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Research Article
Revision and molecular phylogeny of the genus Gastrobothrus (Coleoptera: Staphylinidae: Pselaphinae)
expand article infoYeon-Jae Choi‡§, Richard A. B. Leschen|, Jong-Seok Park§
‡ Kyungpook National University, Sangju, Republic of Korea
§ Chungbuk National University, Cheongju, Republic of Korea
| Manaaki Whenua – Landcare Research, Auckland, New Zealand
Open Access

Abstract

We conducted a taxonomic revision of the New Zealand members of Gastrobothrus sensu novo (the former “Gastrobothrus-group”) consisting of the genera Eupinogitus Broun, 1921, Gastrobothrus Broun, 1882, and Physobryaxis Hetschko, 1913. Based on the results of a molecular phylogenetic analysis, a new synonymy is established for the following taxa: Gastrobothrus Broun, 1882 (= Eupinogitus Broun, 1921, syn. nov. = Physobryaxis Hetschko, 1913, syn. nov.). New combinations are proposed as well: Gastrobothrus inflatus, comb. nov. (= Physobryaxis inflatus (Sharp, 1874)) and Gastrobothrus sulcipennis, comb. nov. (= Eupinogitus sulcipennis Broun, 1921). The five previously described species are redescribed and 14 new species are described: Gastrobothrus angustus sp. nov., G. biligo sp. nov., G. directus sp. nov., G. giganteus sp. nov., G. grossus sp. nov., G. lunatus sp. nov., G. occultus sp. nov., G. pilosus sp. nov., G. planiventris sp. nov., G. pumilus sp. nov., G. quadrus sp. nov., G. retusus sp. nov., G. spinifer sp. nov., and G. trigonus sp. nov. An identification key to all formally described and unnamed species within the genus and high-resolution images of diagnostic characters for each are provided.

Key words

Ant-loving beetles, Brachyglutini, Crosby’s code, Multi-gene phylogeny, New combination, New Zealand, Synonym, Systematics

1. Introduction

New Zealand is renowned for its diverse and often discordant biota, particularly its high level of insect endemism (Buckley et al. 2015). Notably, the New Zealand pselaphine fauna lacks known species of the supertribes Batrisitae and Clavigeritae (Klimaszewski et al. 1996; Parker and Grimaldi 2014; Nomura and Leschen 2015). In contrast, the majority of indigenous genera and species, particularly those of Faronitae, are distributed across New Zealand (Park and Carlton 2011, 2013, 2014a, 2014b, 2015a, 2015b, 2015c, 2015d, 2015e). Regarding Goniaceritae, only one tribe, Brachyglutini, is known to exist in New Zealand (Klimaszewski et al. 1996). Thomas Broun, a Scottish soldier and entomologist, made significant contributions to our understanding of Brachyglutini by describing numerous species from New Zealand (Broun 1880, 1881, 1893, 1921).

More recently, Shen and Leschen (2018, 2019, 2020) and Shen et al. (2020a, 2020b) have revised and analyzed the phylogenetic relationships within the tribe Brachyglutini of New Zealand. However, their phylogenetic inference primarily focused on relationships within the genus Eupines King (Shen et al. 2020a) and suggested the existence of the “Gastrobothrus-group”. This clade includes the type species of the genera Gastrobothrus Broun, Physobryaxis Hetschko, Eupinogitus Broun, and an unidentified species of the genus Rybaxis Saulcy. Their results indicated moderately to poorly supported nodes due to the limited number of known species and the gene loci employed.

In this study, we describe 13 and one new species collected on North Island and South Island, respectively. Furthermore, we conducted a comprehensive phylogenetic analysis using a dataset comprising three mitochondrial genes (COI, COII, and 16S) and three nuclear genes (18S, 28S, and wingless). This analysis included both new and type species to validate the monophyly of the “Gastrobothrus-group” and the necessity of taxonomic changes in its three genera. Hereafter, we use “Gastrobothrus sensu novo” to refer to the monophyletic Gastrobothrus-group (former Gastrobothrus + Eupinogitus + Physobryaxis).

2. Material and methods

2.1. Taxon sampling

We sampled 12 species within Gastrobothrus sensu novo and representatives from allied genera, Anabaxis foveolata (Broun, 1880) and Simkinion bimanum Park & Pearce, 1962 as outgroup taxa from New Zealand (Table S1).

2.2. DNA extraction and PCR

Specimens for DNA extraction were stored in 95% EtOH at –80°C to minimize degradation. Genomic DNA was extracted using the Qiagen DNeasy® Blood & Tissue kit following the manufacturer’s protocol with the modification of using 50 µL of AE buffer. The oldest specimen that yielded informative genomic DNA in this study was collected in 2008 (extracted in 2023). Partial sequences of three mitochondrial genes and three nuclear genes were targeted and amplified. The PCR mixture consisted of 1 µL genomic DNA, 7 µL distilled water, 10 µL 2× Lamp Taq PCR Master Mix (BioFACT™), 1 µL forward primer, and 1 µL reverse primer. Examined genes and primers are detailed in (Table S2). The remaining exoskeletons after extraction were mounted on glass slides to examine detailed morphological characters such as male genitalia.

2.3. Gene Sequencing

Sanger sequencing was commercially performed by Macrogen (Daejeon, Republic of Korea), and both forward and reverse sequences were used to generate consensus sequences. The newly generated sequence data were BLASTed against GenBank (www.ncbi.nlm.nih.gov/genbank). Note that Rybaxis sp. from the dataset of Shen et al. (2020a) was excluded due to identification credibility issues.

2.4. Phylogenetic analyses

Consensus sequences were aligned using MAFFT v7.490 under the L-INS-i algorithm (Katoh and Toh 2008) for all loci, and less informative sites were removed with GBLOCKS 0.91b (Castresana 2000) only for ribosomal RNA gene loci. PartitionFinder 2.1.1 (Lanfear et al. 2017) selected the best-fit partitioning scheme and substitution models. Protein-coding genes (COI, COII, and wingless) were partitioned for each codon position, while ribosomal RNA genes (16S, 18S, and 28S) were treated as single partitions. The following substitution models were selected: (i) 16S — GTR + G; (ii) 18S(2) and 18S(3) — GTR + G; (iii) 28S — GTR + I + G; (iv) position 1 of COI and position 1 of COII — GTR + G; (v) position 2 of COI — GTR; (vi) position 3 of COI — GTR + G; (vii) position 2 of COII — GTR + G; (viii) position 3 of COII — GTR + G; (ix) position 1 of wingless — GTR + G; (x) position 2 of wingless — GTR; (xi) position 3 of wingless — GTR + G.

Bayesian inference (BI) was conducted using MrBayes 3.2.7a (Ronquist et al. 2012) with two independent runs of four chains for 5,000,000 generations, sampling trees every 1,000 generations with a heating parameter of 0.2, and discarding the first 25% of trees as burn-in. The starting tree was randomly selected. For all partitions the “ratepr” parameter was set to “variable” (File S1).

Maximum likelihood (ML) analysis was performed using IQ-TREE 2.0.3 (Minh et al. 2020) with default settings and 1,000 ultrafast bootstrap replicates, applying the same partitioning scheme as in the BI analysis.

2.5. Morphological study

A total of 104 specimens from the New Zealand Arthropod Collection (NZAC, Auckland, New Zealand) and 13 specimens from the Auckland War Memorial Museum (AMNZ, Auckland, New Zealand) were examined. The depository of each specimen is indicated in parentheses, and the specimens are expected to be returned to their original collections after examination and description. Specimens were observed using Leica M80 and DM1000 LED optical microscopes. To study detailed characters and male genitalia, structures were mounted on slides using Euparal solution. Images were generated using Sony ILCE-7RM3 mirrorless camera and stacked with Zerene Stacker v. 1.04. The length of the holotypes was determined by measuring the head, pronotum, elytra, and abdomen in lateral view (Fig. 1). The width of body parts was measured according to the widest point. Specimen label data for the holotypes were transcribed verbatim, and data for other specimens were standardized for consistency. Terminology and nomenclature used follow Chandler (2001) and Lawrence et al. (2011). We employ the words “tergites” and “sternites” when referring to the abdominal segments. The area codes and their localities of the North Island and South Island of New Zealand are based on fig. 2 in Crosby et al. (1998), which was subsequently re-drawn (Figs 2, 3).

Figure 1. 

Measuring method. Sum of head, pronotum, elytra, and abdomen length. — Abbreviations: HL = Head length; PL = Pronotum length; EL = Elytra length; AL = Abdomen length.

Figure 2. 

Distribution map of the genus Gastrobothrus Broun, 1882, sensu novo in the North Island of New Zealand. Two-letter codes represent standard geographic regions (Crosby et al. 1998). Colored symbols indicate collection localities. Note that G. buckleyi is excluded from this map as it is endemic to the Three Kings Islands, off the northern coast of the mainland.

Figure 3. 

Distribution map of the genus Gastrobothrus Broun, 1882, sensu novum in the South Island of New Zealand. Two-letter codes represent standard geographic regions (Crosby et al. 1998). Colored symbols indicate collection localities.

3. Results

3.1. Phylogenetic analyses

The final concatenated molecular dataset is 3,915 bp, comprising 16S (369 bp), 18S (672 + 517 bp), 28S (559 bp), COI (685 bp), COII (630 bp), and wingless (483 bp). GenBank accession numbers for all sequences used are provided in Table S1. Both topologies of the resulting BI (Fig. 4) and ML (File S2) trees indicate that the Gastrobothrus sensu novo, comprising the former genera Eupinogitus Broun, Gastrobothrus Broun, and Physobryaxis Hetschko is a monophyletic sister group to the outgroups with strong support values (posterior probability, PP = 1; bootstrap, BS = 100). However, they exhibit topological incongruence regarding the branching order. Clades C and D are recovered as sister groups in the ML analysis, whereas their relationship remains unresolved in the BI analysis. Between the two analyses, the BI tree was selected as the preferred topology because most clades are strongly supported. Each clade is recovered as monophyletic with strong PP values. Among the clades, relationships within clade A are poorly resolved.

Figure 4. 

Phylogenetic tree of the genus Gastrobothrus Broun, 1882, sensu novo reconstructed by Bayesian inference. Right shows gular characters of members grouped into each clade, outgroup, and species that were not included in the dataset.

A transverse pronotal sulcus crossing the antebasal foveae, elytral striae, and subhumeral foveae are the primary diagnostic features used to distinguish the genus Gastrobothrus from allied genera. If we adhere to the previous diagnosis by Nomura and Leschen (2006) or the identification key by Shen et al. (2020a), some of our new species would be classified into the former genera Eupinogitus, Gastrobothrus and Physobryaxis. However, the molecular phylogeny constructed in this study reveals that the Gastrobothrus sensu novo itself is monophyletic. While members within clade A possess a pronotal sulcus, G. pumilus sp. nov. lacks this trait. Additionally, the clade comprising G. sharpi + G. planiventris sp. nov. is sister to G. giganteus sp. nov., which exhibits a distinct pronotal sulcus. Clade B comprises two species, previously recognized as E. sulcipennis and the newly described G. spinifer sp. nov., both of which share thoracic characters such as a pronotal sulcus and elytral striae. Given these incongruences in character states, we consider that the pronotal sulcus has evolved independently, preventing the separation of members within Gastrobothrus sensu novo into distinct genera. Based on our molecular phylogeny, we propose a new synonymy for the genera Eupinogitus and Physobryaxis under the genus Gastrobothrus. Members within this genus are characterized by gular carinae that gradually broaden toward the apex (Fig. 4). Gastrobothrus sp. 15–17, which will be described after collecting additional specimens, also exhibit this trait. In contrast, G. buckleyi exhibits sharply broadening gular carinae, similar to outgroup taxa such as Anabaxis foveolata and Simkinion bimanum. Superficial differences in morphology and distribution prompt a reconsideration of the taxonomic position of G. buckleyi relative to its congeners. The inclusion of this species and taxa from Brachyglutini across Australasia and elsewhere in future dataset could potentially lead to a revision of the generic classification we propose here.

3.2. Taxonomy

Family Staphylinidae Latreille, 1802

Subfamily Pselaphinae Latreille, 1802

Supertribe Goniaceritae Reitter, 1882

Tribe Brachyglutini Raffray, 1904

Subtribe Brachyglutina Raffray, 1904

Gastrobothrus Broun, 1882, sensu novo

Eupinogitus Broun, 1921, syn. nov.

Physobryaxis Hetschko, 1913, syn. nov.

Type species.

Bryaxis abdominalis Broun, 1880

Diagnosis.

Gular carinae gradually broadened toward apex (except for G. buckleyi); antennomere 5 slender and elongate; elytra sharply narrowed at base; leg with swollen femur; abdominal sternite V or VI often bearing spatula(e).

3.2.1. Key to the species of the genus Gastrobothrus (based on males)

1 Body longer than 3.5 mm (Figs 9A, 23A) 2
1’ Body shorter than 3.2 mm 3
2 Protibia with spine at middle level; abdominal spatula on sternite VI (Fig. 9D) G. giganteus sp. nov.
2’ Protibia lacking spine; abdominal sternite VI simple (Fig. 23B) Gastrobothrus sp. 15
3 Vertexal foveae setose (Figs 5B, 6B, 8B, 10B, 11B, 13B, 14B, 15B, 16B, 19B, 20B, 21B, 22B) 4
3’ Vertexal foveae asetose (Figs 7B, 12B, 17B, 18B) 17
4 Each elytron with sutural stria, discal stria, and subhumeral fovea (Figs 20C, 21D) 5
4’ Elytra without stria or subhumeral fovea 6
5 Spine on leg acute (Fig. 20F, G, I); abdominal sternites IV–VIII depressed (Fig. 20D) G. spinifer sp. nov.
5’ Spine on leg minute (Fig. 21G, H, J); abdominal sternites not depressed (Fig. 21E) G. sulcipennis (Broun, 1921) comb. nov.
6 Abdominal sternites V–VIII extremely concave and sternite VI with pair of spines projecting toward midline (Fig. 19E) G. sharpi (Broun, 1880)
6’ Abdominal sternites normal or slightly depressed 7
7 Pronotum with transverse sulcus spanning antebasal foveae (Figs 5C, 6C, 11D, 14C, 22C, 23C, 23F) 8
7’ Pronotum without sulcus 14
8 Abdominal sternite V with a pair of spatulae (Figs 6D, 23F) 9
8’ Abdominal sternite VI with single spatula (Figs 5D, 11D, 14D, 22E, 23D) 10
9 Abdominal sternites IV–VIII medially depressed; spatulae distantly situated (Fig. 6D) G. biligo sp. nov.
9’ Abdominal sternites not depressed; spatulae closely situated (Fig. 23D) Gastrobothrus sp. 17
10 Abdominal spatula wider than median sclerite of sternite IX (Figs 14D, 22E) 11
10’ Abdominal spatula not wider than median sclerite of sternite IX 12
11 Abdominal spatula triangular (Fig. 22E) G. trigonus sp. nov.
11’ Abdominal spatula rounded (Fig. 14D) G. pilosus sp. nov.
12 Metatibia with apical trichome (Figs 5H, 11I) 13
12’ Metatibia without apical trichome Gastrobothrus sp. 16
13 Abdominal spatula narrower than median sclerite of sternite IX (Fig. 11E) G. inflatus (Sharp, 1874) comb. nov.
13’ Abdominal spatula as wide as median sclerite of sternite IX (Fig. 5D) G. angustus sp. nov.
14 Leg enlarged, mesotibia with acute apical spine G. grossus sp. nov.
14’ Leg slender, tibia lacking spine or bearing minute spine 15
15 Aedeagal endophallus with armature exceeding penis (Figs 8F, 15E) 16
15’ Aedeagal endophallus with armature not exceeding penis (Fig. 13F) G. occultus sp. nov.
16 Apex of aedeagal armature medially curved (Fig. 15E) G. planiventris sp. nov.
16’ Aedeagal armature straight (Fig. 8F) G. directus sp. nov.
17 Body overall shiny G. ignotus (Broun, 1881)
17’ Body not glossy, at least moderately covered with setae 18
18 Mesotibia with apical spine or tubercle (Figs 7H, 12G, 17G, 18G) 19
18’ Mesotibia lacking apical spine 22
19 Abdominal sternite V with single spatula (Figs 12D, 17D, 18D) 20
19’ Abdominal sternite V without spatula G. buckleyi Théry & Leschen, 2013
20 Median spine on protibia acute (Figs 12F, 17F) 21
20’ Median spine on protibia blunt (Fig. 18F) G. retusus sp. nov.
21 Abdominal spatula transverse and rounded (Fig. 12D) G. lunatus sp. nov.
21’ Abdominal spatula narrow and lingulate (Fig. 17D) G. quadrus sp. nov.
22 Metatibia with apical trichome (Fig. 16H) G. pumilus sp. nov.
22’ Metatibiae without trichome G. platynotus (Broun, 1893)

3.2.2. Species descriptions

Gastrobothrus angustus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 5

Type material.

N = 10, 7♂♂ 3♀♀. — Holotype. ♂ point-mounted: “New Zealand: ND, Warawara St Forest, nr Hauturu Stm Campsite, 7-/Dec/2008, 35.22S 173.17E, sifting rotten wood and leaf litter, RL1372, R. Leschen, T. Buckley & D. Seldon // NZAC04291730”. — Paratypes. 1♂ point-mounted (NZAC04261726): “NEW ZEALAND: BP, Waimana Valley, II-1937 // A. E. Brooke Collection”. 1♂ point-mounted (NZAC04261720): “NEW ZEALAND: CL, Kopu Road, Coromandel Peninsula, 17-III-1967, leaf litter, R. A. Cumber”. 1♂ point-mounted (NZAC04261719): “NEW ZEALAND: ND, Kohuronaki 270 m, 24-XI-1982, litter and decayed wood 82/122, G. Kuschel”. 1♂ point-mounted (NZAC04261721): “NEW ZEALAND: ND, Mangamuka Summit, 13-XII-1976, litter 76/102, V. A. May”. 2♀♀ point-mounted (NZAC04291732; NZAC04291733): “NEW ZEALAND: ND, Puketi Forest, Manginagina Scen. Reserve (nr. HQ, Nature Walk), 18-III-2011, 35°12'40.80"S 173°47'36.20"E 320 m, sifting litter in mixed deciduous forest with Kauri, A. Schomann, J. Pedersen”. 2♂♂ DNA extracted and slide-mounted (NZAC, code absent) and 1♀ point-mounted (NZAC04291731): “NEW ZEALAND: ND, Warawara St Forest, nr Hauturu Stm Campsite, 7-XII-2008, 35.22S 173.17E, sifting rotten wood and leaflitter, RL1372, R. Leschen, T. Buckley & D. Seldon”.

Figure 5. 

Gastrobothrus angustus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus. G Mesotibia and tarsus. H Metatibia and tarsus; trichome [red arrow]. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word angustus (“narrow”, masculine) that refers to the relatively narrower pronotum than that of other species.

Diagnosis.

Pronotum with transverse sulcus (Fig. 5C); acute spatula on abdominal sternite VI in male (Fig. 5D); apical trichome on metatibia in male (Fig. 5H).

Description.

Color of body brown, elytra yellowish-brown. MALE: Measurements: Length 2.31 mm, width 0.89 mm (Fig. 5A). Head: Length 0.50 mm and width 0.51 mm (Fig. 5B); vertex sparsely covered with long setae, vertexal foveae moderate and setose; eye moderately developed, shorter than temple; ocular-mandibular carina present; gular carinae gradually broadened toward apex and not swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket weakly developed; antenna unmodified overall, 11-segmented and antennal club consisting of three antennomeres; scape and pedicel cylindrical; 3–7 longer than wide and broadened apically; 8 shortest; 9–10 broadened apically; 11 longer than 9+10. Prothorax: Length 0.48 mm and width 0.54 mm (Fig. 5C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae less setose, transverse sulcus across foveae. Elytra: Length 0.59 mm and width 0.87 mm (Fig. 5C), and broadened markedly, widest 2.5 times as wide as base; basal fovea absent. Hindwing absent. Legs: Long and slender. Protibia and mesotibia with minute apical spine (Fig. 5F, G). Metatibia with apical trichome (Fig. 5H). Abdomen: Length 0.74 mm and width 0.89 mm. Sternite VI bearing acute triangular median spatula (Fig. 5D), as wide as median sclerite of sternite IX. Aedeagus: 0.46 mm long (Fig. 5E); parameres narrowed apically, with median spine and three setae; penis trullate. — FEMALE: Tibial spines and abdominal spatula absent.

Distribution.

North Island: BP, CL, ND (Fig. 2).

Habitat.

Leaf litter and rotten wood from mixed deciduous forest with kauri.

Remarks.

Adults of this species are close to G. inflatus (Sharp, 1874) comb. nov. in having moderate body size and leg but can be distinguished by an acute large abdominal spatula (Figs 5D, 11E).

Gastrobothrus biligo Choi, Leschen & Park, sp. nov.

Figures 2, 4, 6

Type material.

N = 2, 2♂♂. Holotype. ♂ point-mounted: “NEW ZEALAND, ND, Mt Horokaka 518 m, Tangihua Ra., 16 Aug 1977, J. S. Dugdale // litter 77/97 // N.Z. Arthropod Collection, NZAC, Private Bag 92170, AUCKLAND, New Zealand // NZAC04261728”. Paratype. 1♂ DNA extracted and slide-mounted (NZAC, code absent): “NEW ZEALAND: ND, Radar Bush, Te Paki Rec Res., 8-XII-2015, 34°28'26.9"S 172°45'47.1"E, sifting moss, RL1852, R. Leschen, T. Buckley, D. Seldon, J. Tizzard & L. Walker”.

Figure 6. 

Gastrobothrus biligo Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus. G Mesotibia and tarsus. H Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is a combination of the Latin words bi- (“having two parts”) and ligo (“hoe”, masculine) that refers to the paired spatulae on abdominal sternite V.

Diagnosis.

Pronotum with transverse sulcus (Fig. 6C); paired abdominal spatulae and depression on sternites IV–VIII (Fig. 6D); enlarged femur.

Description.

Color of body reddish-brown, elytra yellowish-brown. MALE: Measurements: Length 3.01 mm, width 1.04 mm (Fig. 6A). Head: Length 0.47 mm and width 0.52 mm (Fig. 6B); vertex sparsely covered with long setae, vertexal foveae enlarged and setose; eye moderately developed, situated anterior to level of vertexal foveae; ocular-mandibular carinae present; gular carinae gradually broadened toward apex and not swollen ventrally (Fig. 4). Frontal fovea absent; frontal rostrum lacking and depressed medially; postantennal cavity shallow. Antennal socket moderately developed; antenna unmodified overall, 11-segmented and antennal club consisting of three antennomeres; scape and pedicel cylindrical; 3–7 longer than wide and broadened apically; 8 shortest; 9–10 broadened apically; 11 longer than 9+10. Prothorax: Length 0.64 mm and width 0.67 mm (Fig. 6C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae less setose, transverse sulcus across foveae. Elytra: Length 0.87 mm and width 1.05 mm (Fig. 6C), and broadened markedly, widest 2.4 times as wide as base; basal fovea absent. Hindwing reduced. Legs: Robust. Protibia and mesotibia with prominent spine on apical (Fig. 6F, G). Mesotrochanter acutely extended ventrally. Metatibia simple (Fig. 6H). Abdomen: Length 1.03 mm and width 1.04 mm (Fig. 6D). Sternite V bearing two triangular basolateral spatulae curved medially. Aedeagus: 0.59 mm long (Fig. 6E); parameres enlarged apically, with short and long setae; penis ovoid; endophallus composed of robust armature and short armatures at base. — FEMALE: Unknown.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter and moss.

Remarks.

Adults of this species are similar to those of G. giganteus sp. nov. in having robust leg and short parameres and enlarged penis of aedeagus (Figs 6E, 9E) but can be distinguished by protibia without median spine (Figs 6F, 9F) and paired spatulae on the abdominal sternite V (Figs 6D, 9D).

Gastrobothrus buckleyi Théry & Leschen, 2013

Figures 4, 7

Material examined.

N = 3, 2♂♂ 1♀. Paratype. 1♂ card-mounted (NZAC04261509): “NEW ZEALAND: TH, Three Kings Is., Tasman V, 2-Dec-1983, litter 83/117, J. C. Watt”. Nontypes. 1♂ 1♀ point-mounted (NZAC04261640; NZAC04261641): NEW ZEALAND: TH, Three Kings Is., Tasman V, 2-XII-1983, litter 83/117, J. C. Watt.

Figure 7. 

Gastrobothrus buckleyi Théry & Leschen, 2013. A Habitus, dorsal view. B Head, dorsal view. C Head, lateral view. D Thorax. E Abdominal sternites. F Aedeagus. G Protibia and tarsus. H Mesotibia and tarsus; apical tubercle [red arrow]. I Metatibia and tarsus; trichome [red arrow]. Scale bars: 1 mm (A), 0.1 mm (B–I).

Diagnosis.

Frontal fovea present (Fig. 7B); gular carinae rounded at base, sharply broadened toward apex (Fig. 4) and acutely swollen ventrally (Fig. 7C). mesotibia bearing apical tubercle (Fig. 7G) and metatibia with apical trichome in male (Fig. 7H); abdominal sternite IX partially concealed by transparent lobe on sternite VIII (Fig. 7E); aedeagal endophallus robust and trifid at apex (Fig. 7F).

Description.

See Théry and Leschen (2013).

Distribution.

North Island: TH (not on mainland).

Habitat.

Leaf litter and rotten wood.

Gastrobothrus directus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 8

Type material.

N = 1, 1♂. Holotype. ♂ point-mounted: “NEW ZEALAND: ND, Cape Reinga, Toputupotu Bay, 4-V-1999, 34'26S 172'42E, Berlesate 406, R. Leschen, G. Hall, R. Hoare // NZAC04261631”.

Figure 8. 

Gastrobothrus directus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Pronotum. D Elytra. E Abdominal sternites. F Aedeagus. G Protibia and tarsus. H Mesotibia. I Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–I).

Etymology.

The specific epithet is the Latin word directus (“laid straight”, masculine) that refers to the endophallic armature of this species.

Diagnosis.

Protibia with minute apical spine (Fig. 8G). abdominal spatula absent; aedeagus with straight endophallic armature (Fig. 8F).

Description.

Color of body light brown, elytra yellowish-brown. MALE: Measurements: Length 2.33 mm, width 0.82 mm (Fig. 8A). Head: Length 0.44 mm and width 0.45 mm (Fig. 8B); vertex covered with long setae, vertexal foveae markedly setose; eye well-developed, shorter than temple; ocular-mandibular carina absent; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket well-developed; antenna unmodified overall, 11-segmented and antennal club consisting of three antennomeres; scape cylindrical and longer than wide; pedicel cylindrical; 3–4 broadened apically; 5 longer than wide; 6–7 identical; 8 shortest; 11 as long as 9+10. Prothorax: Length 0.49 mm and width 0.52 mm (Fig. 8C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose. Elytra: Length 0.55 mm and width 0.79 mm (Fig. 8D), and broadened posteriorly, widest 2.1 times as wide as base; basal fovea absent. Legs: Long and straight. Pro- and mesotibiae with minute apical spine (Fig. 8G, H). Metatibia plain (Fig. 8I). Abdomen: Length 0.85 mm and width 0.74 mm (Fig. 8E), lacking median spatula. Aedeagus: 0.6 mm long (Fig. 8F); parameres broadened apically and notched on apex, bearing apical seta; penis ovoid; endophallic armature straight. — FEMALE: Unknown.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter.

Remarks.

Adult of this species is very close to those of G. planiventris sp. nov. but can be distinguished by the endophallus with straight armature (Figs 8F, 15E).

Gastrobothrus giganteus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 9

Type material.

N = 6, 6♂♂. Holotype. ♂ point-mounted: “New Zealand: ND, Waima Forest, Trk. nr. Taita Rd, 19/Feb/2011, 35 31'51.40S 173 26'31.70E (±260 m) 220 m, sifting litter in mixed deciduous forest, A. Schomann & J. Pedersen // ZMUC collection // NZAC04291734”. Paratypes. 1♂ DNA extracted and slide-mounted (NZAC, code absent): “NEW ZEALAND: ND, Puketi Forest, Manginangina Scen. Reserve (nr. HQ, Nature Walk), 18-III-2011, 35°12'40.80"S 173°47'36.20"E 320 m, sifting litter in mixed deciduous forest, A. Schomann & J. Pedersen // ZMUC collection”. 1♂ point-mounted (NZAC04261715): “NEW ZEALAND: ND, Waimatenui, 10-III-1962, litter, R. A. Cumber”. 2♂♂ point-mounted (AMNZ35571; AMNZ35572): “NEW ZEALAND: ND, Waipoua Forest 100 m, 13-XI-2000, ex bush litter on roadside, NZMS 260 O06 622165, S. E. Thorpe”. 1♂ point-mounted (NZAC04261716): “NEW ZEALAND: ND, Waipoua SF, Wairau Smt 390 m, 14-IV-1980, sifting litter 80/51, J. C. Watt”.

Figure 9. 

Gastrobothrus giganteus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus; median spine [red arrow]. G Mesotibia and tarsus. H Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word giganteus (“gigantic”, masculine) that refers to the exceptionally large body size of this species.

Diagnosis.

Large body size; setose frontal fovea on head (Fig. 9B); pronotum with transverse sulcus (Fig. 9C); triangular spatula on abdominal sternite VI (Fig. 9D); acute spine on middle of protibia (Fig. 9F).

Description.

Color of body reddish-brown, head and pronotum darker. MALE: Measurements: Length 3.61 mm, width 1.44 mm (Fig. 9A). Head: Length 0.64 mm and width 0.64 mm (Fig. 9B); vertex sparsely covered with long setae, vertexal foveae enlarged and less setose; eye moderately developed, situated anterior to level of vertexal foveae; ocular-mandibular carinae not complete; gular carinae gradually broadened toward apex and not swollen ventrally (Fig. 4). Frontal fovea distinct, frontal rostrum lacking and depressed medially; postantennal cavity shallow. Antennal socket moderately developed; antenna long and unmodified overall, 11-segmented and antennal club consisting of three antennomeres; scape and pedicel cylindrical; 3–7 elongated; 8 shortest; 9–10 broadened apically; 11 as long as 9+10. Prothorax: Length 0.80 mm and width 0.87 mm (Fig. 9C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose, transverse sulcus across foveae. Elytra: Length 1.01 mm and width 1.44 mm (Fig. 9C), and broadened markedly, widest 2.8 times as wide as base; basal fovea absent. Hindwing reduced. Legs: Robust and long. Protibia with acute median spine (Fig. 9F). Mesotrochanter acutely extended at ventral (Fig. 9D). Metatibia simple (Fig. 9H). Abdomen: Length 1.16 mm and width 1.43 mm (Fig. 9D). Sternite VI bearing large triangular median spatula, as wide as median sclerite of sternite IX. Aedeagus: 0.97 mm long (Fig. 9E); parameres broadened and bifid apically, with two short setae; penis ovoid; endophallus composed of irregular armatures. — FEMALE: Unknown.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter from mixed deciduous forest.

Remarks.

Adults of this species are similar to those of G. pilosus sp. nov. in having dark brown pronotum with a prominent sulcus (Figs 9C, 14C) but can be easily distinguished by the large body size (Fig. 9A), distinct frontal fovea and weakly setose vertexal foveae (Fig. 9B), and protibia with an acute spine at middle level (Fig. 9F).

Gastrobothrus grossus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 10

Type material.

N = 1, 1♂. Holotype. ♂ point-mounted: “NEW ZEALAND: ND, Tangihua Range, N. Auckland, 1800–2000 m, 27-X-1936, E. Fairburn // A. E. Brookes Collection // NZAC04261625”.

Figure 10. 

Gastrobothrus grossus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Pronotum. D Elytra. E Abdominal sternites. F Aedeagus. G Protibia and tarsus. H Mesotibia and tarsus. I Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–I).

Etymology.

The specific epithet is the Latin word grossus (“stout”, masculine) that refers to thick femora and tibiae of this species.

Diagnosis.

Femur and tibia stout, protibia with triangular apical spine (Fig. 10G), mesotibia with acute apical spine (Fig. 10H); abdominal sternites lacking spatula; aedeagal parameres curved medially (Fig. 10F).

Description.

Color of body dark brown, elytra brighter. MALE: Measurements: Length 2.45 mm, width 0.92 mm (Fig. 10A). Head: Length 0.44 mm, width 0.48 mm (Fig. 10B); vertex sparsely covered with long setae, vertexal foveae setose; eye well-developed, shorter than temple; ocular-mandibular carina absent; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent; frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket gently protruded; antenna long and unmodified overall, 11-segmented and club consisting of three antennomeres; scape cylindrical and longer than wide; pedicel as long as antennomere 3; 3 broadened apically and as long as 4; 5 elongated; 6–7 identical; 8 shortest; 9 slightly swollen at middle level; 11 almost as long as 9+10. Prothorax: Length 0.49 mm and width 0.57 mm (Fig. 10C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose, transverse sulcus absent. Elytra: Length 0.63 mm and width 0.92 mm (Fig. 10D), and broadened markedly, widest 2.6 times as wide as base; basal fovea absent. Legs: Robust and long. Pro-, meso-, and metatibiae with acute spine apically (Fig. 10G–I). Abdomen: Length 0.89 mm and width 0.90 mm (Fig. 10E). Sternites IV–VIII depressed medially, abdominal spatula absent. Aedeagus: 0.6 mm long (Fig. 10F); parameres stubby and curved medially, apex enlarged, inward and outward setae on apex; penis ovoid and well-sclerotized basally. — FEMALE: Unknown.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Unknown.

Remarks.

Adult of this species is similar to those of G. lunatus sp. nov. in having robust femur and tibia (Figs 10A, 12A) but can be distinguished by the absence of median spine on the protibia and abdominal spatula.

Gastrobothrus ignotus (Broun, 1881)

Figure 2

Bryaxis ignotus Broun, 1881; Eupines (Byraxis) ignotus [combination and subgeneric assignment by Nomura and Leschen (2006)]; Gastrobothrus ignotus [combination by Shen and Leschen (2020)]

Diagnosis.

Dorsal side of body smooth, rarely punctate and nearly asetose; metatibiae bearing apical trichome.

Redescription.

Body overall shiny, hardly covered with setae. Color of body dark and reddish-brown color, tarsus brighter. Head: Wider than long, bearing asetose frontal fovea and vertexal foveae; eye shorter than temple. Antenna unmodified overall, 11-segmented and antennal club consisting of three antennomeres; non-club antennomere short. Pronotum: Wider than long, median antebasal fovea punctiform and asetose, lateral antebasal foveae setose. Elytra: Wider than long, hardly punctate and nearly asetose. Legs: Slender and long. Metatibia with apical trichome. Abdomen: As wide as elytra. Tergite III largest; paratergites III–IV thick.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Unknown.

Remarks.

The collection locality of the holotype representing Bryaxis ignotus Broun, 1881 is Manaia, Whangarei Harbour, which overlaps that of paratypes representing G. retusus sp. nov. (Mair Park, Whangarei). Nonetheless, this species may be distinguished from G. retusus sp. nov. by the absence of protibial spine on middle. The holotype of this species was not examined for this paper. The brief description is based on the image provided in Shen and Leschen (2019).

Gastrobothrus inflatus (Sharp, 1874), comb. nov.

Figures 2, 4, 11

Bryaxis inflata Sharp, 1874; Physa inflata [combination by Raffray (1890)]; Physobryaxis inflata [combination by Hetschko (1913)]; Gastrobothrus inflata [misplacement by Hudson (1923, 1934)]; Physobryaxis inflatus [gender change in specific epithet by Shen and Leschen (2019)]

Material examined.

N = 18, 13♂♂ 5♀♀. 1♂ 1♀ point-mounted (NZAC04261732; NZAC04261733): NEW ZEALAND: AK, Bethells, Matuku Res., 25-VIII-1992, moss 92/47, G. Hall. 1♀ point-mounted (AMNZ66250): NEW ZEALAND: AK, Mataitai Forest S. A., III-2002, pitfall D13, A. Warren. 1♂ DNA extracted and slide-mounted (NZAC, code absent): NEW ZEALAND: AK, Waitakere Ranges Reg. Park, Kauri Grove Trk., 2-II-2011, 36.964526°S, 174.512425°E (±250m), 300 m, deciduous forest with Kauri, sifted litter & handpick, Scho002, A. Schomann and J. Pedersen. 1♂ point-mounted (NZAC04261723): NEW ZEALAND: BP, Lottin Point Road, Waenga Bush, 27-I-1993, litter, J. I. Townsend. 1♂ point-mounted (NZAC04261722): NEW ZEALAND: BP, Papatea, 25-I-1993, litter 93/1, J. S. Dugdale. 1♂ point-mounted (NZAC04261724): NEW ZEALAND: BP, Te Koau, Hovells, Watching Dog Track 360 m, 31-I-1993, litter 93/11, J. S. Dugdale. 2♂♂ point-mounted (AMNZ26786; AMNZ26788): NEW ZEALAND: CL, Cuvier I., Old Radar Station, 200 m, 10-XI-1999–18-XI-1999, grassy clearing, yellow pan trap L8177, J. W. Early, S. E. Thorpe. 1♀ point-mounted (AMNZ29721): NEW ZEALAND: CL, Cuvier I., Pumphouse Stream, 120 m, 4-IV-2000, wet rotting nikau fronds on ground L8232, R. F. Gilbert. 1♂ point-mounted (AMNZ31070): NEW ZEALAND: CL, Cuvier I., Radar Clearing, 200 m, 29-III-2000–5-IV-2000, forest edge, yellow pan trap L8241, J. W. Early, R. F. Gilbert. 1♂ point-mounted (NZAC04261738): NEW ZEALAND: CL, Great Barrier I, Mt Hobson 620 m, 23-III-1978–28-III-1978, litter 78/134, ESNZ Field trip. 1♂ point-mounted (NZAC04261729): NEW ZEALAND: RI, Ohakune, 20-III-1969, R. A. Cumber. 1♂ 1♀ point-mounted (NZAC04261730; NZAC04291729): NEW ZEALAND: WO, Kawhia, 19-III-1959, R. A. Cumber. 1♂ slide-mounted (NZAC04261731): NEW ZEALAND: WO, Mahoenui, Gribbons Road, 26-VI-1977, sifted litter 77/78, N. H. Mancer. 1♂ 1♀ point-mounted (NZAC04261743; NZAC04261744): NEW ZEALAND: WO, Waitomo, 24-V-1983, decayed wood and litter 83/56, G. Kuschel.

Figure 11. 

Gastrobothrus inflatus (Sharp, 1874), comb. nov. A Habitus, dorsal view. B Head, dorsal view. C Head, lateral view. D Thorax. E Abdominal sternites; spatula [red arrow]. F Aedeagus. G Protibia and tarsus. H Mesotibia and tarsus. I Metatibia and tarsus; trichome [red arrow]. Scale bars: 1 mm (A), 0.1 mm (B–H).

Diagnosis.

Pronotum with transverse sulcus (Fig. 11C); pro- and mesotibia with minute apical spine in male (Fig. 11G, H); metatibia with apical trichome in male (Fig. 11I); abdominal sternite VI with fine median spatula in male (Fig. 11E).

Redescription.

Color of body reddish-brown, elytra and appendages brighter. MALE: Measurements: Length 2.61–2.79 mm, width 0.96–1.17 mm (Fig. 11A). Head: Length 0.55–0.61 mm, width 0.52–0.56 mm (Fig. 11B, C); vertex sparsely covered with long setae, vertexal foveae setose; eye shorter than temple; ocular-mandibular carinae present; gular carinae gradually broadened toward apex and gently swollen ventrally (Figs 4, 11C). Frontal fovea faint, frontal rostrum lacking and depressed medially; postantennal cavity absent. Antennal socket moderately developed; antenna unmodified overall, 11-segmented and antennal club consisting of three antennomeres; scape cylindrical; pedicel parallel; antennomeres 3–5 elongate; 6–7 subequal in length; 8 shortest, slightly longer than wide; 9–10 nearly symmetrical; 11 largest. Prothorax: Length 0.53–0.56 mm and width 0.60–0.63 mm (Fig. 11D). Pronotal median antebasal fovea small and asetose, lateral antebasal foveae setose, transverse antebasal sulcus prominent. Elytra: Length 0.68–0.72 mm and width 0.94–1.11 mm (Fig. 11D), and broadened markedly, widest 2.4–2.6 times as wide as base; basal fovea absent. Legs: Slender and long. Protibia straight with tiny apical spine (Fig. 11G). Mesotibia straight with tiny apical spine (Fig. 11H). Metatibia straight with apical trichome (Fig. 11I). Abdomen: Length 0.80–0.92 mm and width 0.96–1.17 mm (Fig. 11E). Tergites IV–VII narrowed in order, bearing basolateral foveae; IV with mediobasal foveae. Sternites III–IV with mediobasal foveae; IV with basolateral foveae; V–VIII lacking fovea; VI with mediobasal spatula small and triangular; median sclerite of IX somewhat triangular, wider than abdominal spatula. Aedeagus: 0.76 mm (Fig. 11F); parameres small, blunt apex with two apical setae and single subapical seta on each paramere; penis guttiform; endophallus not sclerotized. — FEMALE: Eye less developed; fore leg and mid leg unmodified.

Distribution.

North Island: AK, BP, CL, RI, WO (Fig. 2).

Habitat.

Leaf litter and moss from deciduous forest with kauri. Rotten fronds of nikau on ground.

Remarks.

The collection locality of the holotype representing Bryaxis inflata Sharp, 1874 is Auckland. Although several specimens examined this study were collected from BP and CL where two paratypes of G. angustus sp. nov. were originated, they are distinguished from each other by size of abdominal spatula (Figs 5D, 11D).

Gastrobothrus lunatus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 12

Type material.

N = 5, 5♂♂. Holotype. ♂ point-mounted: “NEW ZEALAND: ND, Waimatenui, 10 Mar 1962, R. A. Cumber // litter // R. A. Cumber Collection // N.Z. Arthropod Collection, NZAC, Private Bag 92170, AUCKLAND, New Zealand // NZAC04261608”. Paratypes. 2♂♂ point-mounted (NZAC04261597; NZAC04261607): “NEW ZEALAND, ND, Waimatenui, 10-III-1962, litter, R. A. Cumber”. 2♂♂ card-mounted (NZAC04261601; NZAC04261602): “NEW ZEALAND: ND, Waipoua Forest, 24-XI-1994, leaf litter, J. T. Nunn”.

Figure 12. 

Gastrobothrus lunatus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus; median spine [red arrow]. G Mesotibia and tarsus. H Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word lunatus (“crescent”, masculine) that refers to the half-moon-shaped abdominal spatula of this species.

Diagnosis.

Acute spine at middle level of protibia (Fig. 12F); transversely elliptic spatula on abdominal sternite V (Fig. 12D).

Description.

Color of body brown. MALE: Measurements: Length 2.76 mm, width 1.01 mm (Fig. 12A). Head: Length 0.43 mm and width 0.50 mm (Fig. 12B); vertex moderately covered with long and short setae, vertexal foveae asetose; eye moderately developed, shorter than temple; ocular-mandibular carinae absent; gular carinae gradually broadened toward apex and not swollen ventrally (Fig. 4). Frontal fovea faint, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket gently protruded; antenna long and unmodified overall, 11-segmented and antennal club consisting of three antennomeres; scape cylindrical and longer than wide; pedicel as long as antennomere 3; 3 broadened apically and as long as 4; 5 elongated; 6–7 identical; 8 shortest; 11 almost as long as 9+10. Prothorax: Length 0.55 mm and width 0.60 mm (Fig. 12C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose, transverse sulcus absent. Elytra: Length 0.69 mm and width 0.98 mm (Fig. 12C), and broadened markedly, widest 2.7 times as wide as base; basal fovea absent. Legs: Robust and long. Protibia with moderately acute spine on middle (Fig. 12F). Mesotibia with minute apical spine (Fig. 12G). Metatibia plain (Fig. 12H). Abdomen: Length 1.09 mm and width 1.01 mm (Fig. 12D). Sternite V bearing median spatula, as wide as gap between mediobasal foveae on sternite IV (Fig. 12D). Aedeagus: 0.84 mm long (Fig. 12E); parameres slender and curved internally, apex enlarged, inward and outward setae on subapical; penis elongated ovoid and well-sclerotized basally; endophallus composed of fine armatures pointed anteriorly. — FEMALE: Unknown.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter.

Remarks.

Adults of this species are similar to those of G. quadrus sp. nov. in having robust femur and a prominent spine on the middle of protibia (Figs 12F, 17F) but can be distinguished by a wide elliptic spatula on the abdominal sternite V (Fig. 12D).

Gastrobothrus occultus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 13

Type material.

N = 4, 4♂♂. Holotype. ♂ point-mounted: “NEW ZEALAND: AK, Lynfield, 4-XII-1974, G. Kuschel // NZAC04261577”. Paratypes. 2♂♂ point-mounted (NZAC04261580; NZAC04261622): “NEW ZEALAND: ND, Kauri Sanct., Omahuta SF, 16-VII-1974, litter 74/40, G. Kuschel”. 1♂ point-mounted (NZAC04261638): “NEW ZEALAND: ND, Waipoua, Waikohatu Stm, 30-X-1985, sifted litter 85/49, R. C. Craw”.

Figure 13. 

Gastrobothrus occultus Choi, Leschen & Park, sp. nov. A Habitus, dorsal habitus. B Head. C Pronotum. D Elytra. E Abdominal sternites. F Aedeagus. G Protibia and tarsus. H Mesotibia. I Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–I).

Etymology.

The specific epithet is the Latin word occultus (“concealed”, masculine) that refers to completely reduced median antebasal fovea on the pronotum of this species.

Diagnosis.

Median antebasal fovea on pronotum barely visible (Fig. 13C); abdominal spatula absent (Fig. 13E).

Description.

Color of reddish-brown, elytra and appendages brighter. MALE: Measurements: Length 2.02 mm, width 0.75 mm (Fig. 13A). Head: Length 0.39 mm and width 0.41 mm (Fig. 13B); vertex sparsely covered with long setae, vertexal foveae setose; eye well-developed, as long as temple; ocular-mandibular carina absent; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket well-developed; antennae unmodified overall, 11-segmented and club consisting of three antennomeres; scape cylindrical and longer than wide; pedicel cylindrical; 3–4 broadened apically; 5 longer than wide; 6–7 identical; 8 shortest; 11 as long as 9+10. Prothorax: Length 0.39 mm and width 0.45 mm (Fig. 13C). Median antebasal fovea on pronotum completely reduced, lateral antebasal foveae setose. Elytra: Length 0.51 mm and width 0.75 mm (Fig. 13D), and markedly broadened posteriorly, widest 2.6 times as wide as base; basal fovea absent. Legs: Long and stout. Protibia with blunt spine apically (Fig. 13G). Mesotibia with minute spine apically (Fig. 13H). Metatibia plain (Fig. 13I). Abdomen: Length 0.73 mm and width 0.74 mm (Fig. 13E), lacking median spatula. Aedeagus: 0.54 mm long (Fig. 13F); parameres skinny and curved outward at apex, bearing apical short seta and subapical long seta; penis fusiform; endophallus composed of two straight armatures. — FEMALE: Unknown.

Distribution.

North Island: AK, ND (Fig. 2).

Habitat.

Leaf litter.

Remarks.

Adults of this species is similar to those of G. planiventris sp. nov. but may be distinguished by smaller lateral antebasal foveae and barely visible median antebasal fovea on the pronotum (Fig. 13C) and the short aedeagal parameres (Fig. 13F).

Gastrobothrus pilosus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 14

Type material.

N = 3, 1♂ 2♀♀. Holotype. ♂ DNA extracted and slide-mounted (NZAC, code absent): “New Zealand: ND, Waipoua Forest, Yakas Trk. (ca 1km of Tane Mahuta), 19/Mar/2011, 35 36'27.80S 173 32'6.30E (±450 m), sifting litter, A. Schomann & J. Pedersen // ZMUC collection”. Paratypes. 1♀ point-mounted (NZAC04261671): “NEW ZEALAND: ND, Omahuta SF, 23-IV-1975, liverworts and mosses from tree bases and ground 75/142, J. S. Dugdale”. 1♀ point-mounted (NZAC04291735): “NEW ZEALAND: ND, Waipoua Forest, Yakas Trk. (ca 1km of Tane Mahuta), 19-III-2011, 35°36'27.80"S 173°32'6.30"E, sifting litter, A. Schomann & J. Pedersen // ZMUC collection”.

Figure 14. 

Gastrobothrus pilosus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus. G Mesotibia and tarsus. H Metatibia and tarsus; trichome [red arrow]. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word pilosus (“hairy”, masculine) that refers to the head coarsely covered with long yellow setae.

Diagnosis.

Shallow punctiform depression on frons (Fig. 14B); moderate trichome on apical of metatibia (Fig. 14H); semicircular spatula on abdominal sternite VI in male (Fig. 14D).

Description.

Color of body brown, head and pronotum darker. MALE: Measurements: Length 2.62 mm, width 1.10 mm (Fig. 14A). Head: Length 0.46 mm and width 0.56 mm (Fig. 14B); vertex moderately covered with long setae, vertexal foveae moderate and markedly setose; eye well-developed, shorter than temple; ocular-mandibular carina present; gular carinae gradually broadened toward apex and not swollen ventrally (Fig. 4). Frontal fovea shallow, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket moderately developed; antenna short and unmodified overall, 11-segmented and club consisting of three antennomeres; scape cylindrical and longer than wide; pedicel cylindrical; 3–4 broadened apically; 5 elongated; 6–7 broadened apically; 8–10 wider than long and projected internally; 8 shortest; 11 longer than 9+10. Prothorax: Length 0.62 mm and width 0.65 mm (Fig. 14C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose, transverse sulcus across foveae. Elytra: Length 0.72 mm and width 1.02 mm (Fig. 14C), and broadened markedly, widest 2.4 times as wide as base; basal fovea absent. Hindwing absent. Legs: Robust and long. Protibia and mesotibia with minute apical spine (Fig. 14F, G). Metatibia with moderate apical trichome (Fig. 14H). Abdomen: Length 0.82 mm and width 1.10 mm (Fig. 14D). Sternite VI bearing wide semicircular median spatula, wider than median sclerite of sternite IX. Aedeagus: 0.77 mm long (Fig. 14E); parameres slender and narrowed apically, with two short setae and one long seta; penis fusiform; endophallus composed of paired triangular armatures. — FEMALE: Tibial spines and abdominal spatula absent.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter. Liverwort and moss from tree bases and ground.

Remarks.

Adults of this species are similar to those of G. inflatus (Sharp, 1874) comb. nov. in having setose vertexal foveae (Figs 11B, 14B) but can be easily distinguished by a large semicircular spatula on the abdominal sternite VI (Fig. 14D).

Gastrobothrus planiventris Choi, Leschen & Park, sp. nov.

Figures 2, 4, 15

Type material.

N = 12, 12♂♂. Holotype. ♂ point-mounted: “NEW ZEALAND: WO, Waharoa, 4 Jan 1992, P. A. Maddison, Litter 92/1 // NZAC04261629”. Paratypes. 1♂ point-mounted (AMNZ27144): “NEW ZEALAND: AK, Auckland 20 m, Western Springs, 4-I-2000, litter under pine trees, NZMS 260 R11 642805, S. E. Thorpe”. 1♂ card-mounted (NZAC, code absent): “NEW ZEALAND: AK, Auckland, 26-VI-1932, Kauri gully, C. E. Clarke”. 1♂ point-mounted (NZAC04261575): “NEW ZEALAND: AK, Lynfield, 19-III-1977, G. Kuschel”. 1♂ card-mounted (NZAC, code absent): “NEW ZEALAND: AK, Papakura, 17-I-1942, A. Richard”. 1♂ card-mounted (NZAC04261627): “NEW ZEALAND: AK, Papakura, 19-X-1935, E. S. Gourlay”. 1♂ card-mounted (NZAC04261574): “NEW ZEALAND: BP, Tui Rd at gate, Mt Te Aroha, 19-XI-2005, moss from drainage ditch, J. T. Nunn”. 1♂ point-mounted (NZAC04261630): “NEW ZEALAND: ND, Waipoua SF, 100 m, Kauri Ricker TK, 13-IV-1980, sifted litter 80/49, J. C. Watt”. 2♂♂ DNA extracted and slide-mounted (NZAC, code absent): “NEW ZEALAND: ND, Warawara St Forest, nr Hauturu Stm Campsite, 7-XII-2008, 35.22S 173.17E, sifting rotten wood and leaf litter, RL1372, R. Leschen, T. Buckley, and D. Seldon”. 1♂ card-mounted (NZAC04261636): “NEW ZEALAND: TK, Patea Walk, Egmont MP, 3-I-1990, 1543, J. T. Nunn”. 1♂ point-mounted (NZAC04261581): “NEW ZEALAND: WO, Piripiri, Caves Reserve, 25-V-1986, leaf litter, CL 480, C. L. Lyal”.

Figure 15. 

Gastrobothrus planiventris Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites. E Aedeagus. F Protibia and tarsus. G Mesotibia and tarsus. H Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is a combination of the Latin words planus (“flat”, masculine) and ventris (“abdomen”, masculine) that refers to the unmodified abdomen.

Diagnosis.

Abdominal spatula absent (Fig. 15D); aedeagus with elongate curved endophallic armature (Fig. 15E).

Description.

Color of body brown, elytra yellowish-brown. MALE: Measurements: Length 2.53 mm, width 0.97 mm (Fig. 15A). Head: Length 0.45 mm and width 0.51 mm (Fig. 15B); vertex moderately covered with long setae, vertexal foveae markedly setose; eye moderately developed, shorter than temple; ocular-mandibular carina indistinct; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket well-developed; antenna unmodified overall, 11-segmented and club consisting of three antennomere; scape cylindrical and longer than wide; pedicel rectangular; 3–4 broadened apically; 5 longer than wide; 6–7 identical; 8 shortest; 11 as long as 9+10. Prothorax: Length 0.57 mm and width 0.61 mm (Fig. 15C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose. Elytra: Length 0.63 mm and width 0.91 mm (Fig. 15C), and broadened markedly, widest 2.4 times as wide as base; basal fovea absent. Hindwing absent. Legs: Long and slender. Protibia and mesotibia with minute apical spine (Fig. 15F, G). Metatibiae plain (Fig. 15H). Abdomen: Length 0.88 mm and width 0.97 mm (Fig. 15D), lacking median spatula. Aedeagus: 0.61 mm long (Fig. 15E); parameres broadened apically with apical seta; penis ovoid; endophallus composed of short armature and long armature curved medially. — FEMALE: Unknown.

Distribution.

North Island: AK, BP, ND, TK, WO (Fig. 2).

Habitat.

Leaf litter under pine tree. Moss from drainage ditch. Rotten wood.

Remarks.

Adults of this species are similar to those of G. sharpi (Broun, 1880) in having hairy body and thick paratergites III (Figs 15A, 19A) but can be distinguished by abdominal sternites without modification (Fig. 15D).

Gastrobothrus platynotus (Broun, 1893)

Figure 2

Bryaxis platynota Broun, 1893; Eupines platynota [combination by Hudson (1923)]; Eupines (Eupines) platynota [subgeneric assignment by Nomura and Leschen (2006)]; Gastrobothrus platynotus [combination by Shen and Leschen (2020)]

Diagnosis.

Rectangular head; slender and simple leg.

Redescription.

Color of head–pronotum moderately dark brown, elytra–abdomen and legs brighter and reddish-brown. Head: Wider than long, bearing asetose vertexal foveae; eye shorter than temple. Antennae composed of 11 antennomeres and club consisting of three antennomeres. Pronotum: Wider than long, sparsely covered with long setae, median antebasal fovea punctiform and asetose, lateral antebasal foveae setose. Elytra: Wider than long, sparsely punctate and hardly setose. Legs: Slender, without modifications. Abdomen: As wide as elytra, moderately covered with long setae. Tergite III as long as IV; paratergites III–IV thin.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Unknown.

Remarks.

The holotype of this species was not examined for this paper. The brief description is based on the image provided in Shen and Leschen (2019). Note that the distribution of Hawkes Bay (HB), was in error in Shen and Leschen (2019) and it should be corrected to Northland, the location of northern Wairoa.

Gastrobothrus pumilus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 16

Type material.

N = 7, 3♂♂ 4♀♀. Holotype. ♂ point-mounted: “New Zealand, ND, Waipoua Forest, Hill south of DoC HQ, Ngãpuhi, Ngãti Whãtua, Te Roroa, 5/Feb/2006, 35.65090 S 173.55724 E, sifting litter & dead wood, RL1102, T. Buckley and R. Hoare // NZAC04291724”. Paratypes. 1♀ point-mounted (AMNZ84815): “NEW ZEALAND: AK, Atuanui, Mt Auckland, I-2002, pitfall B25, A Warren”. 1♂ DNA extracted and slide-mounted (NZAC, code absent): “NEW ZEALAND: AK, Waitakere Ra., Kauri Grove TK, 20-X-2017, 36.965000°S, 174.503167°E, sifted leaf litter, JWS0021, Z. McGrath and J. Shen”. 1♀ point-mounted (NZAC04261598): “NEW ZEALAND: ND, Mt Horokaka 518 m, Tangihua Ra, 16-VIII-1977, litter 77/97, J. S. Dugdale”. 1♀ point-mounted (NZAC04261619): “NEW ZEALAND: ND, Te Paki, Coastal Reserve, Wirihi Swamp, 19-X-1982, litter 82/98, C. F. Butcher”. 1♀ point-mounted (NZAC04261618): “NEW ZEALAND: ND, Te Paki, Radar Bush, 12-I-2007–12-II-2007, Native bush., pitfall trap 5, O. Ball”. 1♂ point-mounted (NZAC04261617): “NEW ZEALAND: ND, Te Paki, Radar Bush, 13-IV-2007–14-V-2007, Native bush., pitfall trap 3, O. Ball”.

Figure 16. 

Gastrobothrus pumilus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus. G Mesotibia and tarsus. H Metatibia and tarsus; trichome [red arrow]. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word pumilus (“dwarf”, masculine) that refers to the small body size of this species.

Diagnosis.

Metatibia with apical trichome (Fig. 16H); abdominal sternite V with shallow triangular spatula (Fig. 16D).

Description.

Color of body reddish-brown. MALE: Measurements: Length 1.76 mm, width 0.69 mm (Fig. 16A). Head: Length 0.33 mm and width 0.36 mm (Fig. 16B); vertex sparsely covered with short setae, vertexal foveae setose; eye moderately developed, shorter than temple; ocular-mandibular carina not complete, ventrolateral margin extending beneath eye; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket barely developed; antenna short and unmodified overall, 11-segmented and club consisting of three antennomere; scape asymmetrical and as long as wide; pedicel ovoid; 3–5 broadened apically; 6 narrowed; 7 trapezoidal; 8–10 transverse; 8 shortest; 11 longer than 9+10. Prothorax: Length 0.39 mm and width 0.40 mm (Fig. 16C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose. Elytra: Length 0.52 mm and width 0.66 mm (Fig. 16C), and broadened markedly, widest 2.7 times as wide as base; basal fovea absent. Hindwing absent. Legs: Slender. Protibia with fine apical spine (Fig. 16F). Metatibia with long apical trichome (Fig. 16H). Abdomen: Length 0.52 mm and width 0.69 mm (Fig. 16D). Sternite V bearing wide triangular median spatula, wider than median sclerite of sternite IX. Aedeagus: 0.39 mm long (Fig. 16E); parameres slender and plain with two setae; penis ovoid; endophallus with two pairs of longitudinal armatures. — FEMALE: Tibial spines and abdominal spatula absent.

Distribution.

North Island: AK, ND (Fig. 2).

Habitat.

Leaf litter and rotten wood.

Remarks.

Adults of this species are similar to those of G. buckleyi Théry & Leschen, 2013 in having small body size but can be distinguished by absence of frontal fovea, setose vertexal foveae (Fig. 16B), setose lateral antebasal foveae on pronotum (Fig. 16C), and spatula on the abdominal sternite V (Fig. 16D).

Gastrobothrus quadrus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 17

Type material.

N = 8, 4♂♂ 4♀♀. Holotype. ♂ point-mounted: “New Zealand: ND, Waipoua Forest, Yakas Trk. (ca 1 km of Tane Mahuta), 19/Mar/2011, 35 36'27.80 S 173 32'6.30 E 340m, sifting litter, A. Schomann & J. Pedersen // ZMUC collection // NZAC04291725”. Paratypes. 1♂ point-mounted (AMNZ35069): “NEW ZEALAND: ND, Aranga, Trounson Kauri Park, 1-XI-2000, litter in forest remnant, NZMS 260 O07 695087, S. E. Thorpe”. 1♀ point-mounted (NZAC04261610): “NEW ZEALAND: ND, Waipoua Forest, Yakas Track, 6-II-1995, litter, bracket fungus & Rhopalostylis sapida bases, J. Klimaszewski”. 1♂ DNA extracted and slide-mounted (NZAC, code absent) and 1♀ point-mounted (NZAC04291726): “NEW ZEALAND: ND, Waipoua Forest, Yakas Trk. (ca 1 km of Tane Mahuta), 19-III-2011, 35°36'27.80"S 173°32'6.30E" 340m, sifting litter, A. Schomann & J. Pedersen // ZMUC collection”. 1♀ point-mounted (NZAC04261600): “NEW ZEALAND: ND, Waipoua SF, 290 m, Waikohatu Bridge, 14-IV-1980, litter 80/53, J. C. Watt”. 1♀ point-mounted (NZAC04261599): “NEW ZEALAND: ND, Waipoua SF, Toatoa Grove, 25-XI-1980, plants and litter 80/117, G. Kuschel”. 1♂ point-mounted (NZAC04261596): “NEW ZEALAND: ND, Waipoua, Waikohatu Stm, 30-X-1985, sifted litter 85/49, R. C. Craw”.

Figure 17. 

Gastrobothrus quadrus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus; median spine [red arrow]. G Mesotibia and tarsus. H Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word quadrus (“square”, masculine) that refers to the angulate head of this species.

Diagnosis.

Acute spine on middle of protibia (Fig. 17F); ligulate spatula on abdominal sternite V (Fig. 17D).

Description.

Color of body reddish-brown, head to pronotum darker. MALE: Measurements: Length 2.46 mm, width 0.95 mm (Fig. 17A). Head: Length 0.47 mm and width 0.50 mm (Fig. 17B); vertex sparsely covered with short and long setae, vertexal foveae asetose; eye moderately developed; ocular-mandibular carina indistinct, ventrolateral margin extending beneath eye; gular carinae gradually broadened toward apex and not swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket moderately developed; antenna unmodified overall, 11-segmented and club consisting of three antennomere; scape cylindrical, almost twice as long as pedicel; pedicel as long as antennomere 3; 3 broadened apically and slightly longer than 4; 4 shortened; 5 elongated; 6–7 identical and 1.6 times as long as 8; 8 shortest; 11 almost as long as 9+10. Prothorax: Length 0.52 mm and width 0.60 mm (Fig. 17C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose with short setae. Elytra: Length 0.71 mm and width 0.95 mm (Fig. 17C), and broadened markedly, widest 2.4 times as wide as base; basal fovea absent. Hindwing absent. Legs: Robust and long. Protibia with minute apical spine and acute spine on middle level (Fig. 17F). Mesotibia with apical spine (Fig. 17G). Abdomen: Length 0.76 mm and width 0.94 mm (Fig. 17D). Sternite V bearing narrow median spatula. Aedeagus: 0.58 mm long (Fig. 17E); parameres slender and curved internally, apex stubby with subapical spine, setae on apical and subapical; penis ovoid and well-sclerotized basally, with paired armatures; endophallus with three asymmetrical armatures curved outward. — FEMALE: Thoracic characters overall skinny; legs without modifications; abdominal spatula absent.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter in forest remnant. Bracket fungus in nikau.

Remarks.

Adults of this species are similar to that of G. platynotus (Broun, 1893) in having asetose vertexal foveae (Fig. 17B) but can be distinguished by the angular temple (Fig. 17B).

Gastrobothrus retusus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 18

Type material.

N = 10, 4♂♂ 6♀♀. Holotype. ♂ point-mounted: “New Zealand: ND, Puketi Forest, Manginagina Scen. Reserve (nr. HQ, Nature Walk), 18/Mar/2011, 35 12'40.80S 173 47'36.20E (±250 m) 320 m, mixed deciduous forest with Kauri, litter, A. Schomann & J. Pedersen // ZMUC collection // NZAC04291727”. Paratypes. 1♂ 3♀♀ card-mounted (NZAC04261603; NZAC04261604; NZAC04261605; NZAC04261606): “NEW ZEALAND: ND, Mair Park, Whangarei, 7-V-1944, leaf mould, E. Fairburn // A. E. Brookes Collection”. 1♂ point-mounted (NZAC04261609): “NEW ZEALAND: ND, Omahuta Forest, Kauri Sanctuary, 31-I-1995, sifted litter Agathis australis, J. Klimaszewski”. 1♀ point-mounted (NZAC04261588): “NEW ZEALAND: ND, Omahutu SF, 18-III-1978, litter, S. B. Peck”. 1♂ DNA extracted and slide-mounted (NZAC, code absent) 1♀ point-mounted (NZAC04291728): “NEW ZEALAND: ND, Puketi Forest, Manginagina Scen. Reserve (nr. HQ, Nature Walk), 18-III-2011, 35°12'40.80"S 173°47'36.20"E (±250 m) 320 m, mixed deciduous forest with Kauri, litter, A. Schomann & J. Pedersen // ZMUC collection”. 1♀ point-mounted (NZAC04261612): “NEW ZEALAND: ND, Te Paki, Radar Bush, 13-IV-2007–14-V-2007, Native bush., pitfall trap 3, O. Ball”.

Figure 18. 

Gastrobothrus retusus Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites; spatula [red arrow]. E Aedeagus. F Protibia and tarsus; median spine [red arrow]. G Mesotibia and tarsus. H Metatibia. Scale bars: 1 mm (A), 0.1 mm (B–H).

Etymology.

The specific epithet is the Latin word retusus (“blunt”, masculine) that refers to the shape of tibial spine and spatula.

Diagnosis.

Blunt spine on middle of protibia (Fig. 18F); short ligulate spatula on abdominal sternite V (Fig. 18D).

Description.

Color of body reddish-brown, head to pronotum slightly darker. MALE: Measurements: Length 2.06 mm, width 0.86 mm (Fig. 18A). Head: Length 0.39 mm and width 0.46 mm (Fig. 18B); vertex sparsely covered with short and long setae, vertexal foveae moderate and asetose; eye moderately developed, as long as temple; ocular-mandibular carina not complete; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity faint. Antennal socket gently protruded; antenna unmodified overall, 11-segmented and club consisting of three antennomere; scape cylindrical and elongated; pedicel slightly longer than antennomere 3; 3 broadened apically and slightly longer than 4; 4 cylindrical; 5 elongated; 7 slightly longer than 6; 8 shortest; 11 almost as long as 9+10. Prothorax: Length 0.44 mm and width 0.52 mm (Fig. 18C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose with short setae. Elytra: Length 0.59 mm and width 0.85 mm (Fig. 18C), and broadened markedly, widest 2.6 times as wide as base; basal fovea absent. Hindwing absent. Legs Slender and long. Protibia with blunt spine on middle level and fine apical spine (Fig. 18F). Mesotibia with apical spine (Fig. 18G). Abdomen: Length 0.64 mm and width 0.86 mm (Fig. 18D). Sternite V bearing short and narrow median spatula. Aedeagus: 0.61 mm long (Fig. 18E); parameres short slender and apex weakly stubby, seta on subapical; penis ovoid and well-sclerotized basally; endophallus consisting of fine irregular armatures. — FEMALE: Legs without modifications and abdominal spatula absent.

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter from mixed deciduous forest with kauri.

Remarks.

Adults of this species are close to G. quadrus sp. nov. in having asetose vertexal foveae (Figs 17B, 18B) and a lingulate abdominal spatula (Figs 17D, 18D). However, the spine on the protibia is blunt (Fig. 18F) and the abdominal spatula is shorter (Fig. 18D) than that of the latter species.

Gastrobothrus sharpi (Broun, 1880)

Figures 2, 4, 19

Bryaxis sharpi Broun, 1880; Bryaxis abdominalis Broun, 1880; Gastrobothrus abdominalis [combination by Broun (1882)]; Gastrobothrus sharpi [synonymized by Shen et al. (2020a)]

Material examined.

N = 12, 3♂♂ 9♀♀. 1♀ point-mounted (AMNZ91972): NEW ZEALAND: AK, Auckland City, Symonds St Cemetery, Grafton Gully side, 24-XII-2003, sedges in bush, S. E. Thorpe. 1♀ point-mounted (NZAC04261576): NEW ZEALAND: AK, Lynfield L, 19-IV-1981, Dysoxylum mould, G. Kuschel. 1♀ point-mounted (NZAC04261578): NEW ZEALAND: AK, Lynfield, 6-XII-1975, blind weevil site, G. Kuschel. 1♀ point-mounted (NZAC04261579): NEW ZEALAND: AK, Lynfield, 24-VII-1976, Cortaderia, G. Kuschel. 1♂ DNA extracted and slide-mounted (NZAC, code absent): NEW ZEALAND: AK, Mt. William Scenic Reserve, 26-I-2011, 37°12'55.10"S 175°1'42.10"E 150 m, sifting litter in mixed forest, R. Anderson. 1♀ point-mounted (NZAC04261500): NEW ZEALAND: ND, Headquarters Rd, beside river, Waipoua Forest, 16-XII-1976, litter 76/110, V. A. May. 2♀♀ point-mounted (AMNZ151662; AMNZ151663): NEW ZEALAND: ND, Kapowairua, 25-III-2018–27-III-2018, pan trap in damp bush, L30478, J. W. Early. 1♀ card-mounted (NZAC04261626): NEW ZEALAND: ND, Mair Park, Whangarei, 12-XII-1944, leaves, E. Fairburn // A. E. Brookes Collection. 1♂ point-mounted (NZAC04261501): NEW ZEALAND: ND, Mangamuka Summit 400 m, 13-XII-1976, litter 76/106, V. A. May”. 1♂ 1♀ point-mounted (NZAC04261502; NZAC04261503): “NEW ZEALAND: ND, Waipoua SF 100 m, 17-III-1978, under Agathis australis log on bank, S. Peck & J. Peck // S. & J. Peck Collection.

Figure 19. 

Gastrobothrus sharpi (Broun, 1880) A Dorsal habitus. B Head, dorsal view. C Head lateral view. D Thorax. E Abdominal sternites. F Aedeagus. G Protibia and tarsus. H Mesotibia and tarsus. I Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–H).

Diagnosis.

Prominent median depression on abdominal sternites VI–VIII and pair of basolateral projections on VII toward midline (Fig. 19D).

Redescription.

Color of body dark brown, maxillary palpi and elytra yellowish-brown. MALE: Measurements: Length 2.86–3.02 mm, width 1.15–1.31 mm (Fig. 19A). Head: Length 0.53–0.55 mm and width 0.52–0.55 mm (Fig. 19B, C); vertex moderately covered with long setae, vertexal foveae moderate and setose; eye longer than temple; ocular-mandibular carina indistinct; gular carinae gradually broadened toward apex and gently swollen ventrally (Figs 4, 19C). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity absent. Antennal socket slightly developed; antenna 11-segmented and club consisting of three antennomeres; scape cylindrical; pedicel parallel; antennomeres 3–4 short, subequal in length; 5 elongate, as long as scape; 6–7 subequal in length; 8 shortest, wider than long; 9–10 asymmetrical; 11 largest. Prothorax: Length 0.56–0.60 mm and width 0.65–0.71 mm (Fig. 19C). Pronotal median antebasal fovea punctiform and asetose, lateral antebasal foveae setose. Elytra: Length 0.74–0.85 mm and width 1.08–1.18 mm (Fig. 19C), and broadened markedly, widest 2.6–2.7 times as wide as base, basal fovea absent. Legs: Robust and long. Protrochanter acutely pointed at ventral, profemur swollen, protibia curved inward with tiny apical spine (Fig. 19F). Mesotibia with minute apical spine (Fig. 19H). Abdomen: Length 1.02–1.03 mm and width 1.15–1.31 mm (Fig. 19E). Tergites IV–VIII subequal in length and bearing basolateral foveae; V broader than IV. Sternites III barely visible; IV short at middle; middle of V depressed, bordering IV; middle of VI acutely descending posteriorly; VII acutely ascending posteriorly at middle, basolateral spines medially protruded; VIII acutely ascending posteriorly at middle, lateral concealed by VII. Aedeagus: 0.65 mm (Fig. 19F); parameres thick, gently curved inward and notched at apex, single short seta and single long seta on each paramere; penis ovoid; endophallus composed of single sclerotized armature rightward, bifid basal one-third and longer branch situated longitudinally. — FEMALE: Eye less developed; legs simple; abdomen narrow, tergite VII concealed by VI in dorsal view, sternites unmodified.

Distribution.

North Island: AK, ND (Fig. 2).

Habitat.

Leaf litter from mixed forest. Logs of kauri. Sedges in bush. Fungus in Dysoxylum.

Gastrobothrus spinifer Choi, Leschen & Park, sp. nov.

Figures 3, 4, 20

Type material.

N = 8, 5♂♂ 3♀♀. Holotype. ♂ point-mounted: “New Zealand: DN, Leith Valley, Leith saddle track, 18/Dec/2017, 45.802799S,170.490550E, sifted broad leaf litter, JWS043, J. Shen & J. Nunn // NZAC04291717”. Paratypes. 1♂ DNA extracted and slide-mounted (NZAC, code absent) 1♂ 1♀ preserved in ethanol (NZAC, code absent) 1♀ point-mounted (NZAC04291718): “NEW ZEALAND: DN, Leith Valley, Leith saddle track, 18-XII-2017, 45.802799S,170.490550E, sifted broad leaf litter, JWS043, J. Shen & J. Nunn”. 1♂ 1♀ preserved in ethanol (NZAC, code absent): “NEW ZEALAND: DN, Otago peninsula, Sandymount, 18-XII-2017, 45°89'13"S, 170°67'22"E, sifted tussock litter, JWS042, J. Shen & J. Nunn”. 1♂ preserved in ethanol (NZAC, code absent): “NEW ZEALAND: DN, Ross creek reservoir, 17-XII-2017, 45°84'96"S, 170°49'41"E, sifted Kanuka and Schefflera litter, JWS041, J. Shen & J. Nunn”.

Figure 20. 

Gastrobothrus spinifer Choi, Leschen & Park, sp. nov. A Habitus, dorsal view. B Head. C Thorax. D Abdominal sternites. E Aedeagus. F Protibia and tarsus; apical spine [red arrow]. G Mesotibia and tarsus; apical spine [red arrow]. H Metatibia and tarsus; trichome [red arrow]. I Protrochanter and femur; spine [red arrow]. J Mesotrochanter and femur. K Metatrochanter and femur. Scale bars: 1 mm (A), 0.1 mm (B–K).

Etymology.

The specific epithet is the Latin word spinifer (“spiny”, masculine) that refers to the spines on legs of this species.

Diagnosis.

Pronotum bearing transverse sulcus (Fig. 20C); each elytron with sutural stria, discal stria, and subhumeral fovea (Fig. 20C); robust femur (Fig. 20I–K); acute spine on protrochanter, pro- and mesotibiae (Fig. 20F, G, I); concave abdominal sternites IV–VIII (Fig. 20D).

Description.

Color of body reddish-brown. MALE: Measurements: Length 2.58 mm, width 0.91 mm (Fig. 20A). Head: Length 0.51 mm and width 0.46 mm (Fig. 20B); vertex moderately covered with short setae, vertexal foveae setose with short setae; eye moderately developed; ocular-mandibular carina distinct, ventrolateral margin extending beneath eye; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity shallow. Antennal socket moderately developed; antenna long and unmodified overall, 11-segmented and club consisting of three antennomeres; scape and pedicel cylindrical; 3–7 elongated; 8 shortest; 9–10 broadened apically; 11 as long as 9+10. Prothorax: Length 0.46 mm and width 0.55 mm (Fig. 20C). Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose with short setae. Elytra: Length 0.65 mm and width 0.91 mm (Fig. 20C), and broadened markedly, widest 2.1 times as wide as base; each elytron with two basal foveae, sutural stria, discal stria, and subhumeral fovea. Hindwing absent. Legs: Robust and long. Protrochanter with acute spine ventrally projecting (Fig. 20I), protibia with prominent apical spine (Fig. 20F). Mesotrochanter with blunt spine ventrally projecting (Fig. 20J), mesotibia with prominent apical spine (Fig. 20G). Metatrochanter with blunt spine ventrally projecting (Fig. 20K), metatibia with apical trichome (Fig. 20H). Abdomen: length 0.96 mm and width 0.91 mm (Fig. 20D). Sternites IV–VIII depressed medially. Aedeagus: 0.62 mm long (Fig. 20E); parameres curved and each paramere with transverse lobe at apex, with two short setae; penis ovoid; endophallus composed of paired armatures. — FEMALE: Eye slightly smaller; legs slender and simple; concavity on abdominal sternites absent.

Distribution.

South Island: DN (Fig. 3).

Habitat.

Broad leaf litter. Litter of tussock grass, kanuka, and Schefflera.

Remarks.

Adults of this species are close to those of G. sulcipennis (Broun, 1921) comb. nov. in having sulci on elytra (Figs 20C, 21D) but can be distinguished from by following the combination of the characters: larger body size; sharply projected protrochanter (Fig. 20I); prominent spine on pro- and mesotibiae (Fig. 20F, G); depression on abdominal sternites IV–VIII (Fig. 20D).

Gastrobothrus sulcipennis (Broun, 1921), comb. nov.

Figures 3, 4, 21

Eupinogitus sulcipennis Broun, 1921; Eupinogitus picescens Broun, 1921 [impliedly synonymized with Eupinogitus sulcipennis by Nomura and Leschen (2006)]

Material examined.

N = 3, 2♂♂ 1♀. 1♂ 1♀ point-mounted, 1♂ DNA extracted and slide-mounted (NZAC04291715; NZAC04291716; NZAC, code absent): NEW ZEALAND: FD, Road to Percy Saddle, 24-I-2018, 45.6066S 167.341767E 380 m, sifting litter among sedges, RL2021, R. Leschen, S. Tassell.

Figure 21. 

Gastrobothrus sulcipennis (Broun, 1921), comb. nov. A Habitus, dorsal view. B Head, dorsal view. C Head, lateral view. D Thorax. E Abdominal sternites. F Aedeagus. G Protibia and tarsus. H Mesotibia and tarsus. I Metatibia and tarsus; trichome [red arrow]. J Protrochanter and femur. K Mesotrochanter and femur. L Metatrochanter and femur; spine [red arrow]. Scale bars: 1 mm (A), 0.1 mm (B–K).

Diagnosis.

Pronotum bearing transverse sulcus (Fig. 21D); each elytron with sutural stria, discal stria, and subhumeral fovea (Fig. 21D); tiny acute spine on mesotrochanter (Fig. 21K); apical trichome with metatibia (Fig. 21I).

Redescription.

Color of body light reddish-brown, appendages yellowish-brown. MALE: Measurements: Length 1.94 mm, width 0.78 mm (Fig. 21A). Head: Length 0.42 mm and width 0.39 mm (Fig. 21B); vertex moderately covered with short setae, vertexal foveae moderate and setose with short setae; eye slightly shorter than temple; ocular-mandibular carina distinct, ventrolateral margin extending beneath eye; gular carinae gradually broadened toward apex and gently swollen ventrally (Figs 4, 21C). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity distinct. Antennal socket barely visible; antenna 11-segmented and club consisting of three antennomeres; scape cylindrical; pedicel parallel; antennomeres 3–4 short, subequal in length; 5 slightly elongate; 6–7 subequal in length; 8 shortest, longer than wide; 9–10 asymmetrical; 11 largest. Prothorax: Length 0.41 mm and width 0.49 mm (Fig. 21D). Pronotal median antebasal fovea punctiform and asetose, lateral antebasal foveae setose with short setae. Elytra: Length 0.61 mm and width 0.78 mm (Fig. 21D) and broadened markedly, widest 2.2 times as wide as base; each elytron with two basal foveae, sutural stria, discal stria, and subhumeral fovea. Legs: Moderately slender. Protrochanter bluntly pointed at ventral profemur slightly swollen (Fig. 21J), protibia straight with apical spine (Fig. 21G). Mesotrochanter sharply pointed at ventral, mesofemur slightly swollen (Fig. 21K), mesotibia straight with apical spine (Fig. 21H). Metatibia with apical trichome (Fig. 21I). Abdomen: Length 0.50 mm and width 0.77 mm (Fig. 21E). Tergites IV–VII subequal in length and narrowed in order, bearing basolateral foveae; IV with mediobasal foveae. Sternite III with basolateral foveae; IV with mediobasal foveae; V–VIII lacking foveae. Aedeagus: 0.54 mm (Fig. 21F); parameres lanciform bearing three setae on each. — FEMALE: Antennae less elongate; lateral antebasal fovea on pronotum smaller; mesotrochanter lacking spine.

Distribution.

South Island: FD, OL (Fig. 3).

Habitat.

Broad leaf litter. Litter among sedges.

Gastrobothrus trigonus Choi, Leschen & Park, sp. nov.

Figures 2, 4, 22

Type material.

N = 2, 2♂♂. Holotype. ♂ point-mounted: “NEW ZEALAND: BP, Lake Okataina, 9-X-1995, litter 95/12, M-C Larivière & A Larochelle // NZAC04261725”. Paratype. 1♂ point-mounted (NZAC04261727): “NEW ZEALAND: TO, Minginui SF, 28/Jul/1977, litter 77/88, J. S. Dugdale”.

Figure 22. 

Gastrobothrus trigonus, sp. nov. A Habitus, dorsal view. B Head. C Pronotum. D Elytra. E Abdominal sternites; spatula [red arrow]. F Aedeagus. G Protibia and tarsus. H Mesotibia and tarsus. I Metatibia and tarsus. Scale bars: 1 mm (A), 0.1 mm (B–I).

Etymology.

The specific epithet is the Latin word trigonus (“three-angled”, masculine) that refers to the triangular abdominal spatula of this species.

Diagnosis.

Pronotum bearing transverse antebasal sulcus (Fig. 22B); metatibia with apical trichome (Fig. 22I); abdominal sternite VI with large triangular spatula (Fig. 22E).

Description.

Color of body reddish-brown, elytra and appendages yellowish-brown. MALE: Measurements: Length 2.5 mm, width 0.93 mm (Fig. 22A). Head: Length 0.58 mm and width 0.51 mm (Fig. 22B), vertex moderately covered with long setae, vertexal foveae markedly setose; eye well-developed, shorter than temple; ocular-mandibular carina not complete; gular carinae gradually broadened toward apex and gently swollen ventrally (Fig. 4). Frontal fovea absent, frontal rostrum lacking and depressed medially; postantennal cavity vague. Antennal socket well-developed; antennae unmodified overall, 11-segmented and club consisting of three antennomeres; scape cylindrical and longer than wide; pedicel cylindrical; 3–4 broadened apically; 5 longer than wide; 6–7 identical; 8 shortest; 11 as long as 9+10. Prothorax: Length 0.51 mm and width 0.57 mm (Fig. 22C), regularly covered with long setae. Pronotal median antebasal fovea reduced and asetose, lateral antebasal foveae setose, transverse antebasal sulcus prominent. Elytra: Length 0.61 mm and width 0.93 mm (Fig. 22D), broadened posteriorly, widest 2.4 times as wide as base; regularly covered with long setae; basal fovea absent. Legs Long and slightly stout. Protibia with blunt apical spine (Fig. 22G). Mesotibia with minute apical spine (Fig. 22H). Metatibia with apical trichome (Fig. 22I). Abdomen: Length 0.80 mm and width 0.93 mm (Fig. 22E). Sternite VI bearing triangular spatula twice as wide as median sclerite of sternite IX. Aedeagus: 0.63 mm long (Fig. 22F); parameres swollen apically, each bearing two short apical setae and one long subapical seta; penis ovoid; endophallic armatures dagger-like. — FEMALE: Unknown.

Distribution.

North Island: BP, TO (Fig. 2).

Habitat.

Leaf litter.

Remarks.

Adults of this species are similar to those of G. inflatus comb. nov. but can be distinguished by the larger abdominal spatula (Fig. 22E).

3.2.3. Unplaced Specimens

The following species are included in the dichotomous key to facilitate future identification. However, they remain unnamed due to the unavailability of aedeagal characters. Although their overall morphology provides sufficient diagnostic features, the dissection and formal description of these specific specimens are deferred due to their extreme fragility. We prioritized the preservation of their morphological integrity over invasive dissection until fresh material becomes available.

Gastrobothrus sp. 15

Figures 2, 4, 23A, B

Material examined.

N = 1, 1♂. 1♂ point-mounted (NZAC04261717): NEW ZEALAND: ND, Waipoua, Tane Mahuta, 29-X-1985, sifted litter 85/47, C. Lyal, R. C. Craw.

Figure 23. 

Gastrobothrus spp. 15–17. A, C, E Habitus, dorsal view. B, D, F Habitus, ventral view. A, B Gastrobothrus sp. 15. C, D Gastrobothrus sp. 16. E, F Gastrobothrus sp. 17. Scale bars: 1 mm.

Diagnosis.

Body length 3.51 mm, head to pronotum black, elytra to abdomen and leg highly dark brown (Fig. 23A); frontal fovea present; pronotal sulcus present (Fig. 23A); protibia lacking spine; abdominal spatula absent (Fig. 23B).

Distribution.

North Island: ND (Fig. 2).

Habitat.

Leaf litter.

Remarks.

This specimen is similar to G. giganteus sp. nov. but differs from it by the absence of spine on protibia and abdominal spatula (Fig. 23B).

Gastrobothrus sp. 16

Figures 2, 4, 23C, D

Material examined.

N = 1, 1♂. 1♂ point-mounted (NZAC04261712): NEW ZEALAND: ND, Omahuta SF, 23-IV-1975, liverworts and mosses from tree bases and ground 75/142, J. S. Dugdale.

Diagnosis.

Body length 2.22 mm, light brown color (Fig. 23C); dorsal side sparsely covered with long setae (Fig. 23C); head and pronotum narrow (Fig. 23C); pronotal sulcus present (Fig. 23C); mesotibia with minute apical spine; abdominal sternite V with small spatula (Fig. 23D).

Distribution.

North Island: ND (Fig. 2).

Habitat.

Liverwort and moss from tree bases and ground.

Remarks.

This specimen is most similar to G. angustus sp. nov. but differs from it by the absence of trichome on metatibia and the smaller blunt abdominal spatula (Fig. 23D).

Gastrobothrus sp. 17

Figures 2, 4, 23E, F

Material examined.

N = 1, 1♂. 1♂ point-mounted (NZAC04261714): NEW ZEALAND: ND, Omahuta SF, 23-IV-1975, liverworts and mosses from tree bases and ground 75/142, J. S. Dugdale.

Diagnosis.

Body length 2.03 mm, light yellowish-brown color (Fig. 23E); dorsal side sparsely covered with long setae (Fig. 23E); pronotum bearing transverse sulcus (Fig. 23E); metatibia with apical trichome; abdominal sternite V with closely paired spatulae (Fig. 23F).

Distribution.

North Island: ND (Fig. 2).

Habitat.

Liverwort and moss from tree bases and ground.

Remarks.

This specimen is most similar to Gastrobothrus sp. 16 but differs from it by the smaller abdominal spatulae separated into two fragments (Fig. 23F). Note that the coloration of this specimen might be faded.

4. Discussion

It is premature to reconstruct the biogeographic history of the genus Gastrobothrus without further field sampling and molecular data, particularly given our conservative approach of refraining from naming or assigning unplaced specimens to valid species.

That said, most members of the genus Gastrobothrus sensu novo are restricted to the North Island, apart from the sister species G. spinifer sp. nov. and G. sulcipennis comb. nov. Within the North Island fauna, one species (G. inflatus comb. nov.) is recorded just south of the Taupo Line (Ellis et al. 2015) and another (G. trigonus sp. nov.) occurs in the Bay of Plenty and Tongariro. The remaining species exhibit high endemism in Northland (10 species) or have broader distributions encompassing this region (three species in Auckland and Northland, and two widespread species), alongside one species strictly endemic to the Three Kings Islands (G. buckleyi). The remarkable diversity in Northland is likely linked to the presence of Pliocene Islands (Strogen et al. 2023) and subsequent Pleistocene climatic and geomorphological fluctuations (e.g., Ellis et al. (2015), Marske and Boyer (2024)). Should the unplaced specimens eventually be validated as distinct species, the recognized species richness in Northland will further increase.

Status of G. buckleyi

Although our molecular phylogeny did not include Gastrobothrus buckleyi Théry & Leschen, morphological evidence indicates that this species deviates significantly from the diagnosis of the genus Gastrobothrus Broun, sensu novo. The prominent ventral protrusion and the distinct widening pattern of the gular carinae imply that this species should be provisionally excluded from the genus in future taxonomic revisions.

5. Data availability

Sequence data used in this study are all available via GenBank. The accession numbers, primers used, and collection information are provided in Table S1 and Table S2.

6. Acknowledgements

We thank Ruby B. Moore and John W. Early (Auckland War Memorial Museum, New Zealand) for the loan of valuable specimens. This work was supported by a grant from the National Research Foundation of Korea (NRF) grant funded by the Korean government (MSIT) (RS-2024-00334183). RABL was funded by Strategic Science Investment Funding for Crown Research Institutes from the Ministry of Business, Innovation and Employment’s Science and Innovation Group; permits were issued by the Department of Conservation, New Zealand.

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Supplementary materials

Supplementary material 1 

Tables S1, S2

Choi Y-J, Leschen RAB, Park J-S (2026)

Data type: .docx

Explanation notes: Table S1. GenBank accession numbers of taxa included to dataset. — Table S2. Primers used in this study.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 2 

Files S1, S2

Choi Y-J, Leschen RAB, Park J-S (2026)

Data type: .zip

Explanation notes: File S1. Nexus file including molecular data for Bayesian inference executed by MrBayes. — File S2. Phylogenetic tree reconstructed by Maximum likelihood.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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