Corresponding author: Helena Shaverdo (
Academic editor: Mariano Michat
Nine new species groups of
Shaverdo H, Surbakti S, Warikar EL, Sagata K, Balke M (2019) Nine new species groups, 15 new species, and one new subspecies of New Guinea diving beetles of the genus
This paper introduces nine new species groups of
The present work is based on material from the following collections:
Our methods follow those described in detail in our previous articles (
The keys are based mostly on the male characters. In many cases, females cannot be assigned to species due to similarity of their external and internal structures (for female genitalia see
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Here, we provide general diagnostic characters for all representatives of the groups, which can be used to separate them from some of the previously studied groups. To complete diagnoses, special diagnostic characters for each group, mainly based on shape of the median lobe and shape and setation of the parameres, are provided below, before the species treatments.
– beetles small or medium-sized (
– habitus elongate to oval, in most species oblong-oval; with rounded pronotal and elytral sides, body outline continuous;
– pronotum short, trapezoidal, with posterior angles not drawn backwards;
– pronotum and elytra without striae or strioles;
– antennomeres not modified, simple;
– male protarsomeres 1–3 not expanded laterally;
– male protarsomere 4 cylindrical, narrow, with a large, hook-like to thin, long, slightly curved anterolateral seta;
– male protarsomere 5 not modified, long and narrow, sometimes slightly concave ventrally;
– median lobe of aedeagus with continuous outline in ventral and lateral view;
– ventral sclerite of median lobe more or less deeply divided apically.
All treated species groups (except for the monotypic
This group is characterised by extremely fine, inconspicuous dorsal punctation, pronotum with distinct lateral bead; median lobe of aedeagus without setation, simple, with rounded apex in ventral view; apexes of ventral sclerites of median lobe almost equal; paramere with distinct dorsal notch and large, long subdistal part with numerous strong setae, proximal setae more or distinct.
Indonesia: Papua Province: Pegunungan Bintang Regency, Aipomek,
Habitus and colouration
For complete description, see
The species shows variability within and between populations in shape of the apex of the median lobe, which can be shorter or more elongate (Figs
In the area of its distribution,
Indonesia: Papua Province: Pegunungan Bintang Regency and Papua New Guinea: Sandaun Province (Fig.
This group is characterised by relatively dense and coarse dorsal punctation; pronotum with distinct lateral bead; median lobe of aedeagus without setation, with apex thick, short, pointed and strongly curved downwards in lateral view; apexes of ventral sclerites of median lobe almost equal; paramere with distinct notch on dorsal side, subdistal part relatively large, rounded, with dense and strong setae, proximal setae inconspicuous.
Papua New Guinea: Hela Province, Hedamali, ca.
The species can be distinguished from the species co-occurring in the same area (
Papua New Guinea: Hela Province, Koroba area (Fig.
The species is named after Koroba Village. The name is a noun in the nominative singular standing in apposition.
This group is characterised by fine and sparse dorsal punctation; pronotum without lateral bead; median lobe of aedeagus without setation, simple; in lateral view, apex thick, short and slightly curved downwards, its minuscule tip curved upwards; apexes of ventral sclerites of median lobe slightly unequal: left one slightly longer that right one; paramere without dorsal notch, evenly tapering to distal part, with numerous small spines and without long setae.
Papua New Guinea: Sandaun Province, Ofektaman,
From most species co-occurring in the same area (
Papua New Guinea: Sandaun Province (Fig.
The species is named after Mekil Village where most specimens of the species were found. The name is an adjective in the nominative singular.
This group is characterised by fine and sparse dorsal punctation; pronotum with narrow lateral bead; median lobe of aedeagus without setation, evenly curved, rather thin, lateral margins thickened proximally; in lateral view, its apex elongate, slightly thickened and rounded, in ventral view, median lobe broad proximally and distinctly narrowed in distal half, its apex bluntly pointed; apexes of ventral sclerites of median lobe almost equal; paramere slightly concave on dorsal side, its subdistal part with dense, strong setae, proximal setae weaker, less distinct.
Papua New Guinea: Morobe Province, Garaina,
From the species co-occurring in the same area (
Papua New Guinea: Morobe Province (Fig.
Map of the eastern part of New Guinea showing distribution of the species of the monotypic groups.
The species is named after Morobe Province, the only province of
The representatives of this group are characterised by fine to coarse dorsal punctation; pronotum with distinct lateral bead; median lobe of aedeagus without setation, simple, broadly pointed; apexes of ventral sclerites of median lobe almost equal; paramere evenly tapering to apex, proximal setae often longer and more distinct that subdistal.
Papua New Guinea: Madang Province, Akameku - Brahmin, Bismarck Range,
From the species co-occurring in the same area (from
Papua New Guinea: Madang Province, Bismarck Range (Fig.
The species is named after Akameku Village. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Madang Province, Finisterre Range, Damanti,
For complete description, see
The species shows variability in size, colouration, how strongly impressed dorsal punctation and, more seldom, microreticulation, and slightly in shape of the apex of the median lobe (Fig.
From the species co-occurring in the same area (
Papua New Guinea: Madang, Simbu, Eastern Highlands, Morobe and Gulf Provinces (Fig.
Papua New Guinea: Central Province, Woitape,
Colouration of the specimens from Woitape distinctly paler, yellowish; the specimen form Wagau much darker, brown.
From the nominotypical subspecies, it can be distinguished by shinier dorsal surface, shorter setae of male protarsomere 4, and by apex of the median lone elongate, thinner, with slightly enlarged tip. The further study is necessary to confirm the status of this taxon, which seems to replace the nominotypical subspecies in the Papuan Peninsula.
Papua New Guinea: Morobe and Central Provinces (Fig.
The subspecies is named after Herzog Mts., where the subspecies was the first time discovered. The name is an adjective in the nominative singular.
Indonesia: Papua Province: Pegunungan Bintang Regency, Borme, ca.
1 male “IRIAN JAYA Zentralmassive
For complete description, see
The species can be distinguished from the species co-occurring in the same area (
Indonesia: Papua Province: Pegunungan Bintang Regency, Borme (Fig.
Papua New Guinea: Morobe Province, Aseki, Oiwa (a village about 100 km to the west of Bulolo),
From the species co-occurring in the same area (from
Papua New Guinea: Morobe Province (Fig.
The species is named after Oiwa Village. The name is a noun in the nominative singular standing in apposition.
Indonesia: Papua Province, Pegunungan Bintang Regency, south from Ok Sibil, tributary Digul River
The species is very similar to
Indonesia: Papua Province, Pegunungan Bintang Regency, Ok Sibil area (Fig.
The species is named after Ok Sibil River. The name is an adjective in the nominative singular.
1 | Beetle dorsally submatt, with strong and dense punctation and strongly impressed microreticulation (Fig. |
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– | Beetle dorsally shiny, often with very weak punctation, invisible on elytra, and weakly impressed microreticulation |
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2 | Anterolateral seta of male protarsomere 4 hook-like, large, strongly curved (Fig. |
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– | Anterolateral seta of male protarsomere 4 thin, long, slightly curved, equal to or smaller than more laterally situated large setae |
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3 | Median lobe with subparallel sides and short apex in ventral view; in lateral view, apex elongate. Setae of paramere uniform, distinct (Fig. |
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– | Median lobe more or less evenly tapering to apex in ventral and lateral views. Proximal setae of paramere usually longer, sometimes also much stronger, than subdistal |
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4 | Beetle usually smaller, |
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– | Beetle usually larger, |
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5 | Beetle with stronger elytral microreticulation and more distinct punctation, duller (Fig. |
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– | Beetle with weaker elytral microreticulation and usually invisible elytral punctation, shinier (Fig. |
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Habitus and colouration
Map of the eastern part of New Guinea showing distribution of the species of the
This group is characterised by fine to coarse dorsal punctation; pronotum with distinct lateral bead; median lobe of aedeagus without setation; in ventral view, with distinctly concave apex forming two apical lobes; in lateral view, apex tip prolongated into characteristic “nose”; apexes of ventral sclerites of median lobe almost equal or slightly unequal; paramere without distinct notch but slightly concave on dorsal side, its subdistal part with dense, strong setae, proximal setae inconspicuous.
Papua New Guinea: Morobe Province, Aseki, Oiwa, ca.
From the species co-occurring in the same area (from
Papua New Guinea: Morobe Province (Fig.
The species is named after Aseki Village. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Morobe Province, Herzog Range, Wagau (Vagau), ca.
For complete description, see
In the area of its distribution,
Papua New Guinea: Eastern Highlands and Morobe Provinces (Fig.
Papua New Guinea: Central Province, Kailaki,
Shape of apex of the medial lobe varies. In some specimens, especially from Myola, it is not clearly truncate in lateral view but very slightly concave and, due to that, the “nose” is more distinct.
Papua New Guinea: Central Province (Fig.
The species is named after Kailaki Village. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Central Province, Kokoda Track,
Papua New Guinea: Central Province (Fig.
The species was mistaken for
1 | Beetle dorsally submatt, with strong and dense punctation and strongly impressed microreticulation (Fig. |
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– | Beetle dorsally shiny, with very weak punctation, often invisible on elytra, and weakly impressed microreticulation (e.g., Fig. |
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2 | Anterolateral seta of male protarsomere 4 thin, slightly curved, equal to or smaller than more laterally situated large setae (Fig. |
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– | Anterolateral seta of male protarsomere 4 hook-like, large, strongly curved |
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3 | Apical lobes of median lobe weak, truncate in lateral view, “nose” indistinct (Fig. |
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– | Apical lobes of median lobe strong, rounded in lateral view, “nose” distinct (Fig. |
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Habitus and colouration
Map of the eastern part of New Guinea showing distribution of the species of the
This group is characterised by fine and sparse dorsal punctation; pronotum with very narrow lateral bead; median lobe of aedeagus with or without setation, very broad, robust, with sides strongly thickened; in ventral view, almost parallel-sided, with slight median constriction; apexes of ventral sclerites of median lobe very unequal: right one much longer than left one; paramere slightly concave on dorsal side and with long and dense subdistal and inconspicuous proximal setae.
Papua New Guinea: Madang Province, Baiteta,
For complete description, see
In the area of its distribution,
Papua New Guinea: Madang and Eastern Highlands Provinces. The species is known from numerous specimens from the central and wertern part of Madang and from northern part of Eastern Highlands (Fig.
Papua New Guinea: Central Province, Mafulu, ca.
The specimen from Telefomin (Sandaun Province), 1 male “PAPUA, Selminumtem [Selminum Tem, 45 km SWS Telefomin, ca. 5°S; 141°15'E], W.Sepik d. P.Beron leg.”, “
Papua New Guinea: Central and National Capital District Provinces (Fig.
Papua New Guinea: Simbu Province, Crater Mountain, Haia, ca.
Papua New Guinea: Simbu and Eastern Highlands Provinces (Fig.
The species is named after Wara Hulene Village where one of the paratype was collected. The name is an adjective in the nominative singular.
1 | Median lobe robust, lateral sides strongly thickened and apex much broader in lateral view and deeply concave in ventral view (Fig. |
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– | Median lobe slender, lateral sides more weakly thickened and apex thinner in lateral view and slightly concave in ventral view (e.g., Fig. |
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2 | Beetle brownish, usually smaller, |
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– | Beetle piceous, usually larger, |
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Habitus and colouration
This group is characterised by more or less coarse and dense dorsal punctation; pronotum with narrow lateral bead; median lobe of aedeagus without setation, broad, robust, sides slightly thickened; in ventral view, it broadened medially or subdistally; apexes of ventral sclerites of median lobe almost equal; paramere with distinct dorsal notch and subdistal part well developed, with long and dense subdistal and inconspicuous proximal setae.
Papua New Guinea: Sandaun Province, Mianmin,
In the area of its distribution,
Papua New Guinea: Sandaun Province (Fig.
The species is named after Mianmin Village. The name is an adjective in the nominative singular.
Indonesia: Papua Province: Pegunungan Bintang Regency, Bime, ca.
2 females “IRIAN JAYA: 11.9.1993 Bime – Calab Gebiet, Bime, 1400m”, “leg. M. Balke (12) ca.
For complete description, see
In the area of its distribution,
Indonesia: Papua Province: Pegunungan Bintang Regency. The species is known only from the type material, i.e., Borme – Bime – Emdoman area (Fig.
1 | Beetle smaller, |
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– | Beetle larger, |
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Map of the eastern part of New Guinea showing distribution of the species of the
This group is characterised by extremely fine and sparse dorsal punctation; pronotum with distinct lateral bead; median lobe of aedeagus simple; in lateral view, slightly or more strongly curved, apex slightly curved downwards and bluntly pointed; in ventral view, apex bluntly pointed and often twisted sidewards; apexes of ventral sclerites of median lobe almost equal; paramere slightly concave on dorsal side, subdistal setae strong and dense, proximal setae usually inconspicuous.
Papua New Guinea: Simbu Province, between Supa and Haia Villages (Airstrips), ca.
See under
Papua New Guinea: Simbu Province, Crater Mountain area (Fig.
Papua New Guinea: Morobe Province, E Pindiu, Kobau, ca.
Papua New Guinea: Morobe Province (Fig.
The species is named after Kobau Village. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Central Province, Moroka, Kailaki, Wareaga,
From species of the
Papua New Guinea: Central Province (Fig.
The species name derives from Latin
Papua New Guinea: Simbu Province, between Supa and Haia Villages (Airstrips), ca.
See for
From the species co-occurring in the same area (
Papua New Guinea: Simbu and Eastern Highlands Provinces, Crater Mountain area (Fig.
The species is named after Haia Village. The name is a noun in the nominative singular standing in apposition.
1 | Beetle larger, |
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– | Beetle smaller, |
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2 | Beetle colourful, with reddish head and bicoloured elytra: yellowish at shoulders and brownish distally; smaller, |
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– | Beetle dark brown, with paler sides of pronotum and head anteriorly; larger, |
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3 | Apex of median lobe shorter and thicker, with right lateral margin slightly concave (Fig. |
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– | Apex of median lobe longer and thinner, its lateral margins straight (Fig. |
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Habitus and colouration
Map of the eastern part of New Guinea showing distribution of the species of the
We are grateful to Dr H. Schillhammer (Vienna) for the habitus photographs and to Prof. D. Bilton and C.R. Turner (Plymouth) for a linguistic review of earlier versions of the manuscript. Fieldwork was supported by the UK Darwin Initiative project “Training the next generation of
Specimens were exported with permission from the Department of Environment and Conservation (DEC, Port Moresby), or loaned from
Financial support for the study was also provided by the FWF (Fonds zur Förderung der wissenschaftlichen Forschung – the Austrian Science Fund) through a project P 24312-B17 to Helena Shaverdo. Michael Balke was supported by the German Science Foundation (DFG BA2152/11-1, 11-2, 19-1, 19-2).