A new species of Fraus, Walker 1856 (Lepidoptera: Hepialidae) from the south-west of Western Australia

Abstract

The species Fraus extremapodus sp. n. from Western Australia is described, illustrated and compared with the sympatric F. basidispina Nielsen and Kristensen, 1989 and F. polyspila (Meyrick, 1890). With a wingspan of 39 mm it is one of the largest species of Fraus Walker, 1856 and, compared to wing length, it has the longest legs of any Australian hepialid.

Full text

A NEW SPECIES OF FRAUS WALKER, 1856 (LEPIDOPTERA:
HEPIALIDAE) FROM THE SOUTH-WEST OF WESTERN AUSTRALIA
ETHAN P. BEAVER and MICHAEL D. MOORE
South Australian Museum, Adelaide, South Australia, 5000
ethan.beaver@live.com.au
Abstract
The species Fraus extremapodus sp. n. from Western Australia is described, illustrated and compared
with the sympatric F. basidispina Nielsen and Kristensen, 1989 and F. polyspila (Meyrick, 1890).
With a wingspan of 39 mm it is one of the largest species of Fraus Walker, 1856 and, compared to
wing length, it has the longest legs of any Australian hepialid.
Key words: taxonomy, homoneurous, short-range endemic, mesic zone
Introduction
Fraus Walker, 1856 is a diverse, endemic genus of primitive ghost moths with 25
species found across much of southern and eastern Australia. They are among the
smallest of the Hepialidae, and certainly the smallest in Australia with a wingspan
between 15 and 35 mm. Like many genera in this family, Fraus are often conned
to areas of higher rainfall, though a few species are known from drier semi-arid
environments such as mallee or dry heathland particularly in South Australia and
Western Australia.
Fraus are generally considered Hepialidae s. lat. or ‘primitive’ Hepialidae due to
the presence of a proboscis vestige in some species, the lack of pseudoempodial
setae on the pretarsus, less connection between the trulleum and juxta, and an
unbroken ventral row of spines on segment VII of the pupa (Nielsen and Scoble
1986; Nielsen and Kristensen 1989). Fraus was removed from Hepialidae s. str.
by Nielsen and Scoble (1986) along with the similar genera Gazoryctra Hübner,
Antihepialus Janse and Afrotheora Nielsen and Scoble, a determination followed
by Grehan (2012) and Simonsen and Kristensen (2017), who found that those
latter genera together form a monophyletic clade, however were unable to resolve
the relationship between that clade, Fraus, and the Hepialidae s. str. Nielsen and
Kristensen (1989) postulated that the monophyly of Hepialidae s. str. could not
be supported without retention of Fraus as Hepialidae s. lat. Reiger et al. 2015
found evidence in the form of a molecular phylogenetic analysis of the basal
lineages of Lepidoptera which supported the monophyly of Hepialidae s. lat. only
for Gazoryctra. Further work is required to determine if the ‘primitive’ hepialid
genera warrant subfamilial status (J. Grehan pers. comm.).
Fraus are easily distinguished from other comparably small Australian Hepialidae
such as Jeana Tindale by comparison of wing venation. In Fraus, Rs1 shares a
stalk with Rs2, and Rs3 + Rs4 also share a common stalk in both wings compared
with Jeana which has typical ‘oxycanine’ venation (Meyrick, 1890; Dumbleton,
1966), where Rs3 branches o Rs1 + Rs2, as well as major dierences in male
valvae, which though simple parallel-sided in Jeana, are often elaborate or club-
shaped in Fraus, and with the retention of the proboscis in some species of the
latter, which Jeana lack. Oncopera Walker, though of a comparable size, are
Australian Entomologist 47 (1): 39–50, 27 March 2020 39

of much heavier build and, except for O. brachyphylla Turner, have elaborate
metatibial androconia in the males, which Fraus lack. All Fraus share the
combined features of: elongate elliptical scales with attened or serrate apexes,
and the females with a patch of hair-scales at the hindwing basal area.
A succinct account of the short taxonomic history of Fraus is recorded by Nielsen
and Kristensen (1989), who added 17 new species to the existing eight in their
landmark revision, and recognised two synonyms. Fraus extremapodus sp. n. is
the rst new Fraus species to be described since Nielsen and Kristensen (1989),
bringing the total number of recognised species to 26.
Methods
Material examined for this study for morphological comparison other than that which
is listed within the systematic component: F. pelagia (Turner): 1 ♂, lectotype, 1 ♀
paralectotype (in ANIC); F. simulans (Walker): 105 specimens: 63 ♀, 33 ♂ (in SAMA),
7 ♂, 2 ♀ (in EPBC). F. basidispina Nielsen and Kristensen: 8 paratypes: 2 ♂, 6 ♀, non-
types: 1 ♀, 1 ♂, (all in SAMA). F. pteromela (Lower): 1 ♂ lectotype, 1 ♀ paralectotype,
non-types: 43 ♂ 38 ♀ (in SAMA), 11 ♂, 5 ♀ (in EPBC). F. polyspila (Meyrick): 119 ♂, 33
♀, (in SAMA), 10 ♂, 4 ♀ (in EPBC). F. quadrangula Nielsen and Kristensen: 10 ♂, 1 ♀
paratypes, non types: 2 ♂, 5 ♀ (in SAMA).
Other material examined primarily for comparison of wing scale structure and/or leg ratios
(see discussion), all in SAMA: Gorgopis libania (Stoll): 3 ♂ 1 ♀. Eudalaca stictigrapha
(Hampson): 2 ♂ paratypes. Gazoryctra wielgusi Wagner and Tindale: paratypes: 11 ♂.
Jeana delicatula Tindale: 1 ♂ holotype, 1 ♀ paratype, 1 ♂ paratype. non-types: 4 ♂, 9 ♀.
Abantiades argentata (Tindale): 1 ♂ holotype, 1 ♀ paratype. Aenetus lewinii (Walker):
3 ♂. Oncopera intricata Walker: 1 ♂. Elhamma australasiae (Walker): 1 ♂. Oxycanus
australis Walker: 1 ♂. Zelotypia stacyi Scott: 1 ♂. Anomosetidae: Anomoses hylecoetes
1 ♂ . Prototheoridae: Prototheora petrosema: 1 ♂.
Dissection: Genitalia: The abdomen was removed and placed in 20% KOH
solution and heated over a ame for 6–8 minutes before being cleaned with
distilled water, then stored in ethanol in a labelled vial kept in a tray adjacent
to the pinned specimen in SAMA. Legs: the legs in Figs 12–17 were removed
from the set specimens and placed in 20% KOH solution for 15 minutes at
ambient room temperature, then were cleared of scales using a ne brush and
are stored in a separate vial in the same manner as the genitalia. Genitalia were
compared directly to both other dissections and with set specimens, and also with
those gured in Nielsen and Kristensen (1989: 176–185, Figs 306–363), which,
although slide mounted, still allow for appropriate morphological comparison
with the non-slide mounted dissections performed for this study.
Dissections were imaged in an alcohol solution 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 camera, with a Canon EF 100 mm 1:2.8 Macro lens and Canon Speedlite
Transmitter and Speedlite 430 EX11 ashes. The images were then stacked using
the Zerene stacker software programme.
Diagrams of adult wing venation were drawn freehand by tracing images of the
40 Australian Entomologist

wings, the resulting drawings were then compared with the set specimens to
ensure accuracy. Terminology for genitalia and wing morphology follows Nielsen
and Kristensen (1989), and Simonsen (2018). Informal species-groups referred to
in the text follow Nielsen and Kristensen (1989).
Fig. 1. Male Fraus extremapodus sp. n., holotype.
Figs 2–3. Male Fraus spp: (2) F. polyspila; (3) F. basidispina, paratype.
1
2 3
47 (1): 39–50 (2020) 41

Abbreviations: General: SAMA—South Australian Museum, Adelaide. ANIC—
Australian National Insect Collection, Canberra. WAM—Western Australian
Museum, Perth. KGC—research collection of Kees Green, Melbourne. EPBC—
research collection of E.P. Beaver, Adelaide.
Morphology: FW—forewing. HW—hindwing. For male genitalia (Figs 3–11):
br—basal rim of pseudotegumen. ip—intermediate plate. jx—juxta. pa—
pseudoteguminal arm. pst—pseudotegumen. sac—saccus. scc—sacculus. s8—
eighth sternite. tl—tergal lobe. tp—twin processes. tr—trulleum. vi—apodemal
vinculum. vl—valvae. For legs (Figs 12–17): fm—femur. ta—tarsi. ti—tibia.
Systematics
Hepialidae Stevens, 1829, s. lat., sensu Nielsen and Kristensen 1989
Fraus extremapodus sp. n.
(Figs 1, 4–8, 13–15, 19–21)
Material examined. Holotype ♂, WESTERN AUSTRALIA: Walpole,
34°58’35.4”S 116°42’38.9”E, 22 April 2017, M.D. Moore / specimen no. 17083,
H025, dissection no. EPB-05 (in WAM).
Paratypes: 1 ♂: same data as holotype except specimen no. 17084, H026, (in
SAMA). 2 ♂: Keystone SF, Walpole, WA, S34.57.58°, E116.41.12°, 50 m elev.,
22–28 April 2017, Kees Green (in KGC). 1 ♂: nr Albany, Western Australia,
34.52.427°S, 117.49.984°E, 1–2 May 2000, M. Court. ex. Light trap, Eucalyptus
globulus plantation #4, 1 / BOLD id 10ANIC-09640 / ANIC Database No. 31
036927 (in ANIC).
Diagnosis. Fraus extremapodus is distinctive and not easily confused with
any other species. It is distinguished from the males of all other Fraus species
by the following external characteristics: antennae liform; absence of wing
patterns; extremely long legs, midleg almost the length of the forewing; wing
venation distinctive, Sc in FW divided, CuP to margin, 2A present in HW; scales
at wing margin dentate. There are distinctive dierences in male genitalia, such
as the presence of piliform scales on the valvae, although otherwise they bare
general similarities to the Western Australian F. basidispina, F. polyspila, and the
Tasmanian F. pelagia (see Discussion).
Description. Male. Head: Antennae 8 mm, to middle of costa, slender,
liform, pale brown, scape sub-rectangular, pedicel depressed ovoid, 33 or 34
agellomeres. Eyes moderate, smaller than head capsule, widely spaced dorsally.
Scales on frons and vertex dense and grey. Labial palps two segmented, posterior
palpomere short, ovoid, anterior palpomere elongate, narrow. Proboscis absent.
Thorax: pro- and mesothorax dorsally and ventrally covered in light grey scales
and interspersed with brown scales on ventral surface. Legs very long, all subequal
(typical for Fraus), (Figs 13–15) prothoracic leg 16 mm, mesothoracic leg 17 mm,
metathoracic leg 14.5 mm, all with short grey scales on femur, tibia and tarsus,
epiphysis absent, tibia dorsally convex medially in pro and midleg, concave in
metaleg. Wings: FW: length 18.5 mm, expanse 39 mm. Narrow, elongate and
42 Australian Entomologist

subtriangular. Costa straight. Apex rounded. Wing venation (Fig. 21) Sc divided,
CuP to margin, 2A absent, otherwise typical for Fraus. HW: Narrow, elongate,
subtriangular. Costa slightly concave centrally. Tornus rounded. Wing venation
(Fig. 21): Sc not divided, 2A present. Wings nely scaled, semi-translucent.
Dorsal FW ground colour light grey, without pattern. Scales along costa and
basal area slightly darker. Fringe with elongate dark grey scales from apex to
jugum, longest at jugum, scales serrate, apex with 6–9 teeth. Jugum long. Patch
of long black scales between jugum and thorax. Ventral surface with piliform
scales basally. HW dorsal surface light grey, darker along costa. Fringe as for
FW. Ventral surface as above. Basal area covered with white piliform scales.
Abdomen: long, narrow, uniformly dark grey interspersed with brown scales.
Tergum 1 sub-rectangular, length/width ratio 0.89, posterior margin with central
indentation. Sternum 8 (Fig. 5) sub-trapezoidal, posterior margin with deep
central indentation, anterior margin rounded, sclerotised at posterior corners, less
sclerotised medially and anteriorly.
Male genitalia (Figs 4; 6–8). Apodemal vinculum broad and short, with paired
posterior projections rounded and smooth. Saccus subtriangular, conuent with
apodemal vinculum. Intermediate plate highly sclerotised, roughly triangular
with anterior apex pointed. Basal rim of pseudotegumen broad, rounded, posterior
margin of pseudotegumen simple, sclerotised, smooth. Ventral pseudoteguminal
arm membranous. Twin processes very short, rounded. Juxta broad and rectangular,
slightly membranous towards trulleum. Trulleum elongate, highly sclerotised,
positioned inferior to pseudotegumen, hook-like projection extending between
valvae towards saccus. Valvae large, straight, posterior margin slightly concave,
large triangular sclerotised projection at sacculus. Posterior margin clothed with
long setae, anterior margin with both setae and piliform scales.
Female Unknown.
Etymology. The species is named in reference to its extremely long legs, longest
for any Fraus spp. both by actual and proportional length. A suggested common
name is ‘Long-legged primitive ghost moth’.
Phenology. Flight time is recorded from the 22 April until 2 May, during or on
the same night as rainfall. The species is apparently localised, uncommon to light,
and is rarely collected. Some Fraus species such as F. polyspila (Meyrick, 1890)
or F. pteromela (Lower, 1892) can occur at light in huge numbers, however F.
extremapodus has so far only been encountered singularly or in pairs.
Distribution. Specimens are known only from the high rainfall region of south-
western Western Australia, where it has been taken from Walpole and near Albany
(Figs 22, 23).
Biology. The adults are nocturnal and have been taken at MV and UV light at
night, during rainfall. This species occurs in tall mixed Eucalyptus diversicolor
and E. jacksonii woodland and in E. marginata woodland, in a thick band of
remnant roadside vegetation near a cleared area, while the specimen from near
Albany was taken from a Eucalyptus globulus plantation. Little is known of the
47 (1): 39–50 (2020) 43

biology of most Fraus species, however one species F. simulans Walker, 1856 is
univoltine with concealed larvae that are nocturnal grass foliage feeders of both
native and exotic species (Hardy, 1973). Further study is required to corroborate
the life history of the other Fraus spp., including that of F. extremapodus.
Discussion
Various aspects of adult morphology are unique to this species, and warrant
discussion and comment in comparison with other species.
Regarding male genitalia structure, F. basidispina (Figs 3, 9–12) have similarly
long valvae to F. extremapodus, however are distinguished by the presence of two
to three short spines on the sacculus; the short, ovoid trulleum; vinculum without
posterior projections; pseudotegumen with elongate, spine-like pseudoteguminal
arm, twin process longer, narrower; intermediate plate uniformly narrower with
apex rounded, not pointed. The male F. polyspila (Fig. 2) (for genitalia of F.
polyspila see Nielsen and Kristensen 1989: 184, 185, Figs 356–359) are similar in
the broad sense by way of the trulleum and vinculum shape however the valvae are
considerably broader, without anterior triangular structure; pseudotegumen with
large centero-ventral spine and prominent ventral arm, which F. extremapodus
lacks. The male genitalia of F. pelagia (see Nielsen and Kristensen 1989: 185,
Figs 362, 363) is similar by way of general pseudotegumen and trulleum shape,
Figs 4–8. Male genitalia and sternite 8 of Fraus extremapodus sp. n. holotype male,
dissection ID EPB-05: (4) posteroventral view; (5) sternite 8; (6) posterodorsal view; (7)
ventral 3-quarter view; (8) lateral view. For abbreviations see ‘Methods’ section.
4 6
5
7 8
44 Australian Entomologist

however dier in that the valvae are distinctly club-shaped, pseudoteguminal
arms prominent and spine-like, and the vinculum narrow. Despite the supercial
similarities in some aspects of male genitalia, F. basidispina, F. polyspila, and
F. pelagia are all distinguished by way of the external characteristics previously
listed, and F. basidispina with the addition of the presence of a proboscis in that
species, which is absent in F. extremapodus. A further characteristic of note is that
tergum 1 is very similar in shape to F. quadrangula, also a Western Australian
species, though there is a slightly higher length/width ratio in F. quadrangula
(0.93) whereas it is 0.89 in F. extremapodus, the two species are otherwise
dissimilar in many respects, particularly by way of male genitalia, where F.
quadrangula has distinctively short valvae and a globose trulleum (see Nielsen
and Kristensen 1989: 177, 178, Figs 316–319).
There are at least three scale types on the wings of F. extremapodus, the most
common type is the ‘simulans’ type illustrated in Nielsen and Kristensen (1989:
Fig. 70 for F. pelagia) where the scales are long, slender and with an acute apex,
however some of the wing scales are the more typical Fraus type where they
are elongate with apical margin almost straight with only minute dentation,
characteristic of Fraus spp. A third scale type that has not been previously recorded
Figs 9–12. Male genitalia of Fraus basidispina male, dissection ID EPB-02: (9)
posteroventral view; (10) posterodorsal view; (11) 3-quarter view; (12) lateral view.
For abbreviations see ‘Methods’ section. Note that sternite 8 could not be located for
comparison.
9 10
11 12
47 (1): 39–50 (2020) 45

in other Fraus spp. but is present in F. extremapodus is best described as an extreme
condition of the typical Fraus type, where the apical dentations are elongate (Fig.
10), this scale type is conned to the fringe of both fore and hindwings of the
new species. This third type is present at the wing fringe in other Hepialidae s.
lat. such as Gazorycta and in Hepialidae s. str. such as Gorgopis and Eudalaca
as well as in Anomoeses (Anomosetidae) and Prototheora (Prototheoridae). This
was also recorded by Nielsen and Kristensen (1989) who noted that wing-scale
Figs 13–18. Legs of Fraus spp, (13–15): Fraus extremapodus sp. n.; (13) prothoracic leg;
(14) mesothoracic leg; (15) metathoracic leg. (16–18) Fraus basidispina (16) prothoracic
leg; (17) mesothoracic leg; (18) metathoracic leg. For abbreviations see ‘Methods’ section.
Scale bar applies to all gs.
13 14 15
16 17 18
46 Australian Entomologist

Figs 19–20. Close-up of key characteristics of male F. extremapodus sp. n.; (19) liform
antennae; (20) Scales at the margin of the forewing, note the dentate apex (black arrow).
Fig. 21. Wing venation diagram of male Fraus extremapodus sp. n.
1919 20
21
47 (1): 39–50 (2020) 47

apical margins of Hepialidae s. str. are similar, as is that of Antihepialus, while the
same scale apex type was recorded in Afrotheora by Nielsen and Scoble (1986)
and by Simonsen and Kristensen (2017) for Paratheora Kristensen & Simonsen,
Palaeoses Turner, and Genustes Issiki & Stringer. Another common scale shape
in Hepialidae s. str. is short, ovoid scales which are present in Abantiades and
Aenetus while Jeana interestingly have slender scales with an acute apex of the
same shape as the F. simulans group (see methods for all material examined). The
presence of the prominently dentate scale margins within F. extremapodus and
in several other unrelated genera is likely plesiomorphic. Details of wing scale
structure such as presence/absence of microtrichia could not be observed with
the available equipment, however may be of additional future research interest.
Sexual dimorphism is present in Fraus spp., with the females larger and usually
lighter marked or coloured compared with the males. If this pattern of dimorphism
is similar in F. extremapodus then the female may be larger than F. basidispina,
which with a wingspan of 43 mm is currently recognised as the largest Fraus
species. Additionally, all known female Fraus have a patch of hair-scales present
(often weakly) at the base of the hindwing, a character unique to Fraus, however
further collecting is required to discover the female of F. extremapodus and check
if this feature is present.
Unusually, the wing venation of F. extremapodus is dierent to that of other
Fraus spp., which are otherwise mostly consistent in terms of wing vein number
(Nielsen and Kristensen 1989: Figs 60–62). In the forewing, CuP continues to
the wing margin in F. extremapodus whereas it ends in the basal area in other
Fraus species. Additionally, this new species has two anal veins in the hindwing,
whereas other Fraus species only have a single anal vein. Some Fraus species
such as F. nanus have 2A greatly reduced in the forewing, while others lack it
entirely (Nielsen and Kristensen 1989: Figs 61, 62), this new species belongs
to the latter group. Vein Sc in the forewing is very nely stalked and S1 does not
extend to the wing margin. Although other Fraus spp. do not have Sc divided,
other genera are known to have species with both typical and divided conditions
such as Endoclita Felder (Zhu et al. 2004).
Male Fraus generally have bipectinate antennae with long, narrow rami except
for the Western Australian F. pilosa, which can be loosely termed ‘unipectinate’
in that it only has a single, broadly rounded lobe circumscribing the antennal
agellomeres. Fraus extremapodus is further unique within Fraus as the antennae
lack rami. However, it is not unusual within Hepialidae for extreme infrageneric
variation to exist in male antennae structure. Abantiades Herrich-Schäer for
example has species with short, singular rami such as A. aphenges (Turner),
bipectinate species such as A. mysteriella Simonsen, and several species with
complex, tri-forked rami such as in A. atripalpis (Walker) (see Simonsen, 2018
for further comparisons). There is also great infrageneric variation in antennae
structure within the large genus Oxycanus Walker, with species with bipectinate
(O. silvanus Tindale) to quadripectinate (O. byrsa (Ptzner)) structure, though
this may be problematic as the monophyly of Oxycanus is yet to be tested (Beaver
et al. 2020). Nielsen and Kristensen (1989) list Dalaca Walker and Callipielus
48 Australian Entomologist

Butler as genera with species that exhibit both liform and bipectinate conditions,
and noted that other genera with the dual condition were known to them.
The legs are noteworthy, when compared to forewing length this species has the
longest legs of any Australian hepialid. By comparing the midleg, which is usually
the longest leg, and the forewing, the males of other Fraus species generally have
a leg-to-wing ratio of 0.62–0.80. For example: F. simulans: 0.72, F. pteromela:
0.67, F. polyspila: 0.80, F. basidispina: 0.62, while in F. extremapodus it is
0.91. For comparison of other Australian Hepialidae s. str. genera: Abantiades
argentata: 0.53, Aenetus lewinii: 0.68, Zelotypia stacyi 0.35, Oncopera intricata:
0.58, Jeana delicatula: 0.66, Elhamma australasiae: 0.55, Oxycanus australis:
0.50. Archaeoaenetus Simonsen could not be examined but is expected to be
similar to that of Aenetus.
The lack of a distinct (by hepialoid standards) proboscis vestige suggests that F.
extremapodus may be close to the simulans group of species, which is the only
species group of Fraus to lack the proboscis. As previously noted, F. extremapodus
is similar to F. basidispina in the biloba group mainly by way of male genitalia
where both species appear closer to each other (regarding valvae and general
pseudotegumen shape) than to F. biloba, however that species and F. basidispina
are both bipectinate and have a developed tibial epiphysis. The absence of a
distinct proboscis vestige, shortened apical palpomere, and the absence of an
epiphysis on the foretibia of F. extremapodus would suggest this species is close to
or within the simulans s. str. group sensu Nielsen and Kristensen (1989), however
the dierences in wing scale structure, antennae, and wing venation presents a
serious diculty with assigning this species to any of the informal species groups
suggested in that publication and as such this new species is excluded from such
grouping.
Acknowledgements
We thank Marianne Horak, Ted Edwards and You Ning Su (Canberra, ACT) for
Figs 22–23. (22) Map of Western Australia; (23) close-up of the south-western corner with
the distribution records of Fraus extremapodus sp. n.
22 23
47 (1): 39–50 (2020) 49

enabling examination of specimens in the ANIC, and Kees Green (Melbourne,
Victoria) for examination of the specimens in his private collection. Ben Parslow
(SAM, Adelaide) is thanked for his kind assistance with imaging techniques, and
John Grehan (McGuire Center, Florida, USA) for helpful discourse in relation
to this genus. Some material examined for this study was collected under DEW
scientic licence no. Y26844-2 to EPB. Travel for this research was funded
by an Australian Lepidoptera Research Endowment grant to the authors; and a
Lirrabenda grant to MDM in 2017. We are thankful to Thomas Simonsen and
Max Moulds for critically reviewing the manuscript.
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