Undarobius, a new genus of cavernicolous weevils (Curculionidae: Entiminae: Leptopiini) from the Undara Lava Caves in north-eastern Australia, with an overview of anophthalmic and microphthalmic Australian Curculionidae

Abstract

Undarobius gen. n., a new genus of cavernicolous weevils with two new species, U. howarthi sp. n. and U. irvini sp. n., is described from the Undara Lava Cave system in north-eastern Queensland, Australia. These are the first cavernicolous weevils to be described from Australia, and U. howarthi is a new addition to the rich arthropod fauna of Bayliss Cave. Undarobius weevils are relatively large in size (4.0–5.5 mm long), anophthalmic and apterous with a robust, flattened body and long legs. The genus has affinities with Leptopiini, but its placement in the tribe is uncertain. We also provide a list of the known anophthalmic and microphthalmic weevils in Australia, spanning 65 species classified in 20 genera, eight tribes and about seven subfamilies and found in diverse hypogean habitats, mainly leaf litter but also soil, beach sand, subterranean aquifers and mosses.

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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by R. Anderson: 27 Jul. 2021; published: 18 Aug. 2021 207
Zootaxa 5023 (2): 207–222
https://www.mapress.com/j/zt/
Copyright © 2021 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5023.2.2
http://zoobank.org/urn:lsid:zoobank.org:pub:32EA32BC-0E25-4BF9-86CB-A5C881CB5BCD
Undarobius, a new genus of cavernicolous weevils (Curculionidae: Entiminae:
Leptopiini) from the Undara Lava Caves in north-eastern Australia, with an
overview of anophthalmic and microphthalmic Australian Curculionidae
HERMES E. ESCALONA1* & ROLF G. OBERPRIELER1,2
1Australian National Insect Collection, CSIRO, G.P.O. Box 1700, Canberra, ACT 2601, Australia.
2
�
Rolf.Oberprieler@csiro.au; https://orcid.org/0000-0002-1837-580X
*Corresponding author.
�
hermes.escalona@csiro.au; https://orcid.org/0000-0003-3487-9474
Abstract
Undarobius gen. n., a new genus of cavernicolous weevils with two new species, U. howarthi sp. n. and U. irvini sp. n.,
is described from the Undara Lava Cave system in north-eastern Queensland, Australia. These are the first cavernicolous
weevils to be described from Australia, and U. howarthi is a new addition to the rich arthropod fauna of Bayliss Cave.
Undarobius weevils are relatively large in size (4.0–5.5 mm long), anophthalmic and apterous with a robust, flattened
body and long legs. The genus has affinities with Leptopiini, but its placement in the tribe is uncertain. We also provide
a list of the known anophthalmic and microphthalmic weevils in Australia, spanning 65 species classified in 20 genera,
eight tribes and about seven subfamilies and found in diverse hypogean habitats, mainly leaf litter but also soil, beach
sand, subterranean aquifers and mosses.
Key words: Curculionidae, Entiminae, Leptopiini
Introduction
The Undara Lava Cave system in north-eastern Queensland is a natural wonder of Australia and epitomises the
complex geological and biological past of the continental landscape. The tube system was formed about 190000
years ago by activity of the Undara Volcano. As the vast lava flow that erupted from the volcano cooled and
solidified, it formed at least 60 tubes and caves (Webb et al. 1982, Atkinson & Atkinson 1995, Stephenson et al.
1998, Whitehead 2010), which are now marked above ground by lush green vegetation contrasting with the open
eucalyptus forest of the area (Fig. 37).
The longest lava tube of this system, the 1.3 km long Bayliss Cave, has one of the highest diversity of
apparent troglobites known in the world (ca. 24 species), despite its high levels of carbon dioxide. The Bayliss
Cave invertebrate fauna includes eyeless cockroaches, planthoppers, beetles, spiders and myriapods, among others
(Howarth 1988, Howarth & Stone 1990, Culver & Sket 2000, Stone 2010, Stone et al. 2012). One of these taxa
is a peculiar weevil, which has been recorded previously (Howarth 1988, Howarth & Stone 1990, Clarke 2010,
Stone 2010, Stone et al. 2012) but remained undescribed. Another closely similar species has been found in the
smaller Taylor Cave (Howarth 1988, Clarke 2010, Stone 2010) and probably yet another in Barkers Cave (Clarke
2010). The species from Bayliss and Taylor Caves were collected in 1985 and 1986 and reported in the literature
as ‘Rhytirhininae n. gen. sp. 1 and sp. 2’, respectively, as then identified by E. C. Zimmerman (Howarth 1988), but
the specimens from Barkers Cave, found in 1989, were just recorded as ‘undetermined weevils’ (Clarke 2010). No
further weevils appear to have been found in the Undara Lava Cave system since. Our efforts to trace the location
of the Barkers Cave specimens have so far been unsuccessful, and we therefore here describe only the genus and the
two species from Bayliss and Taylor Caves.
Although these are the first Australian cavernicolous weevils to be described, they are not the first to be found,
as a single fragmented specimen (missing all legs and the antennal funicles) of a different eyeless genus and species

ESCALONA & OBERPRIELER
208 · Zootaxa 5023 (2) © 2021 Magnolia Press
was recovered from Kelly Hill Cave on Kangaroo Island in 1982 (Table 1, Fig. 6). A weevil has also been reported
from the karst Bullita Cave system in the Northern Territory (Moulds & Bannink 2012), but, having been found in the
Entrance Zone of the Dingo Cave, it was deemed to be a non- troglobitic species and may not even be a cavernicole.
In the last two decades, several taxa of small anophthalmic weevils have been collected from subterranean calcrete
aquifers in the Pilbara and Yilgarn regions of Western Australia, one of them illustrated by Guzik et al. (2010), and
numerous other anophthalmic and microphthalmic weevils have been extracted from soil, leaf litter and similar
substrates around the country in recent times. We here present a brief overview of this fauna (Table 1) and propose
a concept of microphthalmy that should aid a more consistent assessment of eye reduction in weevils as adaptation
to life in conditions of low light intensity, including caves.
Material and methods
General morphological terms and descriptions follow Marvaldi et al. (2014) and Oberprieler & Zimmerman (2020).
The classification scheme follows Pullen et al. (2014). Selected specimens were macerated in KOH, dissected
and stained with chlorazol black, rinsed in water and temporarily stored in glycerine on slides. Photographs were
taken using a BK Lab Imaging system, , and a Leica M205 digital camera attached to a microscope and edited with
Adobe® Photoshop®. Body length is measured from the anterior margin of the pronotum to the apex of the elytra in
lateral view. Label data of the types are cited verbatim, with a slash (/) denoting different lines on a label and a double
slash (//) different labels on the pin of a specimen. Codens used for specimen repositories are: ANIC—Australian
National Insect Collection, CSIRO, Canberra, Australia; QMB—Queensland Museum, Brisbane, Australia.
Taxonomy
Undarobius Escalona & Oberprieler gen. n.
(Figs. 9–36)
http://zoobank.org/urn:lsid:zoobank.org:act:442F6299-8BD7-4FB4-8DF4-B9EFF52B5EED
Type species, by present designation: Undarobius howarthi Escalona & Oberprieler sp. n.
Diagnosis. Body longer than wide, length 4.0–5.5 mm, somewhat flattened, covered with sparse to dense, appressed,
fluted squamae and sparse, short, erect to semi-erect setae. Eyes absent (Fig. 19). Rostrum dorsally not separated
from head by groove (sulcus); frons indistinct; epistome short, apical margin bisinuate (Fig. 21). Antennal scrobes
broadly open and poorly defined (Fig. 19). Scapes long, reaching apical third of prothorax in repose. Prothorax with
anterior margin laterally without ocular lobes or vibrissae. Wingless. Tibiae mucronate; metatibiae without corbels
or bevels. Tarsal claws free.
Description. General appearance. Body longer than wide, length 4.0–5.5 mm, compact, robust, broad,
flattened, surface without ridges, tubercles or spines (Figs. 9–13). Rostrum and legs relatively long. Integument pale
ferrugineous (reddish-brown) to dark brown, sparsely to densely covered with appressed, pale yellowish-brown,
fluted squamae (scales) and sparse, short, erect to suberect, truncate or acute setae.
Head. Rostrum (Fig. 19) ca. 3 × longer than head, oval in cross-section. Epistome slightly projecting, short,
apex asymmetrically emarginate medially, base with two epistomal setae. Frons (Fig. 19) indistinct, slightly domed,
triangular, surface with microsculpture near apex, otherwise smooth. Epifrons (Fig. 21) with indistinct but deep
median groove in anterior third (between antennal sockets), asulcate in basal two thirds. Antennal sockets (Figs. 21)
large, not reaching rostral apex, closed laterally and open dorsally. Scrobes (Fig. 19) lateral, shallow, broad, short
(not extending to base of rostrum), borders poorly demarcated. Eyes absent, location marked by small glabrous area
(Fig. 19). Antennae (Fig. 22) long, inserted near rostral apex; scapes slightly shorter than funicle and club; funicles
7-segmented, segment 1 shorter than 2, expanded apicad, 3–7 subequal, moniliform. Mouthparts imperfectly
adelognathous, with maxillary stipites exposed next to base of prementum (Fig. 20); mandibles (Fig. 15) robust,
externally paucisetose, with four long and two smaller setae, small subtriangular scar of deciduous cusp present in
ventral position; maxillae (Fig. 16) with single apical lobe and 3-segmented palpus, segments 1 and 2 transverse,

UNDAROBIUS, A NEW GENUS OF CAVERNICOLOUS WEEVILS Zootaxa 5023 (2) © 2021 Magnolia Press · 209
2 half as long as 1, 3 conical and surrounded by digitiform sensilla; prementum (Figs. 14, 20) slightly enlarged,
subhexagonal, flat, asetose, labial palpi 2-segmented, ligula absent.
Thorax. Prothorax (Fig. 24) about as long as wide, narrowing apicad from apical third, apically slightly
constricted, forming a slight ring ventrally, anterolateral margins without ocular lobe or fringe of fine setae;
pronotum slightly convex. Procoxal cavities (Fig. 23) circular, medially contiguous, broadly interrupting prosternal
process. Mesocoxal cavities circular (Fig. 26), laterally closed by meso- and metaventrites, separated by about 0.7
× their width. Mesanepisterna and mesepimera fused but suture between them distinct. Elytra slightly longer than
their joint width, medially fused but suture visible, fine, sides evenly rounded, with 10 complete striae of small,
widely spaced punctures; interstriae flat, with single row of sparse, erect, spatulate setae. Metanepisterna fused
to metaventrite but metanepisternal suture present as deep groove through most of their length; metepimera not
exposed. Scutellar shield small, triangular, slightly convex, denuded. Wingless. Legs. Coxae with conspicuous,
long stiff seta directed ventrad; procoxae (Fig. 23) globular, medially contiguous. Trochanters obliquely truncate,
without single long seta. Femora (Figs. 28–30) slender, subcylindrical, conspicuously inflated in distal third. Tibiae
long, terete, subcylindrical; protibiae (Fig. 28) slightly bent inwards apically, meso- and metatibiae (Figs. 29–30)
straight; tibial apex with small mucro surrounded by scattered setae, without spurs, without bevel or corbel; tarsi
with segment 1 longer than wide, enlarged apicad, 2 moniliform, 3 deeply lobed, claws simple, free, divergent.
Abdomen. Ventrites (Fig. 27) jointly subtriangular, 1 and 2 connate but suture between them distinct, arcuate,
3–5 free; 1 in middle ca. 1.6 × longer than 2, with broad, subtruncate intercoxal process, 2 ca. 2 × longer than
3 and 4, these subequal in length, 5 longer than 3 and 4; sexual dimorphism slight, in male ventrites 1 and 2
less convex and 5 shorter, more broadly rounded. Male terminalia. Aedeagus (Figs. 31–32) of pedal type; penis
tubular, narrowing apicad, apex roundly acuminate, temones (apodemes) shorter than body, attached to its base
in dorsal position; tegmen without parameres; endophallus without flagellum or internal sclerite; sternite VIII
with hemisternites forming one plate, sternite IX with basal plate divided and subtriangular, with long apodeme
(spiculum gastrale). Female terminalia. Ovipositor (Fig. 34) short, weakly sclerotised, without baculi (rods); distal
gonocoxites cylindrical, with apical third setose, apex with short setose stylus; spermatheca (Fig. 35) sclerotised,
hook-shaped, without distinct nodulus, ramus or collum, spermathecal gland sac-like, about as long as spermatheca,
duct short; sternite VIII (Fig. 33) with broad, spatulate plate and long thin spiculum ventrale.
Derivation of name. The name of the genus is derived from the locality of the two known species, the Undara
Volcanic National Park, and the Greek noun bios, meaning life; its gender is masculine. The name Undara is an
Ewamian Aboriginal word meaning ‘long way’.
Comments. Elwood Zimmerman (1912–2004) deemed these weevils to be a new genus of the former subfamily
‘Rhytirhininae’ (Howarth 1988, Howarth & Stone 1990, Stone 2010, Stone et al. 2012), but their possession of
deciduous mandibular cusps (the break planes clearly visible as flat scars) and adelognathous mouthparts (albeit
imperfectly) demonstrates that they belong in the subfamily Entiminae (sensu Marvaldi et al. 2014) rather than
Cyclominae (sensu Oberprieler 2010, 2014), in which the Rhythirrinini are now included as a tribe. In the key to
the tribes of Australian Entiminae of Oberprieler & Zimmerman (2020), Undarobius runs to the large and poorly
constituted tribe Leptopiini, but it shares no obvious characters with any known leptopiine genus in Australia and
appears to occupy an isolated position in this tribe. No other anophthalmic leptopiines are known from Australia,
although reduced eyes occur in the small genus Howeocis Lea (a relative of the large terricolous genus Mandalotus
Erichson) and in several other similar leaf-litter species. Undarobius currently comprises two species, but others
may yet be discovered in other caves of the Undara Lava Cave system and perhaps in other cave systems in northern
Queensland.
Undarobius has a most unusual shape for a cavernicolous weevil, being broad and flat with long legs, almost
spider-like. Other anophthalmic cave weevils, in particular entimines such as the various Palaearctic Otiorhynchini
(see, e.g., Osella & Zuppa 1998, Hlavac 2011, Hlavac & Skuhrovec 2016, Bello et al. 2021) and Laparocerini (e.g.,
Machado 2011a–b, Machado et al. 2017) are usually shaped like their epigean relatives or narrow and elongate with
normal-sized legs. The peculiar shape of Undarobius suggests a specialised life style, possibly of crawling along
the walls and even roofs of caves or among the root tresses hanging from the cave roofs, but no observations of the
habits of the species are available so far.

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210 · Zootaxa 5023 (2) © 2021 Magnolia Press
FIGURES 1–8. Selected Australian anophthalmic and microphthalmic Curculionidae. 1—Hexonomys reginalis Kuschel (SL
0.9 mm); 2—Myrtonymus moorei Kuschel (SL 0.9 mm); 3—Notiomimetes pascoei Wollaston (SL 1.3 mm); 4—Wollastonicis
minutus Lea (SL 1.6 mm); 5—Tasmanica sp. (SL 1.7 mm, Tas, Kettering); 6—Listroderini undetermined genus (SL 3.8 mm, SA,
Kelly Hill Cave, Kangaroo Island, ANIC); 7—Cryptorhynchini undetermined genus (SL 2.7 mm, WA, Mt. Trafalgar, ANIC);
8—Halorhynchus caecus Wollaston (SL 1.6 mm).
Key to the species of Undarobius
1. Elytral setae short, apically widened and truncate; spermatheca as in Fig. 35 . . . . . . . . . . . . . . . . . . . . . . . . U. howarthi sp. n.
- Elytral setae longer, apically narrow and acute; spermatheca as in Fig. 36 ............................. U. irvini sp. n.

UNDAROBIUS, A NEW GENUS OF CAVERNICOLOUS WEEVILS Zootaxa 5023 (2) © 2021 Magnolia Press · 211
Undarobius howarthi Escalona & Oberprieler sp. n.
(Figs. 9–10, 12, 14–35)
http://zoobank.org/urn:lsid:zoobank.org:act:E53C4BEB-FA92-4141-AD93-CB3405870D4A
Diagnosis. Body length 4.0–5.5 mm. Elytral setae short, apically widened and truncate. Punctures on pronotum and
elytra fine, sparse, distance between punctures greater than puncture sizes. Body 1.5 × longer than wide, rostrum 2.1
× longer than wide. Prothorax and elytra 1 × and 1.1 × as long as wide, respectively. Spermatheca as in Fig. 35.
Description. General appearance. Body length 4.0–5.5 mm, 1.5 × longer than wide. Squamae sparse on pronotal
and elytral discs but denser laterally, easily abraded (Figs. 9–10); pronotal and elytral setae short, apically widened
and truncate. Head. Rostrum 2.1 × longer than wide and 1.6 × longer than head in lateral view (Fig. 19), oval in
cross-section. Prementum as in Fig. 20. Position of eyes indicated by small glabrous area (Fig. 19) and in cleared
specimens by small mark (probably remnant of an ommatidium). Antennae. Scapes 0.8 × shorter than funicle and
club. Funicles with segment 1 longer than 3. Thorax. Pronotum as long as wide, with sparse fine punctures, distance
between them larger than puncture size. Hypomera behind procoxae 1.5 × longer than coxal width. Elytra 1.1 × as
long as wide, disc barely convex, setae short, apically widened and truncate. Legs. Femora (Figs. 28–30) inflated in
distal third. Metatibial apex with reduced or well developed mucro. Tarsi with segment 1 shorter than 3–5. Abdomen
as in Fig. 27. Male genitalia. Penis as in Figs. 31–32. Female genitalia. Spermatheca as in Fig. 35.
Material examined (14 ex.). HOLOTYPE, ♂: QLD Undara NP / Bayliss Cave / 700m 15 Jun 1985 / F.G.
Howarth, F.D. Stone / J. Bresnan BY66 // On cornflakes bait / Cave sections The Wall & / Duckunder // HOLOTYPE
/ Undarobius howarthi / Escalona & Oberprieler 2021 (ANIC). PARATYPES: 2 ex. (1 ♀), same data as holotype
(ANIC); 6 ex. (2 ♂♂) “QLD 18º 13’S x 144º 36’E / Bayliss Cave Undara NP / 12 Jun 1986 / F. Howarth & S.
Robson” (QM); 5 ex. (1 ♂, 1 ♀) “QLD 18º 13’S x 144º 36’E / Bayliss Cave Undara NP / 31 May 1986 / F. Howarth
& D. Irwin” (QM); 1 ♂, “QLD Undara NP / Bayliss Cave / 22 May 1985 / F.D. Stone, F.G. Howarth / D. Irwin
BY8B / [dissected]” (ANIC); all labelled “PARATYPE / Undarobius howarthi / Escalona & Oberprieler 2021”.
Derivation of name. The species is named for the biospeleologist Francis G. Howarth (Hawaii Biological
Survey, Hawaii, U.S.A.), one of the discoverers of this exceptional cave inhabitant as well as of other animals of the
Undara Lava Cave system.
Comments. All specimens were collected in a deep section of Bayliss Cave known as The Wall, which is about
650 m from the entrance and part of the “stagnant-air” or “bad-air” zone as it has a high concentration of carbon
dioxide (Howarth 1988). The weevils were collected at bait put out on the cave floor, the holotype and one paratype at
cornflakes and the others at pieces of sweet potato tuber (Ipomoea batatas) (see Fig. 2D in Howarth & Stone 1990 and
Fig. 11 in Stone 2010). Bayliss Cave contains abundant organic material, mainly bat guano and root tresses (Howarth
& Stone 1990). Stone et al. (2012) considered these weevils to be part of the cave tree-root community, functioning
as primary consumers of the roots penetrating into the cave from above (identified as Brachychiton, Eucalyptus and
Ficus). As entimine larvae generally feed on roots in the soil, it is possible that those of U. howarthi do so as well,
although they may also live among the root tresses above the cave floor or even in the roof of the caves.
Undarobius irvini Escalona & Oberprieler sp. n.
(Figs. 11–13, 36)
http://zoobank.org/urn:lsid:zoobank.org:act:2BC48169-7DBE-4E82-94EB-1A72AD5116EB
Diagnosis. Body length 4.0 mm. Elytral setae longer, apically narrow and acute. Body 1.8 × longer than wide,
rostrum 1.9 × longer than wide. Prothorax and elytra 0.9 × and 1.2 × as long as wide, respectively. Spermatheca as
in Fig. 36.
Description. General appearance. Body length 4.0 mm, 1.8 × longer than wide. Squamae sparse on pronotal
and elytral discs but denser laterally; scape, pronotum and elytra with long, apically narrow and acute setae. Punctures
mostly fine, distance between them greater than puncture sizes. Head. Rostrum 1.9 × longer than wide and 1.6 ×
longer than head in lateral view, oval in cross-section. Position of eyes indicated by small glabrous area. Antennae.
Scapes 0.8 × shorter than funicle and club. Funicles with segment 1 longer than 3. Thorax. Pronotum 0.9 × as long
as wide with sparse fine punctures. Hypomera behind procoxae 1.5 × longer than coxal width. Elytra 1.2 × as long as
wide, disc barely convex, setae long, apically narrow and acute. Legs. Metatibial apex with reduced mucro, visible
at high magnification. Female genitalia. Spermatheca as in Fig. 36.

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Material examined (1 ex.). HOLOTYPE, ♀: AUSTRALIA: N E Qld / Mt. Garnet, Undara, 1000 m / 20 May
1985 / F. G. Howarth, D. Irvin // Yarramulla Sta / Taylor Cave // HOLOTYPE / Undarobius irvini / Escalona &
Oberprieler 2021 (ANIC).
FIGURES 9–16. Undarobius habitus and mouthparts. 9–10, 12—U. howarthi sp. n. (SL 3.8 mm, ♂) 9–10: dorsal, 12: lateral
aspect; 11, 13—U. irvini sp. n. (SL 3.8 mm, ♀) 11: dorsal, 13: lateral aspect; 14– 16—U. howarthi sp. n. ♀, mouthparts, 14:
labium, ventral view, 15: left mandible, dorsal view, 16: right maxilla, ventral view.

UNDAROBIUS, A NEW GENUS OF CAVERNICOLOUS WEEVILS Zootaxa 5023 (2) © 2021 Magnolia Press · 213
FIGURES 17–25. Undarobius howarthi sp. n., ♂, structural details. 17—head, dorsal view; 18—head, ventral view; 19—head,
lateral view; 20—mouthparts, ventral view; 21—apex of rostrum, dorsal view; 22—left antenna, dorsal view; 23—prothorax,
ventral view; 24—prothorax, dorsal view; 25—elytra, dorsal view.

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FIGURES 26–36. Undarobius species, structural details. 26–34 U. howarthi sp. n., 26—pterothorax and elytra, ventral view;
27—ventrites, ♂, ventral view; 28—foreleg; 29—middle leg; 30—hindleg; 31—penis, dorsal view; 32—penis, lateral view;
33—sternite VIII, ♀, ventral view; 34—ovipositor, ventral view; 35—spermatheca; 36—U. irvini sp. n., spermatheca.

UNDAROBIUS, A NEW GENUS OF CAVERNICOLOUS WEEVILS Zootaxa 5023 (2) © 2021 Magnolia Press · 215
FIGURE 37. Satellite image with panoramic view of Undara Volcano and North and Northwest Lava Tubes. Source Google
Earth, 7 April 2021.
Derivation of name. The species is named for the speleologist Douglas Irvin (Chillagoe Caving Club), collector
of the only known specimen.
Comments. This species is so far known from a single female, collected from the deep zone of Taylor Cave
(Howarth 1988). It is noticeably smaller and narrower than the females of U. howarthi, and it can also be distinguished
by its longer and sharp-pointed elytral setae. Its spermatheca is narrower and has a longer and more sharply angled
cornu, although additional specimens will need to be studied to assess its variability.
Overview of microphthalmic and anophthalmic weevils in Australia
Weevils (Curculionoidea) are the largest group of phytophagous beetles and have diversified particularly well in
vegetated ecosystems, but they have also colonised subterranean habitats such as soil, sand, humus and caves, to
which they are most notably adapted by the reduction to complete loss of their eyes. Morrone & Hlavac (2017)
listed 85 genera and 485 species of soil-dwelling weevils with reduced eyes worldwide. Seven of these genera
and eleven of the species occur in Australia, but the list is not comprehensive as many other microphthalmic and
anophthalmic taxa are known to occur in the country, both described and undescribed (see Table 1, Figs. 1–8).
Additional described microphthalmic species were omitted from the list either because their eye size has not been
adequately noted in their descriptions (e.g., given only as ‘small’, Lea 1926a 1927) or because relevant literature
was overlooked, such as Lea (1926b) for Howeocis microps (Lea) and Zimmerman (1993) for Baeosomus Broun (as
Bryocatus Broun). A few undescribed anophthalmic species have also been recorded in the literature, i.e. the ones
described herein (Howarth 1988, Howarth & Stone 1990, Clarke 2012, Stone 2010, Stone et al. 2012) as well as a
small weevil collected from a subterranean calcrete aquifer in the Pilbara region of Western Australia and illustrated
by Guzik et al. (2010). Most of these are only represented in collections and have yet to be studied in detail.
An additional problem in compiling lists of weevils with reduced eyes is that microphthalmy is never properly
defined. Reduced eye size is implicitly compared with “normal” size and deemed to involve a decrease in the number
of ommatidia, but it is never stipulated how this reduction is measured and how many remaining ommatidia denote a
weevil as being microphthalmic. Furthermore, the reduction in the number of ommatidia may not occur evenly but
only in one dimension (typically in length but not in height of the eye), and a decrease in the number of ommatidia
may be negated by an increase in the size of these ommatidia, so that an eye with a few large ommatidia may be

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216 · Zootaxa 5023 (2) © 2021 Magnolia Press
almost as large as a “normal” eye with many smaller (“normal-sized”) ommatidia (e.g., in Geochus). As a decrease
in the number of ommatidia generally appears to be an adaptation to low-light conditions, even if the ommatidia are
enlarged, we here apply a more consistent concept of microphthalmy to the assessment of eye reduction by counting
the number of ommatidia along the length of the eye, in a row (usually slanting) from the middle of the anterior eye
margin to the posterior margin (representing the greatest eye length). When the number of ommatidia in this middle
row is four or less, we deem the weevil to be microphthalmic. The number of ommatidia thus measured is usually
consistent in the species we examined, but in some species it may vary slightly (see Table 1).
In Table 1 we list all the microphthalmic and anophthalmic weevils occurring in Australia as found by us,
together with their current classification, eye length (number of ommatidia across middle), region of occurrence,
habitat and relevant references. The list spans 20 genera (9 of them undescribed) and 65 species (47 undescribed),
distributed over eight tribes and about seven subfamilies. The uncertain number of subfamilies is due to the unresolved
phylogenetic affinities of the Myrtonymini (here provisionally classified in Erirhininae) and the Notiomimetini,
whose previous inclusion in Cyclominae (Oberprieler 2010) is repudiated by the position of Aphela Pascoe in the
phylogeny estimate of Gillett et al. (2018) (a result supported by a similar placement of Aphela and also Psaldus
Pascoe in a larger phylogenomic analysis; in preparation). The phylogenetic relationships and classification of
Geochus Broun and some of the undescribed genera in Table 1 also remain to be investigated more thoroughly.
Psaldus liosomoides Pascoe, included in the list of Morrone & Hlavac (2017), has an eye length of eight ommatidia
and is therefore not a microphthalmic species in our concept of this condition. Note that undescribed genera and
species in ANIC are clearly labelled as identified in Table 1.
Of the species listed in Table 1, 25 (38 %) have no eyes (ommatidia) at all. In many cases such anophthalmy
occurs in all species of a genus, but in some genera there are a few species with one or two ommatidia remaining.
Although these species have been collected in the same habitats as their anophthalmic congeners (generally in leaf
litter), they may occupy niches of slightly higher light intensity (i.e. live in shallower or looser substrates).
The vast majority of these Australian weevils with reduced eyes occurs in habitats on or in the soil, mainly in
leaf litter but also moss cushions and soil layers just beneath the surface. Only eight of the species appear to lead
a cavernicolous life style, three of them in larger caves and the others recovered from the cavities above small
subterranean aquifers. The exploration of such aquifers in Western Australia in recent years (e.g., Humphreys 2001,
Guzik et al. 2010) has yielded many more such weevils, but they remain to be studied and classified.
Subterranean organisms are typically categorised as troglobites (or troglobionts), trogloxenes and troglophiles,
using the Schiner-Racovitza system as amended by Trajano & Carvalho (2017), which applies primarily ecological
criteria, specifically habitat restriction, to distinguish between these categories. In this system, troglobites are
species restricted to habitats of low food availability and permanent darkness and with exclusively hypogean source
populations, trogloxenes also live in habitats of low food availability but are not restricted to permanent darkness
and have source populations in both hypogean and epigean habitats, and troglophiles live in habitats of high food
availability and only visit habitats of total darkness and have source populations only in epigean habitats. Trajano
& Carvalho (2017) discussed this classification system in detail and stressed that categorisation of organisms as
troglobites, trogloxenes and troglophiles cannot be made based on isolated collecting events and simple extrapolation
of apparent morphological adaptations (such as the loss of eyes) and that, with few exceptions, it is not possible to
assign a subterranean organism to any of these categories “with an acceptable degree of confidence after a single
or a few instances of field observation, and especially without a thorough taxonomic study” (Trajano & Carvalho
2017: 13). An additional issue is that the terms troglobite, trogloxene and troglophile intrinsically refer to organisms
living in caves (from the Greek word troglos, meaning cave) and are not actually applicable to those living in more
or less dense soil among fine roots or in deep organic litter, even though they also live in subterranean (hypogean)
habitats of low light or permanent darkness and often display the same morphological adaptations as cavernicolous
species do, such as the loss of eyes and pigmentation. Similarly, cavernicolous species cannot be regarded as soil-
dwellers when they live above the ground surface in their caves.
Among the Australian weevils, these problems are well exemplified by the weevils from the subterranean
aquifers in Western Australia, which have been treated as troglobites and part of the troglofauna of the Pilbara and
Yilgarn aquifer systems (Guzik et al. 2017). It is, however, unknown whether they are restricted to the small cavities
above these aquifers or whether they occur more widely among roots in the soil above and around the aquifers.
The latter is certainly indicated for the undescribed cryptorhynchine species (Table 1), which have closely related
(possibly congeneric) species inhabiting leaf litter in the Kimberley area further north, suggesting that the aquifer

UNDAROBIUS, A NEW GENUS OF CAVERNICOLOUS WEEVILS Zootaxa 5023 (2) © 2021 Magnolia Press · 217
TABLE 1. Known species of anophthalmic and microphthalmic Curculionidae in Australia (eye length measured as number of ommatidia across middle of eye, see text for de-
tails; question mark (?) indicating uncertain family-group placement).
Species Subfamily Tribe Eye length State (region) Habitat References
Hexonymus reginalis Kuschel, 2014
(Fig. 1)
Erirhininae? Myrtonymini 0 Qld. unknown Kuschel (2014), ANIC
Myrtonymus australicus Kuschel,
2014
Erirhininae? Myrtonymini 0 N.S.W. unknown Kuschel (2014), ANIC
Myrtonymus eucalypti Kuschel, 2014 Erirhininae? Myrtonymini 0 W.A. Eucalyptus log litter Kuschel (2014), ANIC
Myrtonymus moorei Kuschel, 2014
(Fig. 2)
Erirhininae? Myrtonymini 0 N.S.W. Eucalyptus litter and
humus
Kuschel (2014), ANIC
Myrtonymus peckorum Kuschel, 2014 Erirhininae? Myrtonymini 0 W.A. Eucalyptus rotten bark Kuschel (2014), ANIC
Baeosomus uvidus (Oke, 1931) Erirhininae Tanysphyrini 4 Vic. moss, lichens, leaf litter ANIC, Oke (1931), Zim-
merman (1993)
Baeosomus spp. n. 1–3 Erirhininae Tanysphyrini 4–5 Tas. leaf litter ANIC
Baeosomus sp. n. 4 Erirhininae Tanysphyrini 4 Vic., S.A. leaf litter ANIC
Baeosomus spp. n. 5–7 Erirhininae Tanysphyrini 4–5 N.S.W. leaf litter ANIC
Baeosomus sp. n. 8 Erirhininae Tanysphyrini 4 A.C.T. leaf litter ANIC
Baeosomus spp. n. 9–10 Erirhininae Tanysphyrini 4 Qld. leaf litter, moss ANIC
Genus n. 1 nr. Baeosomus sp. n. 1 Erirhininae Tanysphyrini 3–4 N.S.W. (Lord Howe Isl.) moss ANIC
Genus n. 1 nr. Baeosomus sp. n. 2 Erirhininae Tanysphyrini 3–4 Tas. moss ANIC
Genus n. 2 nr. Baeosomus sp. n. Erirhininae Tanysphyrini 3 Tas. heath turf ANIC
Notiomimetes pascoei Wollaston,
1873 (Fig. 3)
uncertain Notiomimetini 2–3 W.A., S.A., Vic. beach under seaweed and
drift litter and rock holes
Oberprieler (2010, 2014),
ANIC
Notiomimetes sp. n. uncertain Notiomimetini 3–4 W.A., S.A. beach under seaweed and
drift litter
ANIC
Wollastonicis minutus Lea, 1909
(Fig. 4)
uncertain Notiomimetini 3 W.A. beach under seaweed and
drift litter
Oberprieler (2010, 2014),
ANIC
Wollastonicis sp. n. uncertain Notiomimetini 2 S.A. beach under seaweed ANIC
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TABLE 1. (Continued)
Species Subfamily Tribe Eye length State (region) Habitat References
Geochus howensis Lea, 1927 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) Lea (1927), Pullen et al.
(2014), ANIC
Geochus sp. n. 1 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Geochus sp. n. 2 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Geochus sp. n. 3 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Geochus sp. n. 4 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Geochus sp. n. 5 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Geochus sp. n. 6 Phrynixinae? 4 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Geochus sp. n. 7 Phrynixinae? 3 N.S.W. (Lord Howe Isl.) leaf litter (leaf mines?) ANIC
Mandalotina atranotata Oke, 1931 Cyclominae Listroderini 4–5 Vic. leaf litter, moss ANIC
Mandalotina sp. n. Cyclominae Listroderini 4 Tas. leaf litter, moss ANIC
Tasmanica myrmecophila Lea, 1900 Cyclominae Listroderini 0 Tas. leaf litter
(under stone in ant nest)
Lea (1900, 1906), ANIC
Tasmanica sp. n. 1 (Fig. 5) Cyclominae Listroderini 0 Tas. leaf litter ANIC
Tasmanica sp. n. 2 Cyclominae Listroderini 0 A.C.T., N.S.W. leaf litter ANIC
Tasmanica sp. n. 3 Cyclominae Listroderini 0 W.A. (Stirling Rg.) leaf litter ANIC
Genus n. 1 nr. Tasmanica sp. n. 1 Cyclominae Listroderini 0 N.S.W. leaf litter ANIC
Genus n. 1 nr. Tasmanica sp. n. 2 Cyclominae Listroderini 0 N.S.W. leaf litter ANIC
Genus n. 1 nr. Tasmanica sp. n. 3 Cyclominae Listroderini 0 N.S.W. leaf litter ANIC
Genus n. 1 nr. Tasmanica sp. n. 4 Cyclominae Listroderini 2 Qld. leaf litter ANIC
Genus n. 2 nr. Tasmanica sp. n. Cyclominae Listroderini 0? W.A. (Pilbara) subterranean aquifer Guzik et al. (2010, Fig. 2e)
Genus n. nr. Reyesiella sp. n. 1 Cyclominae Listroderini 2 N.S.W. leaf litter?
(Nothofagus bark)
ANIC
Genus n. nr. Reyesiella sp. n. 2 Cyclominae Listroderini 1 Tas. leaf litter ANIC
Genus n. nr. Reyesiella sp. n. 3 Cyclominae Listroderini 1 Tas. turf (Poa)ANIC
Gen. & sp. n. (Kelly Hill Cave)
(Fig. 6)
Cyclominae Listroderini 0 S.A. (Kanga- roo Isl.) cave floor?
(ca. 45 m below ground)
ANIC
......continued on the next page

UNDAROBIUS, A NEW GENUS OF CAVERNICOLOUS WEEVILS Zootaxa 5023 (2) © 2021 Magnolia Press · 219
TABLE 1. (Continued)
Species Subfamily Tribe Eye length State (region) Habitat References
Howeocis microps (Lea, 1926) Entiminae Leptopiini 4 N.S.W. (Lord Howe Isl.) leaf litter Lea (1926b), ANIC
Howeocis nodipennis (Lea, 1926) Entiminae Leptopiini 4 N.S.W. (Lord Howe Isl.) leaf litter ANIC
Howeocis setosus Lea, 1926 Entiminae Leptopiini 4 N.S.W. (Lord Howe Isl.) leaf litter Lea (1926a), ANIC
Genus n. cf. Howeocis sp. n. 1 Entiminae Leptopiini 4 Vic. leaf litter ANIC
Genus n. cf. Howeocis sp. n. 2 Entiminae Leptopiini 4 Vic. leaf litter ANIC
Genus n. cf. Howeocis sp. n. 3 Entiminae Leptopiini 4 Vic. leaf litter ANIC
Genus n. cf. Howeocis sp. n. 4 Entiminae Leptopiini 4 Qld. leaf litter ANIC
Undarobius howarthi
Escalona & Oberprieler, h.o.
Entiminae Leptopiini? 0 Qld. lava caves h.o., ANIC
Undarobius irvini
Escalona & Oberprieler, h.o.
Entiminae Leptopiini? 0 Qld. lava caves h.o., ANIC
Genus n. 1 sp. n. 1 (Fig. 7) Molytinae Cryptorhynchini? 1 W.A. (Kimberley) leaf litter ANIC
Genus n. 1 sp. n. 2 Molytinae Cryptorhynchini? 0 W.A. (Kimberley) leaf litter ANIC
Genus n. 1 sp. n. 3 Molytinae Cryptorhynchini? 0 W.A. (Kimberley) leaf litter ANIC
Genus n. 1 sp. n. 4 Molytinae Cryptorhynchini? 0 W.A. (Kimberley) leaf litter ANIC
Genus n. 2 sp. n. 1 Molytinae Cryptorhynchini? 0 W.A. (Yilgarn) subterranean aquifer ANIC
Genus n. 2 sp. n. 2 Molytinae Cryptorhynchini? 0 W.A. (Yilgarn) subterranean aquifer ANIC
Genus n. 2 sp. n. 3 Molytinae Cryptorhynchini? 0 W.A. (Yilgarn) subterranean aquifer ANIC
Genus n. 2 sp. n. 4 Molytinae Cryptorhynchini? 0 W.A. (Yilgarn) subterranean aquifer ANIC
Halorhynchus caecus Wollaston, 1873
(Fig. 8)
Cossoninae Onycholipini? 0 W.A., S.A., Vic. beach sand among roots
(of spinifex grass)
Lea (1906),
Oberprieler (2010), ANIC
Halorhynchus geniculatus Lea, 1900 Cossoninae Onycholipini? 0 S.A. beach sand among roots
(of Atriplex saltbush)
Lea (1900, 1906),
Oberprieler (2010), ANIC

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species may also occur from leaf litter on the soil surface throughout the soil column down to the aquifers (probably
along roots, in which their larvae may develop) and that they are then soil-dwellers rather than cavernicoles. As
cautioned by Trajano & Carvalho (2017), it is not feasible to categorise these species as troglobites based on single
sampling events from cavities above aquifers and without a thorough taxonomic study; if they can be considered
as troglofauna at all, they may at best be trogloxenes or troglophiles. Much more extensive sampling of soil strata
above the aquifers is needed to assess this.
The categorisation of Undarobius as troglobitic (Howarth & Stone 1990, Clarke 2010, Stone 2010) is equally
precarious in the absence of data from concerted collecting efforts throughout the Undara cave system. Although
these weevils have so far only been found in the deep zones of the caves, attracted to bait put out in specific locations,
it is not known where the adults and larvae normally live, whether they may occur in or move through other zones
as well and whether they may also (or mainly) occur among root masses above (outside of) the caves. Osella &
Zuppa (1998) posited that no weevils can be considered to be troglobitic because of their phytophagous life style
and the general absence of living plants in caves. However, in caves with permanent roots penetrating through the
roof, such as the Undara lava caves, it should be possible for rhizophagous weevils to become true troglobites. If the
Undarobius species can be shown to be restricted to the deep zones of the Undara lava caves, they would be the first
troglobitic weevils recorded from Australia and the only other troglobitic beetles besides a few species of Carabidae
recognised as such (Moore 1964, Lawrence & Ślipiński 2013). For the moment, however, it is pertinent to treat U.
howarthi and U. irvini only as troglomorphic cavernicolous species.
Acknowledgements
We sincerely thank Debbie Jennings (ANIC, CSIRO) for the preparation of the illustrations used in this paper.
Thanks to Christine Lambkin, Susan Wright and Karin Koch (QMB) for their assistance during a visit by HE and
loan of specimens. Frank Howarth (Hawaii, U.S.A.) kindly provided additional information on collection methods
and habitat. Peter Bannink (Chillagoe Caving Club) assisted with the location of caves in the Undara System.
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