Exocelina damantiensis (Balke, 1998)

Shaverdo, Helena, Sagata, Katayo & Balke, Michael, 2016, Taxonomic revision of New Guinea diving beetles of the Exocelinadanae group, with the description of ten new species (Coleoptera, Dytiscidae, Copelatinae), ZooKeys 619, pp. 45-102 : 53-63

Taxon classification Animalia Coleoptera Dytiscidae

6. Exocelina damantiensis (Balke, 1998) Figs 9-10, 11, 12, 13-14, 15, 16, 31

Copelatus (Papuadytes) damantiensis Balke 1998: 314; Nilsson 2001: 76 (catalogue).

Papuadytes damantiensis (Balke, 1998): Nilsson and Fery 2006: 56 (comb. n.).

Exocelina damantiensis (Balke, 1998): Nilsson 2007: 33 (comb. n.); Toussaint et al. 2014: Supplementary Figs 1-4, Tab. 2.

Copelatus (Papuadytes) inornatus Balke 1998: 316, not Copelatus inornatus Sharp, 1882; Nilsson 2001: 77 (catalogue); syn. n.

Copelatus (Papuadytes) madangensis Balke 2001: 362 (nom.n. for Copelatus (Papuadytes) inornatus Balke, 1998).

Exocelina madangensis (Balke, 2001): Nilsson 2007: 34 (comb. n.); Toussaint et al. 2014: Supplementary Figs 1-4, Tab. 2.

Copelatus (Papuadytes) patepensis Balke 1998: 317; Nilsson 2001: 77 (catalogue); syn. n.

Papuadytes patepensis (Balke, 1998): Nilsson and Fery 2006: 56 (comb. n.).

Exocelina patepensis (Balke, 1998): Nilsson 2007: 34 (comb. n.); Toussaint et al. 2014: supplementary figs 1-4, tab. 2.

Copelatus (Papuadytes) rivulus Balke 1998: 318; Nilsson 2001: 77 (catalogue); syn. n.

Papuadytes rivulus (Balke, 1998): Nilsson and Fery 2006: 56 (comb. n.).

Exocelina rivulus (Balke, 1998): Nilsson 2007: 34 (comb. n.).

Type locality.

Papua New Guinea: Madang Province, Finisterre Range, Damanti, 05°53'26.5"S; 145°57'50.6"E.

Type material studied.

Exocelina damantiensis . Holotype: male "Stn. No. 37", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120", " Copelatus damantiensis sp.n. Balke des. 1997" [red], “Holotypus” [red] (BMNH). Note: "Stn. 387" in the original description is obviously a type error. Paratypes: 2 males, 11 females "Stn. No. 37", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 16 males, 16 females "Stn. No. 38", "NEW GUIN EA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 3 males, 7 females, 26 exs. "Stn. No. 39", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 2 males, 1 female, 14 exs. "Stn. No. 61", "NEW GUINEA: Madang Dist., Finisterre Mts. Budemu c. 4000 ft. 15-24.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 1 male, 1 female, 5 exs. "Stn. No. 62", "NEW GUINEA: Madang Dist., Finisterre Mts. Budemu c. 4000 ft. 15-24.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 2 males, 1 female, 13 exs. "Stn. No. 73", "NEW GUINEA: Madang Dist., Finisterre Mts. Budemu c. 4000 ft. 15-24.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 3 males, 11 exs. "Stn. No. 74", "NEW GUINEA: Madang Dist., Finisterre Mts. Budemu c. 4000 ft. 15-24.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 6 males, 8 females "Stn. No. 78", "NEW GUINEA: Madang Dist., Finisterre Mts. Moro.C.5550ft. 30.x.-15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 4 males, 6 females, 18 exs. "Stn. No. 82", "NEW GUINEA: Madang Dist., Finisterre Mts. Moro.C.5550ft. 30.x.-15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 2 males, 5 exs. "Stn. No. 83", "NEW GUINEA: Madang Dist., Finisterre Mts. Moro.C.5550ft. 30.x.-15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 2 males, 3 exs. "Stn. No. 89", "NEW GUINEA: Madang Dist., Finisterre Mts. Moro.C.5550ft. 30.x.-15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). 5 males "Stn. No. 95", "NEW GUINEA: Madang Dist., Nr. Sewe, c.5,300 ft. 15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH, NHMW). All these specimens are with red paratype labels "Paratypus Copelatus damantiensis sp.n. Balke des. 1997" [red].

Exocelina madangensis . Paratypes: 4 males, 3 females with the same label as the holotype, except for "Paratypus Copelatus inornatus sp.n. Balke des. 1997" [red] (NHMW).

Exocelina patepensis . Holotype: male "Stn. No. 126", "NEW GUINEA: Morobe Dist., Lae-Bulolo Rd., Patep Ck., 28.xii.1964.", "M.E. Bacchus. B.M. 1965-120", “Holotypus” [red], " Copelatus patepensis sp.n. Balke des. 1997" [red] (BMNH). Paratypes: 4 males, 1 female with the same label as the holotype, except for "Paratypus Copelatus patepensis sp.n. Balke des. 1997" [red] (NHMW).

Exocelina rivulus . Holotype: male "IRIAN JAYA, 12.8.1992 Zentralmassiv, Borme, 140°25'E 04°24'S 900m, leg. M. Balke (8)", “HOLOTYPUS” [red], " Copelatus rivulus Balke des. 1997" [red] (NHMW). Paratypes: 25 males, 15 females with the same label as the holotype (NHMW). 7 males, 5 females "IRIAN JAYA: Borme ca. 140°25'E 04°24'S 950m, 3.9.1993 leg. M. Balke (2)" (NHMW). 12 males, 5 females "IRIAN JAYA Zentralmassiv 140°25'E 04°24'S ", "16.8.1992 Borme, 1000m leg. Balke (15)" (NHMW). 3 males, 5 females "IRIAN JAYA, 4.9.1992 Diuremna - Nalca 139°49'E 04°24'S 1500m, leg. Balke (36)" (NHMW). 38 males, 31 females "IRIAN JAYA, 6.9.1992 Nalca 1700-1800m 139°49'E 04°24'S leg. Balke (38)" (NHMW). 3 males, 1 female "IRIAN JAYA, 7.9.1992 Kono, 1800m 139°47'E 04°21'S, leg. Balke (41)" (NHMW). 55 males, 47 females "IRIAN JAYA, 12.9.1992 Angguruk, 1400m 139 °25'E 04°15'S, leg. Balke (48)" (NHMW). 5 males, 1 female "IRIAN JAYA: Angguruk, 8.10.1993 Angguruk, ca. 1350m", "ca. 139°25'E 04°15'S leg. M. Balke (32)" (NHMW). 1 male "IRIAN JAYA: Borme Tarmlu, 1500m 6.9.1993", "ca. 140°25'E 04°24'S, leg. M. Balke (4-6)" (NHMW). 53 males, 42 females "IRIAN JAYA: 11.9.1993 Bime - Calab Gebiet, Bime, 1400m", "leg. M. Balke (12) ca. 140°12'E 04°20'S " (NHMW). 57 males, 20 females "IRIAN JAYA: 22.9.1993 Bime - Calab Gebiet, Bime, 1400m", "ca. 140°12'E 04°20'S, leg. M. Balke (16)" (NHMW). These females are a mixture of two species: Exocelina damantiensis and Exocelina aipomek (Balke, 1998). 2 males, 3 females "IRIAN JAYA: 28.9.1993 Eme Gebiet Emdoman, 1150m", "ca. 139°55'E 04°14'S, leg. M. Balke (23)" (NHMW). 6 males, 5 females "IRIAN JAYA: 29.9.1993 Eme Gebiet Emdoman, 800m", "ca. 139°55'E 04°14'S, leg. M. Balke (24)" (NHMW). 2 males "IRIAN JAYA: 29.9.1993 Eme Gebiet Emdoman, 800-1000m", "leg. M. Balke (25) ca. 139°55'E 04°14'S " (NHMW). 23 males "IRIAN JAYA: 1.10.1993 Eme Gebiet Okloma, 1500m", "ca. 139°55'E 04°14'S, leg. M. Balke (28)" (NHMW). All these specimens are with red paratype labels "PARATYPUS Copelatus rivulus sp.n. M. Balke des. 1997" or "Paratypus Copelatus rivulus sp.n. Balke des. 1997" [red].

Additional material.

Indonesia: West Papua Province: Teluk Wondama Regency: 3 males, 1 female "IRIAN JAYA: Wandammen Bay, Wondiwoi Mts. Wasior, 300-700 m, 14.I.2001 leg. A. RIEDEL" (NHMW, SMNS, ZSM). 3 females "IRIAN JAYA: Wandammen Bay, Wondiwoi Mts. Wasior, 250-600 m, 4.I.2001 leg. A. RIEDEL" (SMNS). 2 males "Indonesia: West Papua: Wandammen Bay, Wasior, 4-5.I.2001 leg. A. Riedel 2?45.940'S 134?31.738'E " (ZSM). Papua Province: Paniai Regency: 2 males "IRIAN JAYA: Paniai Prov. road Nabire-Ilaga, km 140 4.9.1996, 450 m leg. M. Balke (96 # 13)" (NHMW). 1 male, 5 females "IRIAN JAYA: Paniai Prov. road Nabire-Ilaga, km 160 4.9.1996, 600 m leg. M. Balke (96 # 14)" (NHMW). 1 male "IRIAN JAYA: Paniai Prov. road Nabire-Ilaga, km 165 4.9.1996, 650 m leg. M. Balke (96 # 15)" (NHMW). All these specimens (locs. 13, 14, 15) are with red paratype labels "PARATYPUS Copelatus rivulus sp.n. M. Balke des. 1997" or "Paratypus Copelatus rivulus sp.n. Balke des. 1997" [red] but they are not considered as paratypes because they are not included into the type material of the original description. Intan Jaya Regency: 8 males, 13 female "IRIAN JAYA: Paniai Prov. Kemandoga, Homeyo, Sabisa 1700-1900m, 5.1.1996 leg. A. Riedel" (NHMW, ZSM). Puncak Jaya Regency: 1 male "Indonesia: Papua, Wano Land, red clay creek nr cave, 1100m, 3.ix.2014, nr -3.587955 137.5114945 (Pap024)", "M.Balke 6516" [green text] (ZSM). 12 males, 7 females "Indonesia: Papua, Wano Land, river grey sediment, 980m, 3.ix.2014, -3,587955 137,5114945 (Pap025)" (NHMW, ZSM). 8 males, 1 female "Indonesia: Papua, Wano Land, river ca 15m wide, 930m, 3.ix.2014, -3,587955 137,5114945 (Pap026)" (NHMW, ZSM). 6 males, 3 females "Indonesia: Papua, Wano Land, creek @ jungle helipad, 870m, 4.ix.2014, -3,584077 137,5042947 (Pap027)", two males with an additional labels "M.Balke 6525" and "M.Balke 6526" [green text] (NHMW, ZSM). Puncak Regency: 11 males, 4 females "Indonesia: Papua, Wano Land, below Puluk, 1100m, 2.ix.2014, nr -3.660272 137.5207436 (Pap021)", one of the males with an additional label "M.Balke 6510" [green text] (NHMW, ZSM). Pegunungan Bintang Regency: 15 males, 15 females "IRIAN JAYA Zentralmassiv 140°25'E 04°24'S ", "Kali Takime, 1000m 15.8.1992 leg. Balke (14)" (NHMW). 6 males, 7 females "IRIAN JAYA Zentralmassiv 140°25'E 04°24'S ", "Kali Takime, 900m 18.8.1992 leg. Balke (16)" (NHMW). 19 males, 27 females "IRIAN JAYA Zentralmassiv 140°25'E 04°24'S ", "Kali Takime, 900m 18.8.1992 leg. Balke (17)" (NHMW). All these specimens (locs. 14, 16, 17) are with red paratype labels "PARATYPUS Copelatus rivulus sp.n. M. Balke des. 1997" or "Paratypus Copelatus rivulus sp.n. Balke des. 1997" [red] but they are not considered as paratypes because they are not included into the type material of the original description. 9 females "IRIAN JAYA: 11.9.1993 Bime - Calab Gebiet, Bime, 1400m", "leg. M. Balke (12) ca. 140°12'E 04°20'S " (NHMW).

Papua New Guinea: Sandaun: 3 females "Papua New Guinea: Sandaun, Mianmin, Fak River, 775m, 14.xi.2003, 453 53.00S 141 36 39.40E, K. Sagata (WB17)", one of them with an additional label "DNA M. Balke 678" [green text] (ZSM). 2 females "Papua New Guinea: Sandaun, Mianmin, Fak River, 775m, 15.xi.2003, 453 53.00S 141 36 39.40E, K. Sagata (WB22)" (ZSM). 1 male "Papua New Guinea: Sandaun, Sandaun, Fak River (WB24), 23.x.2003, K. Sagata, DNA M Balke: MB 685", "DNA M. Balke 685" [green text] (ZSM). 2 males, 5 females "Papua New Guinea: Sandaun, Sandaun, Fak River, 775m, 15.x.2003, 4 53 53.00S #, K. Sagata (WB24)" (ZSM). 1 male "Papua New Guinea: Sandaun, Sandaun, Sek River (WB50), 21.x.2003, K. Sagata, DNA M Balke: MB 668", "DNA M. Balke 668" [green text] (ZSM). 3 males, 3 females "Papua New Guinea: Sandaun, Sandaun, Sek River 775m, 13.x.2003, K. Sagata (WB50)" (NHMW, ZSM). 5 males, 1 female "Papua New Guinea: Sandaun, May River, 970m, 19.x.2003, 4 49.779S 141 38.174E, K. Sagata (WB43)", one of the males with an additional label "DNA M. Balke 687" [green text] (NHMW, ZSM). 2 males, 5 females "Papua New Guinea: Sandaun, Wara Uk, - 900m, 14.xi.2003, Not taken, K. Sagata (WB16)", "DNA M. Balke 676", "DNA M. Balke 677" [green text] (ZSM). 1 male "Papua New Guinea: Sandaun, Sandaun, Faklows (WB87), 24.x.2003, K. Sagata, DNA M Balke: MB 656", "DNA M. Balke 656" [green text] (ZSM). 1 male "Papua New Guinea: Sandaun, Sokamin4, 1200m, 19.x.2003, 4 50.845S 141 37.865E, K. Sagata (WB102)", "DNA M. Balke 675" [green text] (ZSM). 2 males, 2 females "Papua New Guinea: Sandaun, MekilWX25, 1718m, 13.x.2003, 4 48.637S 141 38.994E, K. Sagata (WB109)", one of the males with an additional label "DNA M. Balke 669" [green text] (ZSM). 2 males, 1 female "Papua New Guinea: Sandaun, Mianminold, 898m, 20.x.2003, 4 53.419S 141 37.028E, K. Sagata (WB66)", one male additionally with "DNA M. Balke 674" [green text] (ZSM). 1 female "Papua New Guinea: Sandaun, Sandaun, Mianmin (WB75), 9.x.2003, K. Sagata, DNA M Balke: MB 667", "DNA M. Balke 667" [green text] (ZSM). 11 males, 1 female "Papua New Guinea: Sandaun, Mianmin, 670m, 22.x.2008, 4.53.329S 141.35.263E, Ibalim (PNG 189)", one of males with an additional green label "DNA M Balke 3718" (NHMW, ZSM). 69 males, 88 females "Papua New Guinea: Sandaun, Mianmin, 670m, 20.x.2008, 4.53.292S 141.34.118E, Ibalim (PNG 191) (NHMW, ZSM). 22 males, 28 females "Papua New Guinea: Sandaun, Mianmin (river), 990m, 23.x.2008, 4.54.570S 141.35.490E, Ibalim (PNG 192)", one of males with an additional green label "DNA M Balke 3738" (NHMW, ZSM). 8 males, 8 females "Papua New Guinea: Sandaun, Mianmin (pool), 990m, 23.x.2008, 4.54.570S 141.35.490E, Ibalim (PNG 193) (NHMW, ZSM). 51 males, 85 females "Papua New Guinea: Sandaun, Mianmin (river), 1080m, 24.x.2008, 04.55.780S 141.38.185E, Ibalim (PNG 195), some of them with green labels "DNA M Balke" with numbers 3743, 3744, 3779, 3780, 3781, 3782 (NHMW, ZSM). 21 males, 4 females "Papua New Guinea: Sandaun, Mianmin (pool), 1080m, 24.x.2008, 04.55.780S 141.38.185E, Ibalim (PNG 196)", one of males with an additional green label "DNA M Balke 3748" (NHMW, ZSM). 82 males, 82 females "Papua New Guinea: Sandaun, Mianmin (pool), 700m, 21.x.2008, 04.52.858S 141.31.706E, Ibalim (PNG 197) (NHMW, ZSM). 27 males, 43 females "Papua New Guinea: Sandaun, Mianmin (pool), 700m, 21.x.2008, 04.52.858S 141.31.706E, Ibalim (PNG 198) (ZSM). 7 males, 15 females "Papua New Guinea: Sandaun, Mianmin area,>1000m, 23.xii.209, Ibalim & Pius (PNG232)" (ZSM). 9 males, 15 females "Papua New Guinea: Sandaun, Mianmin area,>1000m, 23.xii.2009, Ibalim & Pius (PNG240)" (NHMW, ZSM). 2 males, 2 females "Papua New Guinea: Sandaun, Mianmin area,>1000m, 26.xii.209, Ibalim & Pius (PNG233)" (ZSM). 1 male "Papua New Guinea: Sandaun, Mianmin area,>600m, 13.i.2010, Ibalim & Pius (PNG236)", "DNA M. Balke 4928" [green text] (ZSM). 8 males, 9 females "Papua New Guinea: Sandaun, Mianmin area,>600m, 13.i.2010, Ibalim & Pius (PNG236)" (NHMW, ZSM). 11 males, 7 females "Papua New Guinea: Sandaun, Mianmin area,>600m, 9.i.2010, Ibalim & Pius (PNG237)" (NHMW, ZSM). 7 males, 4 females "Papua New Guinea: Sandaun, Mianmin area,>600m, 6.i.2010, Ibalim & Pius (PNG239)" (NHMW, ZSM). 20 males, 12 females "Papua New Guinea: Sandaun, Mianmin area,>700m, 7.i.2010, Ibalim & Pius (PNG231)" (NHMW, ZSM). Western Province: 27 males, 17 females "Papua New Guinea: Western Province, Tabubil, 600m, 22.vi.2008, 05.15.673S 141.13.738E, Posman (PNG 181)" (NHMW, ZSM). Madang: 1 female "Stn. No. 30", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 1 male, 9 females "Stn. No. 46", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 2 males, 3 females, 15 exs. "Stn. No. 47", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120". These beetles are with paratypes labels "Paratypus Copelatus damantiensis sp.n. Balke des. 1997" but they are not considered as paratypes because they are not included into the type material of the original description (BMNH, NHMW). 2 males, 2 females "Stn. No. 49", "NEW GUINEA: Madang Dist., Finisterre Mts. Damanti 3,550 ft. 2-11.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 2 females "Stn. No. 61", "NEW GUINEA: Madang Dist., Finisterre Mts. Budemu c. 4000 ft. 15-24.x.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 3 females "Stn. No. 82", "NEW GUINEA: Madang Dist., Finisterre Mts. Moro. C. 5550ft. 30.x.-15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 2 males, 1 female "Stn. No. 92", "NEW GUINEA: Madang Dist., Finisterre Mts. Moro. C. 5550ft. 30.x.-15.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 20 males "Papua New Guinea: Madang, Akameku - Brahmin, Bismarck Range, 750m, 25.xi.2006, 05.49.892S 145.24.491E, Balke & Kinibel (PNG 113)" (NHMW, ZSM). 34 males "Papua New Guinea: Madang, Akameku - Brahmin, Bismarck Range, 750m, 25.xi.2006, nr 05.49.307S 145.24.389E, Balke & Kinibel (PNG 114)" (NARI, NHMW, ZSM). 1 male "PAPUA NEW GUINEA Madang Pr. Below Bundi, 500 m, 26IX2002, M Balke (PNG 23), "268 DNA M Balke" [green] (ZSM). 4 males, 11 females "Papua New Guinea: Madang, below Bundi, 500 m, 26.IX.2002 Balke & Sagata (PNG023)" (NHMW, ZSM). 3 males "Papua New Guinea: Madang, Simbai area, 1200m, 10.iii.2007, 05.13.389S 144.37.285E, Kinibel (PNG 152) (ZSM). 2 males "Papua New Guinea: Madang, Simbai area, 1200m, 11.iii.2007, 05.13.333S 144.37.611E, Kinibel (PNG 153) (NHMW, ZSM). Enga: 8 males "Papua New Guinea: Enga, Wapanamanda, 1500m, 6.xii.2006, 05.38.105S 143.55.338E, Balke & Kinibel, (PNG 128)", one of them with an additional green label "DNA M.Balke 1527" (NHMW, ZSM). Western Highlands: 7 males "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)", one of them with an additional green label "DNA M.Balke 1340" (NHMW, ZSM). 1 male "Papua New Guinea: Western Highlands, Lugup River, 1700m, 4.iii.2007, 05.17.237S 144.28.214E, Kinibel (PNG 143)" (ZSM). 7 males "Papua New Guinea: Western Highlands, Above Sendiap, 1400m, 5.iii.2007, 05.19.774S 144.28.307E, Kinibel (PNG 145)", one of them with an additional green label "DNA M.Balke 3314" (NHNW, ZSM). 10 males "Papua New Guinea: Western Highlands, Jimi Valley, above Sendiap Station, 950m, 6.iii.2007, 05.20.587S 144.28.847E, Kinibel (PNG 147) (NHNW, ZSM). Simbu: 1 female "Ibisca Niugini, PNG 6-8.xi.2012 Mount Wilhelm 200m -5,739897251 145,3297424 MW0200 / P0786 Vial 09596" (ZSM). 8 females "Ibisca Niugini, PNG 3-5.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-R-5/8-d10 / Plot 18 / P1238 Vial 15969-CODYTI" (ZSM). 4 females "Ibisca Niugini, PNG 28-30.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-S-7/8-d04 / Plot 15 / P1211 Vial 16189-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 30.x.-1.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-M-3/8-d06 / Plot 13 / P1196 Vial 15980-CODYTI" (ZSM). 1 male "Ibisca Niugini, PNG 31.x.-2.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-D-4/8-d07 / Plot 4 / P1125 Vial 16045-CODYTI" (ZSM). 1 male "Ibisca Niugini, PNG 7-9.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-S-7/8-d14 / Plot 19 / P1248 Vial 15781-CODYTI" (ZSM). 2 males, 1 female "Ibisca Niugini, PNG 3-5.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-K-5/8-d10 / Plot 11 / P1182 Vial 16083-CODYTI" (NHMW, ZSM). 1 female "Ibisca Niugini, PNG 4-6.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-A-6/8-d11 / Plot 1 / P1103 Vial 07195-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 3-5.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-Q-5/8-d10 / Plot 17 / P1230 Vial 16097-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 29 -31.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-J-3/8-d05 / Plot 10 / P1172 Vial 07200-CODYTI" (ZSM). 1 male "Ibisca Niugini, PNG 29-31.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-D-3/8-d05 / Plot 4 / P1124 Vial 07290-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 29-30.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-E-3/8-d05 / Plot 5 / P1132 Vial 07294-CODYTI" (ZSM). 1 male, 8 females "Ibisca Niugini, PNG 27-29.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-F-2/8-d03 / Plot 6 / P1139 Vial 15944-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 31.x.-2.xi.2012 Mount Wilhelm 700m -5,73213905 145,2568207 ", "FIT-MW700-C-4/8-d07 / Plot 3 / P1117 Vial 15664-CODYTI" (ZSM). 1 male, 10 females "Ibisca Niugini, PNG 27-29.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 ", "FIT-MW700-D-2/8-d03 / Plot 4 / P1123 Vial 15972-CODYTI" (ZSM). 2 females "Ibisca Niugini, PNG 27-29.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 ", "FIT-MW700-H-2/8-d03 / Plot 8 / P1155 Vial 15976-CODYTI" (ZSM). 3 females "Ibisca Niugini, PNG 28-30.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 ", "FIT-MW700-M-2/8-d04 / Plot 13 / P1195 Vial 16167-CODYTI" (ZSM). 3 females "Ibisca Niugini, PNG 26-28.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 ", "FIT-MW700-T-1/8-d02 / Plot 20 / P1250 Vial 16254-CODYTI" (ZSM). 1 male, 2 females "Ibisca Niugini, PNG 27-29.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 ", "FIT-MW700-E-2/8-d03 / Plot 5 / P1131 Vial 15937-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 25-27.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 ", "FIT-MW700-D-1/8-d01 / Plot 4 / P1122 Vial 15947-CODYTI" (ZSM). 1 male, 1 female "Ibisca Niugini, PNG 3-5.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1222 Vial 16098" (ZSM). 1 female "Ibisca Niugini, PNG 3-5.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1254 Vial 16105" (ZSM). 1 male "Ibisca Niugini, PNG 5-7.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1247 Vial 16078" (ZSM). 1 male, 4 females "Ibisca Niugini, PNG 28-30.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1243 Vial 16156" (ZSM). 1 male, 7 females "Ibisca Niugini, PNG 28-30.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1235 Vial 16164" (ZSM). 6 females "Ibisca Niugini, PNG 26-28.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1210 Vial 16172" (ZSM). 1 female "Ibisca Niugini, PNG 6-8.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1144 Vial 15649" (ZSM). 1 female "Ibisca Niugini, PNG 30.x.-1.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1220 Vial 15992" (ZSM). 1 female "Ibisca Niugini, PNG 26-28.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1178 Vial 16181" (ZSM). 1 male "Ibisca Niugini, PNG 30.x.-1.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1244 Vial 16285" (ZSM). 5 females "Ibisca Niugini, PNG 28-30.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1179 Vial 16186" (ZSM). 2 females "Ibisca Niugini, PNG 26-28.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1226 Vial 16196" (ZSM). 1 male "Ibisca Niugini, PNG 1-3.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1237 Vial 16231" (ZSM). 2 females "Ibisca Niugini, PNG 1-3.xi.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1213 Vial 16236" (ZSM). 6 females "Ibisca Niugini, PNG 26-28.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1234 Vial 16270" (ZSM). 3 females "Ibisca Niugini, PNG 28-30.x.2012 Mount Wilhelm 700m -5,731960773 145,2521667 MW700 / P1227 Vial 16277" (ZSM). 1 male, 1 female "Ibisca Niugini, PNG 26-28.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-S-1/8-d02 / Plot 19 / P1242 Vial 16118-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 3-5.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-S-5/8-d10 / Plot 19 / P1246 Vial 16092-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 27-29.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-I-2/8-d03 / Plot 9 / P1163 Vial 15933-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 7-9.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-P-7/8-d14 / Plot 16 / P1224 Vial 15796-CODYTI" (ZSM). 5 females "Ibisca Niugini, PNG 27-29.x.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-A-2/8-d03 / Plot 1 / P1099 Vial 15960-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 9-11.xi.2012 Mount Wilhelm 700m", " -5,731960773 145,2521667 FIT-MW700-P-8/8-d16 / Plot 16 / P1225 Vial 16066-CODYTI" (ZSM). 1 female "Ibisca Niugini, PNG 31.x.-2.xi.2012 Mount Wilhelm 1200m", "-5,720873833 145,2694702", "FIT-MW1200-E-4/8-d07 / Plot 5 / P1523 Vial 17348" (ZSM). Simbu/Eastern Highlands: 3 males "Papua New Guinea: Crater Mountain, trek Haia - Wara Sera, 500m, 12IX2002, Balke & Sagata, (PNG 006)" (ZSM). 4 males "Papua New Guinea: Simbu/EHPr. Crater Mountain, Wara Sera Station, 800 m, 14IX2002, Balke & Sagata, (PNG 009)" (NHMW, ZSM). 10 males "Papua New Guinea: Crater Mountain, Wara Sera Station, 800 m, 14IX2002, Balke & Sagata (PNG 010)" (NHMW, ZSM). 1 female "Papua New Guinea: Simbu/EHPr. Crater Mountain, Sera - Herowana, Jau river, 1000 m, 15IX2002, Balke & Sagata (PNG 015)" (ZSM). 2 females "Papua New Guinea: Simbu/EHP, Crater Mountain, Sera - Herowana, Sima river, 1250 m, 15IX2002, Balke & Sagata (PNG 016)" (ZSM). 1 male "PNG Simbu / EHPr. Crater Mountain, Sera - Herowana, Wara Hulene, 1000 m, 16IX2002, Balke & Sagata (PNG 17)", "263 DNA M Balke" [green] (ZSM). 8 males, 5 female "Papua New Guinea: Simbu / EHPr. Crater Mountain, Sera - Herowana, Hulene river, 1000 m, 16IX2002, Balke & Sagata (PNG 017)" (NHMW, ZSM). Eastern Highlands: 1 female "Stn. No. 182", "NEW GUINEA: E. Highland Dist., Purosa Valley, nr. Okapa. 8.ii.1965.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 1 female "Stn. No. 190", "NEW GUINEA: E. Highland Dist., Okapa, c. 5.000ft. 10-11.ii.1965.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 11 males "Papua New Guinea: Eastern Highlands, Akameku - Brahmin, Bismarck Range, 700m, 24.xi.2006, 05.52.754S 145.23.209E, Balke & Kinibel (PNG 109)", one of them with an additional green label "DNA M.Balke 1519" (NHMW, ZSM). 20 males "Papua New Guinea: Eastern Highlands, Akameku - Brahmin, Bismarck Range, 800 m, 24.xi.2006, 05.50.021S 145.24.664E, Balke & Kinibel (PNG 112)" (NARI, NHMW, ZSM). 2 males, 1 female "Papua New Guinea: Eastern Highlands, below Yonki, 850m, 4.iv.2006, 06.11.332S 146.03.052E, Balke & Sagata (PNG 31)", one male additionally with "DNA M.Balke 1311" [green] (ZSM). Morobe: 5 females "Stn. No. 112", "NEW GUINEA: Morobe Dist., Finisterre Mts. Hinggia, c. 2,500ft. 28.xi.1964.", "M.E. Bacchus. B.M. 1965-120" (BMNH). 16 males, 25 females "Papua New Guinea: Morobe, Huon, 1 km E Yakop, 1400m, 14.v.2006, nr 06.10.961S 147.08.204E, Sagata (PNG 74)" (NHMW, ZSM). 74 males, 24 females "Papua New Guinea: Morobe, Huon, Dalasi, 3 km N Yakop, 1900m, 15.v.2006, 06.10.961S 147.08.204E, Sagata (PNG 75)", "DNA M.Balke 1286" [green] (NHMW, ZSM). 1 male "PNG: Huon Peninsula, Morobe Prov., Yus conservation area, 1398m.", "DNA M.Balke 541" [green text] (ZSM). 10 males, 4 female "Papua New Guinea: Morobe, Mindik, 1480m, 10.x.2009, 06.27.335S 147.25.233E, Inaho (03) (PNG 203) (NHMW, ZSM). 6 males "Papua New Guinea: Morobe, Mindik, 1490m, 11.x.2009, 06.27.315S 147.25.166E, Inaho (04) (PNG 204) (NHMW, ZSM). 1 male "PAPUA N.G.: Morobe Prov. Mindik, 1200 - 1500 m, 26.4.1998 leg. A. Riedel" (NHMW). 63 males, 38 females "PAPUA N.G.: Morobe Prov. E Pindiu, Kobau 24.4.1998, 1400 m leg. A. Riedel" (NHMW, ZSM). 3 males, 1 female "Papua New Guinea: Morobe, Penjengjeng, 1200m, 12.x.2009, 06.27.497S 147.29.219E, Inaho (05) (PNG205)", one male additionally with "DNA M.Balke 3822" [green] (NHMW, ZSM). 3 males, 1 female "Papua New Guinea: Morobe, Pindiu, Sulemana, 850m, 15.x.2009, 06.25.169S 147.32.112E, Inaho (08) (PNG 208)" (NHMW, ZSM). 1 male "Papua New Guinea: Morobe, Sattelberg, Maro Creek, 670m, x.2009, ca 06.27.239S 147.42.531E, Inaho (10) (PNG210)", "DNA M.Balke 3826" [green] (ZSM). 3 males, 1 female "Papua New Guinea: Morobe, Sattelberg, Zige River, ca 700m, x.2009, 6 29.233S 147 46.482E, Inaho (12a) (PNG212)" (NHMW, ZSM). 4 males, 3 females "Papua New Guinea: Morobe, Sattelberg, Siki River, ca 700m, 20.x.2009, 6 29.352S 147 46.544E, Inaho (12c) (PNG 214)" (NHMW, ZSM). 15 males, 6 females "Papua New Guinea: Morobe, Huon Pen., Kwapsanek, 850m, 31.iii.2006, 06.34.913S 147.00.526E, Balke & Sagata (PNG 25)", one of males with an additional green label "DNA M.Balke 1315" (ZSM).

Females of doubtful identity.

Indonesia: Papua Province: Pegunungan Bintang Regency: 3 females "IRIAN JAVA: Borme Tarmlu 1500m 6.9.1993", "ca. 140°25'E 04°24'S leg. M. Balke (4-6)" (NHMW). 1 female "IRIAN JAVA: Borme Tarmlu 1500m 6.9.1993", "ca. 140°25'E 04°24'S leg. M. Balke (4)" (NHMW). 2 females "IRIAN JAVA: Borme Tarmlu 1500m 6.9.1993", "ca. 140°25'E 04°24'S leg. M. Balke (6)" (NHMW). These females are a mixture of four species: Exocelina damantiensis , Exocelina ketembang (Balke, 1998), Exocelina aipomek (Balke, 1998), and Exocelina danae (Balke, 1998). 1 male (no genitals), 27 females "IRIAN JAYA: 1.10.1993 Eme Gebiet Okloma, 1500m", "ca. 139°55'E 04°14'S, leg. M. Balke (28)" (NHMW). These females are a mixture of three species: Exocelina damantiensis , Exocelina ketembang , and Exocelina aipomek . 13 females "IRIAN JAYA: 22.9.1993 Bime - Calab Gebiet, Bime, 1400m", "ca. 140°12'E 04°20'S, leg. M. Balke (16)" (NHMW). 2 females "IRIAN JAYA, 24.-26.9.1993 Eipomek [sic!] Gebiet Eipomek [sic!] - Diruemna", "ca. 140°01'E 04°27'S 1800-2600m, leg. M. Balke (21-22)" (NHMW). These females are a mixture of two species: Exocelina damantiensis and Exocelina aipomek . Papua New Guinea: Madang: 15 females "Papua New Guinea: Madang, Akameku - Brahmin, Bismarck Range, 750m, 25.xi.2006, 05.49.892S 145.24.491E, Balke & Kinibel (PNG 113)" (NHMW, ZSM). 25 females "Papua New Guinea: Madang, Akameku - Brahmin, Bismarck Range, 750m, 25.xi.2006, nr 05.49.307S 145.24.389E, Balke & Kinibel (PNG 114)" (NARI, NHMW, ZSM). These females are a mixture of two species: Exocelina broschii (Balke, 1998) and Exocelina damantiensis . 19 females "Papua New Guinea: Madang, Simbai area, 1200m, 10.iii.2007, 05.13.389S 144.37.285E, Kinibel (PNG 152)" (NHMW, ZSM). These females are a mixture of two species: Exocelina broschii and Exocelina damantiensis . 53 females "Papua New Guinea: Madang, Simbai area, 1200m, 11.iii.2007, 05.13.333S 144.37.611E, Kinibel (PNG 153)" (NARI, NHMW, ZSM). These females are a mixture of three species: Exocelina broschii , Exocelina simbaiarea Shaverdo & Balke, 2014, and Exocelina damantiensis . Enga: 10 females "Papua New Guinea: Enga, Wapanamanda, 1500m, 6.xii.2006, 05.38.105S 143.55.338E, Balke & Kinibel, (PNG 128)" (ZSM). These females are a mixture of two species: Exocelina mondmillensis Shaverdo, Sagata & Balke, 2016 and Exocelina damantiensis . 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)" (NARI, NHMW, ZSM). These females are a mixture of three species: Exocelina mondmillensis , Exocelina edeltraudae (Shaverdo, Hendrich & Balke, 2012), and Exocelina damantiensis . 34 females "Papua New Guinea: Western Highlands, Lugup River, 1700m, 4.iii.2007, 05.17.237S 144.28.214E, Kinibel (PNG 143)" (NHMW, ZSM). 9 females "Papua New Guinea: Western Highlands, Above Sendiap, 1400m, 5.iii.2007, 05.19.774S 144.28.307E, Kinibel (PNG 145)" (ZSM). 9 females "Papua New Guinea: Western Highlands, Jimi Valley, above Sendiap Station, 950m, 6.iii.2007, 05.20.587S 144.28.847E, Kinibel (PNG 147) (ZSM). These females are a mixture of two species: Exocelina mondmillensis and Exocelina damantiensis . 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)" (ZSM). 24 females "Papua New Guinea: Eastern Highlands, Akameku - Brahmin, Bismarck Range, 800m, 24.xi.2006, 05.50.021S 145.24.664E, Balke & Kinibel (PNG 112)" (NARI, NHMW, ZSM). These females are a mixture of two species: Exocelina broschii and Exocelina damantiensis .

Diagnosis.

Beetle medium-sized (TL-H 3.7-4.5 mm); uniformly brown to piceous or with paler head, pronotum or only its sides, with or without reddish sutural lines on elytra; shiny, with fine punctation and microreticulation; dorsal punctation on elytra often almost invisible; pronotum with distinct lateral bead; male antennae simple (Fig. 31); protarsomere 4 with large, thick, strongly curved anterolateral hook-like seta; male protarsomere 5 ventrally with anterior band of more than 40 and posterior row of 8 relatively long, thin setae (Fig. 9A); median lobe broad, of characteristic shape in ventral view: broadened subdistally and narrowed apically, with slightly to distinctly concave apex, in lateral view, with curved, slightly elongate and broadly pointed apex, on both lateral sides with fine setae situated on distal part of median lobe under very fine carinas; paramere without notch on dorsal side and dense, long subdistal setae and inconspicuous proximal setae (Fig. 9 B–D). See also the original descriptions in Balke (1998). The species can be easily mixed up with some occurring species: the shiny species of the Exocelina broschii -group, Exocelina broschii and Exocelina mondmillensis , or with Exocelina ketembang and Exocelina aipomek , from which can be reliably distingused only by the shape of the median lobe.

Variability and notes on species delimitation.

Herein, we synonymize with Exocelina damantiensis three species, which were described and treated as representatives of the Exocelina rivulus -group: Exocelina madangensis , Exocelina patepensis , and Exocelina rivulus ( Balke 1998). The main difference between these species was in the shape of the medial lobe, therefore, this character was carefully studied in all available populations and illustrated (in ventral and lateral views) for almost all of them (Figs 10-16). It has been found that the shape of the median lobe varies both within and among populations. It can be slightly (e.g., Figs 10B, D, 11A, C, F, 12D, E, F, 13C) or strongly (e.g., Figs 10A, 12 A–C, H, 13A, B, D) broadened subdistally. Sometimes, the narrower shape might be due to the fact that specimens are teneral (e.g., Fig. 12D) or were treated for SEM (e.g., paratype of Exocelina madangensis , Fig. 12E). The shape can be less and more narrowed apically, the narrower form being characteristic of eastern populations: Simbu, EHL, Madang, and Morobe, but is also found in specimens from Papua, Sandaun, and the Western Province. The less narrowed apically shape is characteristic of specimens from one population in Morobe (Yakop, Fig. 13C), and sometimes both shapes are found in the same population (Tabubil, Western Pr., Fig. 11E, F). The males of the type series of Exocelina patepensis from the Lae–Bulolo region (Morobe) have a median lobe with a more elongate, almost truncate apex (Figs 13D, 16E), but a similar shape can be also observed in some specimens from the other Morobe populations or in some paratypes of Exocelina damantiensis from the Finisterre Range, Madang. A less elongate apex of the median lobe is found in the population from the border region Simbu/EHL (Fig. 15G, H). In short, the shape of the median lobe is not a reliable character to support the earlier recognized species, or to split the present material into several new species or subspecies. The other characters, such as size, coloration, and dorsal punctation, vary little between localities. The beetles are medium-sized, piceous or reddish brown (probably more teneral forms), often with a paler head and pronotal sides, sometimes with reddish sutural lines on the elytra, and are shiny dorsally, with the punctation on the elytra fine, rather distinct or almost invisible. Protarsomere 4 always has a large, thick, strongly curved anterolateral hook-like seta; indication of the "small antero-lateral hook" for Exocelina madangensis in Balke (1998) is probably a mistake.

Thus, the Exocelina rivulus -group is apparently not a complex of several species, but instead a single, very successful polymorphic species Exocelina damantiensis , with the largest geographical range of any New Guinea Exocelina : along the central mountain range from Wandammen Peninsula to Huon Peninsula. Of course, it is not out of the question that further material and study of other aspects (e.g., population genomics) might change this situation.

Distribution and habitats.

Indonesia and PNG. It occurs in the central mountain chain and the mountains of Wandammen and Huon Peninsulas (Fig. 39). There, it is one of the most common and numerous species from 450 m to 1900 m. Usually, it is a dominate species in the biotope and co-occurs with many species, some of which are mentioned under "Females of doubtful identity".