Research Article |
Corresponding author: Jong-Seok Park ( jpark16@tigers.lsu.edu ) Academic editor: Zi-Wei Yin
© 2015 Jong-Seok Park, Christopher E. Carlton.
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:
Park J-S, Carlton CE (2015) Pseudoexeirarthra, a new genus from New Zealand (Coleoptera, Staphylinidae, Pselaphinae), with descriptions of seven new species. ZooKeys 491: 95-118. https://doi.org/10.3897/zookeys.491.9164
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A new endemic genus and seven new species of New Zealand pselaphine staphylinid beetles of the supertribe Faronitae are described as follows: Pseudoexeirarthra Park & Carlton, gen. n. (type species: Sagola spinifer Broun); P. sungmini Park & Carlton, sp. n.; P. kwangguki Park & Carlton, sp. n.; P. youngboki Park & Carlton, sp. n.; P. seiwoongi Park & Carlton, sp. n.; P. parkeri Park & Carlton, sp. n.; P. hlavaci Park & Carlton, sp. n.; P. nomurai Park & Carlton, sp. n. Three species, S. spinifer Broun, S. colorata Broun, and S. puncticollis Broun, are transferred to the genus Pseudoexeirarthra. Six species are synonymized: S. dilucida Broun, S. guinnessi Broun, S. longicollis Broun, S. longula Broun, and S. rectipennis Broun under P. spinifer (Broun); S. insueta Broun under S. colorata (Broun). A lectotype is designated for P. spinifer (Broun). A key, habitus photographs, line drawings of diagnostic characters, and distribution maps are provided for each species.
Taxonomy, biogeography, Faronitae, Faronini, redescription
Sagola Sharp, 1874, the largest genus of the supertribe Faronitae, has been considered to be a paraphyletic assemblage of species (
Approximately four hundred specimens were studied from the Field Museum of Natural History (FMNH), Chicago, IL, USA; Louisiana State Arthropod Museum (LSAM), Baton Rouge, LA, USA; Natural History Museum (NHM), London, United Kingdom; Lincoln University (LUNZ), Lincoln, New Zealand; New Zealand Arthropod Collection (NZAC), Auckland, New Zealand; Auckland Museum, Auckland, New Zealand (AMNZ); personal collection of Donald S. Chandler (DSC), Durham, NH, USA; personal collection of John T. Nunn (JTN), Dunedin, New Zealand.
Holotypes of species described herein are deposited in the New Zealand Arthropod Collection (NZAC), Auckland. Paratype and additional specimen depositions are indicated parenthetically. Specimen label data for types are transcribed verbatim. Data for other specimens are standardized for consistency.
Seven specimens were mounted on permanent slides to aid in observation of internal characters and fine external characters not apparent when using a dissecting microscope. Permanent microscopic slides were prepared using the techniques described by
New Zealand maps were produced by modifying the map of
Each figure of an aedeagus illustrates the organ in dorsal view with the median lobe oriented forward (up on page). Right and left are indicated based on this orientation, not the morphological orientation when inside the body, which would be reversed.
Sagola spinifer Broun, 1895: 75; here designated.
Members of Pseudoexeirarthra can be separated from those of all other faronite genera by the following combination of characters: body length 1.8–2.8 mm; frontal sulcus broad and shallow, reaching level of the midline of eyes (Fig.
Body length 1.8–2.8 mm. Body reddish, antennae, legs, maxillary palpi and elytra paler (Fig.
The generic name refers to the superficial similarity to the genus Exeirarthra Broun.
Members of this genus lack distinct external secondary sexual characters except on abdominal sternite IX. Male sternite IX is fragile, and partially concealed by sternite VIII, rendering it simple and reduced in appearance. Females possess a more robust, heart-shaped or rounded sternite IX that bears two pairs of long setae that are usually visible in ventral view. Female genitalia, including spermathecae, apparently are membranous and were not observed after clearing specimens using 10% potassium hydroxide.
Note. The key is largely based on male genitalia because most specimens are indistinguishable based on the external morphology. Antennal shape and eye size are apparently unique within species. However, apparent variations may result from viewing at inconsistent orientations among specimens. These characters are difficult to interpret consistently when performing identifications, but still useful in comparing types or specimens in series.
1 | Elytra as long as wide (Fig. |
Pseudoexeirarthra kwangguki sp. n. |
– | Elytra longer than wide, abdominal tergite IV with transverse patches of microtrichia | 2 |
2 (1) | Left paramere at least twice as wide as right paramere at midpoint (Fig. |
P. spinifer (Broun) |
– | Let paramere close to as wide as right paramere at midpoint | 3 |
3 (2) | Median lobe of genitalia at least 3 times as wide as either paramere (Fig. |
P. sungmini sp. n. |
– | Median lobe of genitalia at most slightly more than twice as wide as either paramere | 4 |
4 (3) | Parameres broader than median lobe of genitalia (Fig. |
P. puncticollis (Broun) |
– | Parameres narrower than median lobe of genitalia | 5 |
5 (4) | Median lobe of genitalia with acute spine at one-third length (Fig. |
P. nomurai sp. n. |
– | Median lobe of genitalia lacking branch | 6 |
6 (5) | Apical lobe of genitalia triangular (Fig. |
P. seiwoongi sp. n. |
– | Apical lobe of genitalia bluntly rounded | 7 |
7 (6) | Left paramere longer than right (Fig. |
P. parkeri sp. n. |
– | Right paramere longer than left | 8 |
8 (7) | Major apical lobe of genitalia rectangular (Fig. |
P. colorata (Broun) |
– | Major apical lobe of genitalia inverted triangular apically | 9 |
9 (8) | Major apical lobe of genitalia with slightly wider apical margin (Fig. |
P. youngboki sp. n. |
– | Major apical lobe of genitalia with distinctly wider apical margin (Fig. |
P. hlavaci sp. n. |
Sagola spinifer Broun, 1895: 75.
Sagola dilucida Broun, 1914: 157.
Sagola guinnessi Broun, 1911: 502.
Sagola longicollis Broun, 1911: 498.
Sagola longula Broun, 1912: 625.
Sagola rectipennis Broun, 1921: 489.
Lectotype. New Zealand: Waikato (WO): 1♂ (BMNH), glued on rectangular card, “2723.” [white label, handwritten]; “Mount. Pirongia” [white label, printed]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Sagola spinifer” [white label, handwritten]; “LECTOTYPE Pseudoexeirarthra spinifer (Broun) Desig. Park and Carlton 2013” [red label, printed]. *The lectotype designation is required because Broun did not explicitly designate a type specimen, and his comments suggest that three specimens were examined (Broun, 1875: 75). This designation will fix the identity of the species and facilitate its recognition by future workers. Paralectotypes (1 male, 1 female). New Zealand: Waikato (WO): 1♀ (BMNH), glued on rectangular card, “Type” [red label, printed]; “2723.” [white label, handwritten]; “Pirongia” [white label, printed]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Sagola spinifer” [white label, handwritten]; “PARALECTOTYPE Pseudoexeirarthra spinifer (Broun) Desig. Park and Carlton 2013” [yellow label, printed]. 1♂ (NZAC), glued on rectangular card, “Pirongia” [white label, printed]; “var. 2723.” [white label, handwritten]; “S. spinifer antennae.” [white label, handwritten]; “Broun” [white label, handwritten]; “ N.Z. Arthropod Collection, NZAC Private Bag 92710 New Zealand” [yellow label, printed]; “PARALECTOTYPE Pseudoexeirarthra spinifer (Broun) Desig. Park and Carlton 2013” [yellow label, printed].
Holotype of Sagola dilucida: New Zealand: Auckland (AK): 1♀ (BMNH), glued on rectangular card, “Type” [red label, printed]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Epsom. Jany.1912.” [white label, handwritten]; “3520.♂” [white label, handwritten]; “Sagola dilucida” [white label, handwritten]. The original label indicates the specimen is male, but it is female.
Holotype of Sagola guinnessi: New Zealand: Taupo (TO): 1♂ (BMNH), glued on rectangular card, “Type” [red label, printed]; “3373.” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Erua. 5.3.1912.” [white label, handwritten]; “Sagola guinnessi.” [white label, handwritten]. The original label indicates the specimen is female, but it is male.
Holotype of Sagola longicollis: New Zealand: Taupo (TO): 1♀ (BMNH), glued on rectangular card, “Type” [red label, printed]; “3369” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Mahuia. Jany.1911.” [white label, handwritten]; “Sagola longicollis.” [white label, handwritten].
Holotype of Sagola longula: New Zealand: Auckland (AK): 1♀ (BMNH), glued on rectangular card, “Type” [red label, printed]; “15.” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “auckland. N.Z. Lawson” [white label, handwritten]; “Sharp Coll. 1905-313.” [white label, printed]; “Sagola longula.” [white label, handwritten].
Syntype of Sagola rectipennis: New Zealand: Otago Lakes (OL): 1♂ (BMNH), glued on rectangular card, “3997.♂” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Mt. Alfred. 9.2.1914” [white label, handwritten]; “Sagola ♂. rectipennis” [white label, handwritten]. 1♂ (BMNH), glued on rectangular card, “3997.♂” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Mt. Alfred. 9.2.1914” [white label, handwritten]; “Sagola rectipennis” [white label, handwritten]. 1♂ (BMNH), glued on rectangular card, “3997.♂” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Mt. Alfred. 9.2.1914” [white label, handwritten].
(n = 198; 95 males, 103 females). New Zealand: Auckland (AK): 1♂ 1♀, Waitakere Ra, Cascade-Kauri Park, Up. Kauri tr, 170 m, 8 XII 1984–25 I 1985, kauri-podo-hdwd, A. Newton, M. Thayer 680, FIT&window trap (DSC); 1♂, Lynfield, Tropicana dr, 14 VIII 1976, G. Kuschel, litter (NZAC); 1♂ 1♀, Lynfield, 7 IX 1980, G. Kuschel, litter (NZAC); 2♀♀, Woodhill, 27 II 1976, C. F. Butcher, pit trap (NZAC); 1♂, Lynfield, 16 IV 1977, G. Kuschel (NZAC); Bay of Plenty (BP): 1♀, Orete Forest, Te Puia Hut, 230 m, 29 I 1993, R. M. Emberson, litter (NZAC); 1♂1♀, Lottin Pt Rd, Waenga Bush, 24 XI 1992–29 I 1993, R. C. Henderson, Malaise trap (NZAC); 1♂, Lottin Pt Rd, Waenga, 27 I 1993, R. C. Henderson, litter (NZAC); 1♂, Te Koau, Main Ridge, 220 m, 23 IX 1992, J. S. Dugdale, litter (NZAC); 1♂, Te Koau, Twin Puriri’s, 31 I–15 III 1993, R. C. Henderson, pit trap (NZAC); 1♂, Papatea, 13 X–23 XI 1992, G. Hall, pit trap (NZAC); 1♂, Mamaku Ra, 18 I 1972, G. W. Ramsay, litter (NZAC); 1♀, Te Koau, Bush Track, 23 IX 1992, J. S. Dugdale, litter (NZAC); 1♀, Mount Te Aroha, summit, 19 XI 2005, J. Nunn, moss (NZAC); 1♀, Kaimai-Mamaku Forest Park, Mt. Te Aroha summit rd, 450 m, 37°31.43'S, 175°44.01'E, 19 XI 2005, mixed broadleaf forest with many tree ferns, nikau palms, FMHD#2005-016, FIT, A. Newton, M. Thayer, ANMT site 1144 (FMNH); Buller (BR): 5♂♂ 6♀♀, Nelson Lakes NP, Mt. Robert, Speargrass tr, 875 m, 41°49.47'S, 172°48.31'E, 17 XII 2005, Nothofagus forest, FMHD#2005-110, litter, A. Solodovnikov, D. Clarke, ANMT site 1161 (FMNH); 22♂ 12♀♀, (1♂, slide-mounted), Lewis Pass NR, 11.9 km ese Spring Junction, 540 m, 17 XII 1984–21 I 1985, Nothofagus forest, A. Newton, M. Thayer, 715, window trap (FMNH); 8♂♂ 14♀♀, Nelson Lakes NP, Lake Rotoroa, Braeburn tr, 470 m, 16 XII 1984–7 I 1985, Nothofagus forest, A. Newton, M. Thayer 712, window trap (FMNH); 4♂♂ 8♀♀, Nelson Lakes NP, n slope Mt. Robert, Pinchgut tr, 950 m, 14 XII 1984–6 I 1985, Nothofagus forest, A. Newton, M. Thayer 707, window trap (FMNH&DSC); 2♀♀, Nelson Lakes NP, n slope Mt. Robert, Pinchgut tr, 950 m, 14 XII 1984–6 I 1985, Nothofagus forest, A. Newton, M. Thayer 707, litter (FMNH); 1♂2♀♀, Nelson Lakes NP, Lake Rotoiti, St. Arnaud tr, 645 m, 14 XII 1984–6 I 1985, Nothofagus forest, A. Newton, M. Thayer 705, window trap (FMNH); 1♂, Nelson Lakes NP, Mt. Robert Rd, 660 m, 26 XII 1984–6 I 1985, Leptospermum-Nothofagus forest, A. Newton, M. Thayer 722, FIT&window trap (DSC); 1♂, Lewis Pass NR, 13.2 km s Lewis Pass, 650 m, 17 XII 1984–21 I 1985, Nothofagus forest, A. Newton, M. Thayer 713, FIT&window trap (DSC); 2♂♂, Mt. Misery, Ecology Div Stn, 460 m, 24–26 I 1977, J. S. Dugdale, water trap (NZAC); 1♀, Greymouth, Boddytown, 8 II 1984, J.C. Watt, litter (NZAC); 2♂♂, Rd to Mt. Robert, 762 m, Lake Rotoiti, 3 V 1966, J. I. Townsend, moss (NZAC); Central Otago (CO): 1♂ 1♀, Waipori, 610 m, Stony Stream, 2 XI 1979, J. C. Watt, moss (NZAC); 1♀, Piano Flat, Waikaia Forest, 25 IV 2007, washed soil beech forest (JTN); Coromandel (CL): 2♀♀, Cuvier I, Northwest Ridge, 25 II–2 III 1982, G. Hall, malaise trap (NZAC); 1♂, Cuvier I, 25 II–2 III 1982, G. Hall, pit trap (NZAC); 1♂, Great Barrier I, Little Windy Hill, 100 m, 25 II–19 III 2003, K. Parsons, malaise trap (AMNZ); 1♂ 3♀♀, Great Barrier I, Little Windy Hill, 13 XII 2002–17 I 2003, P. Sutton, Malaise (AMNZ); 1♂ 1♀, Cuvier I, Ridge Tr, 100 m, 10–18 XI 1999, J. W. Early, S. E. Thorpe, malaise trap (AMNZ); 1♂, Great Barrier I, Little Windy Hill, 17 I–27 II 2003, K. Parsons, malaise trap (AMNZ); 1♂, Great Barrier I, Little Windy Hill, 220 m, 7 XI–11 XII 2001, P. Sutton, J. Gilbert, malaise trap (AMNZ); 3♀♀, Great Barrier I, Little Windy Hill, 220 m, 11 XII 2001–18 I 2002, P. Sutton (AMNZ); 1♀, Cuvier I, Pumphouse Stream, 120 m, 14 XI 1999, J. W. Early, S.E. Thorpe, litter (AMNZ); 1♀, Great Barrier I, Little Windy Hill, 220 m, 21 II 2001–26 III 2002, P. Sutton, malaise trap (AMNZ); 1♀, Great Barrier I, Little Windy Hill, 220 m, 18 I–21 II 2002, P. Sutton, malaise trap (AMNZ); Fiordland (FD): 2♂♂, Lake Hauroko, Southland, 2 II 1966, J. I. Townsend, moss (NZAC); 1♂, Secretary I, Gut Bay, 24 XI 1981, C.F. Butcher, beech litter and rotten wood (NZAC); 1♂, Fiordland NP, Monowai Lake, 4km sw Monowai, 11–14 III 2010, J. W. Early, yellow pan traps, Nothofagus solandri forest (DSC); 1♂ 1♀, Fiordland NP, Milford Sound rd, Smithy Creek Campground area, 400 m, 44°57.07'S 168°01.16'E, 9 XII 2005, Nothofagus fusca & N. menziesii open forest, FMHD#2005-089, litter, M. Thayer, A. Newton, ANMT site 1170 (FMNH); 1♀, Secretary I, 850 m, 30 XI 1981, C.F. Butcher, pit trap (NZAC); Gisborne (GB): 1♂, Urewera NP, Waikaremoana rd, s end Matanunui Ridge, 720 m, 38°44.40'S, 177°05.81'E, 22 XI–23 XII 2005, mixed broadleaf (incl. Nothofagus fusca)-podocarp, FMHD#2005-028, FIT, M. Thayer, A. Solodovnikov, ANMT site 1149 (FMNH); 1♀, Lake Waikaremoana, 17 I 1972, G. W. Ramsay, litter; 1♀, Urewera NP, at large, #021, 6–8 III 2000, C. Carlton, A. Weir (LSAM); 1♀, Urewera NP, Lake Waikremoana, nr Caravan Park, shoreline toetoe, FIT, 23 III 2000, C. Carlton, A. Weir, #078 (LSAM); Northland (ND): 1♀, Paihia Opua SF, 22 I 1981, G. Kuschel, litter and rotten wood (NZAC); 1♀, Waipoua SF, Waipoua Stm, 70 m, 16–21 III 1978, S. Peck, J. Peck, malaise trap (FMNH); Nelson (NN): 2♂♂ 1♀, 0.6 km e Gowanbridge, 330 m, 18 XII 1984–7 I 1985, Nothofagus forest, A. Newton, M. Thayer 717, FIT&window trap (DSC); 1♂, Kahurangi NP, Cobb Ridge, above Cobb Reservoir, 1050 m, 41°06.35'S, 172°41.66'E, 29 XI–18 XII 2005, Nothofagus forest, FMHD#2005-051, FIT, A. Newton, M. Thayer, A. Solodovnikov, ANMT site 1159 (FMNH); 1♂, Kahurangi NP, Arthur Range, above Flora Saddle, 1000 m, 41°11.35'S, 172°44.46'E, 29 XI–18 XII 2005, Nothofagus forest, FMHD#2005-046, litter, A. Newton, M. Thayer, ANMT site 1156 (FMNH); 1♀, Cobb Ridge, east of Cobb Reservoir, 990 m, 2 I 1985, Nothofagus forest, A. Newton, M. Thayer 728, litter (DSC); 1♂ 1♀, Canaan Harwoods tr, 4 II 1965, L. P. Marchant, litter (NZAC); 1♂, Dovedale, 11 X 1963, J. I. Townsend, litter (NZAC); 2♀♀, Lake Rotoiti, 19 II 1965, L. P. Marchant, moss (NZAC); 1♀, Lake Rotoiti, 27 VII 1965, A. K. Walker, moss (NZAC); Otago Lakes (OL): 1♂ 1♀, 44.5 km nw Wanaka, 350m, Matukituki Valley, 44’29S 168’47E, #079, Nothofagus forest litter, 18 I 1998, C. Carlton, R. Leschen (LSAM); 1♂ 1♀, Makarora Bush, Makarora, 7–9 XI 1997, J. Nunn (JTN); 1♂ 1♀, Paradise, 2 II 1984, J. C. Watt, wood mould (NZAC); 1♂, Upper Makarora, 17 I 1968, F. A. Alack, litter (NZAC); 1♀, 10.5 km nw Glenorchy, Nothofagus forest, 44°47'S, 169°27'E, FIT #143, 19–24 I 1998, C. Carlton, R. Leschen (LSAM); Rangitikei (RI): 1♀, Ruahine Ra, Armstrong Saddle, 1370 m, 26 XII 1983, J. C. Watt, litter (NZAC); 1♀, Ruahine Ra, Triplex, 10 II 1980, C. F. Butcher, litter (NZAC); Marlborough Sounds (SD): 3♂♂ 2♀♀, Wairau V, Schroders Ck, 7 IX 1966, J. I. Townsend, litter 66/295 (NZAC); 1♀, Upper Wairau Valley, 731 m, 5 IX 1966, J. I. Townsend, moss (NZAC); 3♀♀, Head Fabians Valley, 23 X 1963, J. I. Townsend, litter (NZAC); 1♂, Rainbow SF, Connors Ck, 825 m, 21 XII 1981, J. W. Early, sweeping (NZAC); Southland (SL): 2♀♀, Catlins SF Park, 15 II 1982, C.F. Butcher, J. S. Dugdale, litter (NZAC); 1♀, Owaka Glenomaru Reserve, 18 I 1978, S. Peck, J. Peck, litter (FMNH); Taupo (TO): 1♂, Kaimanawa North Forest Park, 850m, 11 III 1978, J. S. Dugdale, moss (NZAC); 1♂, Kaimanawa N Forest Park Saddle, 20 II 1978, J. S. Dugdale, litter (NZAC); 1♀ (slide-mounted), Erua, 27 I 1982, C. F. Butcher, litter and moss (NZAC); 1♀, Erua, 16 XII 1961, G. Kuschel, litter (NZAC); Westland (WD): 1♂, Doughboy Creek, 6 km sw Mahitai, 5 II 1984, J. C. Watt, wood mould (NZAC); 1♀, Hokitika Gorge, 29 I 1978, S. Peck, litter (FMNH); 1♀, Jacksons Bay, 23 IX 1979, A. K. Walker, moss (NZAC); Wanganui (WI): 1♂ 1♀, nr Glow-worm Cave, Table Hill rd nr Apiti, 8 II 1997, J. Nunn (JTN); 1♂, Ashhurst Domain, Ashhurst, 23 X 1998, J. Nunn, litter (JTN); Wellington (WN): 3♂♂ 1♀, Mana Island, 4–6 II 1994, J. Nunn, decayed wood (JTN); 1♂, Tararua Ra, e Basin Logan, 1300 m, 6 XII 1984, R. C. Craw, turf plants (NZAC); 1♂, Pakuratahi Forks, 8 VII 1994, J. Nunn, litter (JTN); 1♀, Tararua Ra, Dundas Hut Ridge, 990 m, 13 II 1985, C. F. Butcher, litter (NZAC); 1♀, n Titahi Bay, Rocky Bay, 28 XII 1980, J. C. Watt, litter and humus (NZAC); 1♀, Tararua Forest Park, Waitewaewae tr, 220 m, 40°51.98'S, 175°15.319'E, 26 XI–21 XII 2005, broadleaf-podo forest, FMHD#2005-034, FIT, A. Newton, M. Thayer, ANMT site 1152 (FMNH); Waikato (WO): 1♂, Pirongia Forest Park, Mahaukura tr, 270 m, 37°58.22'S, 175°06.52'E, 18 XI–27 XII 2005, broadleaf forest, FMHD#2005-009, FIT, A. Newton, M. Thayer, et al., ANMT site 1142 (FMNH).
This species is distinguished from the other species of this genus by the following combination of characters: body length 2.3–2.8 mm; eyes large, as long as temples (Fig.
Length 2.3–2.8 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Mount Pirongia (WO), New Zealand.
Auckland (AK), Bay of Plenty (BP), Buller (BR), Central Otago (CO), Coromandel (CL), Fiordland (FD), Gisborne (GB), Northland (ND), Nelson (NN), Otago Lakes (OL), Rangitikei (RI), Marlborough Sounds (SD), Southland (SL), Taupo (TO), Westland (WD), Wanganui (WI), Wellington (WN), Waikato (WO) (Fig.
Most specimens were collected using malaise, flight intercept, window traps, or by sifting leaf litter in broadleaf, podocarp, hardwood and Nothofagus forests.
Specimens of Pseudoexeirarthra spinifer can be separated from those of the other species by the large eyes, fully developed hind-wings, shapes of antennomeres, and genitalia. The type specimens of Sagola dilucida, S. guinnessi, S. longicollis, S. longula and S. rectipennis share these diagnostic characters. For these reasons, we have placed these names in synonymy with P. spinifer.
Sagola colorata Broun, 1914: 156.
Sagola insueta Broun, 1914: 157.
Holotype. New Zealand: Mid Canterbury (MC): 1♀ (BMNH), glued on rectangular card, “Type” [red label, printed]; “3519.” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “McClennans. 25.3.1912.” [white label, handwritten]; “Sagola colorata.” [white label, handwritten].
Holotype of Sagola insueta. New Zealand: Mid Canterbury (MC): 1♀ (BMNH), glued on rectangular card, “Type” [red label, printed]; “3521.” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Rakaia. 6.7.1912.” [white label, handwritten]; “Sagola insueta” [white label, handwritten].
(n = 23; 21 males, 2 females). New Zealand: Dunedin (DN): 1♂, Nicol Creek, Leith Valley, 25 V 2006, J. Nunn, litter (JTN); 1♂, Vauxhall, 13 IX 2001, J. Nunn, FIT (JTN); 1♂, The Tunnels, Silverpeaks, 2 VI 2001, J. Nunn, moss and litter (JTN); 1♂, Vauxhall, 26 II 2011, 45°54.24'S, 170°31.89'E, 167 m, J. Nunn, washes soil (JTN); 1♂, Town Belt, 26 VII 1997, J. Nunn, decayed wood (JTN); 1♂, Vauxhall, 27 I 2000, J. Nunn, FIT (JTN); Fiordland (FD): 6♂ 1♀, Lake Hauroko, 2 XI 1966, J. I. Townsend, litter (NZAC); 1♂, Hollyford Camp, 10 XII 1966, A. K. Walker, litter (NZAC); Mid Canterbury (MC): 7♂♂, Banks Peninsula, Hay Scenic Res, Pigeon Bay, 25 m, 11 XII 1984–22 I 1985, Podocarp-hdwd forest, A. Newton, M. Thayer 702, window trap (FMNH); 1♂, Craigieburn SF, 850 m, #023, 42°10.8'S, 171°42'E, Nothofagus litter, 9 I 1998, C. Carlton, R. Leschen (LSAM); 1♀, McLellans Bush, Mt. Hutt, 11 I 2008, J. Nunn, on sooty mould on beech bole (JTN).
This species is distinguished from the other species of this genus by the following combination of characters: body length 2.2–2.5 mm; eyes large, two-thirds length of temples (Fig.
Length 2.2–2.5 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
McClennan’s Bush, near Methven (MC), New Zealand.
Dunedin (DN), Fiordland (FD), Mid Canterbury (MC) (Fig.
Specimens were collected using flight intercept traps, and by soil washing or sifting leaf and wood litter.
The type specimen of Sagola insueta shares the diagnostic characters of Pseudoexeirarthra colorata. For this reason, we have placed S. insueta in synonymy with P. colorata.
Sagola puncticollis Broun, 1911: 499.
Holotype. New Zealand: South Canterbury (SC): 1♂ (BMNH), glued on rectangular card, “Type” [red label, printed]; “3370.” [white label, handwritten]; “New Zealand Broun Coll. Brit. Mus. 1922-482.” [white label, printed]; “Timaru. -Wallace.” [white label, hand written]; “Sagola puncticollis” [white label, handwritten].
(n = 34; 29 males, 5 females). New Zealand: Mackenzie (MK): 8♂♀, White Horse Hill, Mt. Cook, 26 X 2009, J. Nunn, litter in podocarp forest (JTN); 1♂, White Horse Hill, Mt. Cook, 25 X 2009, J. Nunn, litter in podocarp grove (JTN); South Canterbury (SC): 18♂♀, Gunns Bush, Waimate, 23 XII 2006, washed soil in broadleaf forest, J. Nunn (JTN); 1♂ 1♀, Orari Gorge SR, Geraldine, 7 VI 2009, J. Nunn, washed soil in broadleaf forest (JTN); 1♀ (slide-mounted), Pioneer Park, Raincliff, 9 VIII 2009, J. Nunn, washed soil in totara and kahikatea forest (JTN); 1♂, Kelceys Bush, Waimate, 20 I 1966, J. I. Townsend, moss; 1♀, Mt. Dalgety, 1737 m, 19 I 1966, G. W. Ramsay, J. I. Townsend, moss (NZAC).
This species is distinguished from the other species of this genus by the following combination of characters: body length 2.3–2.6 mm; eyes one-half lengths of temples (Fig.
Length 2.3–2.6 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Timaru (SC), New Zealand.
Mackenzie (MK), South Canterbury (SC) (Fig.
Specimens were collected by soil washing and sifting moss litter in broadleaf or podocarp forests.
Holotype. New Zealand: Nelson (NN): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “NEW ZEALAND NN Mt Arthur/Flora Sdl Tck, c1400m 28-Nov-05”, “On mossy overhangs by gully. D Clarke, J Nunn”, “HOLOTYPE Pseudoexeirarthra sungmini Park and Carlton des. 2013”. Paratype (1 male): New Zealand: Nelson (NN): 1♂, Cobb, L. Sylvester, 1329 m, 31 III 1969, J. S. Dugdale, litter (NZAC).
This species is named after Dr. Sung Min Boo, Professor of Biology, Chungnam National University (Daejeon, South Korea), world algal systematist and, an enthusiastic supporter of this study.
This species is distinguished from other the species of this genus by the following combination of characters: body length 1.9–2.1 mm; eyes two-thirds length of temples (Fig.
Length 1.9–2.1 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Nelson (NN) (Fig.
Specimens were collected by sifting moss and leaf litter.
Holotype. New Zealand: Stewart Island (SI): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “New Zealand: SI: Table Hill, 366–610m 15 II 1968, G. Kuschel Moss 68/52”, “HOLOTYPE Pseudoexeirarthra kwangguki Park and Carlton des. 2013”. Paratypes (2 females): New Zealand: Stewart Island (SI): 2♀♀ (1♀, slide-mounted), same data as holotype (NZAC).
This species is named for Dr. Kwang-Guk An, Professor of Biology, Chungnam National University (Daejeon, South Korea), freshwater ecosystem specialist, and an enthusiastic supporter of this study.
This species is distinguished from the other species of this genus by the following combination of characters: body length 1.8–2.0 mm; eyes one-half lengths of temples (Fig.
Length 1.8–2.0 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Stewart Island (SI) (Fig.
Specimens were collected by sifting moss litter.
Holotype. New Zealand: Mid Canterbury (MC): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “NEW ZEALAND: MC: Banks Peninsula, Ahuriri Scen. Res., 450m, 43°39.971'S, 172°37.427'E, 3 XII 2005, mixed broadleaf w/emergent podocarp; FMHD#2005-069, berl., leaf & log litter, A. Newton, M. Thayer & A. Solodovnikov; ANMT site 1162”, “HOLOTYPE Pseudoexeirarthra youngboki Park and Carlton des. 2013”. Paratypes (n = 18; 8 males, 10 females): New Zealand: Mid Canterbury (MC): 6♂♂ 6♀♀, same data as holotype (FMHD); 1♂, Banks Peninsula, Ahuriri SR, 450 m, 43°39.971'S, 172°37.427'E, 3–6 XII 2005, mixed broadleaf w/emergent podocarp, FMHD#2005-069, FIT, A. Newton, M. Thayer, ANMT site 1162 (FMHD); 4♀♀, Bank Peninsula, Mt. Sinclair SR, 775 m, 43°42.977'S, 172°51.098'E, 3–16 XII 2005, ridgetop mixed broadleaf w/emergent Podocarpus tatara, FMHD#2005-070, FIT, A. Newton, M. Thayer, ANMT site 1163 (FMNH); 1♂, Prices Valley, 3–24 IV 1981, J. W. Early, yellow pan trap (LUNZ).
This species is named after Dr. Young Bok Cho, curator of Natural History Museum of Hannam University (Daejeon, South Korea), carrion and rove beetles specialist, and an enthusiastic supporter of this study.
This species is distinguished from the other species of this genus by the following combination of characters: body length 1.9–2.2 mm; eyes large, two-thirds lengths of temples (Fig.
Length 1.9–2.2 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Mid Canterbury (MC) (Fig.
Specimens were collected using flight intercept or yellow pan traps, or by sifting moss and leaf litter in broadleaf and podocarp forests.
Holotype. New Zealand: Wellington (WN): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “NEW ZEALAND WN 4 km along Waiotauru Rd. 16/11/91 Tararua FP”, “1159”, “HOLOTYPE Pseudoexeirarthra seiwoongi Park and Carlton des. 2013”. The original label does not record who collected the specimen, but it was collected by J. Nunn. Paratype (1 male): New Zealand: Mid Canterbury (MC): 1♂, Banks Penin., Peraki Saddle Scen Res, 500 m, 11 XII 1984, hdwd-podo.elfin forest, A. Newton, M. Thayer 701, log and leaf litter (FMNH).
This species is named after Dr. Sei-Woong Choi, Professor at Mokpo National University (Mokpo, South Korea), world moth specialist, and an enthusiastic supporter of this study.
This species is distinguished from the other species of this genus by the following combination of characters: body length 1.9–2.1 mm; eyes large, as long as temples (Fig.
Length 1.9–2.1 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Mid Canterbury (MC), Wellington (WN) (Fig.
The paratype was collected by sifting leaf and moss litter.
Holotype. New Zealand: Dunedin (DN): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “NEW ZEALAND DN Rocklands 21 Nov 1981 C.F. Butcher”, “Sweeping tussock nr stream”, “N.Z. Arthropod Collection, NZAC Private Bag 92170 AUCKLAND New Zealand”, “HOLOTYPE Pseudoexeirarthra parkeri Park and Carlton des. 2013”. Paratypes (n = 16; 12 males, 4 females): New Zealand: Central Otago (CO): 2♂♂, Carrick Range, Watts Rock, 1400 m, 11 III 1979, J. C. Watt, litter (NZAC); Mid Canterbury (MC): 1♂ 2♀♀, Bealy Spur, 750 m, 1 VI 1981, C. A. Muir, moss and rotten logs (NZAC); Otago Lakes (OL): 4♂♂, E Matukituki V, 400 m, J. W. Early, 30 I–4 II 1987, yellow pan tap (LUNZ); 1♂, Mt. Aspiring NP, Glacier Burn, 30 I 1987, J. W. Early, 400 m, sweeping Nothofagus forest (LUNZ); Westland (WD): 3♂♂ 1♀ (1♀, slide-mounted), Klondyke Corner, Arthurs Pass, 25 X 1970, D. S. Horning, litter (NZAC); 1♂, Mt. Tuhua, 1067 m, e side of L. Kaniere, 20 X 1984, C. F. Butcher, litter and mats (NZAC); 1♀, Okarito Trig, 150 m, 15 I 1982, J. W. Early, sweeping ferns and kiekie in rimu forest (LUNZ).
This species is named after Dr. Joseph Parker, world pselaphine beetle specialist, and an enthusiastic supporter of this study.
This species is distinguished from the other species of this genus by the following combination of characters: body length 1.8–2.0 mm; eyes large, as long as temples (Fig.
Length 1.8–2.0 mm. Body reddish brown, antennae, legs, maxillary palpi, elytra paler (Fig.
Central Otago (CO), Dunedin (DN), Mid Canterbury (MC), Otago Lakes (OL), Westland (WD) (Fig.
Specimens were collected using yellow pan traps, by sweeping, or by sifting forest litter.
Holotype. New Zealand: Dunedin (DN): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “New Zealand DN Leith Saddle / Leith trig tck, 420m 26 Sep 2011”, “Washed soil sample. temperate cloud forest”, “NZMS 260 144: 173868 430m”, “Voucher specimen “Beetles of Dunedin” project. JT Nunn coll.”, “HOLOTYPE Pseudoexeirarthra hlavaci Park and Carlton des. 2013”. Paratypes (n = 29; 11 males, 18 females): New Zealand: Dunedin (DN): 2♂♂, Swampy Summit, Dunedin, 29 X 2000, J. Nunn, tussock litter (JTN); 1♂, Leith Saddle, Swampy Spur Tck, 14 X 2001, J. Nunn, surface soil (JTN); 1♂, Waitati, 2 VIII 2008, washed soil sample in broadleaf forest (JTN); 1♂, Grahams Bush, Mt Cargill, 6 XII 2002, J. Nunn, FIT (JTN); 1♀, Swampy Summit, Dunedin, 7 I 2000, J. Nunn, shrubbery (JTN); 1♀, Cloud Forest of Leith tr, 30 XI 2003, J. Nunn, moss from tree trunk (JTN); 1♀, Grahams Bush, Mt. Cargill, 9 XII 2001, J. Nunn, podocarp-kamahi-griselinia litter (JTN); 1♀, Careys Creek near Waitati, 12 IV 2008, J. Nunn, washed soil in kanuka forest (JTN); 1♀, Swallow tr, Herbert Forest, 28 VII 2007, washed soil, J. Nunn (JTN); 1♀, Waipori Falls, 17 V 1998, Nothofagus forest litter (JTN); Southland (SL): 3♂♂, Papatowai, 9 II 1989, J. Nunn (JTN); 1♂ 3♀♀, Bog Burn, Waterloo Burn tck, 4 VI 2007, J. Nunn, washed soil in Nothofagus forest (JTN); 1♂, Purakaunui Falls, Catlins, 24 VI 2006, J. Nunn, soil sample in Nothofagus forest (JTN); 1♂, Pourakina River Walk, 27 X 2002, J. Nunn, bracket fungus (JTN); 2♀♀, Princhester Base Hut, Takitimu Forest, 4 VI 2007, J. Nunn, washed soil beech forest, 425 m (JTN); 3♀♀ (1♀, slide-mounted), Blacks Gully, Blue Mtns, 390 m, 2 XII 2006, J. Nunn, washed soil in Nothofagus forest (JTN); 2♀♀, Tautuku, 8 III 1989, J. Nunn (JTN); 1♀, Rakahouka, 2 IX 2007, J. Nunn, washed soil (JTN).
This species is named after Mr. Peter Hlaváč, world pselaphine beetle specialist, and an enthusiastic supporter of this study.
This species is distinguished from the other species of this genus by the following combination of characters: body length 2.3–2.6 mm; eyes large, as long as temples (Fig.
Length 2.3–2.6 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
Specimens were collected mostly by soil washing in Nothofagus and broadleaf forests.
Holotype. New Zealand: North Canterbury (NC): 1♂ (NZAC), aedeagus dissected and mounted in balsam on a clear plastic card, “NEW ZEALAND NC Arthur Pass Kellys Creek, 8 Nov 2005 R Leschen S Nomura”, “ex litter RL1007 42°47'S 171°34'E”, “N.Z. Arthropod Collection, NZAC Private Bag 92170 AUCKLAND New Zealand”, “HOLOTYPE Pseudoexeirarthra nomurai Park and Carlton des. 2013”.
This species is named after Dr. Shûhei Nomura, co-collector of the holotype, world pselaphine beetle specialist, and an enthusiastic supporter of this study.
This species is distinguished from other species of this genus by the following combination of characters: body length 2.0 mm; eyes one-half length of temples (Fig.
Length 2.0 mm. Body reddish brown, antennae, legs, maxillary palpi and elytra paler (Fig.
North Canterbury (NC) (Fig.
The holotype was collected by sifting leaf litter.
We thank James H. Boone, Margaret K. Thayer, Alfred F. Newton (FMNH), John T. Nunn (Dunedin, New Zealand), Richard Leschen (Landcare Research, Auckland, New Zealand), John Early (Auckland Museum, Auckland, New Zealand), John Marris (Lincoln University, Lincoln, New Zealand), and Donald S. Chandler (Durham, New Hampshire, USA) for the loan of valuable specimens and encouragement, and Roger G. Booth (BMNH) for helping with type examinations. Comments of J. T. Nunn and anonymous reviewers guided substantial improvements to the manuscript. This publication is approved by the Director, Louisiana Agricultural Experiment Station as manuscript number 2014-234-18916. This project was funded in part by the Ernst Mayr Travel Grant (Museum of Comparative Zoology, Harvard University) to Jong-Seok Park. This work is/was supported by the USDA National Institute of Food and Agriculture Hatch Project LAB94244.