealand Aenetus virescens (Doubleday, 1843)
(Grehan 1987; Dugdale 1994); the New Caledonian A. cohici Viette, 1961 (Boudinot 1991; Salesne 2010) and for
several eastern and southern Australian species such as A. eximia (Scott, 1869), A. lewinii (Walker, 1856), A. lig-
niveren (Lewin, 1805), A. tindalei Simonsen, 2018, A. scotti (Scott, 1869), A. blackburnii (Lower, 1892) (Beaver &
Grehan 2018), A. moorei Beaver, 2019 (Beaver 2019b) and A. djernaesae Simonsen, 2018 (Kay et al. 2020). Where
known, the early instar larvae feed within the leaf litter on dead wood and polypore fungi for several months as a
‘litter phase’ before entering a ‘transfer stage’ (Grehan 1987) whereupon they ascend the stems of shrubs or trees
within which they bore a tunnel that is concealed by a dense web of excavated sawdust and bark debris and feed
on callous tissue regrowth beneath the feeding web, near to the entrance of the tunnel (Grehan 1988; Beaver &

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450 · Zootaxa 4809 (3) © 2020 Magnolia Press
Grehan 2018). Larval development lasts for at least one year and several species are documented as having two or
more years of development (Grehan 1988; Beaver & Grehan 2018; Kay et al. 2020). Most Australian species occur
in vegetation growing in damp situations such as in close proximity to permanent or ephemeral creeks, swamps,
coastal habitats or areas where soil retains a seasonally very high level of moisture such as temperate woodlands,
wet forests or rainforest; in New Caledonia A. cohici is confined to rainforest environments in both upland and low-
land areas (Salesne 2010), while the New Zealand A. virescens is similarly a rainforest insect (Dugdale 1994). Little
is known of the habitat requirements of the Indonesian and Papua New Guinean species, however Roepke (1935)
lists mountain rainforest for the Moluccan Aenetus sibelae (Roepke, 1935).
The monophyly of Aenetus is supported morphologically; in the male genitalia the distal lobe of the pseudo-
tegumen is broad and ‘ear shaped’; curved valva with one or more hooks and externally, green to blue scales present
in either the male, female, or both (Simonsen 2018). The trulleum is membranous, unique among the West Pacific
higher Hepialidae. Recent taxonomic work on Aenetus, including a revision on the Australian members of the ge-
nus, has led to the discovery of several new species that are externally similar but show species level differences in
genitalia in both sexes and minor, sometimes inconsistent differences in wing morphology (Simonsen 2018, Grehan
et al. 2018, Beaver 2019b).
In this paper we describe three new species from Australia and one from Papua New Guinea. The new Austra-
lian species Aenetus simonseni sp. nov. was previously included under A. thermistis (Lower, 1894) which in turn
was only recently classified as being distinct from A. tegulatus (Pagenstecher, 1888) of Indonesia and New Guinea
(Simonsen 2018, Grehan et al. 2018) and is in the same species group as A. maiasinus sp. nov. from northern West-
ern Australia. The southern Queensland Aenetus trigonogrammus sp. nov. is recognized as being distinct from A.
lewinii. The New Guinea A. albadamanteum sp. nov. from eastern Papua New Guinea has a distinct wing pattern
from all other New Guinean Aenetus species. Based on similarities in adult wing morphology and genitalia it may
fall into a small species group that includes A. hampsoni (Joicey & Noakes, 1914), A. crameri Viette, 1956, and
A. toxopeusi Viette, 1956 of mainland New Guinea; and it may also have some affinity with A. cohici, a species
endemic to New Caledonia. We discuss the probable affinities of these species and aspects of their biogeographic
relationships.
Material and methods
The new species described herein are compared only with species that they either most closely resemble or with
which they are sympatric. Location data are provided as place names and with GPS coordinates where provided
verbatim on the specimen label. Specimens examined in this study are from:
Australian Museum, Sydney, Australia (AM), Research collection of A. Kallies, Melbourne, Australia (AKC),
Australian National Insect Collection, Canberra, Australia (ANIC), Bernice Pauahi Bishop Museum, Hawai’i, Ho-
nolulu, USA (BPBM), Canadian National Collection, Ottawa, Canada (CNC), Research collection of D. A. Lane,
Atherton, Australia (DALC), Research collection of E. P. Beaver, Adelaide, Australia (EPBC), Museum and Art
Gallery of the Northern Territory, Darwin, Australia (MAGNT), Museum Victoria, Melbourne, Australia (MV),
Queensland Museum, Brisbane, Australia (QM), South Australian Museum, Adelaide, Australia (SAMA), Univer-
sity of Queensland Insect Collection, Brisbane, Australia (UQIC).
Other abbreviations: NT—Northern Territory, Australia. NSW—New South Wales, Australia. PNG—Papua
New Guinea. QLD—Queensland, Australia. WA—Western Australia, Australia. VIC—Victoria, Australia.
Specimens examined for this study other than those mentioned within the systematic treatment include the fol-
lowing:
Aenetus thermistis (Lower, 1894)—in total 47 specimens, all Australia, Queensland. 1 ♀ holotype, ‘Mackay,
2511. Hepialus thermistis type, Low., I 14933, SAMA database no. 31-000223’. 1 ♂ A. cyanochlora (syn.) lecto-
type, ‘Mackay, 2512, cyanchlora type, Low., I 14933, SAMA database no. 31-000221’. 1 ♂ A. cyanochlora para-
lectotype, ‘Mackay, 2512, 12, Hepialus cyanochlora Lower, SAMA database no. 31-000222’ (all 3 in SAMA). 1
♀ Charagia walsinghami (syn) holotype, Mt Bartle Frere, K191017 (in AM). 6 ♀, 3 ♂ Kuranda, Qld, F.P. Dodd,
one male with dissection ID 31-016384-EPB (4 ♀, 2 ♂ in SAMA, 2 ♀, 1 ♂ in QM). 2 ♀, 1 ♂, Mackay, Qld, coll.
Lower, male with dissection ID 31-016384. 5 ♀, 4 ♂ Townsville, Qld, F.P Dodd, bred 10-19 Jan 1900 (3 ♀, 2 ♂ in
SAMA, 2 ♀, 2 ♂ in QM). 1 ♀, Atherton, Qld, January 1944. 1 ♀ Cooktown, Qld, dissection ID 31-016407-EPB.

FOUR NEW SPECIES OF AENETUS FROM AUSTRALIA Zootaxa 4809 (3) © 2020 Magnolia Press · 451
1 ♀ no data coll. Rowland Illidge. 1 ♂ Magnetic Island, Qld, Q.A.M. Lea. 1 ♂ Cairns, Qld. 1 ♂ Cairns district,
Qld (all in SAMA). 1 ♀ Shiptons Flat, via Helenvale, N Qld, 30 November 1980, 200m elev., G. Monteith & D.
Cook, emerged 30 December 1985, ex. Glochidion lobocarpum. 1 ♀, 1 ♂, Yeppoon, Qld, 13 November 1924, W.
B. Barnard. 2 ♂, Upper Hall Creek, 12 km WNW Carmila, Qld, 21°52’S x 147°18’E, 350m elev., 03-04 December
1996, G. Monteith, one specimen with dissection no. EPB-QM-002. 2 ♀, ‘Cape York’ Qld, 15 January 1928, W.
B. Barnard (specimens gutted) 1 ♂ same data, with dissection no. EPB-QM-003. 1 ♀ Gordon Ck, Claudie River
Dist, Cape York Pen., Qld, 12°42.78’S, 143°17.97’E, 02 Jan 1995, 50 m, G. and A. Daniels, rainforest MV lamp /
dissection no. EPB-QM-005 1 ♂ Thursday Island, Torres S, QLD, 03 Feb 1928, W.B. Barnard / dissection no. EPB-
QM-004 (all in QM). 1 ♂ West Claudie River, 7 km W road junction, Qld, 12°43’S 143°13’E, 29 Nov 1986, mv
lamp, G. Daniels, M.D. Schneider, dissection no. EPB-ANIC-023. 1 ♀ Yeppoon, QLD, 24 Jan 1993, J. Bugeja, dis-
section no. EPB-ANIC-024 (in ANIC). 1 ♂ 14 January 2019, Herberton Range, 10 km SW of Atherton, Qld, Aust.,
approx 1000 m elev., D.A. Lane, ex pupa from Casuarina torulosa. 2 ♀, 31 December 2015, Diwan, Qld, Aust.,
16°11’27”S 145°24’57”E, E. P & G. P. Beaver (all in EPBC). 6 ♂, 8 ♀, Townsville, Qld, F.P. Dodd. 2 ♀ Kuranda,
Qld, F.P. Dodd. 3 ♀, 4 ♂, Stony Creek, Qld. 1 ♀, Mission Beach, Qld, 09 Feb 2014, L.J. Cookson. 1 ♀, 1 ♂, Cape
York, Qld, W.B. Barnard (all in MV).
Aenetus spp. ‘near’ tegulatus (this is a complex of several species sensu Grehan et al. 2018)—in total 9 speci-
mens. Indonesia: 2 ♀ Little Kei Island, 13-15 Nov. 97, H. Kuhn, SAMA database no. 31-016400 and 31-016408,
with EPB dissection ID. 31-016400-EPB and 31-016408-EPB, 1 ♂ with same data except 19 Nov. 97, E.S. Nielsen
gen slide no. 2407, SAMA database no. 31-016390 (all 3 in SAMA). 2 ♂ and 1 ♀, same data except 28 Oct 97. 1
♀ Sorido, Biak Isl, East Indies, Apr 1945 (all in ANIC). Papua New Guinea: 1 ♀ Kiriwini, Trobriand is, April 1895
(ANIC). Australia, Queensland: 1 ♀ Gabba Island, Torres Strait, N. QLD, 11 Jan 1987, R. B. Lachlan, Dissection
no. EPB-ANIC-17 (in ANIC).
Aenetus splendens (Scott, 1864)—in total 25 ♂, all Australia. Queensland: 1 ♂ Kilarny, Qld, Feb 1940, E.J. Du-
migan. 6 ♂ Brisbane, Qld, one with dissection ID 31-016373-EPB (SAMA). New South Wales: 1 ♂ Bandon Grove,
NSW, M.J. Dowfing, 26 Feb 1962, bred. 1 ♂ Richmond River, NSW, Feb 1936, VJR. 1 ♂ ‘NSW’. 1 m # Newcastle,
NSW. 1 ♂ Tuncurry, NSW, 28 Feb 1919, J. Parkes, additional 4 ♂ with same data except for the dates 25 May 1926;
01 March 1912; 25 Feb 1926; 02 Feb 1920. 1 ♂ Northmead, 07 March 1987, M. Gregg. 1 ♂ Gosford NSW, 26 Feb-
ruary 1943. 6 ♂ no data. (all 17 in AM). 1 ♂ Tooloom Scrub, NSW, 09 Feb 1936, E.J. Dumigan (in SAMA).
Aenetus lewinii (Walker, 1856)—in total 16 ♂, all Australia. New South Wales: 1 ♂ Tuncurry, NSW, 24 Feb
1926, J. Parkes. Additional 2 ♂ with the same data except the dates 02 February 1925; 03 Mar 1926. 1 ♂ Manly,
NSW, 27 Feb 1917. Additional 2 ♂ with the same data except for the dates 29 April 1916; 12 Jan 1911. 1 ♂ no data
except ’03 Feb 1926’ (all 7 ♂ in AM). 1 ♂ Lake Ainsworth, NSW, 28°46’54”S, 153°35’16”E, E. P. Beaver, ex.
Alphitonia excelsa, coll. 01 Jan 2017, pupated 12 Sep 2017, eclosed 16 Oct 2017. 1 ♂ with same data except ‘ex.
Leptospermum sp., coll. 26 Sep 2019, pupated 08 Nov 2019, eclosed 07 Dec 2019’, dissection ID EPB-001 (2 ♂ in
EPBC). Queensland: 5 ♂ Brisbane, Qld, one with coll. Illidge. 1 ♂ Cairns, N. Qld, dissection ID 31-016131-EPB.
1 ♂ no data (all 7 ♂ in SAMA).
Aenetus ligniveren (Lewin, 1805)—in total 34 ♂, all Australia. Queensland: 2 ♂, Brisbane, QLD, R. Illidge,
one with dissection ID 31-016198-EPB. 1 ♂, no data. New South Wales: 1 ♂, Balmain, NSW, 27 Oct 1947. 1 ♂,
Noola via Rylestone, NSW, Jan 1965, N. Blunden. Victoria: 1 ♂ Gippsland, Vic, Jan 1902. 1 ♂ Kewell East, Vic, 15
Oct 1887, J. Hill. 1 ♂, Trafalgar, Vic, 09 Jan 1894. 1 ♂, Kelso, Vic. 1 ♂, Silverband Creek, Grampians, Vic, 25 Dec
1951 N.B. Tindale. 4 ♂, Melbourne, Vic. 2 ♂, Moe, Vic, C.G.L. Gooding, 07 Jan 1938; 11 additional ♂ with differ-
ent date data: 2 ♂ 03 Jan, 2 ♂ 04 Jan, 1 ♂ 08 Jan, 1 ♂ 09 Jan, 1 ♂ Oct 1938, 1 ♂ 28 Nov 1938, 2 ♂ 17 Dec 1921, 1
♂ 31 Dec 1921. 2 ♂ Narracan, Vic, 18 Nov 1902, 4 additional ♂ with different date data: 1 ♂ 17 Dec, 1 ♂ 13 Nov,
1 ♂ 19 Nov, 1 ♂ 24 Nov. South Australia: 1 ♂ 13 Oct 2018, Koorine, LSE of S. Aust, E. P. Beaver, dissection ID
EPB-011. (all 34 in SAMA).
A. bilineatus Beaver, 2019.—in total two ♂, both from ‘New Guinea’. 1 ♂, holotype, ‘New Guinea’ / E. S.
Nielsen gen. slide. 2408 (in SAMA). 1 ♂ ‘Central New Guinea’ in ANIC.
Morphological analysis. Dissections of genitalia were performed as described in Simonsen (2018), by re-
moving 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. Many specimens of two of the new
species described here had been subjected to the now outdated practice of ‘gutting and stuffing’ with cotton wool,
which made dissection and interpretation difficult. Terminology for genitalia and wing venation follows Dugdale

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(1994) and Simonsen (2018). Specimen data are presented with a ‘/’ denoting a second specimen label. Dissections
were imaged in an alcohol solution using a Leica imaging system comprised of a Leica binocular microscope, Leica
DFC 500 camera and the LAS Core software programme. Images of adult specimens were taken using a Canon
EOS camera, with a Canon EF 100 mm 1:2.8 Macro lens and Canon Speedlite Transmitter and Speelite 430 EX11
flashes. The images were then stacked using Zerene stacker software, and placed on 18% grey using Adobe Photo-
shop. Ventral images of adult specimens are provided only for the males of the ‘tegulatus’ group as diagnostic wing
pattern differences are not apparent on the ventral surface of the females of those or the other species. Due to some
shared features of genitalia and wing morphology, the species A. trigonogrammus sp. nov. is compared in diagnosis
with A. splendens, A. lewinii, and with A. ligniveren in Figs 30–33 as these are also the only members of the splen-
dens group to occur in southern Queensland.
Molecular analysis. DNA extraction and PCR amplification were 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 pro-
tocol. We targeted the commonly used mitochondrial (mtDNA) cytochrome c oxidase subunit I (COI) gene which
as successfully delineated between numerous insect species (e.g. Dorey et al. 2019; Grund et al. 2019; Beaver et
al. 2020). PCR products were amplified using the primers LCO1490 (forward) and HCO2198 (reverse) (Folmer et
al. 1994), see Beaver et al. (2020) for full protocols. The PCRs were sent to Macrogen Inc. (Korea) for purification
and sequencing.
All new COI sequence chromatograms (forward and reverse) were manually inspected to resolve ambiguous
base calls and checked for potential contamination using BlastN (NCBI). The resulting sequences were aligned
with other 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 alignment. Nucleotide sequence divergence was calculated using
uncorrected p-distances as implemented in MEGA X (Kumar et al. 2018).
Trees were generated using BEAUTi and BEAST 1.10.4 (Drummond et al. 2002). A separate model of evolu-
tion was applied across each codon partition (1st—GTR + Γ, 2nd & 3rd—GTR + I + Γ), as suggested by PartitionFind-
er v2.1.1 (Lanfear et al. 2016). We used an uncorrelated relaxed molecular clock to calculate trees with empirical
frequency-based priors starting from a random tree and Yule process tree prior and run for 50 million generations,
sampling every 1,000 trees. Convergence and stationarity of model parameters was assessed with Tracer v1.7.1
(Rambaut et al. 2018) with 10% of sampled trees discarded as burn-in, and the maximum credibility tree was gener-
ated using Tree Annotator v1.10.4.
Taxonomy
Family HEPIALIDAE Stephens
Genus Aenetus Herrich-Schäffer, 1855
Aenetus simonseni Beaver & Moore, sp. nov.
(Figs 1–2, 7, 9–11, 18, 21, 24–26, 59–61)
Type specimens: Holotype ♂, ANIC. Paratypes, 10 ♂, 8 ♀, SAMA, ANIC, MAGNT, DALC, UQIC, AKC.
Type locality: East Point, near Darwin, Northern Territory, Australia.
Etymology. This species is named in honour of Danish entomologist Dr Thomas J. Simonsen, in recognition of
his recent work on the Australian Hepialidae. A noun in the genitive case.
Type material. HOLOTYPE, (in ANIC) ♂ East Point, Northern Territory, Australia, 03 December 2018,
12°24’26.4”S 130°49’27.4”E, coll. N. Volpe / Spec. No. 19030 leg removed for tissue storage MD Moore / Dissec-
tion no. EPB-010 / ANIC 31-071235.
PARATYPES: 18 in total. 3 ♀, 1 ♂ (ANIC): 1 ♀, Rimbija Island, Wessel Islands, NT, 11.01S 136.45E, 17 Janu-
ary 1977, E. D. Edwards / Dissection no. EPB-ANIC-18 / ANIC 31-071236. 1 ♂ with the same data except the date
11 January 1977 / dissection no. EPB-ANIC-022 / ANIC 31-071237. 1 ♀ [very poor condition, extreme scale loss]
Black Point, Cobourg Peninsula, NT, 11.09S 132.09E, 30 Jan 1977 E.D. Edwards / Dissection ID. EBB-ANIC-5
/ ANIC 31-071238. 1 ♀ Howard Springs, NT, 10 Nov 1972, I.F.B Common / Genitalia slide H873 / ANIC 31-

FOUR NEW SPECIES OF AENETUS FROM AUSTRALIA Zootaxa 4809 (3) © 2020 Magnolia Press · 453
071239. 1 ♂ (SAMA): ♂ Groote Island , Northern Territory, November 1921, coll. N. B. Tindale / E. S. Nielsen gen
slide no. 2405 / SAMA Database No. 31-016391. 1 ♂ Holmes Jungle, 8 ml. NE of Darwin, NT, 130°56E, 12°24S, 6
December 1972, T. Weir, / 2152 (MAGNT). 1 ♂ 3 ♀ (UQIC): Darwin, NT, E.J. Dumigan, 03-07 February 1964. 2
♂, 1 ♀ (DALC): 1♂ East Point, Darwin, NT bred/larva, 25 November 2002, D. A. Lane, / LM-112 / Leg Taken MD
Moore spec. no. 19321. A further 1 ♂, same data except 29 December 2002. 1 ♀ Darwin, NT, 21 February 1964,
E.J. Dumigan. (MV); 1 ♂ Batchelor, NT, G.F. Hill / T-22494. 1 ♂ P. Darwin, NT, Oct 1908, F.P. Dodd / T-22493. 3
♂ (AKC); 1 ♂ Kakadu NP, Noorlangi Rock, NT, 12°51’37”S 132°49’07”E, larva: 30 March 2009, e.l. 1-15. Dec.
2009, leg. A. Kallies / 398 / MDM specimen number 20022. 1 ♂, same data except 399, MDM specimen number
20023. 1 ♀ same data except 400, MDM specimen number 20024.
Distribution. Aenetus simonseni sp. nov. is widely distributed over the coastal higher-rainfall areas of the Top-
End of the Northern Territory (Fig. 56).
Diagnosis. This is the only Aenetus species and the only member of the Hepialidae known from Top-End
NT. Males of A. simonseni sp. nov. can be distinguished from A. thermistis (Figs 3–4, 8, 12–14, 19, 22, 27–29)
of eastern Queensland by the blue or blue-white hindwings in contrast to broad swathes of pink along the costal,
tornal and inner margin of the hindwing (not so obvious in faded or old specimens). The forewings are generally
more aqua-green than A. thermistis but this is subject to variation. In the forewing, the anterior edge of the medial
line is whiter and posterior edge a darker more defined green. The valvae of the male genitalia are proportionately
larger and the apical, medial and basal hooks are more evenly spaced, with the apex flattened. A thickened portion
of the sacculus is nearly confluent with the basal hook, while in A. thermistis this is represented only as a small
knob. There are also differences in general proportions and shape of the vinculum, where the lateral margin is more
incurved and posterior projections shorter. The intermediate plate has 2-3 deep grooves on the dorsal edge while it
is smooth in A. thermistis. Pseudoteguminal twin processes are shorter, projected farther back on the basal rim. The
disto-posterior margin of the pseudotegumen is broader, and rounder. Sternite 8 in A. simonseni sp. nov. is almost
twice the size of that of A. thermistis, and with the sclerotised region more extensive, posterior projection shorter
and flanked by a small dentation before the postero-lateral corners, which are pointed in this species but rounded
in A. thermistis. Females are also externally similar to A. thermistis, however, A. simonseni sp. nov. has more nu-
merous brown scales on the legs, while the wings are generally ‘cleaner’ and the spots in the centre of the medial
area are smaller—however these are features subject to variation and the most reliable method is dissection of the
genitalia, where the antevaginal lamella differ significantly particularly in the medial area with three indentations
on the posterior margin, whereas there are only two in A. thermistis, which has the apex flattened. Lateral lobes are
smooth, while in A. thermistis they have two indentations. The bursa copulatrix is distinct, with the diverticulum
comparatively shorter than in A. thermistis, and the apex of the corpus bursae, though broad, is less round than in A.
thermistis. Aenetus simonseni sp. nov. can be distinguished from Aenetus nr. tegulatus as examined from Indonesia
and the Torres Strait, Qld (sensu Grehan et al. 2018; see methods) by female genitalia, where in A. nr. tegulatus
the medial area of the antevaginal lamella is broad and with two indentations, closer to that of A. thermistis, and
the bursa copulatrix is distinct, with the diverticulum narrower, pointed at the apex and closer to equal in length to
the corpus bursae, which is further rounded at the apex. Males of A. simonseni sp. nov. are somewhat similar to A.
eximia (Scott) but are distinguished by eighth sternite shape, which is smooth-sided and not anteriorly dentate, and
by general proportions and shape of the valva, in which A. eximia has the medial hook longer. The two species also
differ by the presence of brown scales on the legs and FW costa of A. simonseni sp. nov.. Aenetus sumatraensis
Grehan, Witt & Ignatev, 2018 and A. nr. tegulatus lack scales on the dorsal surface of the antenna, present in both
sexes of A. simonseni sp. nov. and A. thermistis. Differences between A. simonseni sp. nov. and A. maiasinus sp.
nov. are covered under that species.
Description. Male (Figs 1–2). Forewing length: 32–36 mm, hindwing 29–32 mm, expanse: 71–76 mm (n = 7).
Head: Antennae as long as head and slender, filiform, pale brown, covered with fine sensilla chaetica, short flattened
scales from pedicel to mid flagellum, scape rectangular, pedicel globular, 31 flagellomeres. Eyes prominent, larger
than head capsule, larger than in female, almost meeting dorsally. Scales on frons and vertex dense and dark green.
Labial palp three segmented, palpomeres rectangular, middle longest, apical shortest.
Thorax: Pro and mesothorax dorsally and ventrally covered in fine pale green to blueish-green scales and inter-
spersed with pink scales on ventral surface. Legs dark green with brown scales at proximal end of femur, tibia and
tarsi, epiphysis sub-triangular, less than ¼ length of tibia; hind leg with ochreous metatibial tuft of long androconial
scales. Arolium rounded.

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Wings: Forewing: broad and triangular. Costa slightly concave centrally and convex towards apex. Apex point-
ed, wing margin slightly concave toward apex, rounded with gently curved tornus. Wing venation classically hepi-
aline (Dumbleton 1966). Hindwing: broad, triangular with pointed apex, tornus rounded. Dorsal forewing ground
colour light green or aqua-green with an uneven transverse band along medial line from costa to tornus, between
inner margin at Rs2 and A1. Basal-side of line white, proximal dark green lowlights between veins Rs2 and CuA2.
Faint mottled patterns of alternating lighter and darker green across wing surface, extensive near outer wing margin.
Scales fading from light green to aqua-green towards tornus, light pink piliform scales around jugum. Costa light
green with five dark brown bars, final bar distal to medial line. Ventral surface ground color pink with pink-ochre-
ous piliform scales centrally and basally. Patch of green scales present at outer margin between Rs4 and M3. Entire
surface with lustrous iridescent shine not present on dorsal. Hindwing dorsal surface blueish-white to white, pale
blue in fresh specimens, costa and anterior margin marked a mottled green. Fringe dark green. Tornus mottled green;
inner margin with pink piliform scales anteriorly. Apex acutely pointed. Posterior discal cell narrowly triangular,
narrowing towards base, final discal cell rounded. Basal area covered with white or blue piliform scales. Ventral
surface with costa and medial area yellowish-green fading to blue towards tornus, tornus and inner margin pink and
clothed with pink piliform scales into basal area. Apex pink-tipped.
Abdomen: Long, narrow, basally covered in red-pink scales, sometimes white, green at apex. Sternite eight (Fig
21) rectangular, anterior corners rounded, posterior margin heavily sclerotized, with short central tooth, very short
and weakly sclerotised projections midway between posterior corners and central tooth.
Genitalia (Figs 9–11): Apodemal vinculum broad, wider than high in posteroventral view, lateral margin in-
curved sharply, with paired posterior projections sub-rounded and smooth. Saccus sub-triangular, broad, flattened
distally. Intermediate plate ovoid, with three deep grooves ventrally. Basal rim of pseudotegumen narrow, sub-tri-
angular with ‘m’ shaped lateral ridge, dorso- and distoposterior margins of pseudotegumen smooth, rounded and
lightly sclerotized. Twin processes greatly reduced, broad, triangular, pitched back towards intermediate plate. Ven-
tral pseudoteguminal arm present, narrow and pointed, becoming membranous. Valvae (Fig. 18) large, softly sclero-
tized, with three large spines, heavier sclerotisation on spines, valva narrower at proximal end, with sub-square
sacculus. Viewed posteroventrally, valvae curve away from pseudotegumen. Tip of valva ends as spine tapering to
sharp point, oriented vertically away from abdomen. In lateral view (Fig. 11) this spine is positioned at near right
angle, with tip straight. Largest spine from middle of valva, curved, tip pointing anteriorly, inner edge with small
bump. Third spine shortest, curved almost confluent with sacculus margin. Long, fine setae on tip of dorsal surface,
shorter setae present between spines and along inner lateral margin. Juxta broad, flattened and U-shaped. Trulleum
membranous.
Variation. Exact position and shape of medial line on forewing, extent of pink scales on anterior half of abdo-
men, extent of brown scales on forelegs as well as the width of the brown spots along costa is variable. General
ground colour ranges from grass green to greyish blue, with living specimens often appearing brighter than museum
specimens where fading may occur..
Female. (Fig. 7) Forewing length: 31–42 mm, hindwing: 28–33 mm, expanse: 72–84 mm (n = 7). Head: Anten-
nae slightly longer than head to mesothorax, slender, filiform, pale brown, dorsally scaled to middle, scape rectan-
gular, pedicel globular, 28 flagellomeres. Eyes prominent, of roughly same height as head capsule, wider set than
in male. Scales on frons and vertex dense and light green, obscuring base of antenna. Labial palps three segmented,
basal two palpomeres rectangular, apical rounded, middle longest, apical palpomere shortest.
Thorax: Pro- and mesothorax dorsally and ventrally covered in fine light green scales. Patch of dark grey to
brown scales at base of wing. Fore and mid legs covered with dense light green scales, grey to dark brown on tarsus
and anterior apex of tibia. Hind legs light brown, reduced. Foretibia with narrow, triangular epiphysis. Arolium
rounded.
Wings: Forewing: broad and triangular. Costa slightly concave centrally and convex towards apex. Wing ve-
nation classically hepialine. Hindwing: broad, subtriangular, tornus rounded. Dorsal forewing ground colour light
green with darker green mottled pattern consisting of darker green lines of irregular length and angle on all areas
of wing surface between Rs1 and 2A. Two to three cinnamon brown to dark grey spots in centre of wing along
medial line between Rs4 and M3, a smaller spot between CuA2 and 1A along the same line, and sometimes at end
of CuP distal to discal cell. Margins with small cinnamon or dark grey spots between Rs1 and CuA2, fringe dark
brown, light grey-brown at tornus, irregular cinnamon streak between CuA2 and jugum. Costa green with up to 10
cinnamon brown to dark grey spots from discal to medial area, fading to green at apex. Ventral surface cinnamon

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pink, costa as in dorsal, piliform scales basally, entire surface with lustrous iridescent shine. hindwing dorsal surface
uniformly cinnamon pink, unmarked. Ventral surface rust brown, piliform scales basally lustrous as in male.
Abdomen: Elongate, broad, segments covered in dense pinkish red scales basally, light green distally, with mod-
erately long piliform scale-tufts at apex.
Genitalia (Figs 24–26). Dorsal plates broad and high, lightly sclerotised, widest medially, distal ends straight,
setose proximally. Subanal plates short, narrow, horizontally oriented. Antevaginal lamella large, setose in two main
disjunct patches on posterior margin, three indentations on posterior margin, central indentation with longer setae.
Lateral lobes smooth. Ductus bursae elongate and narrow, with diverticulum long, ovoid, apex flattened, corpus
bursae wider distally, sub-ovoid.
Variation: Like all other species in the A. tegulatus group, A. simonseni females have two distinct colour forms,
one with the forewings, thorax, legs and abdomen apex grass green (Fig. 8), and the other colour form with these
areas a dark rusty-brown (Figs 59, 61). The green colour form is more common in collections, and is the basis for
the description.
Biology and phenology. Adult flight time is associated with the wet season, where adult emergence occurs
during or just after rainfall from the 25th of November to the 30th of January. The species has been taken at light in
monsoon rainforests particularly near swamps, creeks, in riparian or coastal situations (Fig. 64) and larvae have
been reared from the stems of Grewia breviflora Benth. (Tiliaceae) (David A. Lane pers. comm).
Remarks. This species is known only from the northern coastal regions and this may reflect a higher rainfall
requirement as suggested by Simonsen (2018: plate 48B). The species is generally widespread in the top-end of the
Northern Territory, where monsoon rainforest can be found in discrete patches within a vast area of 500,000 km2 that
are concentrated and most floristically diverse in the northern coastal regions (Russell-Smith and Lee 1992) where
rainfall is highest. The general dearth of specimens likely reflects an absence of targeted collection rather than a
true absence, as Australian members of this genus are usually uncommon at light and synchronise a short flight time
with specific weather events.
Though specimens have been photographed mating (Figs 59–61), courtship behaviour has not been observed.
The presence of meta-tibial androconial scales of the male, along with the distinctive saccharine scent discernible
from living or fresh male specimens suggests that pheromones play a part in the mate attraction or courtship of Ae-
netus species (Common 1990). Scales with lustrous iridescence are present on entire ventral surface of the forewing
and hindwing of A. thermistis males and females, but both wings only in the female of A. simonseni while these
scales are restricted to the forewing of the male. Comparatively large eyes may indicate a high visual acuity, this
coupled with the dimorphism of iridescent scales on both sexes but particularly so in the female could suggest that
a component of their courtship is visual as well as chemical. The Eurasian Hepialus humuli (Linnaeus) also have
iridescent scales on the wings, as well as hind tibial androconia in the males, and employ a visual courtship strategy
(Anderson et al. 1998). Further dedicated study into the behaviour of Aenetus in general is required to document
their courtship behaviour. Interspecific variation in eye size and degree of dimorphism, as well as variation in andro-
conia extent (with the latter present in both sexes of A. montanus Tindale, 1953 and A. ombraloma (Lower, 1902))
may suggest some level of interspecific variation in courtship behaviour.
Aenetus maiasinus Beaver & Moore, sp. nov.
(Figs 5–6, 15–17, 20, 23)
Type specimens: Holotype ♂, ANIC.
Type locality: Maia Cove, Kimberley Region, Western Australia, Australia.
Etymology. This species is named for the collection locality of the holotype, with the suffix –sinus a Latin noun
that translates to ‘bay’ or ‘cove’. A noun in the nominative singular case.
Type material. HOLOTYPE. (in ANIC): ♂. Maia Cove, Kimberley, WA, December 2017, J & A. Koeyers,
14°12’52.081”S, 126°7’6.431”E, ID no. KCS 18—2083 / dissection no. EPB-ANIC-021 / ANIC 31-071240.
Distribution. Known only from Maia Cove, northern Western Australia (Fig. 56).
Diagnosis. This is the only member of the Hepialidae presently recorded from the tropical north of Western
Australia. Aenetus maiasinus sp. nov. is a distinctive pale species which can be differentiated from A. thermistis
externally by the hindwing entirely white compared with broad swathes of pink and blue along the costal, tornal

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and inner margin of the hindwing in A. thermistis. The forewings are paler than either A. thermistis or A. simonseni,
and the medial line is straighter than in either species. Sternite eight of A. maiasinus sp. nov. has the anterior edge
truncate unlike either species, and has only a single central tooth on the posterior margin, unlike A. simonseni which
has three. Aenetus maiasinus sp. nov. further differs from either species in male genitalia; the apodemal vinculum
is proportionally broader with lateral margin slightly convex, unlike A. thermistis which is straight, or A. simonseni
which is slightly concave. The paired posterior projections are shorter than in A. thermistis, but more pronounced
than in A. simonseni. The saccus is significantly broader with the sulcus set closer to the margin of the apodemal vin-
culum than in either of A. thermistis or A. simonseni. The intermediate plate of A. maiasinus sp. nov. is rectangular,
similar to A. thermistis, however has a single lateral ridge that A. thermistis lacks. The most significant differences
are present in the valvae, where in A. maiasinus sp. nov. the apical spine is proportionally shorter than the middle
and basal spines, somewhat similar to the condition in A. thermistis but the basal spine nearest to sacculus is more
curved, and the sacculus is shorter.
Description. Male (Figs 5–6). Forewing length: 30 mm, hindwing 23 mm, expanse: 62 mm. Head: Antennae as
long as head and slender, filiform, pale brown, covered with fine sensilla chaetica, flagellum without scales; scape
globular, pedicel ovoid, 25–30 flagellomeres. Eyes prominent, larger than head capsule, almost meeting dorsally.
Scales on frons and vertex dense and olive green. Labial palpi three segmented, palpomeres rectangular, basal two
subequal, distal shortest, rectangular with apex blunt.
Thorax: Pro- and mesothorax dorsally and ventrally covered in fine olive green to cream scales and interspersed
with orange scales on ventral surface. Legs olive green with brown scales at proximal end of femur, tibia and tar-
sus; epiphysis triangular, apex rounded, less than ¼ length of femur; hind leg with ochreous metatibial tuft of long
androconial scales. Arolium u-shaped.
Wings: Forewing: broad and triangular. Costa slightly concave centrally and convex towards apex. Apex point-
ed, wing margin straight at apex, outer margin-tornus area rounded with gently curved tornus. Wing venation classi-
cally hepialine. Hindwing: broad, triangular with pointed apex, tornus rounded. Dorsal forewing ground colour light
cream with an even, slightly curved transverse band along medial line from costa to CuA2. Basal side of line paler
cream, fading into ground colour, proximal olive green lowlights between CuA2 and Rs3. Faint mottled patterns of
alternating lighter and darker cream colours across wing surface, extensive near outer wing margin. Pale orange and
cream piliform scales around jugum. Costa light olive green with five dark brown bars, final bar distal to medial
line. Costal-discal area light olive green. Ventral surface overall mottled cream with pale orange piliform scales me-
dially and basally. Patch of green scales present at margin between costa and tornus. Areas with pale orange scales
are lustrous iridescent. Hindwing dorsal surface white, costa, tornus and anterior margin cream. Inner margin with
orange piliform scales anteriorly. Posterior discal cell narrowly triangular, narrowing towards base, final discal cell
narrower, triangular, basally covered with white piliform scales. Ventral with entirely yellowish-green, with whit-
ish-blue scales in discal area.
Abdomen: Long, narrow, basally covered in white scales. Sternite eight (Fig 23) roughly sub-ovoid with anterior
edge truncate, posterior edge unsclerotized, with short central tooth and pronounced lateral corners.
Genitalia (Figs 15–17): Apodemal vinculum broad, slightly wider than long in posteroventral view, lateral mar-
gin slightly convex, with paired posterior projections subtle and sub-rounded. Saccus sub-triangular, broad, flattened
distally. Intermediate plate sub-rectangular, single lateral ridge. Basal rim of pseudotegumen narrow, sub-triangular,
dorso- and distoposterior margins of pseudotegumen smooth, sub-rounded and lightly sclerotized. Twin processes
reduced, blunt, pitched back towards basal rim. Ventral pseudoteguminal arm present, narrow and pointed, becom-
ing membranous. Valva (Fig. 20) lightly sclerotised, with three heavily sclerotised large spines, valva narrower at
proximal end, with subglobular sacculus. Viewed posteroventrally, valvae curve away from pseudotegumen. Tip of
valva ends as apical spine tapering to sharp point, pointing ventrally in habitus away from abdomen. When viewed
laterally (Fig. 17) this spine is positioned at near right angle, with tip straight. Apical spine smallest; mid spine,
curved, tip pointing anteriorly, inner edge with small indentation; basal spine nearest to sacculus shorter than middle
spine, hooked, oriented well away from sacculus. Long, fine setae present on tip of dorsal surface, shorter setae pres-
ent between spines and along inner lateral margin. Juxta broad, flattened and U-shaped. Truellum membranous.
Female. Unknown.
Biology and phenology. Larval biology unknown. The specimen was collected in December, indicating that the
species likely has a northern wet season flight period not unlike other members of the ‘tegulatus’ complex.

FOUR NEW SPECIES OF AENETUS FROM AUSTRALIA Zootaxa 4809 (3) © 2020 Magnolia Press · 457
FIGURE 1–6. Adult male Aenetus spp: 1–2, A. simonseni sp. nov., holotype, 1, dorsal; 2, ventral; 3–4, A. thermistis, Atherton,
QLD, 3, dorsal; 4, ventral; 5–6, A. maiasinus sp. nov., holotype, 5, dorsal, 6, ventral. Scale bars: 10 mm.
FIGURE 7–8. Adult female Aenetus spp. (all dorsal): 7, Aenetus simonseni sp. nov. paratype; 8, A. thermistis, Kuranda, QLD.
Scale bars: 10 mm.

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FIGURE 9–23. Male Aenetus genitalia; ventral, dorsal, lateral; 9–11, A. simonseni sp. nov. holotype, dissection ID EPB-010;
12–14, A. thermistis dissection ID 31-016384-EPB; 15–17, A. maiasinus sp. nov. holotype, dissection ID EPB-ANIC-021; 18–
20, valvae, 18, A. simonseni sp. nov. holotype; 19, A. thermistis, 20, A. maiasinus sp. nov. holotype; 21–23, sternite eight; 21,
A. simonseni sp. nov. holotype; 22, A. thermistis; 23, A. maiasinus sp. nov. holotype; scale bars: 9, 12, 15, 21–23: 1 mm; 18–20:
500 μm. Abbreviations: brp—basal rim of pseudotegumen. ip—intermediate plate. jx—juxta. pt—pseudotegumen. sc—saccus.
scu—sacculus. tp—twin processes of pseudotegumen. tr—trulleum. vi—vinculum. vl—valva.

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FIGURE 24–29. Female genitalia of Aenetus spp, 24–26, A. simonseni sp. nov. paratypes; 24–25, dissection ID EPB-ANIC-
05; 26, dissection ID EPB-ANIC-18; 27–29, A. thermistis dissection ID 31-016407-EPB. Scale bars: 1 mm. Abbreviations:
cb—corpus bursae. db—ductus bursa. div—diverticulum. dp—dorsal plate. lal—lateral area of antevaginal lamella. mal—me-
dial area of antevaginal lamella. sub—subanal plate.
FIGURE 30–33. Male ‘splendens group’ Aenetus spp. that occur in southern Queensland (all dorsal); 30, A. trigonogrammus
sp. nov., holotype; 31, A. lewinii, Brisbane, QLD; 32, A. splendens, Brisbane, QLD; 33, A. ligniveren, Koorine, SA. Scale bar:
10 mm.
Remarks. The dissection is affected by mould which interfered with visibility of some features and imaging
but clear diagnostic differences are nevertheless observable, particularly in the valva and apodemal vinculum. The
species may be found to be more widespread in the high rainfall area of the Kimberley region, as the presumed habi-

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tat of monsoon rainforest totals up to 7000 ha in scattered, isolated patches across 170,000 square km (Kenneally
2018). Lepidoptera are rarely collected from this region due to its general inaccessibility, which likely explains the
absence of available material in major institutions. The single specimen was collected on a boat anchored at the cove
(J. Koeyers pers. comm).
Aenetus trigonogrammus Beaver & Moore, sp. nov.
(Figs 30, 34–36, 40, 42, 62)
Type specimens: Holotype ♂, ANIC. Paratypes: 1 ♂ UQIC.
Type locality: Fig Tree Creek, 27.5 km SW of Gin Gin, Queensland, Australia.
Etymology. The name trigonogrammus is derived from Greek trigono (triangle) and grammi (line) in reference
to the triangular white marking on the male forewing, where the white medial line meets the dorsum. A noun in the
nominative singular case.
Type material. HOLOTYPE. (in ANIC): ♂. Fig Tree Creek (Perry River), 27.5 km SW of Gin Gin, Qld,
25°13’14”S, 151°51’01”E, 27 October 2011, P.J. and Q.A. Hendry / ANIC 31-071241.
PARATYPES. 1 ♂ (UQIC). Toowoomba, Queensland, Australia, E.J. Dumigan, 25 January 1960 / Dissected by
E.P. Beaver, dissection no. UQIC-EPB-01’ (in UQIC).
Distribution. South-eastern Queensland (Fig. 57), where the type specimen was collected 27.4 km SW of Gin
Gin. and a further specimen from Toowoomba.
Diagnosis. Along with the similar ‘splendens’ group species found in Queensland—A. lewinii (Figs 31, 63), A.
ligniveren (Fig. 33), and A. splendens (Fig. 32)—the male of A. trigonogrammus sp. nov. is distinguished from of
the other Queensland Aenetus species such as A. edwardsi Simonsen, A. eximia, A. mirabilis Rothschild, A. scotti,
A. ramsayi (Scott), and A. thermistis by the presence of a narrow white submarginal band from costa to tornus and
baso-tornal patches of white on the forewing. The male of A. trigonogrammus is distinguished externally from the
other ‘splendens’ species by the white triangular patch along the dorsum at the posterior terminus of the submar-
ginal line, and by having the basal triangular marking entirely white. The genitalia are superficially similar only
to A. lewinii (Figs 37–39, 41, 43), but may be distinguished particularly by the pseudotegumen shape, which in A.
trigonogrammus sp. nov. is large and rounded, whereas in A. lewinii it is smaller relative to the basal rim, and sub-
triangular. The valvae are similar to that of A. lewinii, however, they are less strongly sclerotised, and broader cen-
trally, with the area distal to the sacculus narrower. The eighth sternite has sharper anterio-lateral corners while the
apodemal vinculum is broader and with paired anterior projections however the anterior end in A. lewinii is variable
with respect to the presence or absence of paired anterior projections in some specimens. The intermediate plate is
not fused with the basal rim of the pseudotegumen, whereas in A. lewinii it is.
Description. Male (Fig. 30) Forewing length: 24–27 mm, hindwing 18–21 mm, expanse: 50–55 mm. Head:
Antennae longer than head and slender, filiform, densely ciliated, pale brown, 33 flagellomeres, scape cuboid,
pedicel flattened ovoid. Eyes prominent, same height as head capsule, almost meeting dorsally. Scales on frons and
vertex dense and white. Labial palpi three segmented, basal two palpomeres rectangular, apical elliptical, middle
longest, apical palpomere shortest.
Thorax: Pro- and mesothorax dorsally and ventrally covered in white scales and interspersed with olive green
and scales on ventral surface; dorsal prothorax white, base of wing green, posterior end white in lateral view. Legs
white and olive green, with long white piliform scales proximally from middle of tibia and tarsus; dark brown longi-
tudinal line along length of mesotibia; epiphysis short, broad, apex pointed; hind leg white with ochreous metatibial
tuft of androconial scales. Arolium elliptical.
Wings: Forewing broad and triangular. Costa slightly concave centrally and convex towards apex. Apex point-
ed. Wing venation hepialine. Hindwing broad, subtriangular, tornus rounded. Forewing dorsal ground colour olive
green, darker green lines in discal area; an uneven muted white band along medial line from costa to tornus, where
it widens to a white triangle; a second triangle on anal margin in discal area, characteristic of the splendens group of
species, filled white. A short, separate white basal line is present from dorsum near jugum, extending to CuA2 in the
basal area, fusing with discal triangle at CuA2. Costa white, becoming light green at apex. Tornus distal to medial
line suffused with whitish-grey scales between M3 and just beyond CuA2. Ventral hindwing dorsal surface white,
margin at apex light green, margin elsewhere faint bluish green; basal area covered with white piliform scales. Ven-
tral surface as above except basal area bluish-green.

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Abdomen: long, narrow, white piliform scales dorsally and ventrally. Ventral apex with aqua blue scales. White
forked tuft of piliform scales at apex. Sternite eight (Fig. 42) sub-square except strongly concave posterior margin.
Genitalia (Figs 34–36). Apodemal vinculum broad and short, with paired anterior projections rounded. Saccus
uniformly sub-triangular, broad. Intermediate plate laterally compressed, free, with anterior indentation near saccu-
lus. Basal rim of pseudotegumen broad, high in lateral view, disto- and ventroposterior margins of pseudotegumen
broad, rounded, lightly serrate. Ventral pseudoteguminal arm sclerotised. Valva (Fig 40) large, broad at proximal
end, with long, blunt sacculus. Viewed posteroventrally, valvae curve laterally away from pseudotegumen. Apex
of valva rounded, sub-apical area tapering to a sharp point, pointing vertically away from abdomen. When viewed
laterally the apical spine is positioned almost horizontally with tip curving slightly ventrally. A large curving hook
from distal sacculus points vertically down and is more curved than the sub-apical spine. Fine setae sparsely present
between basal hook and apex, with denser patch of hairs between sacculus and hook. Juxta flattened and U-shaped.
Truellum membranous.
Variation. Variable number of darker green lines within the light green baso-discal area; extent of white suffu-
sion beyond medial line. The paratype is smaller than the holotype, with forewing length 24 mm.
Female. Unknown.
Biology and phenology. Larval biology unknown. The holotype (Figs 30, 62) was collected at light during
rainfall (P. Hendry pers. comm.). The site is mostly subtropical Eucalyptus woodland with mixed understorey and
dry vine thicket at the edge of a seasonal creek (Fig 65). Aenetus species, particularly those in the splendens group
of species, are often associated with host plants growing in close proximity to creek-lines, in riparian vegetation
(Beaver 2019b). The two known specimens are from late October to late January.
Remarks. The authors (EPB & MDM) visited southern Queensland in late 2019 but failed to locate further
specimens of this species, which is expected to be highly localised. Only two specimens are known from what is a
heavily cleared and populous area of Australia. Southern Queensland has a significantly high level of Hepialidae
diversity for Australia (Simonsen 2018),—generally within rainforests or wet forest environments which makes the
discovery of this new species unexpected. Aenetus trigonogrammus sp. nov. is allopatric to all other splendens-
group species at the northern site near Gin Gin, however further south at Toowoomba it would appear to overlap in
the general vicinity with A. splendens, A. lewinii and A. ligniveren (Simonsen 2018). The first author has taken each
species in a distinct habitat, and rarely together, suggesting that their range at the local-scale may be closely tied to
habitat preference: A. splendens in subtropical rainforest and rainforest edge, A. lewinii in coastal wallum swamp
and open Melaleuca-Banksia woodland with a diverse heath layer (Beaver & Grehan 2018), and A. ligniveren in
temperate woodland or Eucalyptus wet forest (Beaver 2019b). The main exception being that A. splendens will oc-
casionally occur in temperate woodland very near to A. ligniveren if rainforest elements are present in the general
vicinity. Specimens of those three species were recorded from the Burdekin River, north Queensland by Simonsen
(2018) who remarked that the locality data for these specimens was likely incorrect. Although this may be the case
for A. ligniveren and A. splendens, a further male specimen of A. lewinii is in the SAMA collection and labelled from
Cairns, far north Queensland. The species is also present in central Queensland at the Blackdown Tableland (Simon-
sen 2018) and so may be found to be further sympatric with A. trigonogrammus sp. nov. in southern Queensland.
Aenetus albadamanteum Beaver & Grehan, sp. nov.
(Figs 44–45, 46–49, 51–54)
Type specimens: Holotype ♂, ANIC. Paratypes: 1 ♀ ANIC, 1 ♀ BPBM, 1 ♂ CNC.
Type locality: Mount Kaindi, Morobe Province, Papua New Guinea.
Etymology. The name albadamanteum is from the Latin alba, ‘white’, and adamantem ‘diamond’, referring to
the white, diamond shaped markings present along the medial and sub-marginal lines of the forewing, particularly
in the female. A noun in the nominative singular case.
Type material. HOLOTYPE (in ANIC) ♂. [Papua] New Guinea: NE: [Morobe Province] Mount Kaindi, 2350
m, February 1970, J.H. Sedlacek, Rainforest / Dissection no: EPB-ANIC-019 / ANIC 31-071242.
PARATYPES. 1♀ (in ANIC) Same data as holotype except date as 01 January 1979 / Dissection no: EPB-
ANIC-020 / ANIC 31-071243. 1 ♀ and 1 ♂ in BPBM: ♀ Mt. Kaindi, NG, 2350 m, 23.III.1966 / Light trap, J.L.

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Gressitt / Dissection no: JRG 300. 1 ♂ PNG, Bulldog Road, 7 km s Edie Creek, 2400m, 13-15-xii 1977, UV lt, P.
Herbert & R. Harmsen / Dissection no: JRG208.
Distribution. Known only from Mount Kaindi, Morobe Province, Papua New Guinea (Fig. 58).
FIGURE 34–43. Male Aenetus spp. genitalia, ventral, dorsal, lateral. 34–36, A. trigonogrammus sp. nov. paratype, dissection
ID UQIC-EPB-01; 37–39, A. lewinii, Lennox Head, NSW, dissection ID EPB-001; 40–41, valvae; 40, A. trigonogrammus sp.
nov. paratype; 41, A. lewinii; 42–43, sternite eight; 42, A. trigonogrammus sp. nov. paratype, 43, A. lewinii. Scale bars: 34, 37:
1 mm; 40–43: 500 μm.
Diagnosis. This species is not easily mistaken for any other species that occur in Papua New Guinea. It is super-
ficially similar to A. crameri, A. hampsoni and A. toxopeusi, but without the prominent ‘X’ shaped markings present
between most veins on the forewing of those three species. Furthermore, A. albadamanteum sp. nov. is the only one
of the four species to have two rows of black spots along the medial and submarginal lines in the male. A female of

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A. hampsoni figured in Joicey & Noakes (1914) differs from A. albadamanteum sp. nov. by the presence of numer-
ous bright white ‘X’ shaped markings present in a medial and submarginal line as well as along the wing margin and
basal area. Although females of A. crameri and A. toxopeusi are currently unknown, Viette (1956) figured the male
genitalia of both species and these differ from that of A. albadamanteum sp. nov., particularly in the valvae where
the apex is bulbous, and by the anterior end of the apodemal vinculum, which has only short, close-set anterior
projections, which are much larger and wider-set in A. crameri and absent in A. toxopeusi. Aenetus bilineatus is the
only other New Guinean species which has a well-defined medial and sub-marginal band in the forewing, however
in A. albadamanteum sp. nov. the bands are comprised of a series of disconnected spots, whereas in the former spe-
cies the spots are merged as a continuous line, and the hindwing is very broad and more rounded compared with
the narrow hindwing of A. albadamanteum sp. nov.. Aenetus bilineatus is also smaller, with a male forewing length
of 36 mm. Aenetus albadamanteum sp. nov. is similar to the New Caledonian A. cohici. Grehan (1983) described
and figured the male genitalia of A. cohici (Fig. 50), a species that differs significantly from A. albadamanteum sp.
nov. in the shape of the valvae, which lack the bulbous apex, and the apical teeth proportionally longer. Male wing
pattern in A. cohici is highly variable (Salesne 2010) but includes variations where males have a medial and sub-
marginal line of disconnected spots. Unlike A. albadamanteum sp. nov., which has dark brown spots, the spots of A.
cohici are primarily white with a narrow edge of brown on the outer edge. The new species lacks the knob-shaped
metatibial scent gland found in A. cohici (Grehan 1983, Fig. 1)
Description. Male (Fig. 44). Forewing: 46 mm. Hindwing: 40 mm. Expanse: 83 mm. Head: antennae longer
than head and fine, filiform, pale brown, at least 23 flagellomeres, scape cuboid, pedicel ovoid. Eyes prominent,
wide-set, slightly higher than head capsule. Scales on frons and vertex dense and light green. Labial palps with three
palpomeres, basal two rectangular, third rounded; basal palpomere longest, distal-most shortest.
Thorax: Pro and mesothorax dorsally and ventrally covered in light green scales and interspersed with orange
and yellow scales on ventral surface. Legs uniformly green and light brown, foreleg missing, hind tibia thickened,
with large ochreous metatibial tuft of very long androconial scales characteristic of Aenetus. Arolium u-shaped.
Wings: Forewing broad and triangular. Costa centrally straight and convex towards apex. Apex pointed, wing
venation classically hepialine. Hindwing of similar width to forewing, sub-triangular with sharp acute angle at apex.
Dorsal forewing ground colour light green with alternating dark green lines between all wing veins, and two trans-
verse rows of dark grey spots, one spot between each wing vein; a series of nine semi-circular spots along medial
line between inner margin at 1A and costa, spots between Rs3 and CuA2 with silver centre, and submarginal series
of diamond-shaped dark grey spots between Rs2 and CuA2; Rs3 to M2 with silver centre. Six to seven light brown
bars separated by green along costa with darkest bar medially. Wing margin with three brown spots between each
wing vein, between Rs1 and CuA1, largest spot centrally. Ventral surface plain, spots visible faintly through wing,
faint yellow scales centrally and basally. Hindwing dorsal surface bright white, tornus and inner margin pale green-
ish-cream at margin, fading into white ground area. Sc and Rs1 straight. Apex acutely pointed. Basal area covered
with white piliform scales.
Abdomen: elongate, narrow, scales white. Sternite eight (Fig. 54) subsquare, narrowing slightly posteriorly,
anterior margin tapering to shallow point, posterior margin indented slightly each side of a shallow medial convex
protrusion.
Genitalia (Figs 46–48): Apodemal vinculum broad and short, with paired anterior projections rounded, close-
set. Saccus triangular, broad. Intermediate plate laterally compressed, sub-trapezoidal, free. Basal rim of pseudo-
tegumen broad, high in lateral view, disto- and ventroposterior margins of pseudotegumen broad, sub-triangular.
Ventral pseudoteguminal arm membranous. Valva (Fig. 49) short, broad at proximal end, with short, square saccu-
lus. Viewed posteroventrally, valvae curve laterally away from pseudotegumen. Apex of valva bulbous, with very
short apical spine. When viewed laterally in situ this apical spine is positioned almost horizontally. A large curving
hook from middle of valva points vertically ventrally. A second hook positioned just above the sacculus, less curved
and longer than middle hook. Fine setae present ventrally between sacculus and valva apex. Juxta flattened and U-
shaped. Truellum membranous.
Variation. Paratype male has the diamond-shaped grey spots of the forewing with a more prominent white
centre. Width of valval hooks and juxta variable, level of sclerotisation of pseudotegumen may differ between
specimens.
Female (Fig. 45). Forewing: 86 mm. Hindwing: 57 mm. Expanse: 157 mm. Head: antennae longer than head
and fine, filiform, pale brown, 34 flagellomeres, scape rectangular, pedicel ovoid. Eyes prominent, widely spaced,

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slightly higher than head capsule. Scales on frons and vertex dense and light green. Palps as for male, basal two
partially fused.
Thorax: Pro- and mesothorax dorsally and ventrally covered in light green scales interspersed with orange and
yellow scales on ventral surface. Legs uniformly green and light brown with epiphysis digitiform; arolium as for
male.
FIGURE 44–45. Aenetus albadamanteum sp. nov. (all dorsal); 44, holotype male; 45, paratype female. Scale bar: 10 mm.
FIGURE 46–54. Male and female Aenetus spp. genitalia and sternite eight. 46–48, A. albadamanteum sp. nov. holotype, dissec-
tion ID EPB-ANIC-019, male ventral, dorsal, lateral; 49, A. albadamanteum sp. nov. holotype, valva; 50, male A. cohici, New
Caledonia, ventral (image J.R. Grehan); 51–53, A. albadamanteum sp. nov. paratypes; 51, bursa copulatrix JRG208; 52–53,
dissection ID EPB-ANIC-020; 54, male A. albadamanteum sp. nov. holotype, sternite eight. Scale bars: 46, 54: 1 mm, 49: 500
μm, 52-53: 2 mm.
Wings: Similar to male, hindwing proportionally shorter, forewing broad and triangular. Costa straight centrally
and convex towards pointed apex. Wing venation as for male except 2A present in hindwing. Hindwing narrower,
sub-triangular with sharp angle at apex and tornus. Dorsal forewing ground colour as for male, spots larger and all
with white centre, all diamond-shaped except for semi-circular spots on medial line between Rs4 and M2. Two ir-
regularly shaped grey spots with white centre in discal cell nearest to costa; central and posterior discal cells each

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with a single irregularly shaped white-centred spot; a further single white-centred spot at distal end of CuP, faint
grey spot at middle of CuP. At least three light brown bars separated by green along costa, with darkest bar medial
(costa worn of scales). Outer margin with single brown spot between each wing vein from Rs1 to CuA1. Ventral
surface plain, spots visible faintly through wing, faint yellow scales centrally and basally. Hindwing dorsal surface
pale greenish-yellow, faint pink basally. Green scales irregularly scattered in sub-marginal area to margin. Apex
acutely pointed. Faint pink-cream piliform scales basally.
FIGURE 55. Maximum credibility tree from Bayesian phylogenetic analysis of the mitochondrial DNA COI gene from 7
outgroup taxa and 32 Aenetus sequences from 17 species, including two new Aenetus species. Aenetus thermistis is labelled as
two clades (see Discussion). Posterior probabilities greater than 0.65 are shown at each node, and a * indicates where we have
corrected a GenBank id (see Table S1).

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FIGURE 56–57. Distribution records of Aenetus spp.; 56, the tegulatus complex in Australia, Papua New Guinea, Timor-Leste
and south-eastern Indonesia, with A. simonseni sp. nov. (pink circle), A. maiasinus sp. nov. (white crossed circle), A. thermistis
(black circle), A. tegulatus from Ambon Island (dark grey crossed circle) and at least one undetermined A. tegulatus complex
species (light grey circle) sensu Grehan et al. 2018, modified from Grehan et al. 2018; 57, map of the Australian state of
Queensland with recorded localities of A. trigonogrammus sp. nov..
FIGURE 58. Distribution records of the Aenetus hampsoni species group in New Guinea; pink circle: Aenetus albadamanteum
sp. nov.; dark grey circle: A. crameri; light grey circle: A. toxopeusi; black circle: A. hampsoni.
Abdomen: elongate, narrow, scales green, pinkish basally.
Genitalia (Figs 51–53): Dorsal plates wide, lightly sclerotised, setose, posterior-medial ends narrow, fused
across the median. Subanal plates narrow, dorsal edge with irregular indentations. Antevaginal lamella dorso-ven-
trally high, setose, well sclerotized, particularly along dorsal edge. Dorsal edge medially truncate, lateral lobes
dentate along dorsal margin. No clear distinction between medial and lateral areas. Ductus bursae long and narrow,
diverticulum short, about 1/3 length of corpus bursae, cylindrical, apically tapered to round end, connected to a

FOUR NEW SPECIES OF AENETUS FROM AUSTRALIA Zootaxa 4809 (3) © 2020 Magnolia Press · 467
broadened section of corpus bursae. Corpus bursae elongate, broad, angled slightly before becoming distally wider
and sub-ovoid.
Variation. No significant variation observed.
Biology and phenology. Larval biology unknown. The holotype and two paratype specimens were taken near
the summit of Mt Kaindi (alt. 2362 m), at the recorded elevation of 2350 m Mt Kaindi is predominately lower
montane rainforest dominated by Nothofagus pullei Steenis (Nothofagaceae), with other tree species such as Ac-
ronychia J.R.Forst. & G.Forst. (Rutaceae), Elaeocarpus polydactylus Schltr, Sericolea micans Schltr. (Elaeocar-
paceae), Homalanthus nervosus J.J.Sm. (Euphorbiaceae), and Quintinia macrophylla Hatschb. (Paracryphiaceae).
In the 1990’s this vegetation was reportedly in early-mid succession, having been disturbed by mining and related
activities such as continued clearing and burning (van Valkenburg & Ketner 1994) especially the lower slopes. This
activity has continued and the environment present is now highly degraded including the summit (N. Grimaldi pers.
obs; Fig. 66). The response of this species to this level of disturbance is unknown. A further specimen collected from
south of Edie Creek, which is approximately 10 km SW of Mt Kaindi, was taken at a similar altitude of 2400 m.
All known specimens are from the 13th of December to the 23rd of March, which may indicate a fairly long flight
time.
FIGURE 59–61. Mating pair of Aenetus simonseni sp. nov. from East Point, NT, 29 December 2018; 59, female (brown colour
form) above, male below; 60, close-up anterior view of the male; 61, female, backlit. Fig. 59 courtesy of N. Volpe; Figs. 60–61,
courtesy of C. Henderson.

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FIGURE 62–63. Live habitus of male Aenetus spp., 62, holotype A. trigonogrammus sp. nov. at light sheet; 63, reared A. lewi-
nii, Lennox Head, NSW. Fig. 62 courtesy of P. Hendry.
FIGURE 64. Riparian-associated monsoon rainforest habitat of Aenetus simonseni sp. nov. from Adelaide River, NT. Figure
courtesy of N. Volpe.

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Remarks. The holotype and one paratype have some historic damage by the pest beetle Anthrenus Geoffroy;
the holotype male is less affected though there is some damage to the abdomen, tornal area of the forewings, and
considerable scale loss has occurred across much of the thorax and abdomen of both specimens. Hepialidae are well
known to fade after collection. In life the wing colouration, particularly in the hindwing basal area of the female,
would likely be much brighter.
FIGURE 65. Dry subtropical Eucalyptus woodland and riparian-associated habitat of Aenetus trigonogrammus sp. nov. from
near Gin Gin, Qld. The holotype was taken to light at the shed in centre left. Figure courtesy of P. Hendry.
Molecular analyses
The Bayesian analysis (Fig. 55) reveals a well-supported (PP = 1.0) monophyletic Aenetus clade (compared to
our seven outgroup taxa). Although this is only based on the single mitochondrial COI gene, it does delineate the
two new species, A. simonseni and A. maiasinus from the 15 other Aenetus species in our analysis, which are also
defined by distinct morphological characters (see Taxonomy section above). Supplementary Table S1 details all
specimens used, including GenBank accessions and locality information. Intraspecific sequence divergences, where
we obtained multiple individuals within species, was greatest within A. thermistis with divergences up to 2.1%
(Supplementary Table S2). These are consistent with variation that we would expect within species included here
(e.g. Grund et al. 2019; Beaver et al. 2020; Moore et al. 2020; but see Simonsen et al. 2019: 796–797 for some
potential caveats).
Sequence divergence comparisons between all Aenetus species compared to our outgroup taxa were between
8.4–14.4% (Supplementary Table S2). Comparisons between species reveal interspecific sequence divergences of
up to 9.1% (between A. ramsayi and A. ligniveren). We also observed interspecific sequence divergences that are
close to 3% for A. simonseni sp. nov. and A. maiasinus sp. nov., when compared to each other and either compa-
red to A. thermistis (Supplementary Table S2), which has been observed in other ghost moths (Beaver et al. 2020;
Moore et al. 2020). Although some levels of sequence divergence are low for a number of Aenetus species, Fig. 55
indicates that these are each monophyletic species (PP = 1.0) and correspond to consistent morphological characters
detailed above.

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FIGURE 66. View of Mt Kaindi area, PNG. Environmental degradation via mining activity apparent in centre, while defores-
tation apparent broadly but particularly centre-left. Remnant montane rainforest present lower centre and on distant hill face.
Figure courtesy of N. Grimaldi.
Discussion
Taxonomic and systematic issues
In the revision of Australian Aenetus by Simonsen (2018), A. tegulatus was recognized as a widespread species
across northern and north-eastern Australia, New Guinea, and eastern islands of Indonesia. Subsequent examination
of the female genitalia of A. tegulatus from the type locality (Ambon Islands) revealed a different morphology to
that of the Australian A. tegulatus illustrated by Simonsen (2018). The Australian population was therefore placed
under the re-established name of A. thermistis (Lower, 1894) as having priority over A. cyanochlora (Lower, 1894)
and A. walsinghami (Olliff, 1895) (Grehan et al. 2018). The Australian distribution is now seen to comprise at least
three allopatric species with A. thermistis in eastern Queensland, A. simonseni across the Northern Territory and A.
maiasinus from the Kimberley region of Western Australia (Fig. 56). At this time, the name A. tegulatus can only be
confidently assigned to the holotype locality of Ambon Island in eastern Indonesia. Aenetus sumatraensis, known
only from a single female specimen in northern Sumatra, has the general appearance of the ‘tegulatus’ complex
(Grehan et al. 2018) and is treated as part of this group. The New Guinea ‘tegulatus’ populations may comprise mul-
tiple species and will be the subject of future study. The Australian ‘tegulatus’ group may also include populations
from the northern Torres Strait Islands that are distinct from the three mainland Australian species (D. A. Lane and
E.P. Beaver pers. obs.). Specimens from Cape York and Thursday Island (see methods) have a narrower apodemal
vinculum, but are otherwise identical to A. thermistis further south in Queensland. This is also consistent with the
geographic difference between A. thermistis north of Daintree (Fig. 55 clade ‘A’) and those to the south from the

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Atherton Tableland (Fig. 55 clade ‘B’). These two clades are 1.5-2.1% divergent (see Fig. 55 and Table S2) that
may represent clinal variation in the absence of distinct morphological characters. This is congruent with divergence
within other taxa in North Queensland at the Black Mountain Corridor (Bryant & Krosch 2016).
Based on sequence similarity (Fig. 55) and external morphology, a GenBank (HQ952155) specimen from
Ravensbourne, southern Queensland, listed as A. tegulatus is actually Aenetus scotti (Scott, 1869). The identifica-
tion of another specimen from Brisbane, southern Queensland, listed by Simonsen (2018) is problematic due to
the absence of further specimens from this region or from anywhere south of Byfield, Queensland. Although this
‘Brisbane’ specimen has not been examined by us, the bursae illustrated in Simonsen (2018: Fig. 522) differs from
all other tegulatus-group specimens we have examined and appears to represent an undescribed species of uncertain
provenance.
Almost all specimens of the A. tegulatus complex examined from the Torres Strait Islands and eastern Cape
York Peninsula, Queensland, are historic specimens that were subjected to the now outdated practice of gutting
and stuffing; usually rendering dissection (and therefore species determination) impossible for these specimens.
Monsoon rainforest, coastal and riparian vegetation on the western coast of the Cape York Peninsula, QLD should
be surveyed for Aenetus, as this genus may occur here within the areas that receive above 1000 mm of annual pre-
cipitation. The Cape York region is of further interest as it is geographically situated between two known centres of
Aenetus diversity, the Wet Tropics region of Queensland, and the central New Guinea highlands (Grehan & Mielke
2018).
Aenetus trigonogrammus is identified as belonging to the Australian ‘splendens’ group (sensu Simonsen 2018)
that is characterised by sexually dimorphic adults where the females have forewing elements characterised by broad
areas of brown and green scales; the brown scales are a type-2 bilayer scale covering (Simonsen 2018), often sepa-
rating or surrounding marginal and discal areas of green scales, whereas males are generally light green, yellow (or
light pink in some A. astathes Turner) with a medial white band on the forewing or other white markings, always
with a white triangular marking in the anal margin area of the forewing. This feature is unique within Aenetus, but
there are broadly similar patterns in other Hepialidae such as the western Eurasian genus Triodia Hübner. Species
in the A. splendens group also share three segmented labial palpi where the basal palpomere is only slightly shorter
than the middle palpomere, and have a lyre shaped second sternite with pointed sclerotised anterolateral corners.
Both characteristics are also shared with A. djernaesae Simonsen, A. blackburnii (Lower) and A. dulcis (Swinhoe)
that are not part of the A. splendens group (Simonsen 2018) that currently comprises A. splendens, A. lewinii, A.
astathes, A. ligniveren, A. tindalei, A. moorei, and A. trigonogrammus. The males in this group range from the small-
est of this genus with a FW length of 16 mm for a small male of A. lewinii. The male of A. trigonogrammus is the
largest of this group with a forewing length of 27 mm. Monophyly of the splendens group may be problematic as
the morphological similarities do not correspond fully with COI sequence similarities; however the presence of only
a single gene in the analysis may be generating an erroneous relationship.
A biogeographic review by Grehan & Mielke (2018) demonstrated that many distributions of Hepialidae in
Australasia (both genera and species) corresponded to tectonic and geological structures that were active in the Me-
sozoic. This correlation supports the view that the modern hepialid fauna is not only a Mesozoic remnant, but may
retain some components of their original Mesozoic distributions. The allopatry of A. simonseni, A. thermistis, and
A. maiasinus is consistent with their having originated within their respective ranges by vicariance from an ances-
tor already widespread over the combined range of each species, but only to the north of the McPherson-Macleay
Overlap (MMO) which is a major biogeographic boundary and centre with a complex tectonic history involving
the New England orogeny in the early Mesozoic, terrane accretion, and formation of the Clarence-Moreton basin in
Jurassic-Cretaceous time, and early Cretaceous volcanism (Heads 2014). The MMO is also at or near the northern
distributional boundaries of the other three ‘splendens’ group species in Queensland with the northern limit of the
main distribution of A. lewinii at the MMO along with a disjunct population in central eastern Queensland (Simon-
sen 2018, Fig. 557), the northern boundary of A. ligniveren at the MMO near Brisbane (Simonsen 2018, Fig. 558),
and the northern range of A. splendens just to the north at Fraser Island (Simonsen 2018, Fig. 556).
In both imago size and genitalia morphology A. albadamanteum is similar to the three large species of Indo-
nesian Papua—A. crameri, A. toxopeusi, and A. hampsoni. The presence of two oblique broken transverse bands
on the male and female forewing is also present in some females of A. cohici, and in at least one colour form of
the male (Boudinot 1991; Salesne 2010). This external similarity is paralleled internally by similarities of the bursa
copulatrix where both species have a small diverticulum, a slightly narrowed central corpus bursae, and a wider and

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slightly angled distal corpus. These features distinguish the two species from the ‘tegulatus’ group in Australia (Figs
26, 29), all other Australian Aenetus species (Simonsen 2018, Figs. 493–528), and the New Zealand species Aenetus
virescens (Dugdale 1994, Fig. 217). This division suggests that the phylogenetic affinities of A. cohici are closest to
one or more clades within New Guinea that includes A. albadamanteum and possibly other species that have yet to
be analysed. This closer affinity between the New Caledonian A. cohici and at least one New Guinea species (A. al-
badamanteum) suggests that the origin of A. cohici may not be the result of vicariance by rifting of continental crust
away from Australia, but rather vicariance of an ancestral clade extending north across what is now the Coral Sea.
A comparable biogeographic example is found in the genus Hunga (Chrysobalanaceae) endemic to New Caledonia
and eastern New Guinea (Heads 2014).
Acknowledgments
The authors are grateful to Marianne Horak, Ted Edwards, You Ning Su, Thekla Pleines, and Andreas Zwick (ANIC,
Canberra), Christine Lambkin, Susan Wright and Karin Koch (QM, Brisbane), Derek Smith and Russel Cox (AM,
Sydney), Simon Hinkley, Joseph Schubert and Catriona McPhee (MV, Melbourne) for enabling loan of material
from the collections under their care, and Ben Parslow for allowing access to the collection and assisting greatly
with the molecular component at SAMA. We are also thankful to Peter Hendry (Sheldon, QLD) for kindly provid-
ing the holotype specimen and associated live photograph of A. trigonogrammus; Yi-Kai Tea (AM Sydney) for
assistance with one of the figure plates; Nick Volpe (Darwin, NT) for obtaining the holotype of A. simonseni and
for use of figure 59, Caitlin Henderson (Brisbane, QLD) for providing figures 60–61 and for imaging material in
MAGNT; Nicolas Grimaldi for providing figure 66 and associated observations; David A. Lane (Atherton, QLD)
and Axel Kallies (Coburg, Vic) for providing material for molecular analysis; Mark Heath and John Koeyers (WA)
for information surrounding A. maiasinus; Tony Hiller (Mt Glorious, QLD) John Moss (Capalaba, QLD) and Peter
Hendry are further thanked for kindly showing their private collections (including the Charles King collection) to
EPB & MDM in late 2019. Thomas Simonsen (NHMA, Denmark) is thanked for sharing his expertise with Aenetus.
The first author is indebted to Joseph Schubert (Museum Victoria, Melbourne) and his family for kindly housing me
during my stay in Melbourne in early 2020. Travel for this work as well as funding for the molecular analysis was
supported with an Australian Lepidoptera Research Endowment grant to MDM & EPB in 2019. The authors are
grateful to Thomas Simonsen, Axel Kallies and Carlos Mielke for critically reviewing the manuscript.
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SUPPLEMENTARY FILES
TABLE S1. Specimen data associated with Figure 55 concerning species identification, location of specimen, Genbank
accession number and collection data for all specimens.
TABLE S2. Percentage sequence divergence table associated with Figure 55.