Checklist Journal of Orthoptera Research 2018, 27(2): 119-142 Preliminary study on the diversity of Orthoptera from Kuala Belalong Field Studies Centre, Brunei Darussalam, Borneo MING Kal TAN!, RoOpzAY BIN HAul ABDUL WAHAB? 1 Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Republic of Singapore. 2 Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam, Jalan Universiti, BE1410, Brunei Darussalam. Corresponding author: Ming Kai Tan (orthoptera.mingkai@gmail.com) Academic editor: Juliana Chamorro-Rengifo | Received 1 February 2018 | Accepted 19 March 2018 | Published 29 June 2018 http://zoobank.org/1B247C47-A27B-4C55-9DC7-90201 CE3F278 Citation: Tan MK, Wahab RHA (2018) Preliminary study on the diversity of Orthoptera from Kuala Belalong Field Studies Centre, Brunei Darussalam, Borneo. Journal of Orthoptera Research 27(2): 119-142. https://doi.org/10.3897/jor.27.24152 Abstract The Orthoptera, comprising grasshoppers, crickets, and katydids, is diverse and species rich in tropical Southeast Asia, including the island of Borneo. However, not every part of Southeast Asia is equally well sampled and studied. This includes Brunei Darussalam, specifically at the Kuala Be- lalong Field Studies Centre (KBFSC) within the Ulu Temburong National Park. We present here an annotated and illustrated checklist of Orthop- tera from the primary dipterocarp forest around Kuala Belalong based on three field trips in 2016 and 2017. We provide notes on their taxonomy (including how each species was identified) and natural history of species. In total, 72 species were recorded, representing eight of the 16 monophy]l- etic orthopteran superfamilies. In total, 73.6% of all species recorded were singletons and doubletons, indicating that many species are probably rare. The collection led to the discovery of ten species new to science already published separately, with more expected to be described from pending material and confirmation. More species, including undescribed ones (at least four new species), are expected with continued sampling effort. De- spite the 21 day-long surveying efforts from three trips around KBFSC, we believe that the species list provided here is non-exhaustive and only a preliminary one. Key words checklist, diversity, natural history, Southeast Asia, species richness, taxonomy Introduction Orthoptera is an order of insects consisting of grasshoppers (suborder Caelifera), and crickets and katydids (suborder Ensif- era). These are among the most diverse (around 28,000 species worldwide) and common terrestrial macro-invertebrates (Cigli- ano et al. 2018). There are about 2,000 species from Southeast Asia (Tan et al. 2017a). While some species are notoriously consid- ered as agricultural pests (Willemse 2001), these species represent a small proportion of the overall diversity of orthopterans. In fact, far more species could be found hiding in the pristine forests from Southeast Asia. Southeast Asia is made up of numerous biodiversity hotspots (Myers et al. 2000), yet, the knowledge of orthopteran diversity in this region is not completely understood, with some regions less surveyed than others (Tan et al. 2017a). So, although species new to science are continuously being described from this region, even more can be expected with further intensive surveys into unchar- tered parts of Southeast Asia. On the island of Borneo, pockets of forests remained un- studied for orthopterans. This includes Brunei Darussalam, par- ticularly in the Temburong District (Fig. 1). Within the primary dipterocarp forest of Ulu Temburong National Park lies the Kuala Belalong Field Studies Centre (KBFSC) (Fig. 2A). KBFSC, located along Sungai (= river in Malay) Belalong (Fig. 2B), is under the charge of Universiti (= University, in Malay) Brunei Darussalam. Sungai Belalong forms part of the Sungai Temburong drainage ba- sin which is 1,100 km? and is the third largest catchment in Bru- nei Darussalam. While a huge range of biodiversity surveys were conducted in the field studies center since its opening in 1990, orthopteran diversity around KBFSC remained understudied and overlooked. There was no concerted effort to sample and examine the orthopterans there. In 2016 and 2017, surveys were conducted around the forests of Kuala Belalong with the objectives of discovering species new to science, collecting new material for taxonomic treatments, and making revisions of poorly known or problematic taxa, as well as to present an annotated checklist of Orthoptera from KBFSC. While new species discovery and taxonomic treatments were pub- lished separately in taxonomic journals (Tan and Wahab 2017a, b, Tan et al. 2017c, d), we present here the first annotated and illustrated checklist of Orthoptera from KBFSC based on the col- lections made in 2016 and 2017. We provide references and notes on the taxonomy (including how each species was identified) and natural history of each species, whenever possible. While some of the species could not be promptly identified owing to the insuf- ficient material, we aim to use this publication as a baseline for future work on the taxonomy of Orthoptera from Brunei Darus- salam and Southeast Asia. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 120 Materials and methods Surveys were conducted by the first author (MKT) in the pri- mary lowland and ridge dipterocarp forests (Fig. 2C) surround- ing the KBFSC, Ulu Temburong National Park, Brunei Darussalam (Fig. 1) in September 2016, January 2017, and July 2017. Two main sites were surveyed: Ashton Trail (including the route to Sun- gai Mata Ikan) located behind the field studies center (Fig. 2D), as well as slightly downstream where the Ulu Ulu Resort and Canopy Tower of the Ulu Temburong National Park are located (Fig. 2E). Opportunistic collecting and sweep-netting were conducted dur- ing daytime (0830 to 1500 hours) and/or during night time (1930 to 2300 hours) (Fig. 2F). Light-trapping was also conducted by switching on the corridor lights on the laboratory building facing Sungai Belalong. Whenever possible, acoustic recordings of calling songs were done using the video-recording function of a Canon EOS 500D digital SLR camera or with a digital sound recorder (Olympus WS-750M, frequency response 40-21,000Hz). The specimens were preserved in absolute analytic-grade ethanol and later pinned and dry-preserved. A single hind leg was preserved in absolute analytic-grade ethanol for future molecular work. M.K. TAN AND R.H.A. WAHAB Habitus images were made with a Canon EOS 500D digital SLR camera with a compact-macro lens EF 100mm f/2.8 Macro USM. Close-up images of morphological features (including male genitalia) were done using a Canon EOS 50D digital SLR camera with a macro photo lens MP-E 65mm f/2.8 USM (1-5x). Canon Macro Twin Lite MT-24EX and Canon Macro Ring Lite MR-14EX were used for lighting and flash. Image editing was ac- complished using Adobe Photoshop CC 2014. Scales were giv- en with the images. Acoustic analysis was done using the open source R package WarbleR (Araya-Salas and Smith-Vidaurre 2017) or Sound Ruler software. Specimens were identified by the authors using relevant, published keys and descriptions (see remarks for each species) and by comparisons of voucher specimens and type specimens (sometimes only images available). Where doubtful, the identi- ties of the orthopteran specimens were verified by orthopterists: Andrej V. Gorochov (for Grylloidea), Sigfrid Ingrisch (for Tettigo- nioidea), Tony Robillard (for Eneopterinae), Luc Willemse (for Acridoidea), Josip Skejo (for Tetrigoidea), Josef Tumbrinck (for Tetrigoidea), Sergey Storozhenko (for Tetrigoidea), and Xingbao Jin (for Tettigonioidea). Brunei Darussalam 100 200 300 400km i — Fig. 1. Map of the island of Borneo with the dot indicating the location of KBFSC in Brunei Darussalam. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB Fig. 2. Environment of KBFSC and surrounding habitats: A. View of the KBFSC from Sungai Belalong; B. View of Sungai Belalong from KBFSC; C. Afternoon view of the canopy on the ridge along Ashton Trail; D. Sungai Mata Ikan, a small forest stream near KBFSC; E. Morning view of the canopy at Ulu Temburong National Park; F. Dusk time at the Canopy Tower. Classification and nomenclature of species were based on the Orthoptera Species File (OSF) Online Version 5.0/5.0 (Cigliano et al. 2018). The families, subfamilies, and genera are arranged alphabetically for ease of reference. All specimens collected will eventually be deposited in Institute for Biodiversity and Environ- mental Research (IBER), Universiti Brunei Darussalam and Zoo- logical Reference Collection (ZRC), Lee Kong Chian Natural His- tory Museum (former Raffles Museum of Biodiversity Research), National University of Singapore. Results and discussion Overall, 72 species representing nine families were collected between 2016 and 2017 (Table 1). The species from KBFSC repre- sent eight of the sixteen monophyletic orthopteran superfamilies (sensu Song et al. 2015), thus covering 50% of the lineages in the orthopteran phylogeny. As expected, the monophyletic suborder Ensifera is represented with more species (53 species, 73.6%) than monophyletic suborder Caelifera (19 species, 26.4%). The JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 122 Table 1. A summary of the coverage of orthopteran superfamilies M.K. TAN AND R.H.A. WAHAB and families that were represented in the collection in KBFSC. Ranking Nomenclature Species Richness Ls Order Orthoptera Let: Suborder Caelifera ILE, Superfamily Acridoidea 5 LA Family Acrididae 1.1.2. Superfamily Tetrigoidea 13 1.21 Family Tetrigidae L3l232 Superfamily Trigonopterygoidea 1 isicael Family Trigonopterygidae 125 Suborder Ensifera 12 Superfamily Grylloidea 21 i ya Bal Family Gryllidae 1326722 Family Mogoplistidae L322; Superfamily Gryllotalpoidea 1 12771 Family Gryllotalpidae ibys Superfamily | Rhaphidophoroidea 1 iss Sai | Family Rhaphidophoridae 1.2.4. Superfamily Stenopelmatoidea 5 1.2.4.1 Family Gryllacrididae 1:25: Superfamily Tettigonioidea 25 12:52) Family Tettigoniidae most speciose groups are the Tettigonioidea (34.7%), followed by Grylloidea (29.2%), and Tetrigoidea (18.1%). On the other hand, Trigonopterygoidea and Rhaphidophoroidea are only represented by a single species each. So far, ten species new to science had been described based on material from the collection. From the col- lection, 38 species (52.8%) recorded were singletons, whereas 15 species (20.8%) recorded were doubletons. This represents 73.6% of all species recorded, indicating that many species in the forest are likely to be rare and difficult to collect. More species, including undescribed ones (at least four new species), are expected with continued sampling effort. Despite the 21 day-long surveying ef- forts from three trips, we believe that the species list provided here is non-exhaustive and only a preliminary one. Checklist Order Orthoptera Suborder Caelifera Superfamily Acridoidea Family Acrididae 1. Craneopsis sp. Fig. 3 Remarks.—We tentatively identified this species to be close to C. olivacea Ramme, 1941. Both adults and nymphs of the species were found on Citrus limon at KBFSC. The grasshoppers feed on the leaves, and appear to prefer the older leaves to the younger ones. We identified our species based on the key to species by Willemse (1956) as well as images of type specimens (courtesy of L. Wil- lemse). To date, this genus is endemic to Borneo. The male phallus was previously unexamined. Virtually nothing is known about the life history of these elusive grasshoppers. 2. Perakia sp. Fig. 4A Remarks.—We tentatively identified this species to be near to P. born- eensis Willemse, 1936. This cryptic grasshopper was found on tree branches in the dipterocarp forest. The brown and elongated habitus of the grasshopper enables it to camouflage on branches. Interest- ingly, they were only encountered after the rain, perhaps suggesting that it is a canopy species, brought down to the understory by the rainfall. We identified our species based on the generic key by Wil- lemse (1956). We only have females and can only identify the species name tentatively. Identification was based mainly on color patterns. 3. Traulacris sp. Fig. 4B Remarks.—A female specimen was identified using the generic key by Willemse (1956). The species is very similar to Meltripata Boli- var, 1923 but differs by: head distinctly elevated above level of pro- notum and frontal ridge barely projecting, and female subgenital plate quadrate with a mucronate projection on apical margin. Only two species are currently known from this Bornean-endemic ge- nus, but our specimen differs in coloration from both species, par- ticularly in the hind femur and general body coloration, as well as tegmen length (Willemse 1957). Our specimen may represent an undescribed species, but more material is needed for confirmation. 4. Traulia azureipennis (Serville, 1838) Remarks.—This species was found near the forest edge. It was iden- tified based on the description of the color in Willemse (1957) and comparison with type images from OSF (Cigliano et al. 2018). 5. Traulia borneensis Willemse, 1921 Fig. 4C Remarks.—The two Traulia species were found co-occurring in sim- ilar habitat, although this species can also be found in more for- ested areas of the Canopy Walk. This species was identified based on the description of the color in Willemse (1957) and compari- son with type images from OSF (Cigliano et al. 2018). Superfamily Tetrigoidea Family Tetrigidae Subfamily Cladonotinae 1. Boczkitettix borneensis (Giinther, 1935) Fig. 5A Remarks.—This species was fairly common in the forest floor of Kuala Belalong, usually camouflaging among the leaf litter or for- est trails. We used the key by Tumbrinck (2014) for identification and the material was also verified by J. Tumbrinck and J. Skejo. 2. Epitettix punctatus Hancock, 1907 Fig. 5B, C Remarks.—This species was fairly common in the forest floor of Kuala Belalong, usually camouflaging among the leaf litter or forest trails. We used the key by Tumbrinck (2014) to identify to genus. Subsequently, type images and description were used for comparison. Identification was also verified by J. Tumbrinck and J. Skejo. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 123 Fig. 3. Acrididae: Craneopsis cf. olivacea: A. Male adult; B. Nymph; C, D. Female adult; E. Male genitalia in dorsal (top and bottom left inset), ventral (bottom right inset) views. 3. Potua morbillosa (Walker, 1871) Fig. 5DyE Remarks.—We found this pygmy grasshopper on dead logs and tree trunks along Ashton Trail. We used the key by Tumbrinck (2014) to identify to genus. Subsequently, type images, and description were used for comparison. Identification was also verified by J. Tumbrinck and J. Skejo. Subfamily Discotettiginae 4. Discotettix belzebuth (Serville, 1838) Fig. 5F Remarks.—This species was found on the wooden planks and ground along the trail to the Canopy Tower. We compared our specimens with type images from OSF (Cigliano et al. 2018). Iden- tification was also verified by J. Skejo. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB Hig dh a A Big et er” Fig. 4. Acrididae: A. Perakia nr. borneensis; B. Traulacris sp.; C. Traulia borneensis. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 125 Fe) Fig. 5. Tetrigidae: A. Boczkitettix borneensis; B, C. Epitettix punctatus; D, E. Potua morbillosa; F. Discotettix belzebuth; G, H. Lamellitettigodes contractus; I, J. Pseudoparatettix lineatus. Scale bars: 1 mm. Subfamily Metrodorinae 6. Pseudoparatettix lineatus (Hancock, 1907) Fig. SL, J 5. Lamellitettigodes contractus (Bolivar, 1887) Bigs Gel Remarks.—Unlike most ground-dwelling pygmy grasshoppers, this species was found on foliage of understory plants. This species was Remarks.—A single specimen (a female) collected from the board- identified by J. Tumbrinck. walk en route to the Canopy Tower represents the first record of this species in Brunei Darussalam and Borneo. This species was identified by J. Tumbrinck. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 126 Subfamily Scelimeninae 7. Falconius clavatus Bolivar, 1898 Fig. 6A, B Remarks.—These pygmy grasshoppers were collected near the sandy bank of the Sungai Belalong at KBFSC. We compared our specimens with type images from OSF (Cigliano et al. 2018). Iden- tification was also verified by J. Tumbrinck and J. Skejo. 8. Falconius deceptor Giinther, 1938 Fig. 6C Remarks.—A single female specimen was found attracted to light. We compared our specimen with type images from OSF (Cigliano et al. 2018). Identification was also verified by J. Tumbrinck and J. Skejo. 9. Loxilobus sp. 1 Fig. 6D Remarks.—This pygmy grasshopper was found near small drains near the forest edge at KBFSC. The species was identified by J. Tumbrinck. 10. Loxilobus sp. 2 Fig. GE Remarks.—This pygmy grasshopper was also found near small drains near the forest edge at KBFSC. The species was identified by J. Tumbrinck. 11. Scelimena sp. Fig. 6K G Remarks.—We tentatively identified this species to be near to S. dentiumeris (Hancock, 1907). This large pygmy grasshopper is a powerful flyer that was collected along the sandy river bank along Sungai Belalong at KBFSC. The type specimen of the species was from Brunei Darussalam. We compared our single female speci- men with type images from OSF (Cigliano et al. 2018). 12. Thoradonta nodulosa (Stal, 1861) Fig. 6H, | Remarks.—We compared our single female specimen with type images from OSF (Cigliano et al. 2018). Identification was also verified by J. Tumbrinck and J. Skejo. This is also plausibly the first record of this species in Borneo (Cigliano et al. 2018). Subfamily Tetriginae 13. Coptotettix sp. Fig. 6J Remarks.—This pygmy grasshopper was found near small drains near the forest edge at KBFSC, probably because they are hydro- philic. The species was identified by J. Tumbrinck. M.K. TAN AND R.H.A. WAHAB Superfamily Trigonopterygoidea Family Trigonopterygidae 1. Systella platyptera (Haan, 1842) Fig. 7 Remarks.—We found numerous individuals on the forest under- story near the Canopy Tower. We compared our specimens with images from OSF (Cigliano et al. 2018) for identification. Suborder Ensifera Superfamily Grylloidea Family Gryllidae Subfamily Eneopterinae 1. Cardiodactylus pelagus Otte, 2007 Fig. 8A—D Remarks.—These crickets are sometimes encountered among veg- etation in the understory of the dipterocarp forest. The nymphs can sometimes be found on the forest floor. We compared our specimens with images from Robillard et al. (2013). Identification was verified by T. Robillard. 2. Nisitrus vittatus (Haan, 1844) Fig. 8E-H Remarks.—This species was found abundantly around the station at the forest edge. During the day, the males can be heard calling on plant foliage. We also observed them feeding on flowers (Tan et al. 2017b). We compared this species with other congeners from Malay Peninsula and Borneo with help from T. Robillard. Tenta- tively, we considered the specimens from Kuala Belalong as Nisi- trus vittatus, which can also be found in Malay Peninsula and Su- matra (Robillard and Tan 2013). There are some minor differences in acoustic and morphological characters between our Bornean specimens and those from other parts of Southeast Asia. We need more material from Borneo to better understand the species. Subfamily Gryllinae 3. Gymnogryllus unexpectus trusmadi Gorochov, 2011 Fig. 9A—C Remarks.—The understory can be filled with the loud male calling song of this large cricket during dusk (around 7 pm). The male lives in a burrow and produces loud and high-pitched (peak fre- quency = 4.2 kHz) trilling (mean pulse intervals = 9.1 ms) calls. As they can be quite abundant, the amount of noise produced can be very loud and heard a distance away. Specimens from Kuala Be- lalong resemble the images and description by Gorochov (2011) and identification was verified by A.V. Gorochov. 4. Loxoblemmus sp. Fig. 9D, E Remarks.—This ground cricket was found co-occurring with two species of Velarifictorus among short grasses and herbaceous plants at the Ulu Ulu resort. Despite similarities in habitus, this genus differs from the latter by the head morphology and male genitalia. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 127 Fig. 6. Tetrigidae: A, B. Falconius clavatus; C. Falconius deceptor; D. H, I. Thoradonta nodulosa; J. Coptotettix sp. Scale bars: 1 mm. 5. Mimicogryllus sp. Fig. 9F—I Remarks.—This rare and beautiful black cricket was encountered only once on the forest floor of the dipterocarp forest. A.V. Goro- chov assisted with the identification of this species and this could represent an undescribed species (see Gorochov 1994). However, more material would be necessary before species description can be done. 6. Velarifictorus (Velarifictorus) aspersus aspersus (Walker, 1869) Remarks.—This species is one of the two species of the cosmopoli- tan genus Velarifictorus Randell, 1964 (Modicogryllini) found in Loxilobus sp. 1; E. Loxilobus sp. 2; F, G. Scelimena nr. dentiumeris; Kuala Belalong. Male genitalia resemble that of the species illus- trated in Ingrisch (1998a). This species is widespread in Southeast Asia, and can be found in Thailand, Singapore, and Peninsular Malaysia (Ingrisch 1998a, Tan 2012, Tan and Kamaruddin 2014, 2016, Dawwrueng et al. 2017). Refer to Tan et al. (2017c) for more details on the calling songs and species distribution. 7. Velarifictorus (Velarifictorus) temburongensis Tan et al., 2017 Remarks.—This species was found near river banks, among short grasses and herbaceous plants. It was identified as undescribed and the description was subsequently published in Tan et al. (2017c). This represents the first species of the genus Véelarifictorus to be described from Borneo. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB Fig. 7. Trigonopterygidae: Systella platyptera male adults. Subfamily Landrevinae 8. Duolandrevus (Bejorama) lambir Gorochov, 2017 Remarks.—Prior to the description, we could only tentatively iden- tify the male specimen to be close to D. luzonensis (Tan and Wa- hab 2017a), which was subsequently considered as a new species described from Lambir Hill National Park by Gorochov (2017 a). 9. Duolandrevus (Eulandrevus) kawataredoki Tan & Wahab, 2017 Remarks.—One of the two Landrevinae newly described from the col- lection; this cricket can be commonly heard after dusk in the diptero- carp forest, often while hiding among dead leaves hanging on branch- es. They were also observed to feed on dead leaves. Refer to Tan and Wahab (2017a) for species description and calling song description. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 129 Fig. 8. Gryllidae: Eneopterinae: Cardiodactylus pelagus: A. Male; B. Female; C, D. Male genitalia in ventral and dorsal views respectively; Nisitrus vittatus: E, F Male; G, H. Male genitalia in dorsal and ventral views respectively. 10. Endodrelanva nympha Tan & Wahab, 2017 Remarks.—One of the two Landrevinae newly described from the col- lection; this cricket was found among branches of trees in the diptero- carp forest. Refer to Tan and Wahab (2017a) for species description. Subfamily Podoscirtinae 11. Aphonoides sp. Fig. 10A Remarks.—A single specimen (a female) was collected from a tree trunk in the dipterocarp forest. It resembles congeners of Aphonoides by: inner tympanum open, but outer one obliterated, tegminal Sc with several normal branches, ovipositor with apex rounded, and with large teeth on hind and ventral surfaces (Gorochov 2006). Many species from Borneo were described without the females, so determination of the species name is not possible without the males. 12. Sonotrella (Megatrella) sp. Fig. 10B Remarks.—Our female specimen may belong to Sonotrella (Meg- atrella) optima Gorochov, 2002 or Sonotrella (Megatrella) remota Gorochov, 2002 both known from Borneo. Only the males of both species are known and females are very similar among congeners. Subfamily Pteroplistinae 13. Pteroplistes sp. Fig. 10C, D Remarks. —A single specimen (a female) was spotted camouflaging with the tree trunk of an old tree in the dipterocarp forest. It has characters diagnostic of the genus: large inner and small outer tympana of fore tibia and long ovipositor (Gorochov 2004). Two species have been described from Borneo (i.e. P. borneoensis JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 130 E> —i>— >> — >> — 1 1 17.28 17.3 r 17.24 be Time (s) M.K. TAN AND R.H.A. WAHAB Fig. 9. Gryllidae: Gryllinae: Gymnogryllus unexpectus trusmadi: A. Male; B. Calling song; C. Male genitalia in lateral view; Loxoblemmus sp.: D. Male; E. Female; Mimicogryllus sp.: F Male; G-I. Male genitalia in ventral, dorsal, and lateral views respectively. Gorochov, 2004 and P. lagrecai Gorochov, 2004), neither of which has had the female described (Gorochov 2004). It is therefore difficult to affirm the species name. Subfamily Trigonidiinae 14. Amusurgus or Metiochodes sp.? Fig. 10E Remarks.—Only a single specimen (a female) was collected. The species is characterized by tegmen pubescence, presence of only inner tympanum (very large), and ovipositor with apical valves broader than stem but with acute teeth apically. 15. Amusurgus or Paratrigonidium sp.? Fig. 10F Remarks.— Distinct from all other Trigonidiinae, we, however, only collected females, making it nearly impossible to differentiate them among closely related genera. Tegmen not pubescent and abdomen distinctly humped dorsally. 16. Paratrigonidium sp. Fig. 10G Remarks.—The male can be distinguished from the other Trigoni- diinae above by the developed stridulatory apparatus, and foretib- JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 131 B ree Fig. 10. Gryllidae: Podoscirtinae: A. Aphonoides sp.; B. Sonotrella (Megatrella) sp.; Grylloidea: Pteroplistinae: C, D. Pteroplistes sp.; Trigo- nidiidae: Trigonidiinae: E. Amusurgus or Metiochodes sp.?; F. Amusurgus or Paratrigonidium sp.?; G. Paratrigonidium sp. — JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 132 ia with inner and outer tympana. The male genitalia also resemble that of a typical Paratrigonidium. More specimens will, however, be needed to ascertain the genus and species. This group is probably in need of further taxonomic revision. Family Phalangopsidae Subfamily Phalangopsinae 17. Parendacustes (Minizacla) mulu Gorochov, 2017 Fig. 11A—D Remarks.—Juveniles and adults were found on the tree trunks of old trees in the dipterocarp forest. They were well camouflaged among the ridges of the tree trunk. A.V. Gorochov assisted with the identification of this species which he recently described from Mulu National Park, which is relatively close to KBFSC (Gorochov 201 7b, c). 18. Terrozacla borneo Gorochov, 2014 Fig. 11E-G Remarks.—This species was fairly common in the forest floor of Kuala Belalong, usually camouflaging among the leaf litter or forest trails. T. Robillard assisted with the identification of this species. We compared the genitalia of our specimens with im- ages from Gorochov (2014). This species can be found in Sabah, Sarawak, and now Brunei Darussalam. Subfamily Phaloriinae 19. Phaloria (Papuloria) tristis Gorochov, 2014 Fig. 1IH-K Remarks.—The males produce trilling calling songs after dusk, of- ten on foliage of trees. They are quite common in the dipterocarp forest. The genitalia of our specimens resemble that of the type specimen, described from Mulu National Park (Gorochov 2014). Family Mogoplistidae Subfamily Mogoplistinae 20. Cycloptiloides sp. Fig. 12A Remarks.—These small scaly crickets are usually cryptic but were found hopping around on the forest floor. Only females and nymphs were collected but male genitalia is needed for species identification (Ingrisch 2006). This represents the first record of this genus in Borneo (Ingrisch 2006, Cigliano et al. 2018). Because this genus has rarely been studied (Ingrisch 2006), the species from Brunei could be an undescribed species. 21. Ectatoderus sp. Fig. 12B Remarks.—We tentatively identified this species to be close to E. angusticollis Chopard, 1969. The cricket was found among foliage and branches of understory trees. The key to Southeast Asian spe- cies was used for identification but this genus is likely to be para- phyletic and taxonomic work is needed (Ingrisch 2006). M.K. TAN AND R.H.A. WAHAB Superfamily Gryllotalpoidea Family Gryllotalpidae 1. Gryllotalpa (2) sp. Fig. 12C Remarks.—We recorded the calling songs heard in the evening sporadically across the forest floor, even though we did not man- age to collect any specimen. The calling song is characteristic of congeners from the region. This species tends to call near dead and rotten logs at around 6:45-7pm and the male burrows clearly resemble that of congeners. Superfamily Rhaphidophoroidea Family Rhaphidophoridae 1. Rhaphidophora (2) sp. Fig. 12D-F Remarks.—This group is speciose and can be hard to identify for Southeast Asian representatives (A.V. Gorochoy, in litt.). Superfamily Stenopelmatoidea Family Gryllacrididae 1. Asarcogryllacris (Pseudolarnaca) sp. Fig. 13A-E Remarks.—Our single specimen (a male) resembles the drawing of the habitus of A. genualis (Walker, 1869) from Borneo. The abdomi- nal apex (particularly the medial apical sclerotized processes of ninth abdominal tergite) also resembles that in Gorochov (2005). It clearly differs from the only subcongener, A. (P.) nigroscutata (Brunner von Wattenwyl, 1888) from Java, by color patterns and abdominal apex. 2. Capnogryllacris (?) (Capnogryllacris) sp. Fig. 13F Remarks.—We tentatively identified our female specimen to be close to C. superba (Brunner von Wattenwyl, 1888) using an old key (Murphy 1973). This species is characterized by red hind wings. 3. Melaneremus (2) sp. Fig. 13G-J Remarks.—This is among the most common Stenopelmatoidea that was found in the understory of the Kuala Belalong forest. Un- fortunately, we could only tentatively identify our specimens to this genus using an old key (Murphy 1973). However, our speci- mens are also different in shape of male abdominal apices from described species of the genus. Until a revision of the genera of Asiatic Gryllacridinae is completed, the determination of the spe- cies is problematic (A.V. Gorochoy, in litt.). 4. Phlebogryllacris (2) sp. Fig. 14A—D Remarks.—This stunningly red raspy cricket was found among foli- age in the understory. Without comparison with the type specimen, we could only only tentatively identify our single specimen (a male) to be close to P. venosa (Walker, 1869) as it appears superficially sim- ilar and was described from Sarawak close to Brunei Darussalam. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 133 Fig. 11. Phalangopsidae: Parendacustes (Minizacla) mulu: A. Male; B, C. Male genitalia in dorsal and ventral views respectively; D. Male tegmen; Terrozacla borneo: E. Female; F. Male; G. Nymph; Phaloria (Papuloria) trista: H. Male; I. Female; J, K. Male genitalia in dorsal and ventral views respectively. Scale bar: Imm. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 134 M.K. TAN AND R.H.A. WAHAB i NO Frequency 0 2 Fig. 12. Mogoplistidae, Gryllotalpidae, and Rhaphidophoridae: A. Cycloptiloides sp.; B. Ectatoderus cf. angusticollis; C. Gryllotalpa (?) sp. (frequency in kHz); D. Rhaphidophora (?) sp. male; E, F Male abdominal apex. Scale bars: 1mm. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 135 Fig. 13. Gryllacrididae: A. Asarcogryllacris (Pseudolarnaca) cf. genualis male; B. Face; C. Dorsal view of head and pronotum; D, E. Abdominal apex; FE. Capnogryllacris (Capnogryllacris) cf. superba; G. Melaneremus (?) sp. mating pair; H. Male adult; I. Male stridulatory file; J. Male subgenital plate. Scale bars: 1mm. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 136 M.K. TAN AND R.H.A. WAHAB Fig. 14. Gryllacrididae: A. Phlebogryllacris (?) cf. venosa male; B. Face; C. Male abdominal apex; D. Male subgenital plate; E. Phryganog- ryllacris sp. 5. Phryganogryllacris sp. Fig. 14E Remarks.—We tentatively identified our specimens to this genus using an old key (Murphy 1973) and compared with images of the genus from Gorochov (2005). This genus is characterized by a cup-shaped ninth abdominal tergite, posteriorly with spines pointing downward far apart laterally, and tenth abdominal ter- gite without spine-like process. Superfamily Tettigonioidea Family Tettigoniidae Subfamily Conocephalinae 1. Viriacca modesta Gorochov, 2013 Fig. 1I5A Remarks.—This species was found on the foliage of understory plants in the dipterocarp forest. We used key to genera of Agrae- ciini by Ingrisch (1998b) to identify our specimens. Subse- quently we compared our specimens with species descriptions in Ingrisch (1998b) and Gorochov (2011, 2013). This species was described from Mulu National Park. Subfamily Lipotactinae 2. Lipotactes alienus Brunner von Wattenwyl, 1898 Fig. 15B Remarks.—This katydid is not uncommon in the understory of Kuala Belalong forest. However, curiously, we have yet to encoun- ter the males. Unfortunately, the male of this species has also not been described. Our females correspond to the description by In- grisch (1995) and images from OSF (Cigliano et al. 2018) by color patterns, shapes of ovipositor, and subgenital plate. Subfamily Meconematinae 3. Alloteratura (Alloteratura) belalongensis Tan et al., 2017 Remarks.—This is one of the four new species of Meconematini katydids collected and described from Kuala Belalong. This new species is very similar to A. (A.) karnyi Kastner, 1932 from north- ern Sumatra (Kastner 1932, Gorochov 1998a) and Thailand (see Sanger and Helfert 2000) but could belong to several different subspecies of the same species owing to the similarities in mor- phology (Tan et al. 2017d). We need more material around South- east Asia and study of their population genetics to confirm species limits. Refer to Tan et al. (2017d) for the new species description. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 137 Ampl (V) Scare “. “we “w “ “we . S a “ %, “, “w “ “w “w = o | ! ! 9.9 10 10.2 Time (s) Fig. 15. Tettigoniidae: A. Viriacca modesta; B. Lipotactes alienus; C. Asiophlugis longiuncus; D. Neophisis (Indophisis) cf. longipennis or curvata; Mecopoda elongata: E. Female; F. Calling song. 4. Asiophlugis longiuncus Gorochov, 2013 Fig. 15C Remarks.—This species was found on foliage of the dipterocarp forest understory at night. Usually, a few individuals (mix of adults and nymphs) were found on a single large leaf. Only the female was collected, and was compared with the original description (Gorochov 2013). This species was described from Mulu National Park. 5. Kuzicus mirabilis Tan & Wahab, 2018 Remarks.—This species was encountered on foliage of the diptero- carp forest understory at night. Refer to Tan and Wahab (2018a) for the new species description. 6. Leptoteratura (Rhinoteratura) chela Tan et al., 2017 Remarks.—This species was encountered on foliage of the diptero- carp forest understory at night. Refer to Tan et al. (2017d) for the new species description. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 138 7. Neophisis (Indophisis) sp. Fig. 15D Remarks.—We tentatively identified this species to be close to ei- ther N. longipennis Jin, 1992 or N. curvata Jin, 1992. This species was found fairly frequently on foliage of the dipterocarp forest understory at night. 8. Pulchroteratura huiging Tan et al., 2017 Remarks.—This species was quite frequently encountered on fo- liage of the dipterocarp forest understory at night. The peculiar shape of the abdominal apex led us to consider this to be a com- pletely different genus. Refer to Tan et al. (2017d) for the new spe- cies description. 9. Pseudoteratura (Pseudoteratura) kenuan Tan et al., 2017 Remarks.—Refer to Tan et al. (2017d) for the new species description. Subfamily Mecopodinae 10. Mecopoda elongata (Linnaeus, 1758) Fig. 15E, F Remarks.—This large katydid can be heard calling at night among shrubby bushes. The calling song resembles the train call (see Diwakar and Balakrishnan 2007). Sometimes, they were also at- tracted to light. Subfamily Phaneropterinae 11. Baryprostha bellua Karsch, 1891 Fig. 16A Remarks.—The only specimen (a female) we encountered was found on the forest floor of the dipterocarp forest along Ashton Trail. It is probable that it resides in the canopy but had dropped down to the forest floor. We used the key to the species of Bary- prostha from Ingrisch (1990) and images of the type specimen from OSF (Cigliano et al. 2018) for identification. 12. Casigneta bisinuata Karny, 1926 Fig. 16B Remarks.—This species was found attracted to light, thus appeared around the station. We compared our male specimens with im- ages of type specimens from OSF (Cigliano et al. 2018). 13. Hemimirollia gracilis (Karny, 1925) Fig. 16C Remarks.—Only one male specimen was collected during light trapping. We used the key to genera of Mirolliini by Ingrisch (2011) and compared our specimen with the descriptions of the two other congeners. 14. Holochlora sp. Fig. 16D Remarks.—One male specimen was collected during light trapping. Upon freezing the katydid, a horsehair worm (Nematomorpha) M.K. TAN AND R.H.A. WAHAB emerged from the abdominal apex. This parasite is not uncom- mon among katydids found near the water (i.e. Sungai Belalong). The Nematomorpha was preserved in ethanol. The katydid bears the male diagnostic characters of Holochlora: male tenth abdomi- nal tergite with two apical lobes, area around left stridulatory file strongly swollen (Liu and Kang 2007). 15. Mirollia sp. Fig. 16E Remarks.—We tentatively identified this species to be close to M. abnormis Karny, 1926. One female specimen was collected from the foliage on a tree near the Canopy Tower. We used the key to genera of Mirolliini by Ingrisch (2011) and compared our female specimen with the descriptions. Male specimens are needed to ex- amine the titillators and confirm the species identity. 16. Liotrachela (2) sp. Fig. 16F Remarks.—This species was found attracted to light and we pos- tulate that it is also a canopy species. This species is superficially similar to Holochlora but the tenth abdominal tergite in males does not extend backwards into two lobes (Ingrisch 2002), and it pos- sesses dorsal spines on fore tibia (Webber et al. 2003). We tenta- tively identified the specimens as belonging to this genus although it is not currently known from Borneo (Cigliano et al. 2018). More material is needed for verification. 17. Liotrachela hyalina Karny, 1926 Fig. 16G Remarks.—We compared our male specimen with the original de- scription (Karny 1926) and images of type specimen from OSF (Cigliano et al. 2018): male tenth abdominal tergite, cerci, and subgenital plate resemble our specimen. This species was de- scribed from Java, suggesting that our male specimen represents the first record of this species and genus in Borneo. 18. Phygela sp. Fig. 16H Remarks.—We tentatively identified the species to be close to P. marginata Brunner von Wattenwyl, 1878. One female was col- lected from a light trap. We recognized this species as Phygela owing to the tegmen with Rs and M angled at each cross vein and becoming obliterated in a series of large polygonal cells which occupy the radial field beyond the middle; and presence of a small lateral lobe between the subgenital plate and pileolus (this makes it different from an otherwise similar Pseudopsyra Hebard, 1922). Our specimen appears to differ from the female of P. mar- ginata from Borneo by a more elongated ovipositor and a stouter subgenital plate, but further examination of the three described species from Southeast Asia is needed to confirm species identity. 19. Rectimarginalis sp. Fig. 161 Remarks.—We tentatively identified this species to be similar to R. pro- funda Liu & Kang, 2007. We compared our single specimen (a male) with the descriptions of all species of Rectimarginalis Liu & Kang, 2007 (see Liu and Kang 2007) and images of the type specimen from OSF JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB Fig. 16. Tettigoniidae: Phaneropterinae: A. Baryprostha bellua; B. Casigneta bisinuata; C. Hemimirollia gracilis; D. Holochlora sp.; E. Mirollia cf. abnormis; F. Liotrachela (?) sp.; G. Liotrachela hyalina; H. Phygela cf. marginata; 1. Rectimarginalis nr. profunda; J. Sympaestria lampra. (Cigliano et al. 2018). Although R. profunda is not found in Borneo, our specimen resembles this species in its tenth abdominal tergite, subgenital plate, and genitalia, and is more similar to R. profunda than to its congeners found closer to or in Borneo |[i.e. R. fuscospino- sa (Brunner von Wattenwyl, 1891) and R. ensis (Haan, 1843), respec- tively]. There are some small differences in the male tenth abdominal tergite and genitalia. We require more material for verification since only one specimen was collected from light trapping. 20. Stictophaula bruneii Tan & Wahab, 2017 Remarks.—Only one specimen (a male) was collected during light trapping, perhaps suggesting that this is a canopy species rarely encountered in the forest understory. Extensive work recently on this Southeast Asian genus (see Ingrisch 1994, Gorochov 1998b, Gorochov and Kang 2004, Gorochov and Voltshenkova 2009) and images of type specimen from OSF (Cigliano et al. 2018) allowed JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 140 M.K. TAN AND R.H.A. WAHAB Fig. 17. Tettigoniidae: Pseudophyllinae: A. Olcinia dentata; B. Phyllomimus sp.; C. Tympanophyllum sp. us to confirm that our specimen was undescribed. Full description of the new species and a key to all species can be found in Tan and Wahab (2017b). 21. Sympaestria lampra Hebard, 1922 Fig. 16J Remarks.—This species is among the more common species of or- thopterans attracted to light. They are, however, not encountered during opportunistic surveys in the understory, which is indica- tive of this being a canopy dweller. We used original descriptions (Hebard 1922) and type images from OSF (Cigliano et al. 2018) to identify our specimens. 22. Tapiena paraincisa Tan & Wahab, 2018 Remarks.—Our male specimen, collected during light trapping, resembles Tapiena (Tan et al. 2015). It is however unique in the shape of cercus and tenth abdominal tergite, and also differs from T. incisa Karny, 1923 from Sarawak. Refer to Tan and Wahab (2018b) for the new species description. Subfamily Pseudophyllinae 23. Olcinia dentata de Jong, 1939 Fig. 17A Remarks.—Only one specimen (a female) was collected during light trapping, perhaps suggesting that this is a canopy species rarely encountered on the forest understory. The crenulated cos- tal margin of the tegmina distinguish our specimen from allied genera from the region (such as Sathrophyllia Stal, 1874 and Tegra Walker, 1870) (de Jong 1939, 1960, Bresseel and Vermeersch 2017). We also used the keys by De Jong (1960) and Bresseel and Vermeersch (2017) to identify our specimen. 24. Phyllomimus sp. Fig. 17B Remarks.—This species was found on the understory leaves, and can frequently be heard calling. Upon approaching the katydid, it flattened its body and extended its tegmina to camouflage against the background of the leaf. Patterns on the tegmina resemble epi- phylls that were found commonly on the leaves of the trees in the forest around KBFSC (notice the leaf of Fig. 17B). JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) M.K. TAN AND R.H.A. WAHAB 25. Tympanophyllum sp. Fig. 17C Remarks.—This species was found on the understory leaves. This genus is similar to Chondroderella in habitus and shape of tegmen, but differs by tegmen with Sc and R separated at the base and showing some short cross-veins between them (instead of lying closely appressed). Only a single specimen (a male) was collected but we need both male and female to compare with congeners and identify the species. Acknowledgements The authors are grateful to Andrej V. Gorochov (for Gryl- loidea), Sigfrid Ingrisch (for Tettigonioidea), Tony Robillard (for Eneopterinae and Phalangopsidae), Luc Willemse (for Acrid- oidea), Josip Skejo (for Tetrigoidea), Josef Tumbrinck (for Tetrigoi- dea), Sergey Storozhenko (for Tetrigoidea), and Xingbao Jin (for Tettigonioidea) for their help in identification and verification of the specimens. The collection of material in Kuala Belalong Field Studies Centre, Brunei Darussalam was granted by the Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam (UBD/AVC-RI/1.21.1 [a]). 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