Corresponding author: Lars Hendrich (
Academic editor: M. Michat
The genus
Hendrich L, Shaverdo H, Hájek J, Balke M (2019) Taxonomic revision of Australian
The genus
Most species of
The Australian
All species of Australian
During our recent work on Australasian
This study is based on the examination of 1,682 specimens. Type material of most species, except for
The material used for this study is deposited in the following institutional and private collections:
Beetles were studied with a Leica MZ 12.5 dissecting scope at 10–100×. Habitus photos were made by František Slamka (Bratislava, Slovakia) using a digital photo imaging system and incident light, composed of a Leica DM 2500 M microscope and a Tucsen 5.0 MP camera. The microscope was fitted with Leica HCX PL “Fluotar” 5× and 10× metallurgical grade lenses (Buffington and Gates 2008). Image stacks were aligned and assembled with the computer software Helicon Focus 4.77TM. Male genitalia were studied and figured in wet condition. Aedeagus images were captured by Harald Schillhammer (Vienna, Austria) with a Nikon D4 (in combination with a Novoflex bellows and a Mitutoyo 10/0.25 Apo ELWD) tethered to a PC and controlled with Nikon Camera Control Pro. Resulting image stacks were treated with Zerene Stacker and then post-processed in Adobe Photoshop CS 5.
The descriptive style follows
Abbreviations used in the text are:
Coordinates are given in decimal notation unless cited verbatim from labels. Beside various Australian road maps, we also used Google Earth (
We delineate the species using traditionally employed morphological characters such as shape, structure and setation of the male genitalia; size, shape and colour pattern of the body, shape of the male protibia, as well as features of the dorsal surface sculpture.
The sequence data partly originate from
GenBank accession numbers are
The morphologically delineated species were all retrieved as monophyletic groups in our cox1 DNA sequence tree (Fig.
Small to medium sized (4.2–8.0 mm in Australia), elongate or oblong-oval and more flattened diving beetles, narrowing towards apex. Scutellum visible. Elytra of all Australian species with longitudinal striae being numbered from innermost to outermost (except submarginal stria). Prosternum and its process in same plane or process slightly deflected upwards behind procoxae. Prosternal process short. Metepisternum extended to mesocoxal cavity. Lateral parts of metaventrite (“metasternal wings”) relatively broad at level of mesocoxa and very narrow laterally. Metacoxal processes with an apical cleft; metacoxal lines present. Metafemur ventrally without setigerous row at outer posterior angle. Tarsi with five distinct tarsomeres; male with protarsomeres 1–3 broadly dilated but not forming a round palette, and with four rows of larger adhesive discs bordered by long and thick setae. Metatarsal claws of same length. Median lobe asymmetrical.
“North Australia”.
Habitus and colouration of
Similar to male in habitus. Pro- and mesotarsomeres not broadened, without adhesive setae. Pronotal striae finer and denser.
TL = 5.1–5.25 mm; TL-H = 4.5–4.65 mm; MW = 2.45–2.50 mm.
All specimens studied are rather uniform in shape and colour, and vary only little in body length.
Endemic. The species occurs from the Kimberley region in north-western Australia and in the Arnhemland in the Northern Territory to northern Queensland. In that area, the species is mainly distributed in rainforest pockets of the stone country (Fig.
A lentic species, which occurs in isolated pools of different size in otherwise dry riverbeds of seasonal streams, creeks and rivers. The bottom is consisted of gravel, sand and a layer of rotten plant debris. Occasionally, it occurs in protected bays, at the edge of large (25–50 m width), slow flowing and shallow (up to 20 cm depth) rivers, shaded by old riverine Melaleuca trees. There, the adults can be found among floating roots and organic debris of the paperbark trees. In northern Australia, at Gunlom waterfall, a large series was collected in a small (ca. 50 cm², 10 cm depth), water filled and shaded pothole (Fig.
“Australia [Queensland] (Cape York)”.
Habitus and colouration of
Similar to male in habitus. Protibia simple, not angled basally and only slightly broadened distally; pro- and mesotarsomeres not broadened, without adhesive setae.
TL = 7.2–8.0 mm; TL-H = 6.5–7.15 mm; MW = 3.4–3.5 mm.
All specimens studied are rather uniform but can vary in body length. In approximately half of the studied specimens, stria 2 on elytra is not interrupted and reduced to a few elongated short striae. There is a slight variation in the extension of the ferruginous basal band on elytra.
The species can be separated from all other Australian
The species is widely distributed in the northern half of Australia. Records are from north-western Australia (Kimberley region), Northern Territories and Queensland south to Brisbane and Stradbroke Island (Fig.
In this study, the species is recorded for the first time from southern New Guinea (Indonesia: Papua Province, Merauke Regency and Papua New Guinea: Western Province).
A widely distributed species in tropical northern Australia,
“Australia [Queensland] (Cape York)”.
We were not able to find the type material of
Habitus and colouration of
Habitus and colouration of
Similar to male in habitus. Protibia simple, not angled basally and only slightly broadened distally; pro- and mesotarsomeres not broadened, without adhesive setae.
TL = 6.2–6.3 mm; TL-H = 5.5–5.6 mm; MW = 2.9–2.95 mm.
A dimorphic species. Despite the fact that all specimens studied are rather uniform in habitus and colouration, they vary in extension and number of their elytral striae. Several specimens of both sexes, collected at the same spot at the same time with the main form (NT, Robin Falls), have 11 fully developed elytral striae. (Fig.
On the first view
Endemic. The species is distributed from the Kimberley region in Western Australia, over Northern Territory (Melville Island, Kakadu Area and around Darwin) to coastal Queensland (Cape York Peninsula) south to Townsville (Fig.
The habitat of
“[Australia, Queensland] Lizard Island”.
ANIC, studied but the exact data not noted.
1 ex., “Tasmania, Gladstone, Dorset, A.M. Lea” (SAMA), probably mislabelled.
Habitus and colouration of
Similar to male in habitus. Pro- and mesotarsomeres not broadened, without adhesive setae. Pronotal striae fine and dense.
TL = 6.8–7.8 mm; TL-H = 6.0–6.9 mm; MW = 3.2–3.4 mm.
All specimens studied are rather uniform in shape and size but vary in the extension of the testaceous elytral markings.
The species is close to
The species is widely distributed in the northern half of Australia. Records are from the Northern Territory (inland to the East MacDonnell Ranges), north-western Australia (Kimberley region and the Pilbara), and Queensland south to Brisbane (Fig.
According to the literature,
This is a widely distributed species and one of the most common
“Australia, [Queensland], Rockhampton”.
Habitus and colouration of
Similar to male in habitus. Protibia simple, not angled basally and only slightly broadened distally; pro- and mesotarsomeres not broadened, without adhesive setae. Pronotum basolaterally with numerous short longitudinal strioles. Elytra between striae and side with numerous short longitudinal strioles.
TL = 5.7–6.0 mm; TL-H = 5.3–5.6 mm; MW = 2.7–2.8 mm.
All specimens studied are rather uniform and vary only in body length. In some specimens, the submarginal stria is reduced to few elongate grooves.
The species is close to
The species occurs in tropical and subtropical northern and central Australia, along the east coast south to Brisbane (WA, NT, QLD) (Fig.
Most specimens were collected during or just after the rainy season when the beetles can be collected in seasonal flood meadows along rivers and creeks, shallow roadside ditches and swampy areas. At that time of the year,
Moreguina (St Stephen’s Mission), Central Province, Papua New Guinea Central, 50 m,
Habitus and colouration of
Similar to male in habitus. Protibia not modified. Pro- and mesotarsomeres not broadened, without adhesive setae.
All specimens studied are rather uniform in shape and size but vary a bit in the extension of the testaceous elytral markings.
Based on the characteristic sickle-shaped median lobe, the new species belongs to a difficult complex of species distributed in the Sunda Islands and New Guinea, including
Within other Australian species with six elytral striae,
This species is dedicated in honour of our late colleague Dr Martin Baehr (*10.3.1943, †17.4.2019, Munich, Germany), coleopterist, arachnologist, and others as well as the most knowledgeable authority for Australian ground beetles. The specific epithet is a substantive in the genitive case.
Northern Queensland (Iron Range National Park at Cape York Peninsula) and south-eastern Papua New Guinea (Central Province) (Fig.
Unknown. Most probably, the new species is an inhabitant of temporary lowland rainforest pools. The type specimens were collected in a Malaise Trap and at light.
“Australia, (Champion Bay, Carpentaria, Port Denison, Port Boweii)”.
Habitus and colouration of
Similar to male in habitus. Protibia simple, not angled basally and only slightly broadened distally; pro- and mesotarsomeres not broadened, without adhesive setae.
TL = 5.5–5.8 mm; TL-H = 4.9–5.2 mm; MW = 2.5–2.6 mm.
All specimens studied are rather uniform and vary only in body length.
Endemic. The species is widely distributed in the northern half of Australia. Numerous records are from the Northern Territory, north-western Australia (Kimberley region and the Pilbara), and Queensland south to Brisbane (Fig.
This widely distributed species is the most common
This species is the most common Australian species of the genus in Australian entomological collections.
Habitus and colouration of
Similar to male in habitus. Protibia simple, not angled basally and only slightly broadened distally; pro- and mesotarsomeres not broadened, without adhesive setae.
TL = 5.0–5.7 mm; TL-H = 4.7–5.3 mm; MW = 2.7–2.85 mm.
All specimens studied are rather uniform and vary only in body length and dorsal colouration. In some specimens, sides of elytron and areas between striae often lighter. A single female from Holmes Jungle Park in Darwin is on dorsal surface almost black, matt and with coarse microreticulation and numerous strioles on elytra and pronotum.
The species occurs in the coastal tropical rainforest areas of the Northern Territory and north-eastern Queensland, from northern Peninsula (Lockerbie) to Mackay in the south (Fig.
“[Indonesia, Sumatra] Solok, District of Rawas, Soeroelangoen”.
Not studied. The type specimens should be deposited in the Naturalis Biodiversity Center, Leiden, The Netherlands (former Rijksmuseum van Natuurlijke Historie), but they were not found during a visit of JH. The designation of the lectotype by
Habitus and colouration of
Similar to male in habitus. Protibia simple, not angled basally and only slightly broadened distally; pro- and mesotarsomeres not broadened, without adhesive setae.
TL = 4.2–4.5 mm; TL-H = 3.8–4.1 mm; MW = 1.9–2.0 mm.
All Australian specimens studied are rather uniform and vary only in body length.
The species is similar to
This is the most widespread species in the Indomalayan and Australasian realms. It occurs from Nepal, India, and Sri Lanka over Myanmar, Laos, Vietnam, Thailand, Philippines, Indonesia (
The wide distribution of this species owes to the ability of adaptation to manmade habitats like rice or paddy fields and shallow irrigation ditches (
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Elytron with a submarginal stria and six or more additional discal striae |
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– | Elytron without submarginal stria, but with 11 discal striae. Body colour yellow to pale ferruginous, elytral striae sharply outlined in black (Fig. |
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Elytron with 11 striae |
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– | Elytron with 6–10 |
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Smaller species, TL = 5.1–5.25 mm. Elytron uniformly black without any testaceous basal or apical markings on elytra (Fig. |
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– | Larger species, TL = 6.2–7.8 mm. |
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Elytron with testaceous basal and apical markings (Fig. |
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– | Elytron dark brown to black (Fig. |
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Elytron with 6 striae |
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– | Elytron with more than 6 striae |
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Inner stria of elytron short, less than half length of elytron. TL = 5.7–6.0 mm. Median lobe as in Fig. |
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– | Inner stria of elytron long, at least 2/3rd the length of elytron or longer |
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Elytron dark brown to black, without any testaceous basal marking (Fig. |
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– | Elytron with at least narrow testaceous basal marking (Fig. |
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Body broad, ovoid. TL = 5.0–5.7 mm. Outer stria of elytron complete. Anterior angles of pronotum and base of elytra narrowly testaceous (Fig. |
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– | Body oblong-oval. TL = 6.2–6.75 mm. Outer stria on elytron almost complete. Anterior angles of pronotum, base and tip of elytra testaceous (Fig. |
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Elytron with 8 striae, no striae inwards from 1st row of serial punctures (Fig. |
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– | Elytron with 10 striae, inner striae partially reduced (Fig. |
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Distribution of
Distribution of
Distribution of
Distribution of
Distribution of
Distribution of
Distribution of
Maximum likelihood tree for Australian
Habitat of
Habitat of
Habitat of
Habitat of
Habitat of
Habitat of
Habitat of
We are indebted to the curators C. Reid, T. Weir, C. Taylor, O. Merkl, G. Dully, M. Jäch, C. Burwell, G. Monteith, C. Watts, D. Peggie as well as G. Wewalka for lending type material and specimens, to František Slamka (Bratislava, Slovakia) and Harald Schillhammer (Vienna, Austria) for the excellent photos of habitus and male genitalia, and to Hans Fery (Berlin, Germany) for critical reading an earlier version of the manuscript. The Department of Environment and Conservation in Western Australia is acknowledged for giving permission to conduct scientific research in the Pilbara [Permit numbers: SF 003017 and NE 002348]. The Parks and Wildlife Commission of the Northern Territory, the Australian National Parks & Wildlife Service in Darwin and the Kakadu National Park are thanked for giving permission to conduct scientific research in the Northern Territory (Permit Number: 23929 and RK- 400/ RK- 660). The Department of Environment and Conservation in New South Wales [Scientific Licence No. S12040] and the Department of Sustainability and Environment in Victoria [Research Permit No. 10003840] are acknowledged for giving permission to conduct scientific research in National and State Parks. This work was supported by grants to Michael Balke: Deutsche Forschungsgemeinschaft (BA 2152/4-1, 6-1, 7-1, 11-1 and 11-2), the CLIMAQUA project funded by the Bundesministerium für Bildung und Forschung (BMBF) (grant # 01DR14001 to M. Balke), Lars Hendrich: Deutsche Forschungsgemeinschaft (HE 5729/1-1), Jiří Hájek: Ministry of Culture of the Czech Republic (DKRVO 2019-2023/5.I.a, National Museum, 00023272), and Helena Shaverdo: Fonds zur Förderung der wissenschaftlichen Forschung – the Austrian Science Fund (P 31347-B25). Finally, Lars Hendrich warmly thanks Emma Hendrich (Munich, Germany) and Ingo Weckwerth (Berlin, Germany) for their patience, assistance and enthusiastic encouragement during several field trips.
For full synonymy, see Sheth et al. 2018: 257 and Nilsson and Hájek 2019: 59.
dimorphic species