Corresponding author: Helena V. Shaverdo (
Academic editor: M. Fikácek
The
Especially the New Guinean
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The present work is based on the material from the following collections:
All specimen data are quoted as they appear on the labels attached to the specimens. Label text is cited using quotation marks. Comments in square brackets are ours. We extracted DNA and obtained DNA sequence data for some of the species/specimens, marked with individual DNA extraction numbers (e.g. “256 DNA M. Balke”). All types of the herein described specimens are provided with red labels. The female specimens, identification of which is difficult or sometimes impossible, are included in the type series only when collected with males of respective species and do not show external morphological differences from them. If two or more morphologically similar species collected together (males found together), their females were not included in the types series of the respective species but were mentioned under additional material.
Measurements were taken with a Wild M10 stereomicroscope. The following abbreviations were used: TL (total body length), TL-H (total body length without head), MW (maximum body width), and hw (handwritten). Number of the ventral setae of the male protarsomere 5 is given only for one specimen of each species, which was mounted on a glass slide (see below) for drawing. This character was found not very useful for the species identification since it is possible to make a general statement of the setation pattern (short/long, dense/sparse) but not to count them with certainty at the magnification of normal dissecting scopes. The potential phylogenetic information content of this character will be studied in a further work.
Drawings were made with the aid of a camera lucida attached to a Leica DM 2500 microscope. For detailed study and drawing, antennae, protarsi, and genitalia were removed and mounted on glass slides with DMHF (dimethyl hydantoin formaldehyde) as temporary preparations. The drawings were scanned and edited, using the software Adobe Illustrator CS5.1. Arrangement of the figures follows the species order in the key.
The terminology to denote the orientation of the genitalia (ventral for median lobe and dorsal and external for paramere) follows
This group is newly introduced herein, and the following diagnostic characters of the group are proposed:
- beetles small or middle-sized (TL-H 3.0–5.2 mm);
- habitus oblong-oval (broadest approximately at elytral middle), with rounded pronotal and elytral sides, body outline continuous;
- pronotum short, trapezoidal, with posterior angles not drawn backwards;
- coloration red, reddish-brown to piceous, mainly uniform, sometimes with paler head and pronotum and darker elytra;
- microreticulation and punctation of dorsal suface very fine to strongly impressed, so that beetles shiny to matt dorsally;
- metacoxae and abdominal sternites 2–6 (and 7 in males) with thin, almost longitudinal striae/strioles;
- pronotum and elytra without striae or strioles;
- pronotum with or without lateral bead;
- male antennomeres simple or antennomeres 3–10 differently modified;
- male protarsomeres 1–3 not expanded laterally;
- male protarsomere 4 cylindrical, narrow, with small to large anterolateral hook;
- male protarsomere 5 not modified: long and narrow, without expansion and concavity, ventrally with two sparse rows of relatively short setae;
- median lobe of aedeagus with discontinuous outline in ventral view and sometimes in lateral view;
- ventral sclerite of median lobe more or less deeply divided on the middle, in some species like two long subequal ventral sclerites, because of strong sclerotisation of its lateral sides;
- apical part of median lobe with or without setae;
- paramere with or without notch on dorsal side;
- paramere with long setae occupying whole dorsal side, in most species these setae denser and stronger on subdistal part of paramere and can be of different length and shape.
The main diagnostic character of the group is discontinuous outline of the median lobe of the aedeagus in ventral view and sometimes in lateral view (showed with arrows in
Abbreviations: IN – Indonesia, PNG – Papua New Guinea.
1. |
IN: West Papua: Manokwari |
2. |
IN: West Papua: Manokwari |
3. |
IN: West Papua: Manokwari |
4. |
PNG: Madang |
5. |
PNG: Madang, East Sepik, Enga |
6. |
IN: Papua: Jayawijaya |
7. |
PNG: Sandaun, East Sepik, Madang, Morobe |
8. |
PNG: Madang, Eastern Highlands |
9. |
PNG: Western and Southern Highlands |
10. |
IN: Papua: Nabire, Paniai |
11. |
IN: Papua: Jayawijaya |
12. |
IN: West Papua: Raja Ampat |
13. |
PNG: Morobe |
14. |
IN: Papua: Nabire, Paniai |
15. |
IN: Papua: Nabire, Paniai |
16. |
PNG: Eastern Highlands |
17. |
PNG: Simbu, Eastern Highlands |
18. |
IN: Papua: Nabire, Paniai |
19. |
IN: West Papua: Manokwari |
20. |
IN: Papua: Nabire, Paniai |
21. |
IN: Papua: Nabire, Paniai |
22. |
IN: Papua: Nabire, Paniai |
23. |
IN: Papua: Nabire, Paniai |
24. |
IN: West Papua: Raja Ampat |
25. |
IN: Papua: Nabire, Paniai |
26. |
IN: West Papua: Teluk Wondama |
Indonesia: West Papua Province: Manokwari Regency, Arfak Mts., Tetaho area, Iranmeba.
Beetle middle-sized, dark brown to piceous; pronotum with distinct lateral bead; male antennomeres 3 and 4 evidently larger than other, with external margin expanded (antennomeres triangular, elongated); male protarsomere 4 with small (only slightly larger than more laterally situated large seta), thin, slightly curved anterolateral hook; median lobe with strong submedian constriction in ventral view, apex of median lobe almost truncate in lateral view and broad in ventral view; paramere with notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, and flattened setae. The species is well recognizable by the modified antennae of the males.
Indonesia: West Papua Province: Manokwari Regency. This species is known from the eastern Bird’s head only. Iranmeba and Testega are situated some 20–30 km west of Anggi-Lakes in the Arfak Mountains (
The species is named for friend and most enthusiastic explorer of New Guinea’s entomofauna, Alexander Riedel (Karlsruhe, Germany), who discovered this species. The species name is a noun in the genitive case.
Indonesia: West Papua Province: Manokwari Regency, Arfak Mts., Anggi, Iray,
Beetle middle-sized, reddish-brown to brown; submatt; pronotum with lateral bead; male antennomeres 3–4 strongly enlarged and triangular (3 larger than 4), 5–6 distinctly enlarged, 7 somewhat enlarged; male protarsomere 4 with small (only slightly larger than more laterally situated large seta), thin, slightly curved anterolateral hook; median lobe with strong submedian constriction in ventral view, apex of median lobe elongate in lateral view and narrower in ventral view; paramere with notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, and flattened setae.
Indonesia: West Papua Province: Manokwari Regency. The species is known only from the type locality (
The species is named for the type area, Anggi. The name is an adjective in the nominative singular.
Indonesia: West Papua Province: Manokwari Regency, Arfak Mts., Sijoubreg Village near Mokwam,
Beetle middle-sized, piceous, shiny but with evident dorsal punctation; pronotum with distinct lateral bead; male antennomeres 3–4 strongly enlarged and triangular (3 larger than 4), 5–6 distinctly enlarged, 7 somewhat enlarged; male protarsomere 4 with small, thick (evidently thicker and slightly longer than more laterally situated large seta), slightly curved anterolateral hook; median lobe with strong submedian constriction in ventral view, apex of median lobe elongate in lateral view and narrower in ventral view; paramere with notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, and flattened setae.
Indonesia: West Papua Province: Manokwari Regency. The species is known only from the Arfak Mountains, the eastern part of Bird’s head (
The species is named for the type area, Arfak Mountains. The name is an adjective in the nominative singular.
Papua New Guinea: Madang Province, 1–3 km SE Brahman.
1 male “PAPUA NEW GUINEA Madang Pr. Brahmin, 150 m, 26IX2002, M Balke (PNG 24)”, “273 DNA M Balke” (ZSM). 6 males, 5 females, 7 exs. “Papua New Guinea: Madang, Usino, 260m, 15.iii.2007,
Beetle middle-sized, piceous, with paler pronotum (especially on margins) and head, teneral specimens reddish-brown, submatt; pronotum with distinct lateral bead; male antennomeres simple; male protarsomere 4 with middle-sized, thick, evidently curved anterolateral hook; median lobe short and with extremely strongly discontinuous (broken and curved) outline; paramere with shallow notch on dorsal side and subdistal part elongate, with dense, long, thin setae.
For the complete description see
Papua New Guinea. The species is known only from the Madang Province (
Papua New Guinea: Madang Province, river below Bundi, approximately
Beetle middle-sized, piceous, with paler head, shiny (
For the complete description and illustrations see
Papua New Guinea. The species is known from Madang, East Sepik, and Enga Provinces (
Indonesia: Papua Province: Jayawijaya Regency, Borme, Tarmlu, approximately 04°24'S, 140°25'E.
Beetle small, dark brown to piceous, shiny; pronotum without lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook; median lobe with strong submedian constriction and apex bifid: with small dorsal extension; paramere with notch on dorsal side and subdistal part elongate, with dense, long, thin setae. The species is well recognizable by its characteristic male genitalia.
Indonesia: Papua Province, Jayawijaya Regency. This species is known only from the type locality (
The name refers to the bifid distalodorsal part of the median lobe. The name is an adjective in the nominative singular.
Papua New Guinea: Madang Province, Adalbert Mts., near Keki,
8 females “Papua New Guinea: Madang, Keki, Adalbert Mts., 400m, 29.xi.1994,
Beetle small, piceous, with reddish brown head and pronotum, shiny; pronotum without lateral bead or with weak traces of lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook; median lobe with weak submedian constriction in ventral view; paramere with strong notch on dorsal side, with notch tip sharply pointed, and subdistal part elongate, with upper setae thin and more numerous and lower setae shorter, thick, and flattend. The species is similar to
Papua New Guinea. This species is wide distributed in the Momase Region: Sandaun, East Sepik, Madang, and Morobe Provinces (
The name refers to the village Brahmin where this species was first discovered. The species name is an adjective in the nominative singular.
Papua New Guinea: Eastern Highlands Province, Akameku-Brahmin, Bismarck Range,
Beetles small, dark brown, dorsally less strongly punctate than
Papua New Guinea. The species is known from Madang and Eastern Highlands Provinces (
The species is named after the village Bundi where it was discovered. The name is an adjective in the nominative singular.
Papua New Guinea: Western Highlands Province, Kurumul, 6 km SW Kudjip,
Beetle middle-sized, piceous, submatt; pronotum with distinct lateral bead; male sternite 7 slightly to distinctly concave apically; male antennomeres 3–5 distinctly enlarged, almost equally in size and shape, antennomeres 6–8 enlarged; male protarsomere 4 with middle-sized, slender, evidently curved anterolateral hook; median lobe with very strong submedian constriction and proximal part very broad in ventral view, apex of median lobe pointed and strongly curved downwards in lateral; paramere with distinct notch on dorsal side and subdistal part elongate, with numerous, dense, long, thin setae. The species is well recognizable by the modified antennae of the males and the shape of the median lobe.
Papua New Guinea. The species is known from Western Highlands and Southern Highlands Provinces (
Dedicated to Edeltraud Tötzl, senior author’s mother-in-law: “With my sincere thankfulness. Without your help with my children, my scientific work was not possible during last six years”. The species name is a noun in the genitive case.
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 55th km,
10 females “IRIAN JAYA: Paniai Prov. road Nabire-Ilaga, km 54 10.9.1996, 900m leg. M. Balke (96 # 19)” (NHMW). These females are most likely a mixture of three species:
Beetle small, dark brown, with slightly paler anterior part of head and pronotal sides, shiny; pronotum without lateral bead; male antennomeres 3–10 stout; male protarsomere 4 with large, thick, strongly curved anterolateral hook; median lobe with strong submedian constriction and apical part very broad in ventral view and slightly flattened in lateral view; paramere with notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, and flattened setae.
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
This species is named for the native community (Ekari people) which inhabits the area from which the specimens have been collected. The name is a noun in the nominative standing in apposition.
Indonesia: Papua Province: Jayawijaya Regency, Emdoman, 04°14'S, 139°55'E.
Beetle small, piceous, shiny; pronotum without lateral bead; male antennomeres 3-10 slightly stout; male protarsomere 4 with large, thick, strongly curved anterolateral hook; median lobe with weak submedian constriction in ventral view; paramere with notch on dorsal side and subdistal part elongate, with upper setae thin and less numerous and lower setae long, thick, somewhat flattend, and curved at apex.
Indonesia: Papua Province, Jayawijaya Regency. This species is known only from the type locality (
The species is named for the Eme River, from a tributary of which it has been collected. The name is a noun in the nominative singular standing in apposition.
Indonesia: West Papua Province: Raja Ampat Regency, Waigeo Island, Mountain Nok.
Beetle middle-sized, dark brown to piceous (teneral specimens reddish-brown), shiny; pronotum with lateral bead; male antennomeres 3–7 very slightly enlarged, antennomere 3 slightly more triangular than other antennomeres; male protarsomere 4 with middle-sized, slender, evidently curved anterolateral hook; median lobe with strong submedian constriction in ventral view and truncate apex in lateral view; paramere with notch on dorsal side and subdistal part short and small, with less numerous, relatively short, thick, and flattened setae. The species occurs together with
Indonesia: West Papua Province: Raja Ampat Regency. The species is known only from the type locality (
The species is named for the incredible collector and adventurer, Miss Lucy Evelyn Cheesman who discovered this species. The species name is a noun in the genitive case.
Papua New Guinea: Morobe Province, Herzog Mts., Wagau,
Beetle small, yellowish-red to brown, dorsally strongly punctate, submatt; pronotum with lateral bead; male antennomeres 3–5 enlarged, 6–8 slightly enlarged; male protarsomere 4 with middle-sized, slender, evidently curved anterolateral hook; median lobe with strong submedian constriction and proximal part narrower, apex of median lobe rather broad in lateral view; paramere with shallow notch on dorsal side and subdistal part elongate, with less numerous, long, thin setae.
Papua New Guinea: Morobe Province. The species is known only from the type locality (
The species is named after our friend and colleague Dr. Hans Fery (Berlin). The name is a noun in the genitive case.
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 54th km,
29 females “IRIAN JAYA: Paniai Prov. road Nabire-Ilaga, km 117 Unipo, 2.9.1996, 150m leg. M. Balke (96 # 12)” (NHMW). These females are most likely a mixture of two species:
Beetle small, reddish-brown to dark brown, usually with paler head and pronotal sides; shiny; pronotum without lateral bead; male antennomeres 3–5 distinctly enlarged; male protarsomere 4 with large, thick, evidently curved anterolateral hook; median lobe with very strong submedian constriction, distal and proximal parts equally broad, and symmetrical apex in ventral view; paramere with shallow notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, and flattened setae. The species differs from other ones by the pronotum without lateral bead and modified antennae of the males.
Indonesia: Papua Province. This species is known from Nabire and Paniai Regencies (
The name is derived from the Biak (northern coast of New Guinea) islanders’ word “Irian”, which means “hot island emerging from the sea” and refers to New Guinea. The species name is an adjective in the nominative singular.
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 54th km,
12 females “Indonesia: Papua, Road Nabire-Enarotali KM 60, 640m, 22.x.2011, 03 30.474S 135 42.611E, Uncen (PAP10)”, two of them additionally with labels “DNA M. Balke 4909”, “DNA M. Balke 4913” (NHMW, ZSM). These females most likely belong to
Beetle small, very similar to
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
The Ekari people sometimes called smaller water beetles “kakapupu” (
Papua New Guinea: Eastern Highlands Province, Kainantu, Yoginofi,
Beetle middle-sized, uniformly dark brown or head and pronotum slightly paler; pronotum with distinct lateral bead; male antennomere 3 evidently larger than other; male protarsomere 4 with very small (smaller than more laterally situated large seta), thin, slightly curved anterolateral hook; median lobe with very weak submedian constriction, apex of median lobe almost rounded in lateral view; paramere without notch on dorsal side, with relatively short, sparse, thin setae. The species is well recognizable by its larger size, the modified antennae of the males, and paramere distinctly longer than median lobe.
Papua New Guinea. This species is known from Eastern Highlands Province (
The species is named for an old friend of M. Balke. The name is a noun in the nominative singular standing in apposition.
Papua New Guinea: Simbu/Eastern Highlands Provinces, Crater Mountain, Wara Sera Station,
1 female “Papua New Guinea: Simbu/EHP. Crater Mountain, Sera-Herowana, Hulene river, 1000m, 16IX2002, Balke & Sagata (PNG 017)” (ZSM). 1 male “Papua New Guinea: Crater Mountain, Sera-Herowana, Hulene river, 1000m, 16.IX2002, Balke & Sagata (PNG 017)” (ZSM). 2 females “Papua New Guinea: Crater Mountain, Wara Sera Station, 800m, 14IX2002, Balke & Sagata, (PNG 010)” (NHMW, ZSM). 1 male “PAPUA NEW GUINEA Simbu/EHPr. Crater Mountain, Wara Sera Station, 800m, 14IX2002, Balke & Sagata, (PNG 10)” (NHMW). 1 female “Papua New Guinea: Simbu/EHP, Crater Mountain, Wara Sera Station, 800m, 14IX2002, Balke & Sagata, (PNG 009)” (ZSM).
Beetle middle-sized (TL-H 4.8–5.0 mm, TL 5.1–5.3 mm, MW 2.6 mm), piceous, dull (
For the complete description and illustrations see
Papua New Guinea. The species is known only from Crater Mountain, Simbu and Eastern Highlands Provinces (
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 65th km,
Beetle small, reddish-brown to brown; pronotum with narrow lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook; median lobe slender, with weak submedian constriction in ventral view and elongate apex in lateral view; paramere with strong notch on dorsal side and subdistal part short and large, with long, dense, curved at apex setae.
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
The species is named for “Doc” Ocea Megay, one of our Ekari fellows and most enthusiastic fieldworker who died from a snake bite shortly after we have collected this species. He is buried in Topo. The species name is a noun in the genitive case.
Indonesia: West Papua Province: Manokwari Regency, Arfak Mts., Hatam.
Beetle middle-sized, brown, shiny, with almost invisible dorsal punctation; pronotum with distinct lateral bead; male antennomeres 3–4 strongly enlarged and triangular (3 distinctly larger than 4), 5–6 distinctly enlarged, 7 somewhat enlarged; male protarsomere 4 with small, thin, slightly curved anterolateral hook; median lobe with strong submedian constriction in ventral view, apex of median lobe elongate in lateral view and broader in ventral view; paramere with notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, and flattened setae.
Indonesia: West Papua Province: Manokwari Regency. The species is known only from the lectotype from the Arfak Mountains, the eastern part of Bird’s head (
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 80th km,
Beetle small, very similar to
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
Long time this species was mistaken for
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 80th km,
See in the paragraph of
Beetle small, dark brown, often with paler head and pronotal sides, shiny; pronotum without lateral bead; male antennomeres 3–10 slightly stout; male protarsomere 4 with middle-sized, slender, evidently curved anterolateral hook; median lobe with weak submedian constriction in ventral view and elongate apex in lateral view; paramere with notch on dorsal side and subdistal part short and small, with large brush of thick, somewhat flattend, long, curved at apex setae.
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
This species is dedicated to our old buddy Michael Sopp (Berlin). The species name is a noun in the genitive case.
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 117th km, Unipo, approximately
See in the paragraph of
Beetles small, very similar to
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
The species is named for the type locality. The name is a noun in the nominative singular standing in apposition.
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 80th km,
Beetle small, piceous, shiny; pronotum without lateral bead; male antennomeres 3–10 slightly enlarged; sternite 7 concave; male protarsomere 4 with large, thick, strongly curved anterolateral hook; median lobe long, with very weak submedian constriction and apex narrow in ventral view; paramere large, with strong notch on dorsal side and subdistal part very broad, subquadrate, with dense, long, relatively thick, curved at apex setae. The species is well recognizable by its characteristic male genitalia and concave (also in females) abdominal sternite 7.
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known from the lower Utowa River area and one of its tributaries (
The species is named for Utowa River from which many specimens have been collected. The name is an adjective in the nominative singular.
Indonesia: West Papua Province: Raja Ampat Regency, Waigeo Island, Mountain Nok.
27 females “N.DUTCH NEW GUINEA: Waigeu. Camp Nok. 2,500 ft. iv.1938. L.E.Cheesman. B.M.1938-593.”, one of them additionally with labels “collection 27”, “measured J.Parkin 76” (BMNH). 6 females “N.DUTCH NEW GUINEA: Waigeu.Camp 1.Mt.Nok. 2,500 ft. v.1938. L.E.Cheesman. B.M.1938-593.” (BMNH). These females are a mixture of two species:
Beetle small, reddish-brown, shiny; pronotum with distinct lateral bead; male antennomeres 3–7 very slightly enlarged, antennomere 3 slightly more triangular than other antennomeres; male protarsomere 4 with middle-sized, slender, evidently curved anterolateral hook; median lobe with strong submedian constriction in ventral view and elongate apex in lateral view; paramere with notch on dorsal side and subdistal part short and small, with less numerous, relatively short, thick, and flattened setae.
Indonesia: West Papua Province: Raja Ampat Regency. The species is known only from the type locality (
The species is named in reference to its distribution: Waigeo Island. The name is an adjective in the nominative singular.
Indonesia: Papua Province: Nabire/Paniai Regencies, road Nabire-Enarotali, 55th km,
See in the paragraph of
Beetle small, reddish-brown to brown, with paler head and pronotal sides, shiny; pronotum without lateral bead; male antennomeres 3–10 stout; protarsomere 4 with middle-sized, slender, evidently curved anterolateral hook; median lobe slender, with very weak submedian constriction in ventral view; paramere with notch on dorsal side and subdistal part short and small, with not numerous, relatively short, thick, flattened, slightly curved at apex setae.
Indonesia: Papua Province: Nabire and Paniai Regencies. This species is known only from the type locality area (
Derived from the name of the range, Weyland, at the northern edge of which the type locality is situated. The species name is an adjective in the nominative singular.
Indonesia: West Papua Province: Teluk Wondama Regency, Wandammen Peninsula, Wondiwoi Mts., Wasior,
Beetle small, externally very similar to
Indonesia: West Papua Province: Teluk Wondama Regency. This species is known from the Wondiwoi Mountains of Wandammen Peninsula (
The species is named for the type area, Wondiwoi Mountains. The name is an adjective in the nominative singular.
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 17a and 17b in
1 | Pronotum with distinct lateral bead, broad or narrow | 2 |
– | Pronotum without lateral bead or with weak traces of lateral bead | 15 |
2 | Male antennomeres simple or slightly modified: antennomere 3–7 very slightly enlarged (almost indistinctly), antennomere 3 slightly more triangular than other antennomeres | 3 |
– | Male antennomeres 3–5 evidently enlarged | 8 |
3 | Beetle larger, TL-H: 3.9–5.0 mm, piceous | 4 |
– | Beetle smaller, TL-H: 3.35–4.1 mm, reddish-brown to piceous | 5 |
4 | Beetle larger, TL-H: 4.8–5.0 mm ( |
(17) |
– | Beetle smaller, TL-H: 3.9–4.1 mm ( |
(5) |
5 | Beetle smaller, TL-H: 3.35–3.8 mm ( |
(18) |
– | Beetle larger, TL-H: 3.45–4.1 mm ( |
6 |
6 | Beetle dorsally submatt, with distinct punctation ( |
(4) |
– | Beetle dorsally shiny, with very fine punctation ( |
7 |
7 | Beetle smaller, TL-H: 3.45–3.7 mm, MW: 1.8–2.0 mm ( |
(24) |
– | Beetle larger, TL-H: 3.75–4.1 mm, MW: 1.9–2.2 mm ( |
(12) |
8 | Male antennomeres 3–5 enlarged, rounded, almost equally in size and shape | 9 |
– | Male antennomeres 3 or 3–4 distinctly more modified in shape (triangular) and larger than other antennomeres | 11 |
9 | Male antennomeres 3–5 strongly enlarged ( |
(9) |
– | Male antennomeres 3–5 evidently less enlarged ( |
10 |
10 | Beetle dorsally brightly ferrugineous to castaneous, submatt, with punctation coarse and dense ( |
(13) |
– | Beetle dorsally dark brown, almost shiny, with punctation less coarse and dense ( |
(8) |
11 | Male antennomere 3 much larger than other antennomeres, triangular ( |
(16) |
– | Male antennomeres 3 and 4 much larger than other antennomeres, triangular, beetle smaller, TL-H: 3.7–4.3 mm, MW: 2.05–2.3 mm, of different color, male protarsomere 4 with anterolateral hook thin or thick, slightly curved but larger than more laterally situated large seta, median lobe with stronger median constriction in ventral view and apex different, paramere equal or shorter than median lobe, with notch on dorsal side, setae of subdistal part not numerous, relatively short, thick, and flattened | 12 |
12 | Male antennomeres 3 and 4 more strongly elongated, more equal in size and shape ( |
(1) |
– | Male antennomeres 3 and 4 less elongated, antennomere 3 larger than 4, coloration and elytral punctation different, median lobe with apex elongate in lateral view, paramere broader | 13 |
13 | Beetle dorsally ferrugineous, submatt, with coarse punctuation ( |
(2) |
– | Beetle dorsally brown to piceous, shiny, with distinctly finer punctation, male protarsomere 4 with anterolateral hook thin or thick, median lobe and paramere different | 14 |
14 | Beetle dorsally piceous, with elytral punctation fine but distinct ( |
(3) |
– | Beetle dorsally brown, with elytral punctation almost invisible ( |
(19) |
15 | Male antennomeres 3 and 4 strongly enlarged, 5 less enlarged, and 2, 6–9 slightly enlarged | 16 |
– | Male antennomeres simple or antennomeres 3–10 slightly enlarged (stout) | 17 |
16 | Beetle reddish-brown to brown ( |
(14) |
– | Beetle dark brown to piceous ( |
(26) |
17 | Sternite 7 slightly or strongly concave apically, median lobe long, with very weak submedian constriction and narrow apex in ventral view, paramere large, with strong notch on dorsal side and subdistal part very broad, subquadrate ( |
(23) |
– | Sternite 7 broadly rounded or truncate apically, median lobe distinctly shorter, paramere smaller, with weaker notch on dorsal side and subdistal part small and short or elongate (e.g. |
18 |
18 | Apex of median lobe bifid: with small dorsal extension ( |
(6) |
– | Apex of median lobe not bifid | 19 |
19 | Beetle larger, TL-H: 3.4–3.7 mm, MW: 1.75–1.95 mm ( |
(10) |
– | Beetle smaller, TL-H: 3.0–3.5 mm, MW: 1.6–1.8 mm, apical part of median lobe narrower in ventral view and not flattened in lateral view, paramere with subdistal part small and short or elongate, setation different | 20 |
20 | Median lobe slender, especially its apical part, with very weak submedian constriction in ventral view, paramere with subdistal part small and short, with not numerous, relatively short, thick, flattened, slightly curved at apex setae ( |
(25) |
– | Median lobe more robust, with stronger submedian constriction in ventral view, paramere with subdistal part small and short or elongate, setation different | 21 |
21 | Male protarsomere 4 with middle-sized, slender anterolateral hook ( |
22 |
– | Male protarsomere 4 with large, thick, strongly curved anterolateral hook ( |
23 |
22 | Prosternal ridge evidently rounded and smooth, male antenna stout, median lobe with apex elongate in lateral view and submedian constriction weaker in ventral view, paramere with notch on dorsal side and subdistal part short and small, with large brush of thick, somewhat flattend, long, curved at apex setae ( |
(21) |
– | Prosternal ridge anteriorly evidently less rounded and smooth, male antenna simple, median lobe with apex truncate in lateral view and submedian constriction stronger in ventral view, paramere with shallow notch on dorsal side and subdistal part elongate, with strong tuft of thicker, somewhat flattend, and strongly curved at apex setae ( |
(20) |
23 | Subdistal part of paramere with two kinds of setae: thin upper setae and thick and flattened lower setae, proximal part of paramere with sparse setae ( |
24 |
– | Subdistal part of paramere only with thin setae, proximal part of paramere with dense setae ( |
25 |
24 | Male antennomeres 3–10 stout, median lobe shorter, its apex broader in ventral view and slightly elongate in lateral view, subdistal part of paramere with upper setae less numerous and lower setae long, thick, flattend, and curved at apex ( |
(11) |
– | Male antennomeres simple, median lobe longer, its apex narrower in ventral view and almost truncate in lateral view, subdistal part of paramere with upper setae more numerous and lower setae shorter, thicker, and flattend ( |
(7) |
25 | Male antennomeres simple, median lobe with apex almost truncate in lateral view and submedian constriction stronger in ventral view, paramere with setae of proximal part longer, thicker, distinctly visible ( |
(15) |
– | Male antennomeres 3–10 stout, median lobe with apex elongate in lateral view and submedian constriction weaker in ventral view, paramere with setae of proximal part shorter, thiner, often hardly visible ( |
(22) |
Habitus and coloration.
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 of the
The species are always running water associated, but they do avoid water movement. This appears paradox, but it is a very important microhabitat preference especially in many tropical ecosystems. New Guinean
In general, clay, mossy rocks, as well as presence of rough sand/gravel indicate rich
Habitat of
Habitat of
Habitat of
We are grateful to D. Zimmermann (Vienna) for her help with first steps in using the software Adobe Illustrator, Dr. H. Schillhammer (Vienna) for making habitus photos and habitus plates, and Prof. G.N. Foster (Ayr) for linguistic review of the manuscript.
Fieldwork was supported by Wildlife Conservation Society, PNG Program (now PNG Institute for Biological Research), Goroka, EHP, Papua New Guinea, as well as the PNG Binatang Research Center, Madang, Papua New Guinea. Thanks are especially due to Katayo Sagata, Aloysius Posman, Bangan John, Andrew Kinibel, and Sentiko Ibalim, and also to all other Binatang Boys whose help is greatly appreciated. Recent fieldwork in Indonesia was conducted in the course of a Cendrawasih University (Jayapura, Papua, Indonesia) training course.
Financial support of the study was provided by the FWF (Fonds zur Förderung der wissenschaftlichen Forschung – the Austrian Science Fund) through a project P 24312-B17 to the senior author. Michael Balke was supported by the UK Darwin Initiative and the German Science Foundation (various projects since BA2152/2-1).