Lepanus pygmaeus
Creators: name: Gunter, Nicole L.
name: Weir, Thomas A.
Publisher: name: Zenodo
Year: 2019
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Coleoptera
Scarabaeidae
Lepanus
Lepanus pygmaeus

description: Lepanus pygmaeus (Macleay, 1888)
(Figs. 4 G–H, 7A, 9D, 12C, 13)
Temnoplectron pygmaeum Macleay, 1888: 898; Blackburn 1900: 212; Gillet 1911: 40.
Panelus pygmaeus: Lea 1904: 89; Wilson 1922: 39; Lea 1923: 355; Gillet 1925: 5; Paulian 1935: 119; Carter 1936: 101; Paulian 1939: 70.
Panelus arthuri Blackburn, 1900: 211; Lea 1904: 389 (junior synonym).
Lepanus pygmaeus: Matthews 1974: 112.
Type material examined. Temnoplectron pygmaeum lectotype (here designated): “ N.W. Austr ” / “ Panelus (Temnoplectron) pygmaeus, Macl. Kings Sound. N.W.A ” / “ On permanent loan from MACLEAY MUSEUM University of Sydney ” / “ Syntype ”/ “ANIC Database No. 25-057970” / “ Lectotype Temnoplectron pygmaeum Macleay ♀ top designated by Gunter & Weir 2019” (25-057970 in ANIC); ♀ paralectotype: (on same pin as lectotype) “ N.W. Austr ” / “ Panelus (Temnoplectron) pygmaeus, Macl. Kings Sound. N.W.A ” / “ On permanent loan from MACLEAY MUSEUM University of Sydney ” / “ Syntype ” / “ ANIC Database No. 25-057970” “ Paralectotype Temnoplectron pygmaeum Macleay ♀ bottom designated by Gunter &Weir 2019” (25-057970 in ANIC).
Panelus arthuri lectotype (here designated): “6612 T., N.W.A.” (6612 and N.W.A handwritten in red, and T. handwritten in black) / “Type. H. T.” (round label with red border, typed) / “Blackburn coll. 1910-236” (typed) / “ Panelus arthuri. Blackb.” (handwritten) / “= Panelus pygmaeus Macleay, Det Matthews VI-68 ” (handwritten possibly VII-68 but the second “I” appears crossed out) / “NHMUK011525636” / “ Lectotype Panelus arthuri Blackburn designated by Gunter & Weir 2019” (in BMNH), sex undetermined.
Other material examined. An additional 141 specimens from 35 localities from Northern Territory and Queensland including Groote Eylandt and Torres Strait Islands were examined.
Queensland: Saibai Island, Torres Strait, [9°23'43"S, 142°41'27"E], 17.vii.1975, H. Heatwole (T 169467 in QM); Eet Hill, vicinity, Moa (Banks Island), Torres Strait, [10°09'36"S, 142°17'59"E], 9–13.vii.1977, G.B. Monteith & D.J. Cook (T84016–T 84018 in QM); St Pauls, Moa, (Banks Island), Torres Strait, [10°11'17"S, 142°20'1"E], 14–17.vii.1977, G.B. Monteith & D.J. Cook (T84007–T 84014 in QM); Lake Boronto (= Wincheura), Newcastle Bay, Cape York, [10°46'00"S, 142°33'00"E], 50 m, 21–26.ix.1974, G.B. Monteith (25-015584 x 27 in QM also as T50139–T 50165 in QM); Lake Boronto (= Wincheura), Newcastle Bay, Cape York, [10°46'00"S, 142°33'00"E], 2–26.ix.1974, G.B. Monteith (25-015585 x3, 25 -015586 x2, 25 -015587 x8, 25 -015588 x 1 in QM also as T50166–T50179); 8 km E of Mount Tozer, [12°45'00"S, 143°17'00"E], 8.vii.1986, T.A. Weir (25-057972 x 1 in ANIC); Archer River Xing, 70 km N of Coen, Cape York Peninsula, [13°27'00"S, 142°56'00"E], 17– 18.vii.1975, G.B. Monteith (25-015578 x 1 in QM also as T50109); 18 km W of Port Stewart, [14°07'00"S, 143°31'00"E], 25–28.vi.1976, G.B. & S.R. Monteith (25-015576 x5, 25 -015581 x3, 25 -015582 x8, 25 -017239 x3, 25 -017240 x 2 in QM also as T50102–T50106, T50112–T50115, T50117–T50124); McIvor River Crossing, 40 km N of Cooktown, [15°09'00"S, 145°06'00"E], 15–18.vii.1976, G.B. & S.R. Monteith (25-015580 x 1 in QM also as T50111); Port Douglas, [16°28'58"S, 145°27'53"E], 2 m, 21–23.ii.1984, L. Masner (CMNEN-00030239-00030240 in CMNC); Julatten, [16°34'02"S, 145°21'38"E], 15.x.1978, A. & M. Walford-Huggins (CMNEN-00030224 in CMNC); 8 km N Mareeba, [16°55'28"S, 145°25'17"E], 11–13.ii.1987, H.F. & A.T. Howden (CMNEN-00030246 in CMNC); 8 km N Mareeba, [16°55'28"S, 145°25'17"E], 6.ii.1997, H.F. & A.T. Howden (CMNEN-00030247 in CMNC); 8 km N Mareeba, [16°55'28"S, 145°25'17"E], 8.ii.1997, H.F. & A.T. Howden (CMNEN-00030248 in CMNC); 11 km NW Mareeba, Southedge Tobacco Research Station., [16°58'43"S, 145°21'05"E], 28.xii.1986, H.F. & A.T. Howden (CMNEN-00030241-00030242 in CMNC); Southedge Research Station, 12 km NW Mareeba, [16°58'43"S, 145°21'05"E], 3.ii.1989, H.F. & A.T. Howden (CMNEN-00030243 in CMNC); Southedge Research Station, 12 km NW Mareeba, [16°58'43"S, 145°21'05"E], 26–29.xii.1988, H.F. & A.T. Howden (CMNEN- 0 0 0 30244 in CMNC); 7 km NE of Tolga, [17°09'00"S, 145°31'00"E], iii.1987, R.I. Storey & De Faveri (25- 032635 x 1 in QDPC); 7 km NE of Tolga, [17°09'00"S, 145°31'00"E], 2.ii.1987, R.I. Storey & De Faveri (25- 032637 x 1 in QDPC, 25-058982 x 1 in ANIC); 7 km NE of Tolga, [17°09'00"S, 145°31'00"E], 19.i.1987, R.I. Storey & De Faveri (25- 032636 x 1 in QDPC); 7 km NE of Tolga, [17°09'00"S, 145°31'00"E], 27.i.1987, R.I. Storey & De Faveri (25- 0 58984 x 1 in ANIC); 7 km NE of Tolga, [17°09'00"S, 145°31'00"E], iii.1987, R.I. Storey & De Faveri (25-058985 x 1 in ANIC); Tolga (general), [17°13'00"S, 145°29'00"E], 25.ii.1986, J.D. Brown (25-058983 x 2 in ANIC); Tinaroo Waters, 17°11’02”E, 145°33’07”E, 16–17.ii.2017, K. Ebert et al. (25-065552 x 2 in ANIC); Cardwell Gap, 5 km N, 18°30'6"S, 146°10'56"E, 4–12.ii.1998, G.B. Monteith & D.J. Cook, (T 62200 in QM); 17–20 mi. W Atherton, [17°23'09"S, 145°14'58"E], 17.ii.1975, H.F. & A.T. Howden (CMNEN-00030235 in CMNC); Palm Island, Site 3, [18°44'00"S, 146°35'00"E], 50 m, 20–22.ii.2001, G.B. Monteith (25- 060846 x2, 25 -058986 x 1 in ANIC); Palm Island, Site 3, [18°44'00"S, 146°35'00"E], 20–22.ii.2001, G.B. Monteith (T 111953 –T 111957 in QM); Palm Island, Site 2, [18°44'00"S, 146°36'00"E], 20–21.ii.2001, G.B. Monteith (T 111959 –T 111960, T 111962 – T 111964, T 111966 –T 111967 in QM); Paluma Road, 18°59'24"S, 146°17'42"E, 60 m, 14–16.iii.2012, G.B. Monteith (25-053918 x 1 in ANIC also as COL2404); Balgal Beach, [19°01'00"S, 146°24'00"E], 6.v.1974, R.I. Storey (25-058519 x 1 in ANIC); Northern Territory: 11.26774°S 132.26615°E, near Black Point, Site c002 mushroom paperbark with rainforest understorey, 11°16'04"S, 132°15'58"E, 14.viii.2014, M. Braby (25-065548 in ANIC); West Alligator Mouth, 12°14'0"S, 132°16'0"E, 22–24.vii.1979, G.B. Monteith & D.J. Cook (T 111945 – T 111946 in QM); Kakadu National Park, S Alligator River, Gungaree Rainforest Trail, [12°40'22"S, 132°28'53"E], 12–25.xii.1993, S.B. & J. Peck (CMNEN-00030231 in CMNC); 50 km S Darwin, Berry Spring Park, [12°41'48"S, 131°00'00"E], 4–27.xii.1993, S.B. & J. Peck (CMNEN-00030236-00030238 in CMNC); Nourlangie Creek, 8 km N of Mount Cahill, [12°48'00"S, 132°42'00"E], 21.v.1973, E.G. Matthews (25-057971 x 1 in ANIC); Fish River Station Reserve, Bush Blitz Site B3, 13°49'37"S, 130°42'58"E, 23–26.v.2012, N.L. Gunter (25-057611 x1, 25 - 0 57610 x 1 in ANIC); Litchfield National Park, Pethricks Rainforest, [13°07'40"S, 130°39'43"E], 8–30.xii.1993, S.B. & J. Peck (CMNEN-00030234 in CMNC); Groote Eylandt (General), [14°02'00"S, 136°36'00"E], N.B. Tindale, (25-027815 x 1 in SAMA); The Amphitheatre, Nitmiluk National Park, 14 09'0"S, 132 23'0"E, 16.iii.1995, G.B. Monteith & Janetzki (T 111947 –T 111949, T 111951 in QM); Elsey National Park, 12- Mile Camp, [14°57'00"S, 133°13'00"E], 26–27.vi.2003, G.B. Monteith (T 111934 in QM); Seven Emu, 32 km NNE Pungalina Homestead, Australian Wildlife Conservancy camp, 16°27'53"S, 137°33'17"E, 2–6.vii.2012, N.L. Gunter (25-057710 x1, 25 - 0 57711 x1, 25 - 062104 x1, 25 -057636 x 9 in ANIC); Pungalina Homestead, 16°43'16"S, 137°24'55"E, 26.vi– 8.vii.2012, N.L. Gunter (25-057633 x 1 in ANIC); Pungalina, 9.6 km SSE of Pungalina Homestead, Karns Creek, 16°47'30"S, 137°27'27"E, 26.vi–8.vii.2012, N.L. Gunter (25-057634 x1, 25 -057635 x 2 in ANIC).
Description. Dark brown in colour, sometimes with red humeri. Antennal club grey.
Total length: 2.2–2.6 mm. Lectotype measurements (of Temnoplectron pygmaeum): (♀) Total length 2.2 mm, elytral width 1.5 mm. Lectotype measurements (of Panelus arthuri): (sex not recorded) Total length 2.4 mm, elytral width 1.5mm.
Male: Head: U-shaped between clypeal teeth, which are short and upturned. Margin slightly angulate at clypeogenal suture. Genal angle prominent. Basal carina present and interrupted medially. Dorsal part of eyes moderate in size, separated by an interocular space approximately 16–17 times eye width (32–34: 2); eye canthus not dividing eye. Head 1.4 × wider than long (44:32).
Prothorax: Hypomeral surface finely reticulate with very light punctation, punctures along basal margin slightly elongate and different from the disc. Hypomeral striae about half the length of the hypomeron. Pronotum 1.6 × wider than long (70:44).
Elytra: Surface smooth. Striae 1–6 or 1–5 appearing double with very finely punctate edges. All striae on disc not equal width, striae 5–7 not as wide, striae 6 and/or 7 appearing single (sometimes stria 5 also appears single). Apically, with a short groove extending from end of epipleuron to apex. Elytra 1.2 × wider than long (85:70).
Legs: Front edge of protibiae truncate; apical digit somewhat elongate and sharp; two teeth on outer edge. Inner edge of metatibiae almost straight. Metatarsi with 2nd tarsomere subequal to the 5th. Tarsal claws small and toothed.
Abdomen: Pygidium with wide pear-shape depression containing a somewhat triangular tubercle in upper third. The depression takes up over half of surface and extends under the slightly concave upper edge. Abdominal surface very finely reticulate right across. Segment 6 finely punctate.
Pterothorax: Medial lobe of metaventrite finely punctate and broadly margined between mesocoxae. Mesometaventral suture very slightly arched. Lateral lobe of metaventrite smooth and punctate with setae. Mesoventrite with slight notch anteriorly. Mesoventrite and mesepimeron smooth. Metanepisternum reticulate.
Female: Apical digit absent; apical spur extending straight from front edge. Abdominal ventrite 6 slightly longer than in male.
Distribution. A widespread species known from coast to coast in equatorial latitudes north of about 19°S, the vast majority of specimens have been collected in open forest (Fig. 13). Specimens have been recorded from the IBRA regions Dampierland (DAL), Darwin Coastal (DAC), Pine Creek (PCK), Daly Basin (DAB), Arnhem Coast (ARC), Gulf Coastal (GUC), Cape York Peninsula (CYP), and Wet Tropics (WET) on mainland Australia, and on Groote Eylandt (NT), Saibai and Moa Islands (Torres Strait, Queensland). BIOCLIM modelling indicates appropriate climate for L. pygmaeus exists in Central Kimberley (CEK), Northern Kimberley (NOK), Victoria Bonaparte (VIB), Ord Victoria Plain (OVP), Sturt Plateau (STU), Mount Isa Inlier (MII), Central Arnhem (CEA), Arnhem Plateau (ARP), Gulf Fall and Uplands (GFU), Gulf Plains (GUP), Einasleigh Uplands (EIU), Desert Uplands (DEU), Brigalow Belt North (BBN), Central Mackay Coast (CMC), and South Eastern Queensland (SEQ) IBRA regions. While much of northern Australia remains under sampled, numerous habitat types across CMC, BBN, and SEQ have been extensively surveyed and L. pygmaeus has not been collected in these IBRA regions.
Status of type specimens. Temnoplectron pygmaeum Macleay, 1888. Macleay described this species from one or more specimens collected by Froggatt in the vicinity of King’s Sound, Western Australia and housed in the Macleay Museum; Monteith & Rossini (2017) detailed the Froggatt expedition and history of Macleay’s specimens. In his revision of the genus Lepanus, Matthews (1974) stated that he saw the holotype (sex not recorded) in the Macleay Museum, but when the coleopteran types from that museum were transferred to ANIC, only two female specimens on the one card from that locality and labelled as Panelus (Temnoplectron) pygmaeus Macl., Kings Sound, Western Australia were located. This label may have been added by Macleay Museum’s curator, George Masters, following the transfer of the species to the genus Panelus (Lea 1904). Interestingly, this label appears with added locality details compared to the smaller “N.W. Austr” label and may either be taken from Macleay’s description listing material from the vicinity of King’s Sound or represent a replaced label. These two specimens were presumed to be syntypes by Britton and Stanbury (1981), but there was no label to indicate that they were seen by Matthews. Subsequently, Cassis and Weir (1992) interpreted the incorrect reference to a holotype as a lectotype designation, but, as Matthews did not explicitly state which specimen or give characteristics of the specimen he called holotype, we consider this not to be a valid lectotype designation under Article 74.5 (International Commission on Zoological Nomenclature 1999). The two female specimens have subsequently been remounted on to separate points on the one pin and we hereby designate the female on the upper point with all tarsi intact as the lectotype and have labelled it as such. The female on the lower point with the protarsi missing thus becomes a paralectotype.
Panelus arthuri Blackburn, 1900. Blackburn described this species from one or more specimens from NW Australia given to him by AM Lea and listed as from the Macleay Museum. In his revision of the genus Lepanus, Matthews (1974) stated that he saw the holotype of Panelus arthuri (sex not recorded) in the Macleay Museum, but when the coleopteran types from that museum were transferred to ANIC, no such specimen was found nor was it listed in Britton and Stanbury’s (1982) review of Coleoptera types from Macleay Museum. A literature search revealed that the type was sent to the BMNH (NHM) along with many other Blackburn types in 1910 and after his death in 1912 (Lea 1912). Subsequent contact with that institution resulted in the specimen being located and borrowed by us. Cassis and Weir (1992), following Matthews (1974), listed a holotype (probable). This specimen located in BMNH is certainly the type of Panelus arthuri as evidenced by the characteristic Blackburn labels detailed above (Blackburn marked his types with a “T.” (see Lea 1912)), and by the attached Matthews label indicating it was synonymous with Panelus pygmaeus. Blackburn did not specifically say in his description that there was only a single specimen examined so this specimen is hereby designated as the lectotype of Panelus arthuri Blackburn, 1900. The specimen has been remounted and lacks both protibiae and protarsi, which precludes the sex being determined without dissection, but exhibits enough other characters to show that it conforms to the species concept of Lepanus pygmaeus. Further confirmation of the synonymy of Lea (1904), can be taken from the original justification that “both descriptions were drawn up from specimens taken at King’s Sound by W.W. Froggatt”. There is no reason to question this line of evidence given Lea’s involvement in the transfer of material to Blackburn, Blackburn’s reference that the material was from Macleay Museum, and that Froggatt’s King Sound material was deposited there.
Matthews (1974) examined almost 500 specimens of Lepanus during his revision but only had access to 14 specimens of L. pygmaeus from eight localities. Fortunately, hundreds of specimens belonging to the L. pygmaeus species group have since been collected allowing for rigorous assessment of morphological variation within and between species. Of the material examined by Matthews, the specimen listed from Charters Towers, collected April 1965 by Bornemissza & Yapp (25- 060385 in ANIC) is L. menendezae, while the specimen from Beames Brook, 15 Mi SW Burketown collected May 1972 by Monteith & Monteith (25-058521 in ANIC) is L. guthrieae. Matthews (1974) noted the morphological variation of Queensland specimens of L. pygmaeus and listed diagnostic characters of the species group within the remarks section. Based on these remarks, it is likely he was referring to the variation observed in these specimens from Charters Towers and Beames Brook, or the specimen from Mornington Island, which we did not examine (Matthews 1974).
Specimens from the Northern Territory (Fish River and Pungalina) and Queensland (Paluma) have been sequenced and were included in the phylogenies of Gunter & Weir (2017) and Gunter et al. (2018). These L. pygmaeus specimens formed a clade within a larger clade containing other members of the pygmaeus group. Intraspecific variation is slightly higher than observed for other species of Lepanus, however most other species are more narrowly distributed and other broadly distributed species (e.g., Lepanus australis) also exhibit similarly high levels of intraspecific variation (see Gunter & Weir 2017).
Lepanus pygmaeus has been collected in a variety of habitats including rainforest, heath, open and dry sclerophyll forests, and pasture. Most specimens have been collected in dung baited pitfall traps or at light but a handful of specimens have also been collected in mushroom baited pitfall traps suggesting they may be generalist feeders. Recently collected specimens of L. pygmaeus from Tinaroo Waters on the northern end of the Atherton Tableland were attracted to traps baited with mealworm/cricket carrion. These specimens were collected during a survey that explored bait preferences and interestingly, L. pygmaeus was only collected in the invertebrate carrion traps and not recorded from the traps baited with dung, fruit, mushroom, or vertebrate carrion (K. Ebert & G.B. Monteith, personal communication).
Lepanus andersonorum Gunter & Weir, new species
(Figs. 4C, 5A, 8A, 10A, 14)
Type series. Holotype: ♂ “ 12km SSE Heathlands, 15 Jan 1992 – 26 Jan 1992, Weir,T.A.” [11°51'00"S, 142°38'00"E] / “ human dung trap closed forest” / “ANIC Database No. 25-023701” / “ANIC Image” / “ Lepanus CYP7 det. T.A. Weir 2013” in ANIC” in ANIC. Paratypes: 12 km SSE Heathlands, [11°51'00"S, 142°38'00"E], 15–26.i.1992, T.A. Weir (25-023701 x2 ♂, x1 sex undetermined in ANIC); 12 km SSE Heathlands, [11°51'00"S, 142°38'00"E], 20.i.1992, T.A. Weir & I.D. Naumann (25-023700 x1 ♂, x1 ♀ in ANIC); 12 km SSE Heathlands, [11°51'00"S, 142°38'00"E], 22.iii–25.iv.1992, T. McLeod (25-023703 x1 ♂, x1 ♀ in ANIC); “ABRS Area 2”, Dividing Range, 15 km W of Captain Billy Creek, Cape York Peninsula, [11°40'00"S, 142°45'00"E], 4–9.vii.1975, G.B. Monteith (25-015558 x1 ♀ in QM also as T50228); “ABRS Area 3”, Dividing Range, 15 km W of Captain Billy Creek, Cape York Peninsula, [11°40'00"S, 142°45'00"E], 4–9.vii.1975, G.B. Monteith (25-015556 x1 ♂, 25-015622 x1 ♂, x1 damaged in QM also as T50226, T49910–T49911); “ABRS Area 6”, Captain Billy Creek, Cape York Peninsula, [11°40'00"S, 142°50'00"E], 9–13.vii.1975, G.B. Monteith (25-015557 x1 ♀ in QM also as T50227); Heathlands, [11°45'00"S, 142°35'00"E], 22.i.1992, T.A. Weir & I.D. Naumann (25-023702 x1 ♂ in ANIC); Green Hill Thursday Island, [10°35'00"S, 142°13'00"E], 20–23.v.2003, G.B. Monteith (T169468 x1 ♂ in QM).
Description. Head and pronotum dark brown and elytra light brown. Antennal clubs white.
Total length: 1.8–2.2 mm. Holotype measurements: Total length 2.0 mm, elytral width 1.4 mm.
Male: Head: Broad U-shaped between clypeal teeth, which are short and very slightly upturned. Head reticulate between clypeal suture and anterior edge. Margin slightly angulate at clypeogenal suture. Genal angle rounded. Basal carina absent. Dorsal part of eyes moderate in size, separated by interocular space approximately 10 times eye width (29:3); eye canthus nearly dividing eye. Head 1.25 × wider than long (35:28).
Prothorax: Basal margin slightly angulate. Hypomeral surface smooth with light punctation and long setae. Hypomeral striae about half the length of the hypomeron. Pronotum 1.75 × wider than long (56:32).
Elytra: Surface reticulate at apical half and at the sides. Intervals finely punctate, and with very fine, straight setae longer than the space between punctures. Elytral striae distinct but slightly impressed on apical half. Striae 1– 7 double with very fine, punctate edges. All striae on disc equal width. 8th striae almost extends to the apex of the elytra forming a slight flange. Elytra 1.05 × wider than long (68:65).
Legs: Front edge of protibiae deeply angulate; apical digit somewhat elongate and sharp; two teeth on outer edge. Inner edge of metatibiae almost straight. Metatarsi with 2nd tarsomere subequal to the 5th. Tarsal claws minute and simple.
Abdomen: Pygidium with U-shaped depression containing a faint tubercle that is a raised line but may be absent. Depression occupies medial third of pygidial disc and extends under the angulate upper edge of pygidium. Edges of depression are only moderately sharp, bottom edge is slightly pointed. Abdominal surface reticulate right across. Segment 6 slightly gibbous and finely punctate.
Pterothorax: Medial lobe of metaventrite finely punctate and broadly margined between mesocoxae. Lateral lobe of metaventrite punctate with fine reticulation. Meso-metaventral suture very slightly arched but with a fine short carina extending backwards from middle of meso-metaventral suture as in Fig. 4C. Mesoventrite with slight notch anteriorly. Mesoventrite smooth. Mesepimeron finely reticulate. Metanepisternum reticulate.
Female: As per the L. pygmaeus species group
Etymology. Named for A. Chace and Josie Anderson for their generous support of The Cleveland Museum of Natural History for this research, and encouragement of women pursuing careers in science.
Distribution. Known only from closed forests of northern Cape York Peninsula and Thursday Island (Fig. 14). So far only 15 specimens are known and the species may be more widespread than the CYP bioregion.
Comments. Lepanus andersonorum was previously referred to as “ Lepanus CYP7” under the informal nomenclature sensu Yeates et al. (2011). Specimens have been collected using a variety of methods including dung baited pitfall traps, berlesed leaf-litter and intercept traps.
description: Published as part of Gunter, Nicole L. & Weir, Thomas A., 2019, Revision of Australian species of the dung beetle genus Lepanus (Coleoptera: Scarabaeidae: Scarabaeinae): key to species groups and description of 14 new species from the L. pygmaeus species group, pp. 41-80 in Zootaxa 4564 (1) on pages 50-55, DOI: 10.11646/zootaxa.4564.1.2, http://zenodo.org/record/2588621