Two new endemic species of Abantiades Herrich-Schäffer (Lepidoptera: Hepialidae) from Kangaroo Island, Australia

Abstract

Abantiades penneshawensis Moore & Beaver sp. nov. and Abantiades rubrus Moore & Beaver sp. nov. are described as new. Both species are endemic to Kangaroo Island, and although both are related to species that occur on the Australian mainland and other islands, they are distinguished from those sister and phenotypically similar species by morphology and mtDNA (COI) barcodes. These two new species raise the number of Abantiades species on Kangaroo Island to six, three being endemic, and 45 species in the genus for the whole of Australia. There are now 13 species of Hepialidae (one undescribed) known from Kangaroo Island and we discuss the potential effects of recent catastrophic fire on some distributions.

Full text

ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by C. Mielke: 2 Feb. 2021; published: 7 Apr. 2021 571
Zootaxa 4951 (3): 571–597
https://www.mapress.com/j/zt/
Copyright © 2021 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4951.3.9
http://zoobank.org/urn:lsid:zoobank.org:pub:45D34CAC-0EA1-47AB-A919-CA157741549A
Two new endemic species of Abantiades Herrich-Schäffer (Lepidoptera: Hepialidae)
from Kangaroo Island, Australia
MICHAEL D. MOORE1,3, ETHAN P. BEAVER1,4, ALEJANDRO VELASCO-CASTRILLÓN1,5 & MARK I.
STEVENS1,2,*
1 Biological and Earth Sciences, South Australian Museum, SA 5000, Australia.
2 University of Adelaide, Biological Sciences, SA 5005, Australia.
�
mark.stevens@samuseum.sa.gov.au; https://orcid.org/0000-0003-1505-1639
3
�
michael.moore@samuseum.sa.gov.au; https://orcid.org/0000-0002-8796-3330
4
�
ethan.beaver@live.com.au; https://orcid.org/0000-0002-0613-7046
5
�
a.velascocastrillon@gmail.com; https://orcid.org/0000-0002-3516-6655
* Corresponding author
Abstract
Abantiades penneshawensis Moore & Beaver sp. nov. and Abantiades rubrus Moore & Beaver sp. nov. are described as
new. Both species are endemic to Kangaroo Island, and although both are related to species that occur on the Australian
mainland and other islands, they are distinguished from those sister and phenotypically similar species by morphology
and mtDNA (COI) barcodes. These two new species raise the number of Abantiades species on Kangaroo Island to six,
three being endemic, and 45 species in the genus for the whole of Australia. There are now 13 species of Hepialidae
(one undescribed) known from Kangaroo Island and we discuss the potential effects of recent catastrophic fire on some
distributions.
Key words: DNA barcodes, hepialine, morphology, species divergence, taxonomy
Introduction
Kangaroo Island is a large island off the coast of South Australia, Australia. It is attached to the continental land
mass but glacial retreat in the Permian is thought to have carved a channel between the main continental land mass
and the island itself allowing the ocean to interpose itself between the two. Rises and falls in the ocean levels have
led to periods of separation and continuity in the land pieces (Bourman et al. 2016). Therefore, there have been op-
portunities for species to become isolated on Kangaroo Island and evolve to suit the environments found therein.
Kangaroo Island is the wettest and coldest part of the State and in consequence possesses a unique flora and fauna. It
has already been determined that there is one endemic hepialid. In the present article, two more Abantiades Herrich-
Schäffer species are described for the island raising to three the number of described endemic Hepialid species.
Materials and methods
Molecular and morphological analyses
Here we utilise the mitochondrial (mt) cytochrome c oxidase subunit I (COI) gene to delineate between species.
This gene has been successfully used for this purpose in numerous insects (e.g. Hausmann et al. 2011; Hebert et al.
2013; Dorey et al. 2019; Grund et al. 2019) including recent work on hepialids (Beaver et al. 2020a,b; Moore et
al. 2020a,b). In addition to the two new species described here (see Taxonomy below) our taxon sampling included
other Abantiades species and other ‘hepialine’ genera (see Table 1). The methodology applied here for obtaining
COI gene sequences (DNA extraction, PCR amplification and sequencing) follows Moore et al. (2020b) and were

MOORE ET AL.
572 · Zootaxa 4951 (3) © 2021 Magnolia Press
completed at the South Australian Regional Facility for Molecular Ecology and Evolution (SARFMEE) from a
single leg from each specimen.
All new COI 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
publicly available GenBank and BOLD accessions relevant to the current study (see Table 1) in Sequencher v5.1
(http://www.genecodes.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).
A separate model of evolution was applied across each codon partition in our alignment (1st – HKY + I, 2nd &
3rd – GTR + Γ), as suggested by PartitionFinder v2.1.1 (Lanfear et al. 2016). Trees were generated using BEAUTi
and BEAST v1.10.4 (Drummond et al. 2012; see https://beast.community/index.html). We used an uncorrelated re-
laxed molecular clock to calculate trees with empirical frequency-based priors starting from a random tree and Yule
process tree prior, recommended for species level data (Drummond et al. 2006). Analyses were completed using six
independent runs (to ensure consistent results and stationarity) with each analysis run for 100 million generations,
sampling every 1,000th tree. Post-burnin log and tree files for each run were then combined using LOGCOMBINER
v1.10.4 (Drummond et al. 2012). Convergence and stationarity of model parameters was assessed with Tracer
v1.7.1 (Rambaut et al. 2018; https://beast.community/tracer) with 10% of sampled trees discarded as burn-in, and
the maximum credibility tree was generated using Tree Annotator v1.10.4 (Drummond et al. 2012) before being
visualised and modified using FigTree v1.4.4 (https://beast.community/figtree) and Adobe Illustrator (Adobe Sys-
tems, Inc., San Jose, CA).
All dissections involved sectioning of the relevant specimen, placing the sectioned material in 10% KOH solu-
tion and heating in a boiling water bath for between 5 and 15 minutes depending on the condition of the specimen.
All soft tissue was removed leaving the abdominal tube and the sclerotized parts of the reproductive organs. These
were imaged in an alcohol bath using a Leica imaging system composing a Leica binocular microscope, Leica DFC
500 camera and the LAS Core software programme. Whole specimen images were taken using a Canon EOS cam-
era, with a Canon EF 100 mm 1:2.8 Macro lens and Canon Speedlite Transmitter and Speedlite 430 EX11 flashes.
The images were then stacked using the Zerene Stacker software programme (https://www.zerenesystems.com).
Note that all label references below are verbatim and morphological terms follow Simonsen (2018).
Distribution mapping
We mapped the bushfire area in 2019–20 for Kangaroo Island (Fig. 15) using NatureMaps v. 3.0 (https://data.envi-
ronment.sa.gov.au/NatureMaps/). The map generated in NatureMaps was then used to overlay all known Hepialid
species occurrence records for Kangaroo Island (using occurrence records from ALA (2020)) and combined these
with new records (M.D. Moore and E.P. Beaver, unpubl. data).
Abbreviations
Locations: WA, Western Australia; SA, South Australia; KI, Kangaroo Island.
Institutional Collections (all in Australia): ANIC, Australian National Insect Collection, Canberra, ACT; SAMA,
South Australian Museum, Adelaide, SA; WAM, Western Australian Museum, Perth, WA.
Other Collection: NTPC, Nick Temby Private Collection, Whittlesea, Victoria.
Systematics of Abantiades
Molecular analyses
(see Fig. 1)
The Bayesian analysis for each species (Fig. 1) reveals a well supported (PP = 0.84) monophyletic Abantiades clade
compared to our six outgroups taxa (PP = 1.0). Although this is only based on the single mtDNA (COI) gene, it
does corroborate the morphological distinction of the two new species from the 16 other Abantiades species in our
analysis, which are each also defined by unique morphological characters (see Taxonomy section below). Table 1

TWO NEW ABANTIADES FROM AUSTRALIA Zootaxa 4951 (3) © 2021 Magnolia Press · 573
details all specimens used, including GenBank or BOLD accessions and locality information. For instances where
we obtained multiple individuals within species, we found close intraspecific sequence divergences (Supplementary
Table 1). For example, the greatest sequence divergence observed within any of our Abantiades species varied up
to 1.7%, indicating variation that we would expect within species included here (Supplementary Table 1). These
distances also correspond to those reported elsewhere for Lepidoptera that generally indicate below 3% intraspecific
while above 3% interspecific comparisons using the same COI gene (e.g. Grund et al. 2019; Beaver et al. 2020a,b;
Moore et al. 2020b). However, sequence divergences have been found to vary outside these ranges for some Aban-
tiades species (Simonsen et al. 2019; Moore et al. 2020a,b). We were also interested in the divergence within spe-
cies that we had specimens for from Kangaroo Island and the mainland. For A. marcidus Tindale, 1932 specimens
showed intraspecific variation up to 0.8%, and our outgroup taxon Aenetus tindalei Simonsen, 2018 specimens
showed intraspecific variation up to 0.6%, both consistent with population level divergences and the topology from
Figure 1 (see Supplementary Table 1).
Sequence divergence comparisons among the Abantiades species compared to our outgroup taxa ranged be-
tween 6.7–16% (Supplementary Table 1). Examining between Abantiades species reveal interspecific sequence
divergences of up to 12.7% (e.g. between A. magnificus (Lucas, 1898) and A. aurilegulus Tindale, 1932). Pairwise
comparisons between our new Abantiades species A. rubrus sp. nov. to other Abantiades species ranged between
7.6–11.2%, and A. rubrus sp. nov. groups with the other members of the “labyrinthicus” clade, A. labyrinthicus
(Donovan, 1805) and A. mcquillani Simonsen, 2018 (PP = 1.0; Fig. 1). Pairwise comparisons between our second
new species A. penneshawensis sp. nov. to other Abantiades species ranged between 5.8–12.2% and A. penneshaw-
ensis sp. nov. aligns with A. lineacurva Moore & Edwards, 2014, and place both species in the “lineacurva” clade
(Fig. 1). We also observed several interspecific sequence divergences that are below 3%, for example A. pallida
Simonsen, 2018 and A. tembyi Moore & Beaver, 2020 (2.4%) and A. labyrinthicus and A. cf. mcquillani (2.2%)
(Supplementary Table 1), which has been observed in some ghost moths (e.g. Moore et al. 2020b). Although se-
quence divergence can be low between a number of Abantiades species, each correspond to clear morphological
differences (e.g. Moore et al. 2020b).
Taxonomy
Family Hepialidae Stephens, [1828]
Genus Abantiades Herrich-Schäffer, [1855]
Diagnosis:
Simonsen (2018) produced a new diagnosis for the genus. “A patch of elongate scales at the forewing base and the
‘sensory tubercles’ on S2 are both unique and diagnostic for Abantiades.” He also noted that they have hepialine
(Dumbleton 1966; Dugdale 1994) venation, and monopectinate antennae that lack scales on the flagellum.
Abantiades rubrus Moore & Beaver, sp. nov.
Figs 2a–b, 4a–d
Material Examined: 4 males, 1 female
Holotype ♂. Fig. 2a. Located in SAMA collection: S.A., Western Caravan Park, K[angaroo]. I[sland]., G. Sara
[leg.], 35o30’7.0”S 136o46’25.0”E, 8 April 2019. Spec. No, 19117, Leg removed, for tissue, storage, M.D. Moore.
SAMA Database No. 31-021006.
Paratypes ♂. SAMA, 1 ♂, S.A., Western Caravan Park., K.I., G. Sara, 35o30’7.0”S 136o46’25.0”E, 8 April
2019. Spec. No, 19118, Leg removed, for tissue, storage, M.D. Moore. Dissected E.P. Beaver. SAMA Database No.
31-021007. SAMA, 2 ♂ S.A., Western Caravan Park., K.I., M. Moore, 35o30’7.0’S 136o46’25.0”E, 9 April 2019.
Spec. Nos., 19119, 19120, Leg removed, for tissue, storage, M.D. Moore. SAMA Database Nos. 31-021008, 31-
021009.
♀. SAMA, 1 ♀, Ravine du Corsauer [sic], K. Is., 1.3.86 [interpreted as ‘Ravine des Casoars’, Kangaroo Island,
1.III.1886].

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574 · Zootaxa 4951 (3) © 2021 Magnolia Press
TABLE 1. Specimen data for species identification, location of specimens, GenBank accession numbers and collection data for all specimens listed in Figure 1.
Genbank/
BOLD No.
Specimen
code
Species ID Location Collection date Collector Storing
Institute
Latitude Longitude
Ingroup
(Abantiades)
MK561075 M056 Abantiades marcidus VIC Camperdown 2016 M+M Moore SAMA 38.0 14.0 6.8 143.0 6.0 53.1
MT364913 M057 Abantiades marcidus SA Williamstown 2016 G. Hollamby SAMA 34.0 40.0 6.4 138.0 53.0 15.5
MT364912 G103 Abantiades marcidus SA Kohinoor, Playford
Highway, Kangaroo
Island
30-Apr 2019 D.A. Young SAMA 35.0 53.0 58.0 136.0 53.0 53.6
MT364911 G156 Abantiades marcidus SA Emu Bay, Discovery
Lagoon, Kangaroo
Island
30-Apr 2019 M+M Moore SAMA 35.0 36.0 52.9 137.0 31.0 27.8
MT364915 G087 Abantiades marcidus SA Vivonne Bay,
Kangaroo Island
4-Apr 2019 M+M Moore SAMA 35.0 59.0 3.0 137.0 10.0 37.0
MT364914 G094 Abantiades marcidus SA Mouth Flat Road,
Kangaroo Island
21-Apr 2019 M+M Moore SAMA 35.0 50.0 23.9 137.0 56.0 18.5
MT364909 H013 Abantiades marcidus QLD Goondawindi 2014 N. Temby SAMA 28.0 33.0 4.9 150.0 19.0 22.8
MT052735 H122 Abantiades pallida WA Eucla, Quarantine
Station
7-May 2018 R. Penning SAMA 30.0 49.0 17.8 117.0 29.0 12.3
MT052737 H61 Abantiades tembyi SA 30km EbyN of Swan
Reach
13-May 2017 M. Hura M. Hura Priv.
Coll
34.0 29.0 36.0 139.0 54.0 56.0
MK561106 TJS 13-77 Abantiades
macropusinsulariae
SA Vivonne Bay,
Kangaroo Island
20-Apr 2006 D.A. Young ANIC 36.0 137.2
MK561104 M018 Abantiades
cephalocorvus
SA Goog’s Track 28-May 2016 J. Forrest SAMA 31.0 34.0 11.2 133.0 58.0 3.0
MK561121 M029 Abantiades atripalpis SA Lake Giles, Eyre
Peninsula
24-May 2016 J. Forrest SAMA 33.0 5.0 28.8 136.0 35.0 59.0
MT052712 G51 Abantiades argentata SA Swan Reach 24-May 2018 M. Hura SAMA 34.0 31.0 29.0 139.0 54.0 53.0
MT364908 M061 Abantiades argentata VIC Walpeup Reserve 16-Apr 2015 N. Temby NTPC 35.0 8.0 21.6 142.0 1.0 25.2
ANICM728-10 ANICM728 Abantiades aurilegulus WA 2km WbyS of
Bullabulling
6-May 1984 E.S.Nielsen,
E.D.Edwards
ANIC 31.0 120.5
WALPA4345-13 WALPA4345 Abantiades aurilegulus WA Boxwood Hill,
Chingarrup Sanctuary
1-Mar 2010 P.M. Heath CBG 34.3 118.7
ANICM705 ANICM705 Abantiades leucochiton VIC S.E. Big Desert,
at 15km WSW of
Rainbow
30-Mar 2008 F. Douglas 35.9 142.0
…continued on the next page

TWO NEW ABANTIADES FROM AUSTRALIA Zootaxa 4951 (3) © 2021 Magnolia Press · 575
TABLE 1. (Continued)
Genbank/
BOLD No.
Specimen
code
Species ID Location Collection date Collector Storing
Institute
Latitude Longitude
MT364907 M062 Abantiades
leucochiton
VIC Walpeup Reserve 16-Apr 2015 N. Temby NTPC 35.0 8.0 21.6 142.0 1.0 25.2
MT773570 H20 Abantiades pica VIC Rainbow (also known
from Kangaroo Island)
2014 F. Douglas SAMA 35.9 142.0
MT364901 G089 Abantiades rubrus
sp. nov.
SA Western KI Ca. Pk.,
Kangaroo Island
8-Apr 2019 G. Sara SAMA 35.0 30.0 7.0 136.0 46.0 25.0
MT364902 G091 Abantiades rubrus
sp. nov.
SA Western KI Ca. Pk.,
Kangaroo Island
10-Apr 2019 M. Moore SAMA 35.0 30.0 7.0 136.0 46.0 25.0
MT364900 G088 Abantiades rubrus
sp. nov.
SA Western KI Ca. Pk.,
Kangaroo Island
8-Apr 2019 G. Sara SAMA 35.0 30.0 7.0 136.0 46.0 25.0
MT364903 G090 Abantiades rubrus
sp. nov.
SA Western KI Ca. Pk.,
Kangaroo Island
10-Apr 2019 M. Moore SAMA 35.0 30.0 7.0 136.0 46.0 25.0
MT364904 H023 Abantiades
labrynthicus
NSW Armidale 2017 G. Beaver Ethan
Beaver PC
30.0 27.0 48.4 151.0 40.0 52.6
MT364905 TJS-DNA-
13-144
Abantiades cf.
mcquillani
TAS Mt William NP 29-Feb 2009 G Cocking SAMA 41.500 141.130
MT364923 H100 Abantiades
magnificus
VIC Camp Road, Bunyip State
Forest
30-Jan 2017 K. Green Kees
Green PC
37.987 145.659
MT364922 H001 Abantiades
magnificus
VIC Camp Road, Bunyip State
Forest
30-Jan 2017 K. Green Kees
Green PC
37.987 145.659
MT364917 M045 Abantiades
lineacurva
WA Chingarrup, Boxwood
Hill
25-Apr 2015 P.M. Heath SAMA 34.0 18.0 16.5 118.0 43.0 43.2
MT364918 M069 Abantiades
lineacurva
WA Chingarrup, Boxwood
Hill
25-Apr 2015 P.M. Heath SAMA 34.0 18.0 16.5 118.0 43.0 43.2
MT364916 H031 Abantiades
lineacurva
WA Chingarrup, Boxwood
Hill
29-Apr 2017 M.Moore SAMA 34.0 18.0 16.5 118.0 43.0 43.2
MT773571 M44 Abantiades
lineacurva
WA Dryandra State Forrest,
Narrogin
7-Apr 2015 M+M Moore SAMA 32.0 47.0 7.3 116.0 59.0 1.0
MT364920 G096 Abantiades
penneshawensis sp.
nov.
SA Baudin Cons. Pk,
Penneshaw, Kangaroo
Island
21-Apr 2019 M+M Moore SAMA 35.0 43.0 28.3 137.0 57.0 15.2
MT364919 G097 Abantiades
penneshawensis sp.
nov.
SA Baudin Cons. Pk,
Penneshaw, Kangaroo
Island
24-Apr 2019 M+M Moore SAMA 35.0 43.0 28.3 137.0 57.0 15.2
…continued on the next page

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576 · Zootaxa 4951 (3) © 2021 Magnolia Press
TABLE 1. (Continued)
Genbank/
BOLD No.
Specimen
code
Species ID Location Collection
date
Collector Storing
Institute
Latitude Longitude
MT364921 G098 Abantiades
penneshawensis sp. nov.
SA North Cape Road (439),
Emu Bay, Kangaroo
Island
30-Apr 2019 M+M Moore SAMA 35.0 35.0 58.2 137.0 35.0 40.7
ANICM730-10 ANICM730 Abantiades aphenges NSW CSIRO Exp. Farm,
Wilton
13-Apr 1982 V.J.Robinson ANIC 34.2 150.4
ANICM729-10 ANICM729 Abantiades fulvomarginatus WA L.O`Halloran, Deepdene 24-Apr 1963 V.J.Robinson ANIC 34.3 115.1
Outgroup
MT773572 G79 Aenetus tindalei SA Snake Lagoon,
Kangaroo Island
23-Apr 2019 E. Beaver Ethan Beaver PC 37.0 57.0 26.8 136.0 39.0 32.1
MT773573 G44 Aenetus tindalei SA Gorge Track, Mt. Lofty
Ranges
11-Dec 2018 E. Beaver Ethan Beaver PC 34.0 51.0 35.0 138.0 43.0 37.4
MT773574 H83 Aenetus tindalei SA Montacute, Mt. Lofty
Ranges
16-Feb 2018 E. Beaver Ethan Beaver PC 34.0 51.0 35.0 138.0 43.0 37.4
MT773575 G81 Aenetus tindalei SA Flinders Chase,
Kangaroo Island
25-Jan 2019 E. Beaver Ethan Beaver PC 35.0 56.0 58.3 136.0 40.0 26.5
MT773576 G43 Aenetus tindalei SA Flinders Chase,
Kangaroo Island
20-Dec 2018 E. Beaver Ethan Beaver PC 35.0 56.0 58.3 136.0 40.0 26.5
MT773577 M75 Aenetus djernaesae WA Jurien Bay 18-Apr 2015 M+M Moore SAMA 30.0 20.0 19.7 115.0 4.0 11.6
MN901447 H65 Aenetus blackburnii SA Eyre Pen. (also known
from Kangaroo Island)
12-Apr 2018 M. Moore SAMA 34.0 49.0 4.2 135.0 48.0 31.6
HQ952132 Zelotypia stacyi NSW Otford 25-Feb 1964 V.J. Robinson University of
Maryland
34.2 151.0
MN901446 Oncopora intricata TAS Mt Field 1-Feb 2018 E.P. Beaver SAMA 42.7 146.6
KF491607 Callipielus arenosus Chile Anticura, Puyehue
Osorno
30-Nov 1992 L. Pena University of
Maryland
40.6 73.2
MT773578 G93 Oxycanus occidentalis SA Mouth Flat Road,
Kangaroo Island
21-Apr 2019 M+M Moore SAMA 35.0 50.0 23.9 137.0 56.0 18.5
MN901444 Oxycanus antipoda VIC Anglesea 7-May 2018 N. Temby SAMA 38.4 144.2
MN901445 H17 Elhama australasiae VIC Mt. Disappointment 2017 M. Moore SAMA 37.4 145.1

TWO NEW ABANTIADES FROM AUSTRALIA Zootaxa 4951 (3) © 2021 Magnolia Press · 577
Diagnosis:
Abantiades rubrus sp. nov. is endemic to Kangaroo Island. It is a large, monopectinate “labyrinthicus” styled moth
but where males when fresh, have all wings suffused a russet colour. The male can be identified by its relatively
small triangular shaped pseudoteguminal lobe that has, i) a small sharp angled disto-posterior corner, ii) a posterior
margin that is shorter and steeper than the anterior one, and iii) a rounded obtuse angled disto-posterior process,
iv) the large distal processes of the valva when compared to the pseudoteguminal lobe and v) the shape of sternite
VIII (Fig. 4a–d). Analysis of the mtDNA (COI) will also allow positive identification. For a complete comparison
of the species within this clade, and other Abantiades species likely to be found within the range, see the Remarks
section below.
Description:
Male
Fig. 2a
Head: Frons densely covered in longish grey scales, fronto-clypeus region dark grey with band of much lighter,
grey scales at antenna level. Compound eyes black, slightly smaller than head capsule in anterior view. Labial palps
three segmented, second palpomere longest, third shortest, sub spherical, all densely clothed in longish scales, first
segment sand, second, dark red brown, third, black brown. Antennae wine red in colour, one quarter length costa,
total 59 segments, monopectinate, each ramus lanceolate to elliptical with round blunted apex, concave outer face,
all angled toward distal end, covered in fine, colourless, very short setae.
Thorax: Densely covered in long piliform scales. Anterior portion dark grey, rest of scales sand coloured but
with light grey tips. Forewing; length 39–48 mm. Wing membranes opaque white, costal vein sandy brown others
much lighter brown. Dorsal surface; prominent, discoidal and submarginal marks, both, linear but irregular in shape,
composed short shining white scales, edged and highlighted in dark brown. Anterior to discoidal mark, small area
of dark brown highlighting scales at base of wing. From costa to Rs4 filled with fine grey scrolling lines highlighted
by darker brown scales. Light sand coloured scales fill spaces. Area between dorsum and M3, and from termen to
submarginal white mark, filled with fine grey scrolling lines edged in dark brown. Central wing area filled with red
chestnut scales, most of which are part of thick, broad scrolling lines, with dark brown highlights. Ventral surface:
dorsal markings visible through wing, particularly in apical area and highlighted in dark brown. Apex and termen
with short fine scales producing fine sand coloured scrolling lines, rest of wing covered more sparsely by sand
coloured piliform scales redder along veins. Whole basal area suffused with reddish pink. Hindwing; length 28–35
mm. Dorsal surface, covered in fine cinnamon piliform scales, covered with red/pink suffusion. Ventral surface, as
for forewing including scrolling at apex and termen, rest, sparser covering of long sandy piliform scales with red-
dish suffusion. Legs: Densely covered in scales. Dorsally short scales all legs brown/grey, laterally long pencil like
scales, grey. Ventrally, sand coloured, laterally reddish sandy/grey.
Abdomen: Covered all over in long piliform cinnamon scales. Sternite VIII; Fig. 4d roughly rectangular but all
sides curved. Posterior margin more sclerotised, slightly convex but with central gentle concavity. Laterals margins
concave, anterior margin more so. Posterior corners pointed, anterior corners distinctly rounded.
Genitalia: Fig. 4a–c. Apodemal vincular arms thick, disto-posterior portion thinner, distal portions heavily
sclerotised. Arms meet very broadly producing a broad based “U” shaped saccus. Posterior margin of ventral face
rises gently but has a large, heavily sclerotized distinct square shaped notch centrally. Posterior margin of dorsal
face a gentle shallow “V” shape. Twin tergal processes thick based, lightly sclerotised, long, twice as high as dorso-
posterior corner, linear, pointed apex. Pseudoteguminal lobe heavily sclerotised, roughly triangular. Dorso-posterior
margin, short, concave, dorso-posterior corner, small, angular, protuberant. Disto-posterior margin much longer
slightly concave, irregular. Distal pseudoteguminal process, moderately wide base, rounded. Ventro-posterior margin
slightly longer than disto-posterior margin, sinusoidal but with longer concavity. Pseudoteguminal arms thick round-
ed and re-curved towards juxta. Valvae, “pear” shaped with distal arms equal length of sacculus and almost as broad.

MOORE ET AL.
578 · Zootaxa 4951 (3) © 2021 Magnolia Press
FIGURE 1. Maximum credibility tree from Bayesian phylogenetic analysis of the mitochondrial DNA COI gene of all 36
Abantiades sequences used and 13 outgroup taxa. Included are 18 Abantiades species, including A. rubrus sp. nov. and A. pen-
neshawensis sp. nov. Posterior probabilities greater than 0.7 are shown at each node. An * identifies specimens from Kangaroo
Island, while ** identifies specimens of species known from Kangaroo Island. Details for all sequences including GenBank
accessions and BOLD process IDs are available in Table 1.

TWO NEW ABANTIADES FROM AUSTRALIA Zootaxa 4951 (3) © 2021 Magnolia Press · 579
Female
Fig. 2b
Head: Compound eyes black, slightly smaller than head capsule in anterior view. Labial palps covered in long stiff
scales, three segmented, second palpomere longest, third shortest, stalked, ovate distally. Antennae (damaged: only
one present, broken and re-glued ventrally to body) almost apectinate, with flat vertical distal face and proximal side
rounded. Angled posterior faces with flat profiles.
Thorax: Forewing; length 77 mm, dorsal surface with two large white coloured marks in the discoidal and
submarginal areas, these narrowly highlighted in dark brown. Rest of wing heavily scrolled with linear elements
along dorsum and termen, but many smaller, circular elements in apical and costal regions. Ventral surface sparsely
covered in long sandy piliform scales. Hindwing; 57 mm, dorsal surface, plain but with a reddish hue. Ventrally, as
forewing.
Abdomen: All densely covered in long brown piliform scales.
Genitalia: See remarks.
FIGURE 2. Abantiades rubrus sp. nov., dorsal view. Male, (a) Holotype; SAMA collection: S.A., Western Caravan Park, K.I.,
G. Sara, 35o30’7.0”S 136o46’25.0”E, 8 April 2019. Spec. No, 19117, Leg removed, for tissue, storage, M.D. Moore. SAMA Da-
tabase No. 31-021006. Female, (b) Paratype; SAMA collection: Ravine du Corsauer [sic, interpreted as ‘Ravine des Casoars’],
K. Is., 1.3.86 [interpreted as 1886].
Distribution and biology:
Recorded Localities: Kangaroo Island. A single female specimen was collected at Ravine des Casoars from the
north-west of the island, while the male specimens were all collected approximately 25 km south-east, at Western
Caravan Park (Fig. 15).

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Flight times: Early March and early April.
Larval biology: Unknown.
Habitat: Collected in a developed area of small to moderate sized Eucalypt trees.
FIGURE 3. Abantiades labyrinthicus, dorsal view. Male, (a) SAMA: Victoria; Whittlesea, 37.454889 S, 145.096215 E, 7th
February 2015, N. Temby. SAMA Database No. 31-020398. Female, (b) SAMA: Mt. Buangor, Vic., 1400 ft. at m.v.l., 14 Feb.
1956, I.F.B. Common. SAMA Database No. 31-015722.
Etymology:
‘Rubrus; red’, an eponym reflecting the redness of the specimens distinguishing them from the similar Abantiades
labyrinthicus.

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FIGURE 4. Abantiades rubrus sp. nov. Male genitalia and sternite VIII. (Paratype, S.A., Western Caravan Park., K.I., G. Sara,
35o30’7.0”S 136o46’25.0”E, 8 April 2019. Spec. No, 19118, Leg removed, for tissue, storage, M.D. Moore. Dissected E.P. Bea-
ver. SAMA Database No. 31-021007. (a) lateral view genitalia, (b) ventral view genitalia, (c) dorsal (internal) view genitalia,
(d) sternite VIII.
Morphological structures: am: anterior margin; Brt: basal rim of the pseudotegumen; Dav: distal-posterior arm of vinculum;
Dipm: disto-posterior margin; Dopc: dorso-posterior corner; Dopm: dorso-posterior margin; Dpp: disto-posterior process;
pm: posterior margin; Sba: anterior margin of dorsal face of saccus; Sbf: dorsal face of saccus; Sbp: posterior margin of dorsal
face of saccus; Scu: sacculus; Sfa: anterior margin of ventral face of saccus; Sff: ventral face of saccus; Sfp: posterior margin
of ventral face of saccus; Tpr: dorso-distal twin processes; Va: apodemal vinculum; Vl: valva; Vpa: ventral pseudoteguminal
arm; Vpm: ventro-posterior margin.

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FIGURE 5. Abantiades mcquillani. Male genitalia. (a) lateral view genitalia, (b) ventral view genitalia, (c) dorsal (internal)
view genitalia. (Australia: Devils Elbow Rd, 0.5 km NE of Bass Hwy, 3.2 km E of Montumana, 168m. 40o57.622’S 145o
33.042’E, 19 Jan 2011 K. Hill, D. Marschall.).
Morphological structures: am: anterior margin; Brt: basal rim of the pseudotegumen; Dav: distal-posterior arm of vinculum;
Dipm: disto-posterior margin; Dopc: dorso-posterior corner; Dopm: dorso-posterior margin; Dpp: disto-posterior process;
pm: posterior margin; Sba: anterior margin of dorsal face of saccus; Sbf: dorsal face of saccus; Sbp: posterior margin of dorsal
face of saccus; Scu: sacculus; Sfa: anterior margin of ventral face of saccus; Sff: ventral face of saccus; Sfp: posterior margin
of ventral face of saccus; Tpr: dorso-distal twin processes; Va: apodemal vinculum; Vl: valva; Vpa: ventral pseudoteguminal
arm; Vpm: ventro-posterior margin.
Remarks:
Over time, Abantiades specimens fade to a rather uniform brown colour, and because of this we have kept references
to colour in the description of the female specimen to a minimum. Due to the age and significance of this single
specimen, we have not dissected it.
The males of the three species in the “labyrinthicus” group (Fig. 1) differ in the shape of their sternite VIII and
in the components of the genitalia. Abantiades rubrus sp. nov. and A. mcquillani are most similar and both species
have significantly smaller pseudoteguminal lobes than A. labyrinthicus (see Figs 4a–d, 5a–c, 6a–d). The disto-pos-
terior margin in A. rubrus sp. nov. is straight producing a pseudoteguminal lobe that is triangular with a triangular

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disto-posterior process. In A. mcquillani the disto-posterior process has a deep concavity distally that produces a
distinct posterior facing projection that, with a flat topped pseudoteguminal lobe produces a keel shaped process. In
A. rubrus sp. nov. the anterior margin of the saccus is flatter and hence more “U” shaped and the posterior margin
of the forward face has a wider deeper notch than in A. mcquillani. Compared to the above two species A. labyrin-
thicus (Fig. 6a–d) is significantly different having a much more massive and longer pseudoteguminal lobe, with a
curved and backward raked disto-posterior process. The valva have much thinner distal half, and the twin processes
are much longer with inward curved distal portions. The anterior margin of the saccus is more “V” shaped and the
vincular arms are markedly different being relatively shorter and thicker and of a different form whereas in the other
two species the saccus is less “V” and more “U” shaped and the vincular arms relatively longer and thinner. Sternite
VIII (Fig. 6d) is distinctly different having the most complex shape, with lateral margins that are concave but with
posterior angled processes absent in the other species.
FIGURE 6. Abantiades labyrinthicus. Male genitalia and sternite VIII. (Frank Tetley Park, nr Fred Piper Lookout, Brown Mtn.
nr Bega, N.S.W., 12 Jan 1982, M.S.& B.J. Moulds. SAMA Database No. 31-015733) (a) lateral view genitalia, (b) ventral view
genitalia, (c) dorsal (internal) view genitalia, (d) sternite VIII.
Morphological structures: am: anterior margin; Brt: basal rim of the pseudotegumen; Dav: distal-posterior arm of vinculum;
Dipm: disto-posterior margin; Dopc: dorso-posterior corner; Dopm: dorso-posterior margin; Dpp: disto-posterior process;
pm: posterior margin; Sba: anterior margin of dorsal face of saccus; Sbf: dorsal face of saccus; Sbp: posterior margin of dorsal
face of saccus; Scu: sacculus; Sfa: anterior margin of ventral face of saccus; Sff: ventral face of saccus; Sfp: posterior margin
of ventral face of saccus; Tpr: dorso-distal twin processes; Va: apodemal vinculum; Vl: valva; Vpa: ventral pseudoteguminal
arm; Vpm: ventro-posterior margin.

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FIGURE 7. Abantiades penneshawensis sp. nov., dorsal view. Male, (a) Holotype; SAMA collection: S.A., Baudin Cons. Pk.,
K.I., Penneshaw, M+M Moore, 35o43’28.3”S 137o57’15.7”E, 21 April 2019, Spec. No., 19145, Leg removed, for tissue, storage,
M.D. Moore. Dissected E.P. Beaver. SAMA Database No. 31-021001. Female, (b) Paratype; SAMA collection: S.A., Baudin
Cons. Pk., K.I., Penneshaw, M+M Moore, 35o43’28.3”S 137o57’15.7”E, 21 April 2019, Spec. No., 19160, Leg removed, for
tissue, storage, M.D. Moore. SAMA Database No. 31-021003. Dissected E.P. Beaver.
On Kangaroo Island the large size of A. rubrus sp. nov. will separate it from the related but smaller Oxycanus
Walker, 1856 species and the un-forked antennal pectinations in this species will separate it from the sympatric, tri-
forked A. argentata (Tindale, 1932), A. atripalpis (Walker, 1856) and A. macropusinsulariae Simonsen, 2018, and
the bi-forked and much smaller A. pica (Tindale, 1932). In addition, males can be separated from the other mono-
pectinate species found on the island easily by the aforementioned wing colouring and patterning, as A. marcidus
is a white and grey moth with few scrolling lines and A. penneshawensis sp. nov. is a smaller black coloured moth
with a distinct white line on the forewing and no obvious scrolling lines (see description below).
The females of A. rubrus sp. nov. can be separated from A. penneshawensis sp. nov. on appearance, the latter
being blackish with a large white, sinuous or “T” shaped mark, on the forewing, whereas A. rubrus sp. nov. has two
large marks on the forewing (dull white or brown/grey) which if joined (a common event in A. labyrinthicus) would
not form a sinuous line. The antennae differ too, with A. penneshawensis sp. nov. having plate shaped rami whereas
A. rubrus sp. nov. is almost apectinate.
Dissection may be necessary to discriminate between the females of A. rubrus sp. nov. and A. marcidus, as they
are both large in size, apectinate, have substantial amounts of scrolling lines over their surface and are grey/brown
with two marks on their forewings. The prominences on the flagellum of A. marcidus have more rounded backs than
the flatter ones on A. rubrus sp. nov. but this is difficult to determine without experience.

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The Kangaroo island species is reddish in comparison to the brown eastern species which also has rather yel-
lowish hindwings. This phenotypic difference, along with the abovementioned morphological differences is further
supported by the mtDNA (COI) sequence which also distinguishes this species from A. labyrinthicus and from A.
mcquillani (Fig. 1).
The flight times in Victoria of A. labyrinthicus show that it often emerges around Christmas Day (N. Temby
pers. comm.) and continues to emerge over the intervening months until the end of April. This breeding strategy
is unusual with respect to many other species whose emergence time is limited to a few wet days in autumn (M.D.
Moore and E.P. Beaver, pers. obs.). Nor does A. labyrinthicus seem as tightly bound to rain events as most other
species, as it emerges at times when rainfall cannot be reliably depended upon. What this means with regards to A.
rubrus sp. nov. we do not know but the male specimens collected in 2019 were well in advance of the season break-
ing rains which fell on April 30th and so it too may have a long emergence period, independent of rain, involving
lower numbers at any one time.
Abantiades penneshawensis Moore & Beaver, sp. nov.
Figs 7a–b, 9a–d, 11a, 12a, 13
Material Examined:
Holotype ♂. (Fig. 7a). Located in SAMA collection: S.A., Baudin Cons[ervation]. Pk., K[angaroo].I[sland].,
Penneshaw, M+M Moore [leg.], 35o43’28.3”S 137o57’15.7”E, 21 April 2019, Spec. No., 19145, Leg removed, for
tissue, storage, M.D. Moore. Dissected E.P. Beaver. SAMA Database No. 31-021001.
Paratypes ♀. SAMA, 1♀, S.A., North Cape Road, K.I., Bay of Shoals, M+M Moore, 35o35’58.2”S
137o35’40.7”E, 30 April 2019. Spec. No., 19167, Leg removed, for tissue, storage, M.D. Moore. SAMA Database
No. 31-021002. SAMA, 1♀, S.A., Baudin Cons. Pk., K.I., Penneshaw, M+M Moore, 35o43’28.3”S 137o57’15.7”E,
21 April 2019, Spec. No., 19160, Leg removed, for tissue, storage, M.D. Moore. SAMA Database No. 31-021003.
Dissected E.P. Beaver. SAMA, 1♀, MacGillivray, Kangaroo I. S.A., April 1931. J. Wood.
Diagnosis:
The male of Abantiades penneshawensis sp. nov. is a medium sized moth though small for this genus. They are
black with a prominent white sinuous stripe on the forewing. The females too are dark being black or dark grey also
with prominent, white wing markings, though this mark may be more complex than the white marking of the males
(see Fig. 13 of female specimen).
This species is most externally similar to the Western Australia species A. lineacurva, and dissection is likely
to be needed for both genders to distinguish between the two. In the males there are antennal differences, with the
number of antennal segments on the Kangaroo Island specimen significantly less than in the Western Australian
species (49 versus 57 segments) and at the base of each mid length flagellomere segment the dorsal distal projection
is distinctly longer and more steeply angled with the face of the flagellomere, proscribing a right angle (Fig. 12a),
whereas in A. lineacurva the dorsal distal projection is shorter and the angle with the face of the flagellomere is
much larger (see Fig. 12b). The shape of sternite VIII in A. penneshawensis sp. nov. is unique (Fig. 9d) but amongst
other differences, A. penneshawensis sp. nov. can also be distinguished by the lack of a disto-posterior corner on the
pseudoteguminal lobe (Fig. 9a), whereas in A. lineacurva the corner is large and rounded (Fig. 10a).
The females in the “lineacurva” clade can be distinguished from all other Hepialids by the combination of i)
black/dark grey colouring, ii) monopectinate circular plate like rami on the antennae (Fig. 12c) and iii) the white,
irregular “T” shaped mark on the forewing. To separate A. lineacurva and A. penneshawensis sp. nov. dissection is
needed. Then it can be seen that the shape of the dorsal plate in A. penneshawensis sp. nov. is less regular in form
because the lateral margins are shorter and curved, producing longer rounded laterally displaced lobes than the
shorter, triangular more centrally placed ones in A. lineacurva and that there are differences in antevaginal lamella
where in A. penneshawensis sp. nov., the medial lobe of the antevaginal lamella is relatively taller and thinner and
projects deeper into the central space of the oviduct than in A. lineacurva which is shorter and wider (Fig. 11a–b).
Both A. penneshawensis sp. nov. and A. lineacurva are quickly distinguished from the externally similar A.
moesta (Tindale, 1932) (a WA endemic) by comparison of the antennae, which in the latter species are light coloured
and pseudobipectinate (see Simonsen 2018).

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Analysis of the mtDNA (COI) will also allow positive identification (Fig. 1). For a complete comparison of the
species within this clade, and other Abantiades species likely to be found within the range, see the Remarks section
below.
Description:
Male
Fig. 7a
Head: Frons thickly covered in long grey piliform scales, fronto-clypeus region darker. Compound eyes large,
black, each only slightly smaller than head capsule. Labial palps three segmented, covered in longish stiff grey/
brown scales. Second palpomere longest and straight, first and third similar length, first linear, third subspherical.
Antennae total of 49 antennomeres, one-third length of costa, monopectinate, wine red-brown. At the base of each
flagellomere the dorsal distal projection present and perpendicular with the face of the flagellomere (Fig. 12a). Rami
twice flagellomere height, plate-like with slightly flattened top, not symmetrical, slightly longer towards posterior.
Thickly covered in small, colourless setae, longer on lateral sides. Rami angled towards distal end.
Thorax: Anterior part covered in light grey/white long piliform scales, rest all grey. Forewing; Length 26 mm.
Membrane whitish opaque, veins brownish white picked out with reddish brown scales. Dorsal surface, has numer-
ous white markings but largest and most obvious, a long, widish, sinuous white line that begins at origin of M veins,
runs along M1 and M2 until about middle of wing whence follows M1 to near wing margin, then runs towards
the apex which it fails to meet. Other larger, non-uniform, irregular shaped white marks run from apex making an
irregular “T” shape. A fine white line produced sub-tornally from intervein marks. Other white markings seen in
apical and posterior areas. All white marks are edged dark grey. Dark grey scales also largely fill all the remaining
spaces in anterior half of wing. Along termen the “filler” scales lighter grey and lighter still along the dorsum. Minor
scrolling present along dorsum. Ventral surface, veins thickly covered in longish brown scales, rest thinly covered
in same. Scales longer still towards base and along dorsum. Hindwing; Length 20 mm. Veins brownish, covered in
long thin reddish piliform scales. Dorsal surface, intervein areas thinly covered in long pale straw-coloured scales,
longer at base and along dorsum. Ventral surface, as forewing. Legs: dorsally covered in short dark grey scales,
much longer laterally. Ventrally pale yellow-brown.
Abdomen: Tergites I and II covered in same pale yellow piliform scales present in base of hindwing. Remain-
ing tergites, covered densely in long light grey piliform scales. Ventrally darker, browner scales. Sternite VIII; Fig.
9d, roughly trapezoid in shape. Posterior margin convex but with neat, well defined, sclerotised, central concavity.
Lateral sides straight but with posterior flaring. Anterior margin longest, convex with acute ends and shallow central
concavity.
Genitalia: Fig. 9a–c. Vincular arms relatively short, thick, apodemal vinculum more lightly sclerotised very
thick proximally. Saccus posterior margin of ventral face rising gently to a prominence with a deep square shaped
notch, anterior margin a steep shallow “V” with a very acute, proportionally lengthy tail-like depression. Posterior
margin of dorsal face a shallow “V”. Twin tergal processes longer conical but from broad base, distally curved in-
ward. Basal rim thick posteriorly but narrows quickly as progresses forward. Pseudoteguminal lobe heavily sclero-
tised, minutely toothed along ventro-posterior margin and apex. No dorso-posterior margin or dorso-posterior cor-
ner. Disto-posterior margin straightish from rim but curves outward to produce a large triangular tooth slightly over
half way along its length. From the apex of the triangle straight but then into a deep concavity that rises to a rounded,
narrow based, outwardly curved, serrated disto-posterior process. Ventro-posterior margin sinuous, minutely ser-
rated, pseudoteguminal arms steeply recurved towards juxta. Valvae with very wide blunt distal arms, large relative
to their small rounded base.

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Female
Fig. 7b
Head: Frons densely covered in piliform medium grey scales, fronto-clypeus region darker. Two, large, black,
compound eyes, each only slightly smaller than head capsule. Labial palps three segmented. Second palpomere
longest, linear. First and second clothed in long straight dark grey scales. Third shortest, stalked but spherical end,
clothed shorter light grey scales. Antennae monopectinate, 52 antennomeres, all red ochreous. Rami, distally lean-
ing, lanceolate but with flattened rounded apex, more tear shaped and acute pointed distally. All covered in small,
fine colourless setae.
Thorax: anterior clothed as head, rest darker. Wing membrane, grey, opaque but see through. Forewing; length
44–50 mm. Dorsal surface, many white markings but one thickish, long, sinuous line stretching from the origin of
M vein, along M1 and M2 narrows to follow M1 at about four-fifths distance to termen bends upwards towards apex
which it almost makes. Other white marks posterior to upward turn cause production of vague “T” shape, more ap-
parent in living specimens (Fig. 13). Anterior to main white line in region of cell, basal half thickly edged in dark
grey. Smaller white marks to posterior and anterior of main line. Thick line of white marks sub-tornally that might
be continuous or separated. All white scales somewhat yellowish and all white marks edged narrowly in dark grey
scales. Around apex and anterior half, rest of space filled mainly with dark grey scales, but with a few light grey
areas that produce insignificant scrolling patterns. Posterior to line, the filler scales tend to be lighter in colour. Ven-
tral surface, veins covered in brown/grey piliform scales. Rest sparsely covered in longer scales of the same colour,
getting longer towards base and dorsum. Hindwing; 36–42 mm, veins very pale straw colour. Dorsal surface, veins
covered in reddish/straw coloured piliform scales. Rest sparsely covered in long piliform scales of similar colour.
Ventral surface, as for forewing but slightly darker. Legs: All medium grey, ventrally, lighter.
Abdomen: All densely covered in brown/grey piliform scales.
Genitalia: Fig. 11a. Dorsal plate lightly to moderately sclerotised, split into two long lobes rounded at ends but
with roughly parallel sides due to broad triangular invagination. Invaginated area moderately covered with long
thick setae, more but shorter on face of plate. Antevaginal lamella split into three portions. Two lateral plates, have
shear like inner margin, outer (ventral) margin gentle arc. Outer face shallowly ridged with line of setae along crest.
Medial lobe most heavily sclerotised. Rises steeply to form cone shape but with rounded concave notch on short in-
ner margin, producing a bilobed apex, moderately clothed in longish thick setae. Face of “cone” moderately covered
in less thick setae. Sub-anal plates, small, tear shaped. Inward margins producing point, outer margin seem bilobed,
each rounded. Bursa copulatrix not located.
Distribution and biology:
Recorded Localities: Three localities on eastern coastal Kangaroo Island: Baudin Conservation Park; Penneshaw;
Bay of Shoals; and MacGillivray. MacGillivray is an area comprising a substantial portion of the eastern-central
part of the island; however, it is suspected the specimen would have been taken to the north-east which is near to the
other localities, all of which are coastal (Fig. 15).
Flight times: March and April.
Larval biology: Unknown. Possibly root feeders on Allocasuarina.
Habitat: Collected in and likely closely associated with Allocasuarina woodland.
Etymology:
Named after Penneshaw Primary School, given the students interest in Lepidoptera and their collection of one of the
female specimens used in this work.

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FIGURE 8. Abantiades lineacurva, dorsal view. Male, (a) Holotype; ANIC collection: Western Australia, Kojonup, 18.iv.1960,
Wallace. Female, (b) Paratype; ANIC collection: Nedlands, 21.iv.1964, Wallace.
Remarks:
Differences in Mitochondrial sequencing, morphological structure and its allopatric distribution all support Aban-
tiades penneshawensis sp. nov. as being a valid species.
The mitochondrial DNA (COI) sequencing, with a mean difference of 9.6% clearly clusters A. penneshawensis
sp. nov. separately from A. lineacurva (Fig. 1), and the genitalia structures in both the male and female differ sub-
stantially. In the male, A. lineacurva has a much thinner attachment at the basal rim to the much shorter and different
shaped twin processes than in A. penneshawensis sp. nov. where the attachment is thicker and the twin processes
much taller and pointed. Abantiades lineacurva has a differently shaped pseudoteguminal lobe with a distinct round-
ed dorso-posterior corner, a more distinct triangular point on its posterior margin and a more pointed disto-posterior
process (Figs 9a, 10a). The apodemal vinculum in the two species are quite different. Abantiades penneshawensis
sp. nov. has arms that are proximally very wide and rounded. In A. lineacurva the arms are proximally much thinner,
and less rounded with their broadest point situated more distally producing a saccus that has more area and a more
triangular shape whereas in A. penneshawensis sp. nov. it is long and pointed (Figs 9b–c, 10b–c). Sternite VIII is
different too in the two species. In A. lineacurva it has rounded sides with deep concavities in both the posterior and
anterior margins, whereas in A. penneshawensis sp. nov. it is much squarer with straightish sides and the concavities
in posterior and anterior margins are shorter and much shallower (Figs 9d, 10d).

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Like the males, the females of A. penneshawensis sp. nov. and A. lineacurva are externally similar, and dissec-
tion is required to differentiate the two. In A. lineacurva the lobes of the dorsal plate are distinctly triangular and
symmetrical whereas in A. penneshawensis sp. nov. they are less symmetrical being rounded distally with broad,
round apices. The lateral lobes of the antevaginal lamella whilst similar are deeper in A. penneshawensis sp. nov.
but the medial plate is much taller, narrower, and with a smaller invagination on the inner margin whereas in A.
lineacurva it is broader, proportionally shorter and has a large, wide, “V” shaped central depression (Fig. 11a–b).
On Kangaroo Island the un-forked antennal pectinations in this species will separate it from the sympatric, tri-
forked A. argentata, A. atripalpis and A. macropusinsulariae, and from the bi-forked A. pica, and the rounded plate
shaped rami in both the males and females of A. penneshawensis sp. nov. will distinguish this species from the other
monopectinate, A. marcidus and A. rubrus sp. nov. which in the males have lanceolate rami and where the females
are virtually apectinate.
FIGURE 9. Abantiades penneshawensis sp. nov. Male genitalia and sternite VIII. (Holotype, S.A., Baudin Cons. Pk., K.I.,
Penneshaw, M+M Moore, 35o43’28.3”S 137o57’15.7”E, 21 April 2019, Spec. No., 19145, Leg removed, for tissue, storage,
M.D. Moore. Dissected E.P. Beaver. SAMA Database No. 31-021001. (a) lateral view genitalia, (b) ventral view genitalia,
(c) dorsal (internal) view genitalia, (d) sternite VIII. The red arrow indicates the absence of the dorso-posterior corner on the
pseudotegumen (compared to Fig. 10); and blue arrow indicates the relative thickness of the basal rim of the pseudotegumen
(compared to Fig. 10).
Morphological structures: am: anterior margin; Brt: basal rim of the pseudotegumen; Dav: distal-posterior arm of vinculum;
Dipm: disto-posterior margin; Dopc: dorso-posterior corner; Dopm: dorso-posterior margin; Dpp: disto-posterior process;
pm: posterior margin; Sba: anterior margin of dorsal face of saccus; Sbf: dorsal face of saccus; Sbp: posterior margin of dorsal
face of saccus; Scu: sacculus; Sfa: anterior margin of ventral face of saccus; Sff: ventral face of saccus; Sfp: posterior margin
of ventral face of saccus; Tpr: dorso-distal twin processes; Va: apodemal vinculum; Vl: valva; Vpa: ventral pseudoteguminal
arm; Vpm: ventro-posterior margin.

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FIGURE 10. Abantiades lineacurva. Male genitalia and sternite VIII. (West. Aust.; South Coast, Chingarrup, Boxwood Hill,
34o18’17”S 118o43’43”E, M. Heath; 29th April 2017. Spec. No., 17115. Leg removed for tissue storage M.D. Moore. Dissected
M. Moore 2018. SAMA D atabase No. 31-020334) (a) lateral view genitalia, (b) ventral view genitalia, (c) dorsal (internal) view
genitalia, (d) sternite VIII. The red arrow indicates the presence of the dorso-posterior corner on the pseudotegumen (compared
to Fig. 9); and blue arrow indicates the relative thickness of the basal rim of the pseudotegumen (compared to Fig. 9).
Morphological structures: am: anterior margin; Brt: basal rim of the pseudotegumen; Dav: distal-posterior arm of vinculum;
Dipm: disto-posterior margin; Dopc: dorso-posterior corner; Dopm: dorso-posterior margin; Dpp: disto-posterior process;
pm: posterior margin; Sba: anterior margin of dorsal face of saccus; Sbf: dorsal face of saccus; Sbp: posterior margin of dorsal
face of saccus; Scu: sacculus; Sfa: anterior margin of ventral face of saccus; Sff: ventral face of saccus; Sfp: posterior margin
of ventral face of saccus; Tpr: dorso-distal twin processes; Va: apodemal vinculum; Vl: valva; Vpa: ventral pseudoteguminal
arm; Vpm: ventro-posterior margin.

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FIGURE 11. Female genitalia. Posterior view. (a) Abantiades penneshawensis sp. nov., Paratype; SAMA collection: S.A.,
Baudin Cons. Pk., K.I., Penneshaw, M+M Moore, 35o43’28.3”S 137o57’15.7”E, 21 April 2019, Spec. No., 19160, Leg removed,
for tissue, storage, M.D. Moore. SAMA Database No. 31-021003. Dissected E.P. Beaver. (b) Abantiades lineacurva. Para-
type; ANIC collection: Nedlands, 21.iv.1964, Wallace. The red arrow indicates the lobe on the dorsal plate; and the blue arrow
indicates the medial lobe on the antevaginal lamella.
Morphological structures: Dp: dorsal plate; Lal: lateral lobe of the antevaginal lamella; Mal: medial lobe of the antevaginal
lamella; Sub: sub-anal plate.
FIGURE 12. Male antennae at middle length. (a) Abantiades penneshawensis sp. nov., ventral view, (b) Abantiades lineacurva,
ventral view, (c) Abantiades lineacurva, lateral view.

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FIGURE 13. Photograph of a live female Abantiades penneshawensis sp. nov. 31 March 2011, Penneshaw. Photo David Man-
gham.

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FIGURE 14. Map showing the known distribution across Australia for Abantiades lineacurva, A. labyrinthicus, A. mcquillani,
King Island - A. labyrinthicus. Green fill shows Kangaroo Island where our two new endemic species occur.
FIGURE 15. Distribution on Kangaroo Island for Abantiades rubrus sp. nov., Abantiades penneshawensis sp. nov. and all other
Hepialid species that occur on the island (using occurrence records from ALA (2020) and new records from M. Moore and E.
Beaver, unpubl. data). The ‘year’ of last fire shown, and for 2019 the bushfire extent is shown as red shading (using NatureMaps,
https://data.environment.sa.gov.au/NatureMaps/) (accessed 12/xii/2020).
Discussion
Before further collecting events on Kangaroo Island in Autumn 2019, both of the new species described herein were
represented in SAMA by single female specimens. Abantiades rubrus sp. nov. was represented by a specimen col-

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lected in the extreme north west of Kangaroo Island, South Australia in 1886 and only located when the collection
was databased in 2018, and Abantiades penneshawensis sp. nov. by a specimen collected from the eastern half of
the island in 1931. This latter specimen was examined by M. Moore when he and E. Edwards described and named
A. lineacurva Moore & Edwards, 2014, but because it was a worn female from an “unexpected” location, thus al-
lopatric distribution, its determination and veracity was uncertain, and it was not dissected nor mentioned in the
subsequent paper.
In the phylogeny A. rubrus sp. nov. groups with A. labyrinthicus and A. mcquillani (Fig. 1). Up until 2018,
this clade had only A. labyrinthicus; however, A. mcquillani was added more recently in a taxonomic revision of
the genus (Simonsen 2018). Abantiades labyrinthicus (Fig. 3) is a widespread eastern Australian species found on
the mainland from south-eastern Queensland down and around Australia’s eastern and south-eastern coasts, and in
Tasmania. Abantiades mcquillani is currently known only from Tasmania where according to Simonsen (2018) it
seems sympatric for some of its distribution with A. labyrinthicus.
In the cladogram A. penneshawensis sp. nov. groups with A. lineacurva, which up until now, was its only
member. Abantiades lineacurva has been collected from various locations in Western Australia, distributed between
Gooseberry Hill, near Perth to Boxwood Hill, east of Albany.
The two new Kangaroo Island species described herein and A. macropusinsulariae, raise to three the known
species of the genus Abantiades endemic to Kangaroo Island, suggesting a significant period of isolation.
At present the Backstairs Passage separates Kangaroo Island from the mainland by a 40 m to 80 m deep chan-
nel of water 14 km wide at its narrowest point. Throughout the repeated glacial cycles that have dominated since
the Miocene to the present day (Bourman et al. 2016), there have been several periods where sea-level has dropped
sufficiently enough to expose a land-bridge between the island and mainland. It is thought that the current separa-
tion of the island started at the end of the last glacial maximum 16,000 years ago and attained its current form about
7,000 years ago (Bourman et al. 2016). Based on the level of mtDNA sequence divergences we identified for A.
penneshawensis sp. nov. and A. rubrus sp. nov., and assuming a generalised invertebrate molecular clock rate of
1.5–2.3% divergence per million years that has been widely used (e.g. McGaughran et al. 2010), it is likely that
these moths became isolated on the island during one of these earlier periods of separation in the late Pliocene to
early Pleistocene, and have remained isolated either geographically or ecologically.
By contrast, the Kangaroo Island populations of Aenetus marcidus and A. tindalei (Fig. 1) revealed no signifi-
cant sequence divergence from the populations found on the mainland. This suggests that these species either have
only been recently isolated on Kangaroo Island (see Beaver 2019 for Aenetus tindalei) or that ongoing dispersal
across Backstairs Passage is possible for these two species. Such dispersal may be possible based on the observed
distribution of species such as A. labyrinthicus across six eastern states/territories including Tasmania (separated by
Bass Strait) (Fig. 14), but contrasts with the isolation of the endemic species on Kangaroo Island and A. mcquillani
on Tasmania (see Figs 1, 14). In Australia all the hepialid moth species (with the exception of the early morning
emergence of Aenetus ombraloma (Lower), (E.P. Beaver, pers. obs.)) emerge in the late evening or at night and
observations would suggest that they sit motionless during the day relying on camouflage (external appearance) to
escape predation (especially by birds; M.D. Moore and E.P. Beaver, pers. obs.). In addition, having no mouth parts
evolved for feeding in the imago stage, these moths have very short life spans as adults, even if predation was not a
problem. This suggests that dispersal may be limited. Further study is needed to examine if such island populations
are effectively isolated from their mainland populations and if differing life histories and adaptations to specialised
niches (including host plants) may have had an influence in the speciation of island endemics. Of interest too, are
the factors driving wing patterning in these species and why little change has occurred. In the “labyrinthicus” group
all the currently known species have a virtually identical wing pattern, (See Figs 2, 3) making identifications dif-
ficult without genitalia dissections. For example, in the present study, and in the absence of genitalia examination,
A. penneshawensis sp. nov. (Fig. 7) was originally considered A. lineacurva, (Fig. 8) and Simonsen (2018) only
recently described A. mcquillani from specimens previously believed to belong to A. labyrinthicus. Abantiades has a
much greater species diversity than has been previously recognised (see also Moore & Edwards 2014; Moore 2014;
Simonsen et al. 2019; Moore et al. 2020a,b).
Recent calls to understand and describe the ‘dark taxa’ (e.g. Page 2016) before they go extinct has never been
more relevant in Australia, and this is especially true on Kangaroo Island. Our recent efforts to describe species in
the Hepialidae come at a time when Australia experiences one of its most catastrophic fire seasons on record. Kan-
garoo Island has a mean annual rainfall of 487.7 mm, but 2018 was the third driest (333.9 mm) and 2019 (304 mm)

TWO NEW ABANTIADES FROM AUSTRALIA Zootaxa 4951 (3) © 2021 Magnolia Press · 595
the driest ever recorded (BOM 2020a). During the summer of 2019/2020 catastrophic fires starting in the northwest
of the Island were driven by winds that burnt 50% of the Island, 96% of Flinders Chase National Park and about
70% of the Island’s intact bushland (NPWSA 2020) (Fig. 15). The entire known ranges of A. rubrus sp. nov. and
an as yet undescribed Oxycanus species, and many KI localities of A. pica, A. marcidus, Aenetus tindalei, Aenetus
blackburnii (Lower, 1892) and Oxycanus occidentalis Tindale, 1935 were burnt by this extremely hot fire (see Fig.
15). To what extent these species have survived the fire is not yet known. Their habit of living a subterranean ex-
istence as a large larva could have helped the Abantiades survive the fire, however all of the leaf litter in the burnt
areas have been destroyed leaving thousands of hectares of bare sandy earth. Evidence with Aenetus (Beaver &
Grehan 2018, Kay et. al. 2020) and observations on Abantiades (Nick Temby, pers. com.) suggest the very young
caterpillars require leaf litter to survive the first few months of life. After a fire the adults may still enclose, mate and
lay eggs, however the young larvae may die in greater numbers because of the lack of leaf litter. Of great concern
is that the eucalypt forests of Australia will continue to dry out and remain fire risks threatening the flora and fauna
within them (BOM 2020b).
Acknowledgements
We would like to thank Thomas Simonsen for his help and support and for providing several mtDNA sequences and
Nick Temby for his involvement and preparedness to provide his observations and field work. Thanks to SAMA
staff members, Peter Hudson, Chris Watts, Matt Shaw and Ben Parslow for their ongoing encouragement and ad-
vice, and for access to the SAMA collection. Thanks to friend and colleague Howard Hamon for his ability to turn
raw images into art, and for his constant reviewing of the images and to David Mangham for the use of his image
in this paper. We would also like to thank Greg and Judy Sara, Andy Young and Richard Glatz for their support in
this work and for their collection of specimens and especially to Brianna Manoel and the students in her Year 2,3,4
Composite class at Penneshaw Primary School for their passionate interest and support of the work and for the col-
lection of the first fresh specimen of Abantiades penneshawensis sp. nov. We would also like to thank reviewer Axel
Kallies, for the time and effort he has put into improving this manuscript, and Zootaxa editor Carlos Mielke for his
continuing support. Some of these specimens were collected under permit from the South Australian Department for
Environment and Water. We would sincerely like to thank the South Australian Field Naturalists Lirabenda Grant
for funding much of this research.
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