Research Article |
Corresponding author: Helena Shaverdo ( shaverdo@mail.ru ) Academic editor: Mariano Michat
© 2016 Helena Shaverdo, Katayo Sagata, Michael Balke.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Shaverdo H, Sagata K, Balke M (2016) Description of two new species of the Exocelina broschii-group from Papua New Guinea, with revision and key to all representatives of this species group (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 577: 125-148. https://doi.org/10.3897/zookeys.577.7254
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Two new species of Exocelina Broun, 1886 from Papua New Guinea are described herein: E. mondmillensis sp. n. and E. pseudomarinae sp. n. They are placed into the E. broschii-group based on the shovel/fork-like ventral sclerites of their median lobe. While the former has rather distinct combination of the morphological characters (inconspicuous dorsal punctation, thin apex of the median lobe and ventral sclerite of the median lobe with two tips of different length), the latter is very similar to already described species E. marinae (Shaverdo, Sagata & Balke, 2005). All described species of the group are revised and a key to their identification is provided. Important diagnostic characters (habitus, color, protarsomeres 4–5, median lobes, and parameres) are illustrated. Data on the distribution of all species of the group are given showing that its representatives occur only in Papua New Guinea and most of them are widely distributed in it central part.
Exocelina broschii-group, Copelatinae , Dytiscidae , new species, Papua New Guinea
This paper continues our previous studies on the New Guinea species of the genus Exocelina Broun, 1886 (
Here, we provide a detailed diagnosis of the E. broschii-group, describe two new species, review the known species providing new faunistic data, and present a key to the species and map of their distribution.
We provided electronic resources for the species treated here in the form of species pages, which were automatically created by ZooKeys on the species-id.net portal with the publication of this article. This wiki engine based site provides for example high resolution art work and can be improved interactively should new data become available. By providing these resources, we hope to help creating a more user-friendly, sustainable taxonomy as suggested by
Including the results of this work, 91 Exocelina species are described from New Guinea and 144 worldwide.
The present work is based on the material from the following collections:
BMNH
The
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., “264 DNA M. Balke”). All types of the herein described species are provided with red labels. The female specimens, identification of which is difficult or sometimes impossible, were included in the type series only when collected with males of respective species and did not differ morphologically from them. If two or more morphologically similar species were collected together (i.e., males found together), their females were not included in the types 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 very recent (
Measurements were taken with a Wild M10 stereomicroscope choosing the smallest and the largest specimens within and among the populations. The following abbreviations were used: TL (total body length), TL-H (total body length without head), MW (maximum body width), and hw (handwritten). The number of ventral setae on the male protarsomere 5 is given for only 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, 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 descriptions.
The terminology to denote the orientation of the genitalia (ventral for median lobe and dorsal and external for paramere) follows
The representatives of the E. broschii-group share the following diagnostic characters:
beetles small or middle-sized (TL-H 3.2–4.15 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 brown to piceous, mainly uniform, sometimes with paler head and pronotum and darker elytra;
microreticulation and punctation of dorsal surface very fine to strongly impressed, so that beetles shiny to matt dorsally;
metacoxae and abdominal ventrites 1–5 (and 6 in males) with thin, almost longitudinal striae/strioles;
pronotum and elytra without striae or strioles;
pronotum with lateral bead;
male antennomeres not modified, antennomere 2 larger than antennomere 3;
male protarsomeres 1–3 not expanded laterally;
male protarsomere 4 cylindrical, narrow, with 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 continuous outline in ventral and lateral view;
ventral sclerite of median lobe not deeply divided in the middle, apically forming a shovel/fork-like structure with two apices;
apical part of median lobe with numerous setae;
paramere without notch on dorsal side;
paramere with long setae occupying whole dorsal side.
Representatives of this species group are recorded only from Papua New Guinea (PNG).
Representatives of this species group are recorded only from Papua New Guinea (PNG).
1. | Exocelina broschii (Balke, 1998) | PNG Madang, Eastern Highlands |
2. | Exocelina hintelmannae (Shaverdo, Sagata & Balke, 2005) | PNG: Simbu, Eastern Highlands, Gulf |
3. | Exocelina marinae (Shaverdo, Sagata & Balke, 2005) | PNG: Sandaun, Hela |
4. | Exocelina mondmillensis sp. n. | PNG: Western Highlands, Enga, Madang |
5. | Exocelina pseudomarinae sp. n. | PNG: Hela |
Copelatus (Papuadytes) broschii Balke, 1998: 327;
Papuadytes broschii (Balke, 1998):
Papuadytes broschii (Balke, 1998):
Exocelina broschii (Balke, 1998):
Exocelina undescribed sp. MB1520:
Papua New Guinea: Madang Province, Finisterre Range, Moro, approximately 5°42'47.6"S; 146°03'40.1"E.
Holotype: male “Stn. No. 82”, “NEW GUINEA: Madang Dist., Finisterre Mts. Moro.C.5550ft. 30.x.-15.xi.1964.”, “M.E. Bacchus. B.M. 1965-120”, “Holotypus” [red], “Copelatus broschii sp.n. Balke des. 1997” (BMNH). Paratypes: 4 males, 1 female with the same labels as the holotype, except for “Paratypus Copelatus broschii sp.n. Balke des. 1997” (BMNH,
Madang: 1 male “Stn. No. 82”, “NEW GUINEA: Madang Dist., Finisterre Mts. Moro.C.5550ft. 30.x.-15.xi.1964.”, “M.E. Bacchus. B.M. 1965-120”, “Paratypus Copelatus broschii sp.n. Balke des. 1997” (BMNH) – although this specimen is with the paratype label, it is not included in the type material of the original description in
Females of doubtful identity. Eastern Highlands: 12 females “Papua New Guinea: Eastern Highlands, Akameku - Brahmin, Bismarck Range, 700m, 24.xi.2006, 05.52.754S 145.23.209E, Balke & Kinibel (PNG 109)” (
(original description in
Lateral view of median lobe of (A–E) Exocelina broschii (Balke, 1998) and (F, G) E. hintelmannae (Shaverdo, Sagata & Balke, 2005). A, B Madang, Simbai area, PNG 153 C Madang, Adalbert Mts., Keki, PNG 52 D Madang, Adalbert Mts., below Keki, PNG 53 E Eastern Highlands, Bena Bridge, PNG 164 F, G Simbu/ Eastern Highlands, Crater Mountain, Sera-Herowana, Hulene River, PNG 017. Setae not shown.
Variability: Beetles vary in size, kind of dorsal punctation (Figs
Papua New Guinea: Madang and Eastern Highlands Provinces (Fig.
Papuadytes hintelmannae Shaverdo, Sagata & Balke, 2005: 272.
Exocelina hintelmannae (Shaverdo, Sagata & Balke, 2005):
Papua New Guinea: border Simbu and Eastern Highlands Provinces, Crater Mountain, between Wara Sera Station and Herowana Village, Hulene River, approximately 06°43.4'S; 145°05.6'E.
Holotype: male “264 DNA M Balke” [green], “PNG Simbu / EHPr. Crater Mountain, Sera - Herowana, Wara Hulene, 1000 m, 16IX2002, Balke & Sagata (PNG 17)”, “Holotypus Papuadytes hintelmannae sp.n. des. H. Shaverdo, K. Sagata & M. Balke, 2005” (BMNH). Paratypes: 1 male “256 DNA M Balke” [green], “Papua New Guinea Simbu / EHPr. Crater Mountain, Wara Sera Station, 800 m, 14IX2002, Balke & Sagata (PNG 10)”, “Paratypus Papuadytes hintelmannae sp.n. des. H. Shaverdo, K. Sagata & M. Balke, 2005” (
Simbu/EHL: 1 female “Papua New Guinea: Simbu / EHPr. Crater Mountain, Wara Sera Station, 820m, 14IX2002, Balke & Sagata, (PNG 8)” (
(original description in
Female: Without evident differences in external morphology from males, except for not modified protarsi and abdominal ventrite 6 without striae.
Variability: Beetles vary in shape of the median lobe and its ventral sclerite: mainly shape and length of its left lobe (Fig.
Papua New Guinea: Simbu, Eastern Highlands, and Gulf Provinces (Fig.
Papuadytes marinae Shaverdo, Sagata & Balke 2005: 272.
Exocelina marinae (Shaverdo, Sagata & Balke, 2005):
Papua New Guinea: Sandaun Province, trail from Telefomin to Eliptamin.
Holotype: male “Papua N. G.: Sandaun Prov. Telefomin, 16.–17.5.1998 trail to Eliptamin 1700-1800 m; leg. Riedel”, “Holotypus Papuadytes marinae sp.n. des. H. Shaverdo, K. Sagata & M. Balke, 2005” (
Sandaun: 4 males, 6 females “Papua New Guinea: Sandaun, Mianmin, 670m 20.x.2008, 4.53.292S 141.34.118E, Ibalim (PNG 191)”, two males with additional green labels “DNA M.Balke 3733” and “DNA M.Balke 3734” (NHMW,
Hela: 1 male “Papua New Guinea: Southern Highlands, Tari to Koroba, 1600m, 15.v.2006, 05.46.500S 142.50.000E, Balke (PNG 65)”, “DNA M.Balke 1291” [green] (
(original description in
Female: Without evident differences in external morphology from males, except for not modified protarsi and abdominal ventrite 6 without striae.
Variability: Beetles small to medium-sized: for Mianmin populations: TL-H 3.3–3.55 mm, TL 3.6–3.95 mm, MW 1.75–1.95 mm; for the holotype (Telefomin): TL-H 3.6 mm, TL 4.1 mm, MW 1.9 mm (Fig.
Papua New Guinea: Sandaun and Hela Provinces (Fig.
Papua New Guinea: Western Highlands Province, 5 km SE from Minj, Mondmill, 05°56.80'S, 144°39.90'E.
Holotype: male “Papua New Guinea: Western Highlands, Mondmill, 5 Km SE Minj, small pools near creek, 1741 m, 12.vi.2006, 05.56.801S 144.39.898E, John (PNG 77)” (
Western Highlands: 142 females “Papua New Guinea: Western Highlands, Kurumul, 6 Km SW Kudjip, small stream, 1580 m, 13.vi.2006, 05.53.426S 144.36.600E, John (PNG 78)” (
Beetle medium-sized, brown to piceous, with reddish head and pronotal sides, shiny; median lobe with slightly curved downwards apex but thin in lateral view and ventral sclerite with two unequal apices (left one long, narrow and curved and right one short, broad and more or less rounded). The species is similar to E. marinae and E. pseudomarinae sp. n. in shape of the ventral sclerite of median lobe, but distinctly differs from them in having fine, inconspicuous punctation and weak microreticulation on the dorsal surface and a thin tip of the median lobe. From E. broschii and E. hintelmannae, it can be distinguished by the shape of the median lobe (thin apex) and its ventral sclerite (very long left lobe), and from the former also by finer elytral punctation.
Size and shape: Beetle medium-sized (TL-H 3.5–4.1 mm, TL 3.9–4.5 mm, MW 1.8–2.2 mm), with oblong-oval habitus, broadest at elytral middle. Coloration: Head reddish brown to piceous, usually darker medially and posterior to eyes, sometimes almost uniform; pronotum with brown to piceous medially and reddish brown to brown sides; elytra brown to piceous, usually with narrow reddish sutural lines; head appendages and legs yellowish to reddish, legs distally darker, especially metathoracic legs (Fig.
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 head. Elytra with very sparse and fine punctation, almost invisible. Pronotum and elytra with weakly impressed microreticulation, dorsal surface shiny. Head with microreticulation stronger. Metaventrite and metacoxae distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and very fine and sparse punctation.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, slightly rounded anteriorly. 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 or slightly truncate.
Male: Antenna simple (Fig.
Holotype: TL-H 4.05 mm, TL 4.45 mm, MW 2.15 mm.
Female: Without evident differences in external morphology from males, except for not modified protarsi and abdominal ventrite 6 without striae.
Variability: Beetles vary mainly in shape of the ventral sclerite of the median lobe as shown in Fig.
Papua New Guinea: Western Highlands, Enga, and Madang Provinces (Fig.
The name refers to Mondmill, the type locality, where the species was found in great numbers. The name is an adjective in the nominative singular.
Exocelina undescribed sp. MB1287:
Papua New Guinea: Hela Province, Tari, 05°50.38'S, 142°55.90'E.
Holotype: male “Papua New Guinea: Southern Highlands, Tari (trickle in gardenland), 1700m, 12.v.2006, 05.50.383S 142.55.901E, Balke (PNG 58)”, “DNA M.Balke 1287” [green] (
Beetle medium-sized, brown to dark brown, with reddish head and pronotal sides, submatt; median lobe with apex strongly curved downwards in lateral view and ventral sclerite with two unequal apices (left one long, narrow and curved apically and right one short, broad and more or less strait). The species is similar to E. marinae Shaverdo, Sagata & Balke, 2005 from which distinctly differs in larger size, sparser and finer punctation and weaker microreticulation of the dorsal surface, and strongly curved apex of the median lobe, which is similar to that of E. hintelmannae Shaverdo, Sagata & Balke, 2005. The specimen of E. marinae from Tari-Koroba, though large in size and with the same distribution, has a distinctly stronger sculpture on the dorsal surface and a median lobe with only a slightly curved apex in lateral view and a narrower right lobe of the ventral sclerite. Therefore, it can be easily distinguished from E. pseudomarinae sp. n.
Size and shape: Beetle medium-sized (TL-H 3.55–4.05 mm, TL 3.85–4.4 mm, MW 1.9–2.1 mm), with oblong-oval habitus, broadest at elytral middle. Coloration: Head reddish brown, brown to dark brown medially and posterior to eyes; pronotum with brown to dark brown medially (to piceous – narrow part on disc) and reddish brown sides; elytra brown to dark brown, sometimes with narrow reddish brown sutural lines; head appendages yellowish, legs yellowish to reddish, distally darker, especially metathoracic legs (Fig.
Surface sculpture: Head with very dense punctation (spaces between punctures 1–2 times size of punctures), evidently finer and sparser anteriorly; diameter of punctures smaller than diameter of cells of microreticulation or equal to it. Pronotum and elytra with punctation sparser and finer than on head, but very distinct. Pronotum and elytra with distinct microreticulation, dorsal surface submatt. Head with microreticulation stronger. Metaventrite and metacoxae distinctly microreticulate, metacoxal plates with longitudinal strioles and transverse wrinkles. Abdominal ventrites with distinct microreticulation, strioles, and sparse punctation, coarser on two last abdominal ventrites.
Structures: Pronotum with distinct lateral bead. Base of prosternum and neck of prosternal process with distinct ridge, slightly rounded anteriorly. 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 or slightly truncate.
Male: Antenna simple (Fig.
Holotype: TL-H 4.05 mm, TL 4.4 mm, MW 2.1 mm.
Female: Without evident differences in external morphology from males, except for not modified protarsi and abdominal ventrite 6 without striae.
The name points to similarity of the new species to E. marinae. The name is a noun in the nominative singular standing in apposition.
The key is based mostly on male characters. In many cases females cannot be assigned to a species due to the similarity of their external and internal structures (for female genitalia see Figs 17a and 17b in
1 | Dorsal surface of the body matt, with strongly impressed microreticulation and dense coarse punctation or submatt, with finer microreticulation and punctation (Figs |
2 |
– | Dorsal surface of the body shiny, with distinctly weaker microreticulation and finer punctation (Figs |
3 |
2 | Dorsal surface of body matt, with strongly impressed microreticulation and dense coarse punctation (Fig. |
(3) marinae |
– | Dorsal surface of body submatt, with punctation evidently sparser and finer and microreticulation weaker (Fig. |
(5) pseudomarinae sp. n. |
3 | Dorsal surface with very fine and sparse to moderately fine and dense punctation (Figs |
(1) broschii |
– | Dorsal surface with very fine and sparse punctation (Figs |
4 |
4 | Median lobe with apex broad, strongly curved in lateral view, its ventral sclerite with two more or less equal tips (Fig. |
(2) hintelmannae |
– | Media lobe with apex thin, slightly curved in lateral view, its ventral sclerite with two tips of different length: left one very long and narrow and right one shorter and broader, somehow rounded (Figs |
(4) mondmillensis sp. n. |
We are grateful Dr. H. Schillhammer (Vienna) for the habitus photos and Prof. D. Bilton (Plymouth) for a linguistic review of the manuscript.
Fieldwork was authorized by the PNG Department of Environment and Conservation (now PNG Climate and Environment Protection Authority) and supported by Wildlife Conservation Society, Goroka, EHP, Papua New Guinea, as well as the New Guinea Binatang Research Center, Madang, Papua New Guinea. Thanks are especially due to Aloysius Posman, Bangan John, Andrew Kinibel, and Sentiko Ibalim, and also to all other parataxonomists and paraecologists whose help is greatly appreciated.
Financial support for the study was provided by the FWF (Fonds zur Förderung der wissenschaftlichen Forschung – the Austrian Science Fund) through a project P 24312-B17 to Helena Shaverdo. Michael Balke was supported by the UK Darwin Initiative and the German Science Foundation (DFG).