Academic editor: Mariano Michat
Nineteen new species of
Shaverdo H, Sagata K, Balke M (2018) Introduction of the
Herein, we introduce the new species group of the genus
Including the results of this work, 125 species of
As in most of our previous papers on the genus (
The present work is based on material from the following collections:
All methods follow those described in detail in our previous articles (
Abbreviations:
Species | Distribution | |
---|---|---|
1. | ||
2. | ||
3. | ||
4. | ||
5. | ||
6. | ||
7. | ||
8. | ||
9. | ||
10. | ||
11. | ||
12. | ||
13. | ||
14. | ||
15. | ||
16. | ||
17. | ||
18. | ||
19. | ||
20. | ||
21. | ||
22. | ||
23. | ||
24. |
The diagnostic characters of the group are:
– beetles small or medium-sized (
– habitus elongate to oval, in most species oblong-oval (broadest approximately at elytral midlength); with rounded pronotal and elytral sides, body outline continuous;
– pronotum short, trapezoidal, with posterior angles not drawn backwards;
– coloration reddish to piceous, mainly uniform, sometimes with paler head and pronotum and darker elytra;
– microreticulation and punctation of dorsal surface very fine to strongly impressed, beetles shiny to matt dorsally;
– metacoxae and abdominal ventrites 1–5 (and 6 in males) with thin, almost longitudinal striae/strioles;
– pronotum and elytra without striae or strioles;
– pronotum with or without lateral bead;
– antennomeres not modified;
– male protarsomeres 1–3 not expanded laterally;
– male protarsomere 4 cylindrical, narrow, with anterior angle slightly expanded in some species, with a large, hook-like to thin, long, slightly curved anterolateral seta;
– male protarsomere 5 long and narrow, sometimes slightly concave ventrally;
– median lobe of aedeagus with continuous outline in ventral and lateral view; almost straight or slightly curved in lateral view; in ventral view, almost parallel-sided, often narrowed distally before apex or towards it, or broadened subdistally; apex usually with thickened sides, slightly or distinctly enlarged (“swollen”, often ventrally of shape of a baby pacifier), rounded, truncate, or slightly concave in ventral view;
– ventral sclerite of median lobe more or less deeply divided apically;
– median lobe without setation, in some species with minuscule spines;
– paramere without dorsal notch and with long, dense, thin setae, situated along dorsal margin, subdistal setae usually denser and stronger than proximal ones.
Although the species of the group do not form a monophyletic complex with the distinguished autapomorphic morphological character (Fig.
Phylogenetic relationships of the
Phylogenetically, the group is polyphyletic and includes five different clades, which are partially supported morphologically and contribute to two larger monophyletic complexes: 1) clades I and II plus the
Clade I includes
Clade II is morphologically rather heterogeneous and is comprised of the largest (size) representatives of the group. Three of them,
Clade III includes species without lateral pronotal bead, except for
Clade IV is the most homogeneous and includes the smallest in size species of the group. They are morphologically very similar, and three of them,
Clade V includes two very different species.
Thus, this group, as defined now, is the subject of further study and may be divided into subgroups or even groups as and when additional species are discovered.
Papua New Guinea: Madang Province, Adelbert Mts, Keki to Sewan, 04°41.80'S, 145.25.46'E, 650 m a.s.l.
Papua New Guinea: Madang Province. The species is known only from the type locality (Fig.
The species is named after Adelbert Mountains. The species name is an adjective in the nominative singular.
Papua New Guinea: Southern Highlands Province, Tari, Mt Ambua,
Papua New Guinea: Southern Highlands Province. The species is known only from the type locality (Fig.
The species is named after Mt Ambua. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Sandaun Province, Bewani Mts, approximately
Papua New Guinea: Sandaun Province. The species is known only from the type locality (Fig.
The species is named after the Bewani Mountains. The name is a noun in the nominative singular standing in apposition.
Papua: Nabire Regency, 62 km of road Nabire to Enarotali, ca
1 female “IRIAN JAYA: Paniai Prov. road Nabire-Ilaga, km 65 29.8.1996, 250m leg. M. Balke (96 # 6)” (
For complete description, see
Within the
Papua: Nabire Regency. The species is known only from the area close to the type locality (Fig.
Papua: Jayapura Regency, Cyclops Mts,
Papua: Jayapura Regency. The species is known only from the Cyclops Mountains (Fig.
The species is named after the Cyclops Mountains. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: East Sepik Province, Amboin Patrol Post, Karawari Lodge, ca
For complete description, see
In the area of its distribution,
Within the
Papua New Guinea: East Sepik, Simbu, Western Highlands, and Eastern Highlands Provinces (Fig.
Papua: Pegunungan Bintang Regency, Borme,
For complete description, see
In the area of its distribution,
Within the
Papua: Pegunungan Bintang. The species is known only from the type locality (Fig.
Papua New Guinea: Morobe Province, Herzog Range, Wagau (Vagau), ca
For complete description, see
In the Herzog Range area,
Within the
Papua New Guinea: Morobe Province. The species is known only from the type locality, Wagau in Herzog Range (Fig.
Papua New Guinea: Sandaun Province, Mianmin area, ca
The species has variability in size, coloration and shape of the median lobe. Beetles are small to medium-sized (see the measurements above) and with coloration: from reddish head and pronotum and dark brown elytra to uniformly piceous with reddish brown pronotal sides. Median lobe shows different shape of its apex: in lateral view, it is not curved or differently slightly curved downwards reminding that of
Papua New Guinea: Sandaun Province (Fig.
The species is named for Sentiko Ibalim, one of the great young
Papua New Guinea: Madang, Adelbert Mts, creek near Keki,
Papua New Guinea: Madang and Eastern Highlands (Fig.
species is named after Keki Village. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Enga Province, Kumul Lodge at foot of Mt Hagen,
Papua New Guinea: Enga Province. The species is known only from the type locality (Fig.
The species is named after Kumul Lodge. The name is an adjective in the nominative singular.
Papua New Guinea: Southern Highlands Province, Sopulkul, 30–35 km NE Mendi,
Papua New Guinea: Southern Highlands Province. The species is known only from the type locality (Fig.
The species is named after Mendi Village. The name is an adjective in the nominative singular.
Papua New Guinea: Morobe Province, Menyamya, Mount Inji,
Papua New Guinea: Morobe Province. The species is known only from the type locality (Fig.
The species is named after Menyamya Village. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: East Sepik Province, Amboin Patrol Post, Karawari Lodge.
For complete description, see
See under
Papua New Guinea: Eastern Highlands Province, Wapi Creek, Kimiagomo, Okapa,
3 females “Papua New Guinea: Eastern Highlands, Hano kotu, Kimiagomo, Okapa, 1661m, 11.viii.2006,
Papua New Guinea: Eastern Highlands Province (Fig.
The species is named after Okapa Station. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: East Sepik Province, Lembena,
Papua New Guinea: East Sepik Province (Fig.
The species is named for Pius, a local collector. The species name is a noun in the genitive case.
Papua New Guinea: Madang Province, Wannang,
Papua New Guinea: Madang (Fig.
The species is named “
Papua New Guinea: Simbu Province, Mount Wilhelm,
The species was mistaken for
Papua New Guinea: Madang Province, Akameku-Brahmin, Bismarck Range,
Papua New Guinea: Madang and Simbu Provinces (Fig.
The species name derives from the Latin “pusillus” (small, tiny) to express small size of these beetles. The species name is an adjective in the nominative singular.
Papua New Guinea: Simbu/Eastern Highlands Province, Crater Mountain, Sera – Herowana, Sima River, ca
In absence of the pronotal bead and thin and not hook-like anterolateral seta of the male protarsomere 4,
Papua New Guinea: Simbu and Eastern Highlands Provinces, Crater Mountain. This species is known only from the type locality (Fig.
The species is named after Sima River. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Western Highlands Province, Simbai, Ineng River,
Papua New Guinea: Western Highlands Province, near Simbai (Fig.
The species is named after Simbai area. The name is an adjective in the nominative singular.
Papua New Guinea: Western Highlands Province, Simbai-Jimi,
1 female “Papua New Guinea: Western Highlands, Simbai, 1800–2000m, 25.ii.2007,
Papua New Guinea: Western Highlands Province (Fig.
The species is named after Simbai-Jimi area. The name is a noun in the nominative singular standing in apposition.
Papua: Puncak Regency, south from Iratoi,
Papua: Puncak Regency. The species is known only from the Iratoi area (Fig.
The species is named for our friend Bob Sumoked (Tomohon, Sulawesi). The species name is a noun in the genitive case.
Papua New Guinea: Eastern Highlands Province, Kainantu, Yoginofi,
Papua New Guinea: Eastern Highlands Province (Fig.
The species is named after Yoginofi Village. The name is a noun in the nominative singular standing in apposition.
The key is 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 Figs
1 | Pronotum without lateral bead |
|
– | Pronotum with lateral bead |
|
2 | Male protarsomere 4 with anterolateral seta hook-like, large, strongly curved |
|
– | Male protarsomere 4 with anterolateral seta thin, long, equal or smaller than more laterally situated large setae, slightly curved |
|
3 | Median lobe not or slightly narrowed before truncate or slightly concave apex in ventral view (Fig. |
(4) |
– | Median lobe distinctly narrowed before truncate apex in ventral view (Figs |
|
4 | Apex of median lobe curved downwards, with visible angle on dorsal side in lateral view (Fig. |
(7) |
– | Apex of median lobe not or only slightly curved downwards in lateral view (Fig. |
(9) |
5 | Beetle more oval, broader, with pronotum reddish brown and elytra piceous (Fig. |
(20) |
– | Beetle elongate, narrower, with reddish to reddish brown dorsal coloration (Figs |
|
6 | Dorsal punctation almost invisible on elytra (Fig. |
(14) |
– | Dorsal punctation distinct on elytra (Figs |
|
7 | Dorsal microreticulation more strongly impressed (Fig. |
(10) |
– | Dorsal microreticulation less strongly impressed (Fig. |
(17) |
8 | Beetle reddish brown, more oval. Usually smaller, |
|
– | Beetle reddish brown to piceous, elongate, oblong-oval. Usually larger, |
|
9 | Beetle larger, |
(16) |
– | Beetle smaller, |
|
10 | Median lobe with thinner apex in lateral view, apex narrowed to tip in ventral view (Fig. |
(19) |
– | Median lobe with apex thickened in lateral view, apex not narrowed to tip, broad, differently truncate in ventral view (e.g., Fig. |
|
11 | Median lobe distinctly narrowed distally, with apex roundly truncate in ventral view (Fig. |
(23) |
– | Median lobe not or very slightly narrowed distally, with apex distinctly truncate or slightly concave in ventral view (e.g., Fig. |
|
12 | Apex of median lobe not curved downwards in lateral view (Fig. |
(5) |
– | Apex of median lobe curved downwards in lateral view (Figs |
|
13 | Apex of median lobe narrower in lateral view and slightly concave in ventral view (Fig. |
(3) |
– | Apex of median lobe broader in lateral view and truncate in ventral view (Fig. |
(1) |
14 | Apex of median lobe strait, flatted, and thin apex in lateral view and broadly elongated, lanceolate, impressed in ventral view (Fig. |
(13) |
– | Apex of median lobe of different shape, never so flatted and impressed ventrally, usually thickened in lateral view (e.g., Fig. |
|
15 | Median lobe evenly broad, with rounded, not curved downwards apex in lateral view (e.g., Fig. |
|
– | Median lobe narrowed towards apex, apex pointed or slightly rounded, usually curved downwards in lateral view (e.g., Fig. |
|
16 | Beetle smaller, |
(6) |
– | Beetle larger, |
|
17 | Beetle dark brown, with elytral punctation finer. Median lobe thinner and narrower in lateral view (Fig. |
(8) |
– | Beetle dark brown to piceous, with elytral punctation more distinct. Median lobe thicker and broader in lateral view (Fig. |
(22) |
18 | Beetle more elongate, almost parallel-sided, smaller, |
|
– | Beetle more oval, larger, |
|
19 | Beetle smaller, |
(18) |
– | Beetle larger, |
(21) |
20 | Beetle smaller, |
|
– | Beetle larger, |
|
21 | Apex of median lobe not bent downwards in lateral view, roundly truncate in ventral view (Fig. |
(2) |
– | Apex of median lobe bent downwards in lateral view, rounded in ventral view (e.g., Fig. |
|
22 | Beetle smaller, |
(24) |
– | Beetle larger, |
(15) |
23 | Dorsal punctation and microreticulation weaker (Fig. |
(12) |
– | Dorsal punctation and microreticulation stronger (Fig. |
(11) |
The studied species have the same habitat preferences as those described in
Habitus and coloration
Habitus and coloration
Habitus and coloration
Habitus and coloration
Habitus and coloration
Habitus and coloration
Map of New Guinea showing distribution of the species.
We are grateful Dr. H. Schillhammer (Vienna) for the habitus photos and Clive R. Turner (Plymouth) for a linguistic review of the manuscript.
Fieldwork was supported by the UK Darwin Initiative project “Training the next generation of
The specimens from Mount Wilhelm, Papua New Guinea were collected during the “Our Planet Reviewed Papua-New-Guinea 2012–2013” project. Specimens were exported under the permit # 012297 issued by the Department of Environment and Conservation (DEC, Port Moresby). Sorting and processing of the material was supported by the European Research Council (ERC) grant 669609 to C. Wardhaugh.
Financial support for the study was 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).