Description of 23 new species of the Exocelina ekari-group from New Guinea, with a key to all representatives of the species group (Coleoptera, Dytiscidae, Copelatinae)

Abstract Twenty three new species of Exocelina Broun, 1886 from New Guinea are described herein: Exocelina bewaniensis sp. n., Exocelina bismarckensis sp. n., Exocelina craterensis sp. n., Exocelina gorokaensis sp. n., Exocelina herowana sp. n., Exocelina jimiensis sp. n., Exocelina kisli sp. n., Exocelina ksionseki sp. n., Exocelina lembena sp. n., Exocelina mantembu sp. n., Exocelina michaelensis sp. n., Exocelina pinocchio sp. n., Exocelina pseudoastrophallus sp. n., Exocelina pseudobifida sp. n., Exocelina pseudoedeltraudae sp. n., Exocelina pseudoeme sp. n., Exocelina sandaunensis sp. n., Exocelina simbaiarea sp. n., Exocelina skalei sp. n., Exocelina tabubilensis sp. n., Exocelina tariensis sp. n., Exocelina vovai sp. n., and Exocelina wannangensis sp. n. All of them have been found to belong to the Exocelina ekari-group. An identification key to all known species of the group is provided, and important diagnostic characters (habitus, color, male antennae, protarsomeres 4–5, median lobes, and parameres) are illustrated. Data on the distribution of the new species and some already described species are given.


Introduction
The Exocelina ekari-group, the largest species group of New Guinea Exocelina, was introduced by Balke et al. (2007) and Shaverdo et al. (2012) for 26 species. Listing several diagnostic characters of the group, we proposed a discontinuous outline of the median lobe of the aedeagus as the main diagnostic character of the group (for details, see Shaverdo et al. (2012), p. 4). We have recently detected 23 additional new species of this group, which are described here. Also, E. vladimiri (Shaverdo, Sagata & Balke, 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. Female specimens, identification of which is difficult or sometimes impossible, were included in the type series only when they were collected with males of respective species and did not show external morphological differences from them. If two or more morphologically similar species were collected together (i.e., males found together), their females were not included in the type series of the respective species but were instead mentioned under additional material. Species descriptions are based on the whole type series.
Some of the species treated herein are very similar to each other and, based on low overall genetic divergence, most likely also are of very recent origin (Toussaint et al. 2014). We have used constant morphological differences based on examined series as an indicator of interrupted gene flow and as an operational criterion to delineate biological species. However, we suggest that extensive population genetic work using genomic data might reveal many additional lineages that represent putative species in this highly structured geographic and geological setting.
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), UNCEN (Universitas Cendrawasih, Papua, Jayapura, Indonesia), UNIPA (Universitas Papua, Manokwari, West Papua, Indonesia), 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 to be not very useful for species identification since it is possible to make a general statement on the setation pattern (short/long, dense/sparse) but not to count them with certainty at the magnification of normal dissecting microscopes. 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 Miller and Nilsson (2003). The terminology on the structure of the prosternum follows Larson et al. (2000). Administrative divisions of Indonesia and Papua New Guinea follow information from Wikipedia (2014a-c).

Checklist and distribution of species of the Exocelina ekari-group
Abbreviations: IN -Indonesia, PNG -Papua New Guinea. Only new species are numbered.
Surface sculpture: Head with dense punctation (spaces between punctures 1-3 times size of punctures), evidently finer and sparser anteriorly; diameter of punctures smaller than diameter of cells of microreticulation. Pronotum with much sparser and finer punctation than on head. Elytra with extremely sparse and fine punctation, almost invisible. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum without lateral bead or with weak traces of lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, less rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively broad, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded.
Male: Antenna simple (Fig. 21A). Protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of 11-13 and posterior row of 5-6 short setae (Fig. 21B). Abdominal ventrite 6 with 7-14 lateral striae on each side. Median lobe with weak submedian constriction in ventral view and elongate apex in lateral view (Fig. 21C, D). Upper margin of apex distinctly curved or almost straight in lateral view. Paramere with distinct notch on dorsal side and subdistal part short, rounded, with upper setae almost inconspicuous or rather distinct and lower relatively long, dense, thick, and flattened; setae of proximal part more or less numerous, thin (Fig. 21E).
Female: Without evident differences in external morphology from males, except for abdominal ventrite 6 without striae.
At first, we intended to describe the species with three subspecies as these morphological differences are evident and stable within each region, though insignificant. Finally, we have decided against this, bearing in mind that more material is needed from these regions (especially, from Noiadi) and the regions in-between for a conclusion whether they belong to the different subspecies or maybe even species.
Diagnosis. Beetle medium-sized, dark brown to piceous, with paler clypeus, vertex, and pronotal sides, submatt to matt; pronotum with distinct lateral bead; male antennomeres 3-5 evidently enlarged, with margins more or less rounded, almost equal in size, antennomeres 6 and 7 somewhat enlarged; male protarsomere 4 with medium-sized, slender, evidently curved anterolateral hook-like seta; median lobe with weak submedian constriction, distal part broadened, and apex almost rounded in ventral view and thin, curved, and pointed in lateral view; paramere with shallow notch on dorsal side and subdistal part elongate, with numerous, dense, more or less long, thin setae. The species is similar to E. gorokaensis sp. n., from which differs with duller dorsal surface due to denser punctation and stronger microreticulation, as well as apex of the median lobe rounded in ventral view and narrower in lateral view and paramere with shallow, not distinct notch on dorsal side. From E. vovae sp. n., the species differs with more elongate habitus and apex of the median lobe almost rounded, not distinctly concave in ventral view.
Surface sculpture: Head with very dense, coarse punctation (spaces between punctures 1-2 times size of punctures). Pronotum with punctation finer than on head. Elytra with punctation sparser than on pronotum. Pronotum and elytra with weaker or stronger impressed microreticulation, dorsal surface submatt to matt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and not rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 slightly truncate or concave apically.
Distribution. Papua New Guinea: Eastern Highlands and Simbu Provinces. The species is known mainly from Bismarck Range (Fig. 53).
Etymology. The species is named after Bismarck Range. The name is an adjective in the nominative singular. Diagnosis. Beetle small, piceous, with dark brown head and sides of pronotum; pronotum with lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta; median lobe with submedian constriction in ventral view and strongly elongate apex in lateral view; paramere with notch on dorsal side and subdistal part elongate, with brush of long, dense, thin setae. The species is similar to E. oceai Shaverdo, Hendrich & Balke, 2012, from which differs with darker dorsal coloration and structure of the male genitalia.
Surface sculpture: Head with relatively sparse punctation (spaces between punctures 1-4 times size of punctures), evidently finer and sparser anteriorly; diameter of punctures smaller than diameter of cells of microreticulation. Pronotum with extremely sparse and fine punctation, almost invisible. Elytra without punctation, only with several extremely fine punctures and with punctural rows. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and extremely fine, sparse punctation, almost invisible, only slightly coarser and denser on two last abdominal ventrites.
Structures: Pronotum with lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, less smooth and slightly rounded anteriorly, with anterolateral extensions. Ridge laterally with distinct punctation. Blade of prosternal process lanceolate, relatively broad, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 slightly concave apically.
Female: Without evident differences in external morphology from male, except for abdominal ventrite 6 without striae.
Diagnosis. Beetle medium-sized, piceous, slightly submatt; pronotum with distinct lateral bead; male ventrite 6 slightly to distinctly concave apically; male antennomeres 3-5 distinctly enlarged, almost equal in size and shape, antennomeres 6-8 enlarged; male protarsomere 4 with medium-sized, slender, evidently curved anterolateral hook-like seta; 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 very similar to E. pseudoedeltraudae sp. n., from which differs with slightly shinier dorsal surface, due to weaker punctation and microreticulation, with smaller and less rounded male antennomeres 3-5 (for male antennomeres 3 and 4, ratio width/length: < 0.92) and apex of median lobe broader in lateral view.
Surface sculpture: Head with dense, coarse punctation (spaces between punctures 1-3 times size of punctures). Pronotum with punctation finer, sparser, and more evenly distributed than on head. Elytra with punctation much finer, sparser than on pronotum. Pronotum and elytra with less strongly impressed microreticulation, dorsal surface slightly submatt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, long strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, rounded and smooth anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded, or slightly truncate, or concave apically.
Etymology. The species is named after Goroka, where it was mostly collected. The name is an adjective in the nominative singular. Diagnosis. Beetle small, dark brown, with slightly paler head and pronotum, shiny; pronotum with lateral bead; male antennomeres 3-9 stout, with 4-5 slightly larger than other antennomeres; male protarsomere 4 with large slender, evidently anterolateral hook-like seta; median lobe with very strong median constriction and proximal part very broad in ventral view, apex of median lobe broad, slightly concave in middle and twisted at both sides in ventral view and shortly pointed in lateral view; paramere with distinct notch on dorsal side and subdistal part elongate, with a large brush of long, dense, thin setae; proximal setae almost invisible. The species is similar to E. edeltraudae and E. pseudoedeltraudae sp. n., from which differs with smaller size and stout, not evidently modified, male antennomeres.
Surface sculpture: Head with dense, coarse punctation (spaces between punctures 1-3 times size of punctures). Pronotum with punctation much finer, sparser, and more evenly distributed than on head. Elytra with punctation much finer, sparser than on pronotum, almost invisible. Pronotum and elytra with less strongly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, long strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and slightly rounded anteriorly, without anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded.
Male: Antennomeres 3-9 stout, with 4-5 slightly larger than other antennomeres (Fig. 5A). Protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of 12 setae and posterior row of 4 short setae (Fig. 5B). Abdominal ventrite 6 with 8-10 lateral striae on each side. Median lobe with very strong median constriction and proximal part very broad in ventral view, apex of median lobe broad, slightly concave in middle and twisted at both sides in ventral view and shortly pointed in lateral view (Fig. 5C, D). Paramere with distinct notch on dorsal side and subdistal part elongate, with a large brush of long, dense, thin setae; proximal setae almost invisible (Fig. 5E).
Female: Unknown. Distribution. Papua New Guinea: Simbu/Eastern Highlands Provinces. This species is known only from the type locality (Fig. 53).
Etymology. The species is named after the type locality. The name is a noun in the nominative singular standing in apposition. Diagnosis. Beetle medium-sized, dark brown to piceous, with paler clypeus, vertex, and pronotal sides, slightly submatt; pronotum with distinct lateral bead; male antennomeres 3-5 evidently enlarged, slightly rounded, almost equal in size, external margin of antennomere 5 almost straight, antennomere 6 somewhat enlarged; male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta; median lobe with submedian constriction and apex bluntly pointed, broadened in lateral view; paramere with distinct notch on dorsal side and subdistal part elongate, with numerous, dense, more or less long, thin setae. The species is similar to E. sandaunensis sp. n. and E. simbaiarea sp. n., from which differs with stronger punctation on pronotum, male antennomeres 3-5 smaller and more equal in size, median lobe with stronger submedian constriction in ventral view and more broadened apex in lateral view, subdistal part of paramere with setae more numerous, shorter, and thinner.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and not rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded or slightly truncate apically.
Holotype: TL-H 3.9 mm, TL 4.25 mm, MW 2.15 mm. Female: Antennae simple, abdominal ventrite 6 broadly rounded apically, without striae. Distribution. Papua New Guinea: Western Highlands Province. The species is known only from the area of Jimi River (Fig. 53).
Etymology. The species is named after Jimi River, in the area in which it was collected. The name is an adjective in the nominative singular. Diagnosis. Beetle medium-sized, piceous, with dark brown head and pronotum; pronotum with lateral bead; male antennomere 3 evidently larger than other antennomeres; male protarsomere 4 with very small (smaller than more laterally situated large seta), thin, slightly curved anterolateral hook-like seta; median lobe with evident submedian constriction, apex of median lobe almost rounded in lateral view; paramere without notch on dorsal side, with relatively long and dense subdistal setae and spinelike setae on internal surface. The species is similar to E. knoepfchen and E. ksionseki sp. n. It differs from E. knoepfchen with dorsal surface matt due to stronger punctation and microreticulation, male antennomeres 3-5 larger, and median lobe slender; from E. ksionseki sp. n. with larger size, dorsal surface matt due to stronger microreticulation, male antennomeres 3 smaller and more triangular, male protarsomere 4 with anterolateral hook-like seta smaller than more laterally situated large seta, apex of median lobe more rounded in lateral view, paramere only slightly longer than medial lobe, with less numerous subdistal setae and spine-like setae, and abdominal ventrite 6 less striated.
Surface sculpture: Head with very dense, coarse punctation (spaces between punctures 1-2 times size of punctures), finer and sparser anteriorly; diameter of punctures only slightly smaller than diameter of cells of microreticulation, of some punctures equal to it. Pronotum and elytra with slightly finer and more evenly distributed punctation than on head. Head, pronotum, and elytra with strongly impressed microreticulation. Dorsal surface matt due to strong punctation and microreticulation. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, anteriorly with weak transverse lines and less rounded, without anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded or slightly truncate apically.
Paramere slightly longer than median lobe, without notch on dorsal side, with relatively long and dense subdistal setae, short and sparse proximal setae, and spine-like setae on internal surface (Fig. 17E).
Female: Antennomere 1 as in male or only slightly larger, other antennomeres simple, abdominal ventrite 6 broadly rounded apically, without striae.
Etymology. The species is named for F. Kisl. The species name is a noun in the genitive case. Diagnosis. Beetle medium-sized, piceous, with reddish sides of pronotum and sometimes with reddish head, submatt; pronotum with lateral bead; male antennomere 3 evidently larger than other antennomeres; male protarsomere 4 with very small (only slightly larger than more laterally situated large seta), thin, slightly curved anterolateral hook-like seta; median lobe with very weak submedian constriction, apex of median lobe elongate in lateral view; paramere distinctly longer than median lobe, without notch on dorsal side, with relatively long and dense subdistal setae and numerous spine-like setae on internal surface, proximal setae almost absent. The species is similar to E. knoepfchen Shaverdo, Hendrich & Balke, 2012 except for evidently smaller size, coarse, dense dorsal punctation, beetle submatt, male antennomeres 3 larger, with more rounded external margin, male protarsomere 4 with anterolateral hook-like seta larger than more laterally situated large seta, narrow apical half (in ventral view) of median lobe, with elongate apex in lateral view, and setation of paramere. It differs from E. kisli sp. n., see under E. kisli sp. n.
Surface sculpture: Head with very dense, coarse punctation (spaces between punctures 1-2 times size of punctures), finer and sparser anteriorly; diameter of punctures only slightly smaller than diameter of cells of microreticulation, of some punctures equal to it. Pronotum and elytra with slightly finer and more evenly distributed punctation than on head. Pronotum and elytra with more weakly impressed microreticulation than on head. Dorsal surface submatt due to strong punctation. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and rounded anteriorly, without anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Male: Antennomere 2 very small, stout, antennomere 3 strongly enlarged, evidently larger than other antennomeres, with rounded external margin, antennomeres 4-6 distinctly enlarged, antennomere 7 slightly enlarged ( Fig. 18A). Protarsomere 4 with very small (only slightly larger than laterally situated large seta), thin, slightly curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row (double apically) of 14 short setae and posterior row of 4 short setae (Fig. 18B). Abdominal ventrite 6 with 14-17 lateral striae on each side. Median lobe narrow in apical half and broad in basal one, with weak submedian constriction in ventral view and elongate apex in lateral view (Fig. 18C, D). Paramere distinctly longer than median lobe, without notch on dorsal side, with relatively long and dense subdistal setae and numerous spine-like setae on internal surface (Fig. 18E).
Etymology. The species is named for K. Ksionsek. The species name is a noun in the genitive case. Diagnosis. Beetle small, dark brown to piceous, with paler head or only its anterior part and pronotal sides, shiny; pronotum without lateral bead or with weak traces of lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, evidently curved anterolateral hook-like seta; median lobe with strong submedian constriction in ventral view and almost truncate apex in lateral view; paramere with strong notch on dorsal side and subdistal part elongate but broad, with numerous long, dense, thick, flattened setae, some of them curved at apex; setae of proximal part evident, long. The species is very similar to E. brahminensis Shaverdo, Hendrich & Balke, 2012, from which differs with shape of the median lobe apex and, especially, with setation of the subdistal part of the paramere: it has only thick, flattened setae. From E. mantembu Shaverdo, Hendrich & Balke, 2012, it differs with shape of the median lobe and stronger notch on dorsal side of the paramere.
Surface sculpture: Head with dense (spaces between punctures 1-3 times size of punctures) but fine punctation; diameter of punctures evidently smaller than diameter of cells of microreticulation. Pronotum with much sparser and finer punctation than on head. Elytra with extremely sparse and fine punctation, almost invisible. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum without lateral bead or with weak traces of lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, slightly convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Male: Antenna simple (Fig. 26A). Protarsomere 4 with large, thick, evidently curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of 11 and posterior row of 4 short setae (Fig. 26B). Abdominal ventrite 6 with 6-8 lateral striae on each side. Median lobe with strong submedian constriction in ventral view and almost truncate apex in lateral view (Fig. 26C, D). Paramere with strong notch on dorsal side and subdistal part elongate but broad, with numerous long, dense, thick, somewhat flattened setae, some of them curved at apex; setae of proximal part evident, long (Fig. 26E).
Female: Without evident differences in external morphology from males, except for abdominal ventrite 6 without striae.
Distribution. Papua New Guinea: East Sepik Province. This species is known only from localities near Lembena (Fig. 53).
Etymology. The name refers to the village of Lembena where this species was collected. The name is a noun in the nominative singular standing in apposition. Diagnosis. Beetle small, dark brown, with paler head and pronotum, shiny; pronotum without lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta; median lobe slender, with very weak submedian constriction in ventral view and broadly rounded, elongate apex in lateral view; paramere with shallow notch on dorsal side and subdistal part elongate but broad, with large brush of long, dense, thick, somewhat flattened setae, distal ones longer and curved at apex; setae of proximal part short, thin, almost invisible. The species is similar to E. lembena sp. n., from which differs with shape of the median lobe apex and shallow notch on dorsal side of the paramere. There are only two species on Yapen Island, which can be easily distinguished with size and dorsal sculpture: E. mantembu sp. n., small and shiny, and E. vladimiri, large and matt, as well as using shape and structure of the male genitalia.
Surface sculpture: Head with dense punctation (spaces between punctures 1-3 times size of punctures), evidently finer and sparser anteriorly; diameter of punctures smaller than diameter of cells of microreticulation. Pronotum with much sparser and finer punctation than on head. Elytra with extremely sparse and fine punctation, almost invisible. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum without lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, less rounded anteriorly, without anterolateral extensions. Blade of prosternal process lanceolate, relatively broad, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded or slightly truncate apically.
Male: Antenna simple (Fig. 25A). Protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of 11 short setae and posterior row of 4 short setae (Fig. 25B). Abdominal ventrite 6 with 8-11 lateral striae on each side. Median lobe slender, with very weak submedian constriction in ventral view and broadly rounded, elongate apex in lateral view (Fig. 25C, D). Paramere with shallow notch on dorsal side and subdistal part elongate but broad, with large brush of long, dense, relatively thick, somewhat flattened setae, distal ones longer and curved at apex; setae of proximal part short, thin, almost invisible (Fig. 25E).
Female: Without evident differences in external morphology from males, except for abdominal ventrite 6 without striae.
Distribution. Indonesia: Papua Province: Yapen Islands Regency. This species is known only from the type locality (Fig. 53).
Etymology. The name refers to the region Mantembu where this species was collected. The name is a noun in the nominative singular standing in apposition. Diagnosis. Beetle medium-sized, piceous, with dark brown clypeus and sides of pronotum; pronotum with lateral bead; male antennomeres simple; male protarsomere 4 with large, thin, slightly curved anterolateral hook-like seta; median lobe with submedian constriction in ventral view and elongate and broadly pointed apex in lateral view; paramere with distinct notch on dorsal side and subdistal part elongate, with brush of long, dense, thin setae. Size, dorsal sculpture, and structure of the male genitalia of this species strongly resemble those of E. bismarckensis sp. n., E. gorokaensis sp. n., and E. vovai sp. n., but the species can be easily distinguished from them with its simple, not modified, male antenna.
Surface sculpture: Head with relatively sparse punctation (spaces between punctures 1-4 times size of punctures); diameter of punctures smaller than diameter of cells of microreticulation. Pronotum and elytra with distinct punctation but finer and more evenly distributed than on head. Pronotum and elytra with relatively strongly impressed microreticulation, dorsal surface submatt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and extremely fine, sparse punctation, almost invisible, only slightly coarser and denser on two last abdominal ventrites.
Structures: Pronotum with lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and rounded anteriorly, without anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded.
Male: Antenna simple (Fig. 31). Protarsomere 4 with large, thin, slightly curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of more than 40 and posterior row of 13 thin, moderately long setae ( Fig. 2A). Abdominal ventrite 6 with 15-18 lateral striae on each side. Median lobe with submedian constriction in ventral view and elongate and broadly pointed apex in lateral view (Fig. 2B, C). Paramere with distinct notch on dorsal side and subdistal part elongate, with brush of long, dense, thin setae (Fig. 2D).
Female: Unknown. Distribution. Papua New Guinea: Eastern Highlands Province. This species is known only from the type locality (Fig. 53).
Etymology. The species is named after Mt. Michael where it was collected. The name is an adjective in the nominative singular. Additional material. 26 females with the same label as the holotype (NHMW, ZMS). These females might belong to of two species: E. pinocchio sp. n. and E. brahminensis, therefore, they are not included in the type series.

Exocelina pinocchio
Diagnosis. Beetle small, dark brown to piceous, with paler head and pronotal sides, shiny; pronotum without lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta; in ventral view, median lobe with strong submedian constriction and ventral sclerite apically divided in three parts and in lateral view, with very strongly protruding apex, forming long thin prolongation; paramere with strong notch on dorsal side and subdistal part elongate but broad, with numerous long, dense setae, thinner and shorter distally and thicker, longer, and curved at apex proximally. This species can be easily distinguished from all small, shiny, with simple antennae, and without pronotal bead species (e.g., E. brahminensis or E. lembena sp. n.) with shape of its median lobe.
Surface sculpture: Head with dense (spaces between punctures 1-3 times size of punctures) but fine punctation; diameter of punctures evidently smaller than diameter of cells of microreticulation. Pronotum with much sparser and finer punctation than on head. Elytra with extremely sparse and fine punctation, almost invisible. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum without lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and rounded anteriorly, without anterolat-eral extensions. Blade of prosternal process lanceolate, relatively broad, slightly convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Male: Antenna simple (Fig. 24A). Protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of 12 and posterior row of 4 short setae (Fig. 24B). Abdominal ventrite 6 with 8-10 lateral striae on each side. Median lobe with strong submedian constriction and ventral sclerite apically divided in three parts in ventral view and with very strongly protruding apex, forming long thin prolongation in lateral view (Fig. 24C, D). Paramere with strong notch on dorsal side and subdistal part elongate but broad, with numerous long, dense setae, thinner and shorter distally and thicker, longer, and curved at apex proximally (Fig. 24E).
Female: Without evident differences in external morphology from males, except for abdominal ventrite 6 without striae.
Distribution. Papua New Guinea: Madang Province. This species is known only from the type locality (Fig. 53).
Etymology. The species is named for a fictional character from the book "The Adventures of Pinocchio" by Carlo Collodi because the apex of its median lobe has a prolongation like a "nose". The name is a noun in the nominative singular standing in apposition. Diagnosis. Beetle middle-sized, piceous, with paler pronotum (especially on margins) and head, dorsally with evident punctation, submatt; pronotum with distinct lateral bead; male antennomeres simple; male protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta; 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. The species is very similar to E. astrophallus , except for media lobe without notch on left side and larger and more strongly curved anterolateral hook-like seta.
Surface sculpture: as in E. astrophallus. Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, without anterolateral extensions. Blade of prosternal process lanceolate, rather narrow, strongly convex, with distinct bead and few setae; neck and blade of prosternal process evenly jointed.
Female: Without evident differences in external morphology from male, except for abdominal ventrite 6 without striae.
Distribution. Papua New Guinea: East Sepik Province. This species is known only from localities near Lembena (Fig. 53).
Etymology. This species was mistaken for E. astrophallus due to their external similarity. The name is a noun in the nominative singular standing in apposition. Diagnosis. 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 very similar to E. bifida Shav-erdo, Hendrich & Balke, 2012, except for structure of genitalia: apical lobes slender and more deeply separated, dorsal extension prominent but not deeply cut.
Surface sculpture: Punctation and microreticulation as in E. bifida. Structures: Pronotum without lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, anteriorly less rounded, smooth, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively broad, slightly convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Holotype: TL-H 3.7 mm, TL 4.15 mm, MW 2.0 mm. Female: Without evident differences in external morphology from male, except for abdominal ventrite 6 without striae.
Distribution. Papua New Guinea: Sandaun Province, Mekil. This species is known only from the type locality (Fig. 53).

Exocelina pseudoedeltraudae
Diagnosis. Beetle medium-sized, piceous, submatt; pronotum with distinct lateral bead; male ventrite 6 slightly to distinctly concave apically; male antennomeres 3-5 distinctly enlarged, almost equal in size and shape, antennomeres 6-8 enlarged; male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta; 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 very similar to E. edeltraudae Shaverdo, Hendrich & Balke, 2012, from which differs with less shinier dorsal surface, due to stronger punctation and microreticulation, with larger and more rounded male antennomeres 3-5 (for male antennomeres 3 and 4, ratio width/length: > 1.0) and apex of median lobe narrower in lateral view.
Surface sculpture: Head with dense, coarse punctation (spaces between punctures 1-3 times size of punctures), especially on vertex. Pronotum with punctation finer, sparser, and more evenly distributed than on head. Elytra with punctation finer, sparser than on pronotum. Pronotum and elytra with strongly impressed microreticulation, dorsal surface submatt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, rounded and smooth anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded, or slightly truncate, or concave apically.
Distribution. Papua New Guinea. The species is known only from Hela Province (Fig. 53).
Etymology. In an earlier work (Shaverdo et al. 2012), this species was mistaken for E. edeltraudae. The name is a noun in the nominative singular standing in apposition. Diagnosis. Beetle small, dark brown to piceous, with paler anterior part of head and pronotal sides, shiny; pronotum without lateral bead; male antennomeres 5-10 slightly stout; male protarsomere 4 with large, slender, strongly curved anterolateral hook-like seta; median lobe with submedian constriction in ventral view and elongate apex in lateral view; paramere with notch on dorsal side and subdistal part elongate, with large brush of two kinds of setae: upper setae thin and less numerous and lower setae long, thick, somewhat flattened, and curved at apex; setae of proximal part shorter, thinner, less evident. The species is very similar to E. eme Shaverdo, Hendrich & Balke, 2012 except for more weakly impressed dorsal microreticulation, especially on pronotum, beetle dorsally slightly shinier, as well as for structure and setation of genitalia: median lobe with stronger submedian constriction and symmetrical apex in ventral view; subdistal part of paramere with upper thin setae less numerous making brush smaller.
Surface sculpture: Head with dense punctation (spaces between punctures 1-3 times size of punctures), finer and sparser anteriorly; diameter of punctures smaller than diameter of cells of microreticulation. Pronotum with much finer and sparser punctation than on head. Elytra with very sparse and fine punctation, almost invisible. Head, pronotum, and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum without lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and less rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, slightly convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Male: Antennomeres 5-10 slightly stout (Fig. 27A). Protarsomere 4 with large, slender, strongly curved anterolateral hook-like seta. Protarsomere 5 ventrally with anterior row of 14 and posterior row of 5 short setae (Fig. 27B). Abdominal ventrite 6 with 5-6 lateral striae on each side. Median lobe with submedian constriction and symmetrical apex in ventral view and elongate apex in lateral view (Fig. 27C, D); paramere with notch on dorsal side and subdistal part elongate, with large brush of two kinds of setae: upper setae thin and less numerous and lower setae long, thick, somewhat flattened, and curved at apex; setae of proximal part shorter, thinner, less evident (Fig. 27E).
Female: Without evident differences in external morphology from males, except for abdominal ventrite 6 without striae.
Distribution. Papua New Guinea: Sandaun Province. This species is known only from Mianmin region (Fig. 53).
Etymology. This species was mistaken for E. eme due to their external similarity. The name is a noun in the nominative singular standing in apposition.  E. ketembang Balke, 1998. Diagnosis. Beetle medium-sized, dark brown to piceous, slightly submatt; pronotum with lateral bead; male antennomeres 3-5 evidently enlarged, slightly rounded, antennomeres 3, 4 almost equal in size, antennomere 5 slightly smaller, with external margin almost straight, antennomere 6 somewhat enlarged; male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta; median lobe broad, with very weak submedian constriction in ventral view and thin apex in lateral view, apex with small lateral setae; paramere with notch on dorsal side and subdistal part elongate, with numerous, long, thick, curved at apex setae. This species is similar to E. simbaiarea sp. n., E. tariensis sp. n., and E. jimiensis sp. n., see differences under their diagnoses.
Surface sculpture: Head with dense punctation (spaces between punctures 1-3 times size of punctures), evidently finer and sparser anteriorly; diameter of punctures equal or smaller than diameter of cells of microreticulation. Pronotum with finer, sparser, and more evenly distributed punctation than on head. Elytra with very sparse and fine punctation. Pronotum and elytra with relatively weakly impressed microreticulation, dorsal surface slightly submatt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and not rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lan-ceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 slightly truncate apically.
Etymology. The species is named after Sandaun Province where it was collected. The name is an adjective in the nominative singular. Additional material: 10 females with the same label as the holotype (ZSM), these females might belong to three species: E. simbaiarea sp. n. and two species from the E. broschii-and E. rivulus-groups.

Exocelina simbaiarea
Diagnosis. Beetle medium-sized, blackish brown, with brown head and pronotal sides, slightly submatt; pronotum with lateral bead; male antennomeres 3-5 evidently enlarged, slightly rounded, antennomeres 3, 4 almost equal in size, antennomere 5 slightly smaller, with external margin rounded, antennomere 6 somewhat enlarged; male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta; median lobe with very weak submedian constriction in ventral view and apex relatively short and slightly broadened in lateral view; paramere with notch on dorsal side and side and subdistal part elongate, with numerous, long, thick, curved at apex setae. The species is similar to E. sandaunensis sp. n., except for slightly matter pronotum, more striated abdominal ventrite 6, male antennomere 5 with external margin rounded, and apex of median lobe shorter and broader. See also under diagnosis of E. jimiensis sp. n.
Description. Size and shape: Beetle medium-sized (TL-H 3.65 mm, TL 4.1 mm, MW 1.95 mm), with oblong-oval, broadest at elytral middle. Coloration: Head brown, darker posterior eyes and at middle; pronotum with dark brown disc and brown sides; elytra blackish brown, with reddish sutural lines; head appendages and legs reddish, legs darker distally (Fig. 40).
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and not rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 slightly truncate apically.
Female: Antennae simple, abdominal ventrite 6 without striae. Distribution. Papua New Guinea: Madang Province. This species is known from the type locality (Fig. 53).
Etymology. The species is named after the Simbai area where it was collected. The name is a noun, combination of two words: "Simbai" and "area", in the nominative singular standing in apposition. Diagnosis. Beetle small, broadly oval, piceous, with paler head and pronotum or only with pale anterior part of head and pronotal sides, submatt; pronotum with lateral bead; male antennomeres simple; male protarsomere 4 with medium-sized, slender, slightly curved anterolateral hook-like seta; median lobe with apical discontinuity and deeply concave, bilobed apex in ventral view; paramere without notch on dorsal side, with triangular basal part and thin subdistal part, setae inconspicuous, sparse, thin, and relatively short. This species is similar only to E. vladimiri and probably related to it. In the group, only these two species have outline of the median lobe with apical, not submedial, discontinuity in ventral view and broadly oval habitus. Exocelina vladimiri can be distinguished from E. skalei sp. n. with larger size, absence of the pronotal bead, less concave apex of the median lobe, and paramere setation.
Surface sculpture: Head with dense, coarse punctation (spaces between punctures 1-2 times size of punctures); diameter of some punctures equal diameter of cells of microreticulation. Pronotum and elytra with punctation finer and more evenly distributed than on head but very evident. Pronotum and elytra with evident microreticulation, dorsal surface submatt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, smooth and rounded anteriorly, without small anterolateral extensions. Blade of prosternal process lanceolate, broad, slightly convex, with rounded apex, distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Holotype: TL-H 3.25 mm, TL 3.6 mm, MW 1.9 mm. Female: Without evident differences in external morphology from male, except for abdominal ventrite 6 without striae.
Distribution and habitat. Indonesia: West Papua Province: Kaimana Regency. This species is known only from the type locality (Fig. 53). The species was collected from a small rock pool, without any vegetation (Fig. 54).
Etymology. The species is named for Andre Skale who collected this species, with our sincere thanks for presenting this interesting species for study. The species name is a noun in the genitive case. Diagnosis. Beetle medium-sized, piceous with paler head and pronotum, submatt; pronotum with distinct lateral bead; male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta. The species is similar to E. munaso (Shaverdo, Sagata & Balke, 2005) because of shape of the median lobe (with large lateral folds in ventral view) and paramere (without notch on dorsal side). However, it differs from E. munaso with smaller size, evidently narrower blade of prosternal process, male antennomeres 5-7 evidently enlarged, antennomeres 4, 8, 9 slightly enlarged, medial lobe much narrower, submedian constriction evident in ventral view, apex of median lobe almost rounded and not curved downwards in lateral view, and setae of paramere more numerous.
Surface sculpture: Head with dense punctation (some punctures conjoint or spaces between most of them 1-3 times size of punctures), evidently finer and sparser anteriorly and posteriorly; diameter of punctures equal to diameter of cells of microreticulation. Pronotum with finer, slightly sparser, and more evenly distributed punctation than on head. Elytra with punctuation slightly coarser and denser than on pronotum. Head, pronotum and elytra with strong microreticulation and punctation, dorsal surface submatt. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and rounded anteriorly, without anterolateral extensions. Blade of prosternal process lanceolate, very narrow, convex, with distinct lateral bead and few setae, apex of blade slightly but distinctly bent upwards; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 slightly truncate apically.
Etymology. The species is named after the type locality: Tabubil. The name is an adjective in the nominative singular. Diagnosis. Beetle medium-sized, blackish brown, with brown clypeus and pronotal sides, submatt; pronotum with lateral bead; male antennomeres 3-5 evidently enlarged, with external margin more expanded, antennomeres 3, 4 almost equal in size, antennomere 5 slightly smaller, antennomere 6 somewhat enlarged; male protarsomere 4 with large, slender, slightly curved upwards anterolateral hook-like seta, with pointed apex; median lobe broad, with very weak submedian constriction in ventral view and thin apex in lateral view, apex with small lateral setae; paramere with notch on dorsal side and subdistal part small, elongate, with not numerous, long, thick, almost straight setae. The species is similar to E. sandaunensis sp. n. and E. simbaiarea sp. n., except for more robust habitus, slightly matter pronotum, larger male antennomeres 3-5, with external margin more expanded, pointed and slightly curved upwards anterolateral hook-like seta of male protarsomere 4, subdistal part with setae less numerous and almost straight.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and very slightly rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 slightly truncate apically.
Female: Unknown. Distribution. Papua New Guinea: Hela Province. This species is known only from the type locality (Fig. 53).
Etymology. The species is named after the village of Tari where it was collected. The name is an adjective in the nominative singular. Diagnosis. Beetle medium-sized, dark brown to piceous, with paler clypeus, vertex, and pronotal sides, matt; pronotum with distinct lateral bead; male antennomeres 3-5 evidently enlarged, almost equal in size, antennomeres 5 slightly rectangular, antennomeres 6 and 7 somewhat enlarged; male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta; median lobe with weak submedian constriction and apex evidently concave in ventral view and with apex distinctly pointed in lateral view; paramere with shallow notch on dorsal side and subdistal part elongate, with numerous, dense, more or less long, thin setae. The species is similar to E. gorokaensis sp. n., from which differs with duller dorsal surface due to denser punctation and stronger microreticulation, as well as larger and sometimes less rounded male antennomeres 3-5, paramere with shallow notch on dorsal side, and smaller median lobe, with apex evidently concave in ventral view and distinctly pointed in lateral view. Also it is similar to E. bismarckensis sp. n. from which differs with broader and more oval habitus, less rounded male antennomeres 3-5, narrower median lobe, with apex less rounded and evidently concave in ventral view and stronger pointed in lateral view.
Surface sculpture: Head with very dense, coarse punctation (spaces between punctures 1-2 times size of punctures). Pronotum with punctation finer than on head. Elytra with punctation sparser than on pronotum. Pronotum and elytra with rather strongly impressed microreticulation, dorsal surface matt. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process and neck of prosternal process with distinct ridge, smooth and not rounded anteriorly, with small anterolateral extensions. Blade of prosternal process lanceolate, relatively narrow, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded or slightly truncate apically.
Etymology. The species is named for brother of the senior author, Vladimir (Vova) Shaverdo, with her sincere thanks for his help and interest in her life. Diagnosis. Beetle small, with head and pronotum red to reddish brown and elytra dark brown, shiny; pronotum with narrow, in some specimens indistinct lateral bead; male antennomeres modified: antennomeres 3-5 larger and more rounded than other, antennomeres 6, 7 somehow enlarged; male protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta; median lobe slender, with strong submedian constriction in ventral view and elongate apex in lateral view; paramere with strong notch on dorsal side and subdistal part slightly elongate, broad, with long, dense, relatively thick setae. The species is similar to the complex of the following species: E. edeltraudae, E. pseudoedeltraudae sp. n., E. jimiensis sp. n., E. tariensis sp. n., E. simbaiarea sp. n., and E. sandaunensis sp. n. But it differs from all of them with its smaller size, narrow pronotal lateral bead, less modified male antennomeres, and structure and setation of the male genitalia.
Surface sculpture: Head with dense punctation (spaces between punctures 1-3 times size of punctures), evidently finer and sparser anteriorly; diameter of punctures smaller than diameter of cells of microreticulation. Pronotum with much sparser and finer punctation than on head. Elytra with extremely sparse and fine punctation, almost invisible. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxa distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and fine sparse punctation, coarser and denser on two last abdominal ventrites.
Structures: Pronotum with narrow lateral bead. Some specimens with pronotal lateral bead indistinct and/or reduced at posterior angles. Base of prosternum and neck of prosternal process with distinct ridge, smooth and slightly rounded anteriorly, without anterolateral extensions. Blade of prosternal process lanceolate, relatively broad, convex, with distinct lateral bead and few setae; neck and blade of prosternal process evenly jointed. Abdominal ventrite 6 broadly rounded apically.
Female: Without evident differences in external morphology from males, except for simple antennae and abdominal ventrite 6 without striae.
Distribution. Papua New Guinea. This species is known only from Madang Province (Fig. 53).
Etymology. The name refers to the village of Wannang where this species was first discovered. The name is an adjective in the nominative singular.
Morphological notes. Beetles are smaller (TL-H 3.1-3.55 mm, TL 3.45-3.85 mm, MW 1.7-1.9 mm) than ones from the type locality: IN: Papua, Jayawijaya, Borme, Tarmlu. Also they have a slightly different shape of the median lobe of the aedeagus: the apex is less concave and narrower in ventral view and the right lateral side broader than in E. bifida (Fig. 20C). Differences in the setation of the protarsi and parameres are most likely within in the limits of intraspecific variability (Fig. 20B, E). In order to establish the status of these specimens, additional material from the border area, IN: Papua, Jayawijaya / PNG: Sandaun, is needed.
Distribution. Central part of the New Guinea Island, i.e., Indonesia: Papua Province, Jayawijaya Regency and Papua New Guinea: Sandaun Province.
Distribution. Papua New Guinea. This is one of the most widely distributed species in Papua New Guinea. It is known from the Momase Region: Sandaun, East Sepik, Madang, and Morobe Provinces (Shaverdo et al. 2012) and, now, also from Highlands Region: Eastern Highlands Province. Morphological notes. Most specimens are darker than the holotype, piceous, with dark brown head and pronotal sides. It is obvious that the holotype, with its dark brown dorsal coloration, is a slightly teneral specimen. Some variability (narrower apex in lateral view) in the shape of the apical part of the median lobe is observed.

Exocelina knoepfchen
Distribution. Indonesia: West Papua Province: Manokwari Regency. So far, this species has been known only as its holotype from Arfak Mountains. The present records are an extension of the known distribution of the species to the northwest.
Distribution. Indonesia: Papua Province: Nabire, Paniai, and Jayapura Regencies. So far, this species has been known only from the Nabire-Enarotali region. The present record is an extension of the known distribution of the species to the northeast.

Key to all described species of the Exocelina ekari-group
This key is a modified version of the key to species of the E. ekari-group from Shaverdo et al. (2012). It is based mostly on male characters. In many cases females cannot be assigned to species due to the similarity of their external and internal structures (for female genitalia see figs 17a and 17b in Shaverdo et al. (2005) and fig. 7C in Shaverdo et al. (2013)). Some species are rather similar in external morphology, therefore, in most cases the male genitalia need to be studied for reliable species identifications. Numbers in brackets refer to the order of the new species descriptions above.

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Beetle reddish brown to brown dorsally, median lobe more slender in ventral view, paramere with strong notch on dorsal side and subdistal part short and large (figs 26, 1C-F in Shaverdo et al. 2012 (Fig. 36), median lobe broader in ventral view, paramere with weaker notch on dorsal side and subdistal part elongate ( Fig. 4C-F)  Male antennomeres antennomeres 3-9 stout, with 4-5 slightly larger than other, median lobe with very strong median constriction and proximal part very broad in ventral view, apex of median lobe broad, slightly concave in middle and twisted at both sides in ventral view and shortly pointed in lateral view, subdistal part of paramere elongate, with a large brush of long, dense, thin setae, proximal setae almost invisible (Fig. 5)  Male antennomeres 5-7 evidently enlarged, antennomeres 4, 8, 9 slightly enlarged, male protarsomere 4 with large, slender, evidently curved anterolateral hook-like seta (Fig. 6A, B), median lobe and paramere as in Fig. 6C (Fig. 39), protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta (Fig. 7B), paramere with strong notch on dorsal side and subdistal part slightly elongate and broad (Fig. 7E), median lobe as in Fig. 7C Median lobe with very strong median constriction and proximal part very broad in ventral view, apex of median lobe pointed and strongly curved downwards in lateral view (Figs 8E, 9D), male abdominal ventrite 6 slightly to distinctly concave apically (Fig. 8C)  Male antennomeres 3-5 smaller and less rounded, for antennomeres 3 and 4, ratio width/length: < 0.92 (Fig. 8A), dorsal surface slightly shinier due to weaker punctation and microreticulation (Fig. 41), apex of median lobe broader in lateral view (Fig. 8D)  Beetle dorsally matter, especially pronotum, male antennomeres 3-5 more strongly enlarged, with external margin more expanded, male protarsomere 4 with large, slender, slightly curved upwards anterolateral hook-like seta, with pointed apex, subdistal part of paramere with setae less numerous (Fig. 11)  Male antennomere 5 with external margin rounded (Fig. 12A), abdominal ventrite 6 with 13-14 striae, apex of median lobe shorter (Fig. 12D)  Male antennomere 5 with external margin almost straight (Fig. 13A), abdominal ventrite 6 with 7-10 striae, apex of median lobe longer (Fig. 13D)  Paramere with distinct notch on dorsal side (Fig. 14E). Dorsal punctation finer (Fig. 46)  Habitus more elongate, often with subparallel sides (Fig. 47), apex of median lobe almost rounded in ventral view, with curved part gradually pointed in lateral view (Fig. 15D) (Fig. 48), apex of median lobe not rounded, distinctly concave in ventral view, with curved part more sharply pointed in lateral view (Fig. 16D) (Fig. 49), antennomere 3 smaller, more triangular, median lobe with apex more rounded in lateral view, male protarsomere 4 with anterolateral hook-like seta smaller than more laterally situated large seta, paramere with less numerous subdistal long setae and, especially, internal spines (Fig. 17) (Fig. 50), male antennomere 3 larger, with external margin more rounded, median lobe with apex pointed in lateral view, male protarsomere 4 with anterolateral hook-like seta larger than more laterally situated large seta, paramere with more numerous subdistal long setae and internal spines (Fig. 18) Beetle dorsally piceous, with elytral punctation fine but distinct, male protarsomere 4 with thick anterolateral hook (                    Habitats Shaverdo et al. (2012) provided a summary of habitats for New Guinea Exocelina species. All the species with only one exception (Shaverdo et al. 2013) are running water associated, but avoid the current, i.e., preferred microhabitats are small, quiet backflows, tiny puddles at the edge of streams and creeks, rock holes filled with water ( Fig. 54), and other similar situations. Further information was provided in this wiki site: http://zsm-entomology.de/w/index.php?title=Coleoptera_Fieldwork&oldid=862 West Papua). Permission to conduct fieldwork was provided by Department of Forestry/Nature Conservation in Manokwari and Jayapura. 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 the senior author. Michael Balke was supported by the UK Darwin Initiative and the German Science Foundation (various projects since BA2152/2-1).