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The diversity of Orthoptera in the northwestern parts of Mount Kinabalu, Sabah, Malaysia
expand article infoAmira Aqilah Muhammad‡, Ming Kai Tan§
‡ Universiti Malaya, Kuala Lumpur, Malaysia
§ National University of Singapore, Singapore, Singapore

Corresponding author: Amira Aqilah Muhammad ( miamedulla@gmail.com )

Academic editor: Klaus-Gerhard Heller
© 2025 Amira Aqilah Muhammad, Ming Kai Tan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Muhammad AA, Tan MK (2025) The diversity of Orthoptera in the northwestern parts of Mount Kinabalu, Sabah, Malaysia. Journal of Orthoptera Research 34(1): 95-118. https://doi.org/10.3897/jor.34.128225
ZooBank: urn:lsid:zoobank.org:pub:E468554C-2FB8-421A-B11A-BFD6D126C48B
Open Access

Abstract

The Orthoptera from Borneo require further research since some areas have never been studied, including parts of Mount Kinabalu in Ranau, Sabah. A five-day survey at Marai Parai and Nunuk in October 2023 revealed 32 species of Orthoptera from nine families. There are at least nine new species records for the Mount Kinabalu region and four new species records for the state of Sabah. Twenty-one out of the 32 species collected are singletons, indicating that many species on Mount Kinabalu are rare and of low abundance or collected during the wrong season. Hence, an annotated and illustrated checklist of the Orthoptera of this region is presented with notes on their taxonomy and, where data are available, on their natural history. With extended sampling, it is expected that more species will be discovered than in this preliminary checklist.

Keywords

Checklist, Marai Parai, new record, Southeast Asia, taxonomy

Introduction

Mount Kinabalu is the highest mountain in Borneo, and the habitats in and around this mountain are mega-rich in biodiversity (Nor 2001, Sabah Parks 2022, UNESCO World Heritage Convention 2022). As of May 2024, about 89 species of Orthoptera have been documented to occur at Mount Kinabalu (Cigliano et al. 2024). New species and genera have been recently discovered, including Borneratura atromacula Gorochov, 2022; Depressacca kinabalu Ingrisch, 2020; Subtilodecma bilobata Gorochov, 2022; and Subtilodecma unilobata Gorochov, 2022 (Ingrisch 2020, Gorochov 2022), highlighting that we do not necessarily have a good understanding of this famous mountain’s orthopteran diversity due to the mountain’s vast terrain.

Marai Parai (1652 m.a.s.l.), located on the western slope, is one of the lesser-known areas of Mount Kinabalu (Fig. 1). It is named after the Dusun people’s word for the graminoid vegetation Tetraria pilisepala (Cyperaceae) (Beaman and Beaman 1998), which flourishes in the area. Marai Parai is an ultramafic plateau at the northwest of Mount Kinabalu, Sabah (Fig. 2A) that houses endemic vegetation that includes few species of pitcher plants (Van der Ent et al. 2015) (Fig. 2B). The climate is cold, and the soils are nutrient-poor and waterlogged due to high humidity and frequent rainfall. Kiau Nuluh (880 m.a.s.l.), a village bordering Kinabalu Park in the district of Kota Belud, serves as the starting point of the route to Marai Parai. Surrounding the village is a forest reserve referred to locally as the community forest area of Kiau Nuluh; this extends from the village up to the border of Kinabalu Park, where Nunuk camp (1215 m.a.s.l.) is situated (Fig. 2C, D) (Cooke and Hussin 2014). No published records of Orthoptera from Marai Parai exist (Cigliano et al. 2024), although expeditions have been conducted there as early as 1858 (Gibbs 1914, Anfraix 2005, Barkman et al. 2016, Cross et al. 2022).

Fig. 1. 

Map of Kinabalu Park, Sabah. The dots indicate the location of Marai Parai (blue), Nunuk camp (yellow), and Kiau Nuluh (white) near Mount Kinabalu, Sabah, Malaysia (orange).

Fig. 2. 

Habitats of sampling locations. A. View of the Mount Kinabalu peak from Marai Parai; B. Vegetation along the trail at Marai Parai; C. Vegetation along the trail at Nunuk camp; D. View of Nunuk camp and the surrounding canopy.

In 2023, the first concerted effort to sample orthopterans from Marai Parai was conducted as part of the Marai Parai—West Gurkha Hut Kota Belud Scientific Expedition. As a result, we present here the first annotated and illustrated checklist of Orthoptera from Marai Parai and Nunuk camp. Notes on the taxonomy and natural history of each species are provided whenever possible. While this checklist is likely preliminary, it offers a foundation for future work on the taxonomy of Orthoptera from Mount Kinabalu, Sabah and Southeast Asia.

Material and methods

Sampling.—The first author (AAM) conducted surveys at Marai Parai camp (6.081897°N, 116.520449°E) and Nunuk camp (6.071725°N, 116.512458°E) from 13 October to 17 October 2023. AAM performed opportunistic collecting during the daytime (0830 to 1500 hours) and nighttime (1900 to 2200 hours). Specimens were photographed using an Olympus OM-D E-M5 Mark II interchangeable lens camera with an M.Zuiko Digital ED 60 mm F2.8 macro lens attached.

Specimen preservation and examination of material.—The specimens were first preserved in absolute analytic-grade ethanol before being pinned and dry-preserved. Dissection of the male titillators or phallus was performed by removing them from the abdominal apex of softened specimens before being cleaned using aqueous potassium hydroxide (KOH) and subsequently preserved in glycerine. The morphological features and male genitalia of the specimens were photographed using the same camera and lens listed above and illuminated by external lighting from a Godox Macro Ring Flash ML-150. Additionally, a PULUZ Folding Portable 550LM Light Photo Lighting Studio Shooting Tent Box Kit with a white backdrop was used to enhance the photography. Adobe Photoshop CC 2014 (Adobe Systems Incorporated, San Jose, CA, USA) was utilized for image editing.

Specimen identification.—Both authors identified the specimens using relevant published keys (Willemse 1921, 1933, 1935, Karny 1925a, 1925b Beier 1954, Kevan and Jin 1993, Gorochov 1997, 2002, 2006, 2008a, 2008b, 2014, 2016, 2020, 2021, 2022, Gorochov and Tan 2011, Ingrisch 2011, 2018, Robillard et al. 2023, Tan 2014, Tan and Kamaruddin 2014, Tumbrinck 2014, Tan et al. 2015, 2019, 2021, 2022, Tan and Wahab 2017, 2018, Skejo et al. 2022). The Orthoptera Species File (OSF) Online Version 5.0/5.0 was used for species classification and nomenclature (Cigliano et al. 2024). The identification was further verified by the following orthopterists: Luc Willemse for Acridoidea, Andrej V. Gorochov for Stenopelmatoidea, and Josip Skejo, Karmela Adžić, and Maks Deranja for Tetrigoidea.

Depository.—All specimens collected are planned to be deposited in the Entomological Unit of Sabah Park (EUSP), Kinabalu Park, Ranau, Sabah.

Results and discussion

The five-day sampling period yielded 32 species representing nine families. The species from Marai Parai and Nunuk represent 8 of the 17 monophyletic orthopteran superfamilies (sensu Song et al. 2020), covering 50% of the lineages in the orthopteran phylogeny (Tan and Wahab 2018). The suborder Ensifera is represented by 25 species, which is more than in the monophyletic suborder Caelifera (7 species). The most speciose groups are Tettigonioidea (12 species), followed by Grylloidea (6 species), Tetrigoidea (4 species), and Rhaphidophoroidea (2 species). Eumastacoidea and Gryllotalpoidea are each represented by a single species. Twenty-one of the 32 species (65.6%) collected are singletons, suggesting that many species in this region are rare or of low abundance.

Of the 32 species, only one species (Nisitrus musicus Ingrisch, 1987) was collected from both sampling locations (Marai Parai and Nunuk camp), suggesting that this species is common. Four species were exclusively collected from Marai Parai (Traulacris nr. erecta, Cratioma cf. borneense, Despoina spinosa Brunner von Wattenwyl, 1895, Subtilodecma bilobata) while the other 27 species were collected from Nunuk camp.

Checklist

Suborder Caelifera

Family Acrididae

Subfamily Catantopinae

Traulacris nr. erecta

Fig. 3

Material examined.—EAST MALAYSIA: Sabah • 3♂ 2♀; Ranau, Kinabalu Park, Marai Parai; 14 October 2023; on stem of Tetraria pilisepala plant; leg. A.A. Muhammad, Supani T., Jalter M. & Silver P.; EUSP.

Remarks.—This species is very similar to T. erecta Willemse, 1933, but differs by the anterior and median femora being yellowish green instead of yellowish brown and apical half of hind tibiae of the same color (greenish blue) as the basal half (instead of blackish). It was found within the grassy vegetation of Marai Parai, at approximately 1600 m.a.s.l., but not at Nunuk camp. Male and female specimens were identified using the key in Willemse (1933). Our specimen may represent an undescribed species, but comparison to the type specimen(s) is needed for confirmation. Nevertheless, this finding represents the first record of this species at Mount Kinabalu, which is approximately 170 km west of the type locality (Bettotan [=Betotan], Sandakan).

Fig. 3. 

Traulacris nr. erecta. A–C. Habitus in dorsal, lateral, and frontal view. Scale bars: 2 mm.

Traulia borneensis Willemse, 1921

Fig. 4

Material examined.—EAST MALAYSIA: Sabah • 1♀; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was found on the foliage of understory plants at Nunuk camp. It was identified based on the color description in Willemse (1935) and comparison with type images from OSF (Cigliano et al. 2024). The type locality of this species is unspecified (North Borneo); hence it is impossible to determine it as a first record for this region (Willemse 1921). However, it is probable that this species is widespread in Borneo, as it is also present in Kuala Belalong, Brunei Darussalam (Tan and Wahab 2018).

Fig. 4. 

Traulia borneensis. Scale bar: 10 mm.

Family Chorotypidae

Subfamily Chorotypinae

Chorotypus sp.

Fig. 5

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; at the wall of a building with light source; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Only one individual, attracted to the light source at a building at Nunuk camp, was collected at night. This grasshopper can be found in leaf litter on the forest floor (Tan and Kamaruddin 2014). Identification involved the use of the generic key in Willemse (1930) and comparison with the images in OSF (Cigliano et al. 2024).

Fig. 5. 

Chorotypus sp. Scale bar: 10 mm.

Family Tetrigidae

Subfamily Cladonotinae

Potua cf. morbillosa

Fig. 6A

Material examined.—EAST MALAYSIA: Sabah • 1♂ 2♀; Ranau, Nunuk; 16 October 2023; on rotten logs; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Our specimens are similar to P. morbillosa Walker, 1871, which is commonly found in Borneo, differing slightly by the number of pronotal humps. Unlike P. morbillosa, our specimens have two extra humps in dorsal view (seen as one hump in lateral view) just behind the first hump. Our specimens may represent an undescribed species, but members of this subfamily are known to vary greatly by the shape of their pronotum (Tumbrinck 2014). Hence, more material and alternative analyses are needed for confirmation. This pygmy grasshopper was found on rotten logs along the trail in the community forest area. Morphological identification was verified by J. Skejo, K. Adzic, and M. Deranja and compared to type images in OSF (Cigliano et al. 2024). This finding represents the first official record of this species from Mount Kinabalu and from Sabah. Apart from its type locality in Sarawak, this species has also been recorded in Kuala Belalong, Brunei Darussalam (Tan and Wahab 2018).

Fig. 6. 

Tetrigidae. A. Potua cf. morbillosa; B. Bolivaritettix apterus; C. Disconius shelfordi. Scale bars: 2 mm.

Subfamily Metrodorinae

Bolivaritettix apterus Rehn, 1904

Fig. 6B

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 14 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP • 2♀; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was collected during the day on the foliage of understory plants along the trail in the community forest near Nunuk camp. It is among the most common Tetrigidae species encountered here. Habitus size is rather large (BL > 1.0 mm) for Metrodorinae. The specimens were compared to images in OSF (Cigliano et al. 2024) for identification. This species is widespread in Borneo, with the type locality in Labuan, which is approximately 170 km southwest of Mount Kinabalu.

Subfamily Scelimeninae

Disconius shelfordi Hancock, 1907

Fig. 6C

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; on rotten log; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was found on a rotten log on the forest floor. Unlike Discotettix, this species lacks elevated frontomedial and other highly elevated projections on its pronotum (Skejo et al. 2022). Identification was also verified by J. Skejo, K. Adzic, and M. Deranja. Previously described from Sarawak (Hancock 1907) and Kalimantan (Cigliano et al. 2024), this is the first record of this species for Mount Kinabalu and Sabah.

Discotettix (Discotettix) belzebuth Serville, 1838

Fig. 7

Material examined.—EAST MALAYSIA: Sabah • 2♂ 2♀; Ranau, Nunuk; 16 October 2023; on rotten log; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species is commonly found in the forests of Borneo. In this study, both Disconius shelfordi and D. belzebuth were found co-occurring on a rotten log. This specimen was recognized by its flattened and widened subapical antennal segments and multiple projections on the pronotum. Further identification was done using the description in Skejo et al. (2022).

Fig. 7. 

Discotettix belzebuth. A–C. Habitus in dorsal, frontal, and lateral view. Scale bars: 2 mm.

Suborder Ensifera

Family Gryllidae

Subfamily Eneopterinae

Nisitrus musicus Ingrisch, 1987

Fig. 8

Material examined.—EAST MALAYSIA: Sabah • 1♂ 2♀; Ranau, Kinabalu Park, Marai Parai; 14 October 2023; on foliage; leg. A.A. Muhammad, Supani T., Jalter M. & Silver P.; EUSP • 2♀; Ranau, Nunuk; 16 October 2023; on foliage; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP • 1♀; Ranau, Nunuk; 17 October 2023; on foliage; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Fig. 8. 

Nisitrus musicus. A. Nymph; B. Adult male in dorsolateral view; C. Adult male (right) and female (left). Scale bar: 10 mm.

Remarks.—This species was collected during both the day and night at Marai Parai and Nunuk camp, and was the most common Gryllidae species encountered. During the day, males can be heard calling on plant foliage. The specimens were compared to the descriptions and images in Tan et al. (2021) and Robillard et al. (2023) for identification.

Subfamily Itarinae

Itara sp.

Fig. 9

Material examined.—EAST MALAYSIA: Sabah • 1♀; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Only one female specimen was collected on the foliage of understory plants at Nunuk camp. It resembles the congeners of Itara by the fore tibiae with open outer but slit-like inner tympanum (Gorochov 1997). Determination of the subgenus and species name was not possible because the only descriptions available are of males.

Fig. 9. 

Itara sp. A. Lateral view; B. Dorsal view. Scale bars: 10 mm.

Subfamily Landrevinae

Repapa trusmadi Gorochov, 2016

Fig. 10

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 16 October 2023; in crevices of rotten log; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was collected within the bark crevices of a rotten log at Nunuk camp. During the night, males can be heard calling from these crevices. Species of this genus are similar to those of the genus Duolandrevus Kirby, 1906 in general appearance and in the presence of outer and inner tympana on the fore tibiae, but R. trusmadi differs in their male genitalia by having a long and rather narrow unpaired posteromedian epiphallic lobule, long posterolateral epiphallic lobes with the distal parts curved upwards, and rather long and semitubular rachis (Gorochov 2016). This represents the first record of this species from Mount Kinabalu, with its type locality Mount Trusmadi situated 60 km south.

Fig. 10. 

Repapa trusmadi. A–C. Habitus in frontal, dorsal, and lateral view; D–F. Male genitalia from dorsal, ventral, and lateral view. Scale bars: 10 mm (B, C); 1 mm (D, E, F).

Family Oecanthidae

Subfamily Podoscirtinae

Aphonoides sp.

Fig. 11

Material examined.—EAST MALAYSIA: Sabah • 1♀; Ranau, Nunuk; 17 October 2023; on tree trunk; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—A female specimen was collected from a tree trunk in the forest near Nunuk camp. It resembles congeners of Aphonoides by the inner tympanum open but outer one obliterated, tegminal Sc with several normal branches, ovipositor with apex rounded, and large teeth on hind and ventral surfaces (Gorochov 2006). Determination of the species name is not possible without males since most species from Borneo were described based only on males. Additionally, many species of this genus cannot be distinguished by morphology alone, although no sexual dimorphism is typically observed.

Fig. 11. 

Aphonoides sp. A, B. Habitus in dorsal and lateral view. Scale bars: 10 mm.

Truljalia parvispinosa Chopard, 1930

Fig. 12

Material examined.—EAST MALAYSIA: Sabah • 1♂ 1♀; Ranau, Nunuk; 17 October 2023; at the wall of a building with a light source; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was attracted to the light from a building at Nunuk camp. The specimens were compared to images in OSF (Cigliano et al. 2024) for identification. Male genitalia resemble the description by Gorochov (2002). Apart from its type locality in Sarawak, this species has also been recorded in Mesilau, which is situated about 10 km south of Mount Kinabalu, Sabah (Gorochov 2002).

Fig. 12. 

Truljalia parvispinosa. A. Male; B. Female; C–E. Male genitalia in dorsal, ventral, and lateral view. Scale bars: 1 mm.

Family Phalangopsidae

Subfamily Phalangopsinae

Anemozara (Anemozara) vera Gorochov, 2014

Fig. 13

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 14 October 2023; on forest floor; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was collected during the day on the forest floor along the trail in the community forest area near Nunuk camp. The male genitalia resemble the description by Gorochov (2014). This represents the first record of this species from Mount Kinabalu; its type locality is Mount Trusmadi, which is 60 km south of Mount Kinabalu, Sabah.

Fig. 13. 

Anemozara (Anemozara) vera. A–C. Habitus in frontal, dorsal, and lateral view; D. Male genitalia in dorsal view. Scale bars: 1 mm.

Family Gryllotalpidae

Subfamily Gryllotalpinae

Tribe Gryllotalpini

Gryllotalpa sp.

Fig. 14

Material examined.—EAST MALAYSIA: Sabah • 1 nymph; Ranau, Nunuk; 16 October 2023; in crevice of rotten log; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Only one individual of this species was collected. Found within the crevice of a rotten log, the specimen was identified as a nymph due to the underdeveloped wings and genitalia. Species identification is difficult without an adult or a calling sound recording.

Fig. 14. 

Gryllotalpa sp. A–C. Habitus in postolateral, dorsal, and lateral view. Scale bars: 10 mm.

Family Rhaphidophoridae

Subfamily Rhaphidophorinae

Tribe Rhaphidophorini

Rhaphidophora cf. curta

Fig. 15

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; in rotten tree stump; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Only one male (BL < 20 mm) was collected inside a rotten tree stump at Nunuk camp. This group is speciose, and it can be hard to identify Southeast Asian representatives (A.V. Gorochov in litt.). The habitus closely resembles that of R. curta Gorochov, 2013 described from Mt. Trus Madi [= Trusmadi], but it differs by the abdominal apex. The specimens were compared to images from OSF (Cigliano et al. 2024) for identification.

Fig. 15. 

Rhaphidophora cf. curta. A, B. Male in dorsal and lateral view; C, D. Male abdominal apex in dorsal and ventral view. Scale bars: 10 mm (A, B); 1 mm (C, D).

Rhaphidophora sp.

Fig. 16

Material examined.—EAST MALAYSIA: Sabah • 1♂ 1♀; Ranau, Nunuk; 17 October 2023; in rotten tree stump; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was collected on a rotten tree stump at Nunuk camp, albeit a different stump than that on which Rhaphidophora cf. curta was found. Habitus and size closely resemble that of R. kinabaluensis (BL = ±30 mm), but both differ by male abdominal apex. This group is speciose, and can be difficult to identify for Southeast Asian representatives (A.V. Gorochov in litt.).

Fig. 16. 

Rhaphidophora sp.2. A, B. Male in dorsal and lateral views respectively; C, D. Male abdominal apex in dorsal and ventral view; E, F. Female in dorsal and lateral view; G. Female subgenital plate. Scale bars: 10 mm (A, B, E, F); 1 mm (C, D, G).

Family Gryllacrididae

Zalarnaca cf. teuthroides

Fig. 17

Material examined.—EAST MALAYSIA: Sabah • 1♂ 1♀; Ranau, Nunuk; 17 October 2023; in rotten tree stump; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Only one male specimen was collected on the foliage of understory plants at Nunuk camp. Genus identification was based on descriptions by Gorochov (2008a) and Ingrisch (2018) and verified by A. V. Gorochov. Species identification follows Karny (1925b). This specimen closely resembles Z. teuthroides Karny, 1925b in body length (17 mm), its tawny brown habitus, fore and middle tibiae with five long spines, apical margin of subgenital plate “W” shaped with short and conical cerci, and ninth abdominal tergite with two strongly incurved projections; it differs in tegmina length not twice as long as the whole body. However, comparison to the type specimen is needed to confirm this species identification (Tan 2014, Tan et al. 2019); thus, until a revision of the genera of Asiatic Gryllacridinae is completed, the determination of the species is problematic (A.V. Gorochov, in litt.).This is the first record of this species at Mount Kinabalu and Sabah. The type locality is Mount Dulit, Sarawak, which is more than 400 km southwest of Mount Kinabalu, Sabah.

Fig. 17. 

Zalarnaca cf. teuthroides. A, B. Habitus in dorsal and frontal view; C. Male abdominal apex in dorso-apical view; D. Subgenital plate in ventral view. Scale bars: 10 mm (A); 1 mm (B, C, D).

Caustogryllacris sp.

Fig. 18

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Kinabalu Park, Marai Parai; 15 October 2023; on tree branch; leg. A.A. Muhammad, Supani T., Jalter M. & Frederick G.; EUSP.

Remarks.—Only one male specimen, found on a tree branch, was collected. The specimen resembles members of the genus Caustogryllacris Karny, 1937 by small habitus (BL = 16 mm) with its wings surpassing hind knees, male ninth abdominal tergite globular with the apical area split in the midline by a fine membrane, and a pair of long oval swellings at the apical margin of the ninth tergite that carries a short tooth at the ventral margin (Ingrisch 2018). Until a revision of the genera of Asiatic Gryllacridinae is completed, the determination of the species is problematic (A.V. Gorochov, in litt.).

Fig. 18. 

Caustogryllacris sp. A, B. Habitus in dorsal and frontal view; C. Subgenital plate in ventral view; D. Male abdominal apex in oblique apical view. Scale bars: 10 mm (A); 1 mm (B, C, D).

Monseremus appendiculatus Ingrisch, 2018

Fig. 19

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T., Jalter M. & Frederick G.; EUSP • 1♀; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was collected at night on the foliage of understory plants at Nunuk camp. The specimens were identified using the dichotomous keys of Ingrisch (2018) and Tan et al. (2022). So far, there are only two species known in this genus, and this species is widespread in Sabah. Apart from its type locality in Poring (16 km east of Mount Kinabalu), this species has also been recorded from Mount Silam (220 km southeast of Mount Kinabalu) (Ingrish 2018, Tan et al. 2022).

Fig. 19. 

Monseremus appendiculatus. A–C. Habitus in dorsal, frontal, and lateral view. Scale bars: 10 mm (A); 1 mm (C).

Family Stenopelmatidae

Subfamily Stenopelmatinae

Tribe Siini

Sia bugajus Gorochov, 2021

Fig. 20

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; on the ground; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—One male specimen was collected on the ground at Nunuk camp. The specimen was identified using the keys by Willemse (1933) and Gorochov (2020). So far, this is the only species of this genus recorded from Borneo. It differs from congeners by the shape of the abdominal apex and the subgenital plate (Gorochov 2022). This represents the first record of this species at Mount Kinabalu; its type locality is Crocker Range National Park near Keningau, which is approximately 40 km south of Mount Kinabalu, Sabah.

Fig. 20. 

Sia bugajus. A–C. Habitus in dorsal, lateral, and frontal view; D. Ventral view of male subgenital plate. Scale bars: 10 mm (A, B); 5 mm (C, D).

Family Tettigoniidae

Subfamily Conocephalinae

Tribe Agraeciini

Eumacroxiphus (Eumacroxiphus) caudatus Ingrisch, 1998

Fig. 21

Material examined.—EAST MALAYSIA: Sabah • 1♂ 1♀; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—The titillators, cerci, and subgenital plate of our specimen resemble that of the type specimen (Ingrisch 1998). This species differs from its congeners by the straight apical margin of the male subgenital plate, cerci with dorsal carina ending in an acute angle, and apex of dorsal lobe triangular with apex rounded. This species is widespread in Mount Kinabalu National Park, described also from Sayap Substation (10 km north of Mount Kinabalu) and Poring (16 km east of Mount Kinabalu) (Cigliano et al. 2024).

Fig. 21. 

Eumacroxiphus (Eumacroxiphus) caudatus. A, D. Habitus in lateral and frontal view; B, C. Male abdominal apex in postolateral and posterior view. Scale bars: 1 mm.

Subfamily Lipotactinae

Lipotactes sp.

Fig. 22

Material examined.—EAST MALAYSIA: Sabah • 1 nymph; Ranau, Nunuk; 17 October 2023; on foliage of understory plants; leg. A.A. Muhammad (photograph).

Remarks.—Only nymphs were spotted on the foliage of understory plants along the trail in the community forest area near Nunuk camp and only during the day. Identification used Gorochov (2021), but identification to species is impossible, as adult specimens are needed for comparison and cryptic species exist among these Lipotactes (Tan et al. 2020).

Fig. 22. 

Lipotactes sp.

Subfamily Meconematinae

Asiophlugis borneoensis Jin, 1993

Fig. 23

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 16 October 2023; on underside of foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was found during the night on the underside of the foliage of the understory plants at Nunuk camp. Individuals were found on separate leaves of the same plant or nearby ones. The specimen was compared with images in Gorochov and Tan (2011) and Kevan and Jin (1993). This species was first described from Bundu Tuhan, which is close to the sampling area.

Fig. 23. 

Asiophlugis borneoensis. A, B. Habitus in dorsal and lateral view; C–E. Male abdominal apex in lateral, dorsal, and ventral view. Scale bars: 5 mm (A, B); 1 mm (C, D, E).

Asiophlugis nr. trusmadi

Fig. 24

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 16 October 2023; on underside of foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—Similar to A. borneoensis Jin, 1993, this species was found during nighttime on the underside of foliage of the understory plants at Nunuk camp. This species is similar to A. trusmadi, which has been described from Mt. Trus Madi [= Trusmadi], but differs slightly by the shape of the styli at the base of the subgenital plate. Our specimen may represent an undescribed species, but more specimens are needed for a better understanding of the intraspecific variation. The specimen was compared with images in Gorochov and Tan (2011). The type locality of A. trusmadi is Mount Trusmadi, which is situated 60 km south of Mount Kinabalu, Sabah.

Fig. 24. 

Asiophlugis nr. trusmadi. A, B. Habitus in dorsal and lateral views respectively; C, D. Male abdominal apex in ventral and lateral view. Scale bars: 10 mm (A, B); 1 mm (C, D).

Odonturisca dentata Gorochov, 2022

Fig. 25

Material examined.—EAST MALAYSIA: Sabah • 1♀; Ranau, Nunuk; 17 October 2023; at the wall of a building with a light source; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was attracted to a light source at a building at Nunuk camp. It differs from its congeners by the low and rounded humeral notches of the pronotum and the shape of the female subgenital plate and ovipositor (Gorochov 2022). The type specimen was also described from Mount Kinabalu National Park (Gorochov 2022).

Fig. 25. 

Odonturisca dentata.

Subtilodecma bilobata Gorochov, 2022

Fig. 26

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Kinabalu Park, Marai Parai; 14 October 2023; on underside of foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species was collected during the night on the underside of foliage of the understory plants at Nunuk camp. It can be found co-occurring with Asiophlugis species. There are only two species inthis genus, and both were recorded at Mount Kinabalu National Park (Gorochov 2022), the other being Subtilodecma unilobata Gorochov, 2022. This species differs from its congener by the male cerci being bilobate instead of unilobate apically, the distal portion of the male genital plate narrower, and the distal parts of the male tegmina somewhat shorter (Gorochov 2022).

Fig. 26. 

Subtilodecma bilobata. A, B. Habitus in dorsal and lateral view; C, D. Male abdominal apex in lateral and ventral view. Scale bars: 10 mm (A, B); 0.5 mm (C, D).

Subfamily Phaneropterinae

Mirollia (Hemimirollia) gracilis Karny, 1925

Fig. 27

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; at the wall of a building with light source; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP • 1♂ ; Ranau, Nunuk; 17 October 2023; on foliage of understory plant; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This species is identified as H. gracilis based on the details of the male epiproct, cerci, and subgenital plate: male phallus without large sclerites, at most with small, semi-solid, in situ hidden structures, and the epiproct widening toward apex (Ingrisch 2011). The specimens were collected during nighttime, as they were attracted to a light source on a building at Nunuk camp and also on foliage of an understory plant. Apart from the type locality on Mount Dulit, Sarawak (Karny 1925a), this species has been recorded at Mount Trus Madi [=Trusmadi] (Gorochov 2008b), Sorinsim (Gorochov 2023b), and Kuala Belalong, Brunei Darussalam (Tan and Wahab 2018).

Fig. 27. 

Mirollia (Hemimirollia) gracilis. A, B. Habitus in lateral and frontal view; C, D. Male abdominal apex in ventral and lateral view. Scale bars: 10 mm (A); 1 mm (B, C, D).

Sympaestria acutelobata Brunner von Wattenwyl, 1878

Fig. 28

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 17 October 2023; at the wall of a building with light source; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This specimen is identified as S. acutelobata based on the details of the stridulatory area (shape of the mirror on the right tegmen), although the male abdominal apex is similar (Tan 2014, Tan et al. 2019, 2024a). More material is needed to confirm the identification of this species from this region such that further taxonomic treatment of this genus can be done (Tan 2014, Tan et al. 2019). This species was collected during the night, as it was attracted to a light source on a building at Nunuk camp.

Fig. 28. 

Sympaestria acutelobata. A, B. Habitus in lateral and frontal view; C, D. Male abdominal apex in ventral and ventrolateral view. Scale bars: 10 mm (A); 1 mm (B, C, D).

Tapiena sp.

Fig. 29

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Nunuk; 16 October 2023; on foliage of understory plants; leg. A.A. Muhammad, Supani T. & Jalter M.; EUSP.

Remarks.—This specimen is a member of the genus Tapiena Bolívar, 1906 having a rugose pronotum. The male abdominal apex closely resembles that of T. incisa Karny, 1923 and T. bullata Karny, 1923, but differs by the broad and flat unilobed cerci with serrate apical margin (distinctively bilobate apically with obtuse inner tooth in T. incisa), and subgenital plate with posterior margin more deeply excised in the middle (not excised in T. bullata) (Tan et al. 2015). It appears most similar to Tapiena ?incisa (sensu Tan et al. 2024a) collected from Mount Trus Madi. More material is needed to confirm this species identification in this region. This species was collected during the night on the foliage of understory plants at Nunuk camp.

Fig. 29. 

Tapiena sp. A, B. Habitus in lateral and frontal view; C, D. Male abdominal apex in ventral and ventrolateral view. Scale bars: 10 mm (A); 1 mm (B, C, D).

Subfamily Pseudophyllinae

Tribe Phyllomimini

Cratioma cf. borneense

Fig. 30

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Kinabalu Park, Marai Parai; 15 October 2023; on stem of a plant; leg. A.A. Muhammad, Supani T., Jalter M. & Frederick G.; EUSP.

Remarks.—Only two species of this genus were recorded in Sabah, with the other being Cratioma oculatum Karny, 1926. This species is similar to C. borneense, which has been described from Kina Balu [=Kinabalu] (Beier 1954), by the absence of a circular pattern on the tegmen and denser wing venation than in C. oculatum (Tan et al. 2019). The specimens were compared to images from OSF (Cigliano et al. 2024) for identification.

Fig. 30. 

Cratioma cf. borneense.

Despoina spinosa Brunner von Wattenwyl, 1895

Fig. 31

Material examined.—EAST MALAYSIA: Sabah • 1♂; Ranau, Kinabalu Park, Marai Parai; 14 October 2023; on stem of Tetraria pilisepala; leg. A.A. Muhammad, Supani T., Jalter M. & Silver P.; EUSP.

Remarks.—Only one male specimen was collected from a grassy patch at Marai Parai. This specimen is identified as D. spinosa based on spines on its pronotum, brownish-purple-dotted elytra, and the apex of its hind femora black. Beier (1954) assigned the type specimen collected from Kina-Balu [=Kinabalu] to D. spinosa atrata Beier, 1954, but subspecies identification is impossible due to a lack of material. Comparison with the type specimen of the other subspecies, D. spinosa spinosa Brunner von Wattenwyl, 1895 from Brunei, is also needed.

Fig. 31. 

Despoina spinosa. A–C. Habitus in dorsal, ventral, and lateral view. Scale bars: 10 mm.

Promeca cf. nobilis

Fig. 32

Material examined.—EAST MALAYSIA: Sabah • 1♀; Ranau, Nunuk; 13 October 2023; on foliage of understory plants; leg. Paul Y.I. & Evan Q.S.H.; EUSP.

Remarks.—Only one female was collected during the expedition. This specimen closely resembles P. nobilis, but differs in having dark-colored sternites and the shape of the subgenital plate, possibly due to the artifact of preservation. The specimen was compared with type images from OSF (Cigliano et al. 2024). The type locality of this species is North Borneo (“Nord-Borneo, Waterstradt”), which refers to Sabah state, although the specific locality remains undetermined (Beier 1954, Liana 1999).

Fig. 32. 

Promeca cf. nobilis. A. Habitus in lateral view; B. Ovipositor in lateral view; C. Subgenital plate in ventral view. Scale bars: 10 mm (A, B); 1 mm (C).

Discussion

The orthopteran diversity in and around Mount Kinabalu is not fully understood, although previous expeditions have been conducted there. Very recently, species and genera of katydids and crickets new to science were discovered. These include Holochlorini from Mount Trus Madi (Gorochov 2023a, Tan et al. 2024a); Mirolliini katydids from Mount Trus Madi and Crocker Range (Gorochov 2023b, Tan et al. 2024a); species of Itara and Odontogryllodes crickets and Depressacca katydids, each from Tenompok (Tan et al. 2024b, 2024c, 2024d); and Pendleburyella crickets from Mount Trus Madi and Bukit Hampuan (Tan et al. 2024e). This suggests that continued sampling in other less-explored regions of these vast mountain ranges is needed.

The call for further sampling is corroborated by the new locality records and potential new species found in the five-day sampling effort in Marai Parai and Nunuk presented here. Mount Kinabalu has several types of landscapes at various altitudes, which results in disparate vegetation (Kitayama 1992, Aiba and Kitayama 1999). The difference in habitat composition may influence the orthopteran species assemblages and spatial distribution (Joern 1982, Guido and Gianelle 2001, Ibanez et al. 2013), as reflected in this study in which only one overlapping species was present at both locations. Hence, habitat heterogeneity across this area signifies that more species are likely to be found, especially niche-specific species.

We also believe that many more undescribed species are still awaiting discovery in larger parts of the hyper-diverse Sabah and even Borneo. Mount Kinabalu region has historically been considered one of the more sampled regions in Sabah and Borneo (Tan et al. 2017). It is not a surprise, therefore, that we know even less about the orthopteran diversity in other parts of Sabah and Borneo. Beyond taxonomy, we also know little about the calling songs and natural history of orthopterans in this region (Tan et al. 2023), with most species described only based on morphology. Hence, there is a dearth of knowledge on orthopterans in this region, yet the biodiversity of Borneo is significantly impacted by the negative anthropogenic activities that threaten the existence of these orthopteran species (Miettinen et al. 2011, Ocampo-Peñuela et al. 2020).

Acknowledgments

The authors are grateful to the Board of Trustees of the Sabah Parks, Sabah, Malaysia for organizing the Marai Parai—West Gurkha Hut Kota Belud Scientific Expedition in October 2023. The authors thank Luc Willemse, Andrei V. Gorochov, Josip Skejo, Karmela Adžić, and Maks Deranja for their help in identification and verification of the specimens and for sharing specific references. Many thanks to everyone else involved in the expedition, including Supani Tuboh, Jalter Musin, Veronica Jumil, Geofarry Gunsalam, Paul Yambun Imbun, Assoc. Prof. Dr. Evan Quah Seng Huat, Silver Peter, Johan @ Frederick Gansau, Wahimah Sulaiman, Sinail Dunsul, Martinah Latim, and others for field assistance, necessary equipment, advice, and other valuable support.

References

  • Aiba SI, Kitayama K (1999) Structure, composition and species diversity in an altitude-substrate matrix of rain forest tree communities on Mount Kinabalu, Borneo. Plant Ecology 140: 139–157. https://doi.org/10.1023/A:1009710618040
  • Barkman TJ, Repin R, Sugau JB (2016) The parasitic plant families Loranthaceae and Viscaceae in Sabah, Malaysia. Sandakania 131: 131–169.
  • Beaman JH, Beaman RS (1998) The plants of Mount Kinabalu, 3: gymnosperms and non-orchid mocotyledons. Kota Kinabalu: Natural History Publications (Borneo) in association with Royal Botanic Gardens Kew.
  • Beier M (1954) In Revision der Pseudophyllinen. Instituto Español de Entomología, Madrid, 479 pp.
  • Bolívar I (1906) Fasgonurídeos de la Guinea Española. Memorias de la Real Sociedad Española de Historia Natural 1: 327–377.
  • Brunner von Wattenwyl C (1895) Monographie der Pseudophylliden. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 45: 1–282.
  • Cooke FM, Hussin R (2014) Biodiversity conservation and its social implications: The case of indigenous and community conserved areas in Sabah, Malaysia. Suvannabhumi 6: 3–18.
  • Cross AT, van der Ent A, Wickmann M, Skates LM, Sumail S, Gebauer G, Robinson A (2022) Capture of mammal excreta by Nepenthes is an effective heterotrophic nutrition strategy. Annals of botany 130: 927–938. https://doi.org/10.1093/aob/mcac134
  • Gorochov AV (1997) Partial revision of the subfamily Itarinae (Orthoptera: Gryllidae). Zoosystematica Rossica 6: 47–75.
  • Gorochov AV (2002) Taxonomy of Podoscirtinae (Orthoptera: Gryllidae). Part 1: The male genitalia and Indo-Malayan Podoscirtini. Zoosystematica Rossica 10(2): 303–350. https://doi.org/10.31610/zsr/2001.10.2.303
  • Gorochov AV (2008a) Contributions to the fauna and systematics of the Stenopelmatoidea (Orthoptera) of Indochina and some other territories: VIII. Entomological Review 88: 406–420. https://doi.org/10.1134/S0013873808040040
  • Gorochov AV (2008b) New and little known species of the tribe Mirolliini (Orthoptera, Tettigoniidae, Phaneropterinae) from South-East Asia [in Russian]. Evraziatskii Entomologicheskii Zhurnal 7: 311–321. https://doi.org/10.31610/trudyzin/2008.312.1-2.26
  • Gorochov AV (2014) Classification of the Phalangopsinae subfamily group, and new taxa from the subfamilies Phalangopsinae and Phaloriinae (Orthoptera: Gryllidae). Zoosystematica Rossica 23: 7–88. https://doi.org/10.31610/zsr/2014.23.1.7
  • Gorochov AV (2020) The families Stenopelmatidae and Anostostomatidae (Orthoptera). 1. Higher classification, new and little known taxa. Entomological Review 100: 1106–1151. https://doi.org/10.1134/S0013873820080084
  • Gorochov AV (2021) Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 14. Far Eastern Entomologist 434: 1–25. https://doi.org/10.25221/fee.434.1
  • Gorochov AV (2022) Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 15. Far Eastern Entomologist 459: 1–26. https://doi.org/10.25221/fee.459.1
  • Gorochov AV (2023a) New and little known taxa of the genus Calopsyra Brunner von Wattenwyl, 1891 (Orthoptera: Tettigoniidae: Phaneropterinae) from Indo-Malayan Region. Caucasian Entomological Bulletin 19: 269–275. https://doi.org/10.5281/zenodo.8399608
  • Gorochov AV (2023b) New and little known taxa of the tribe Mirolliini (Orthoptera: Tettigoniidae: Phaneropterinae) from Indo-Malayan region. Proceedings of the Russian Entomological Society, St Petersburg 94: 40–72.
  • Gorochov AV, Tan MK (2011) New katydids of the genus Asiophlugis Gor. (Orthoptera: Tettigoniidae: Meconematinae) from Singapore and Malaysia. Russian Entomological Journal 20: 129–133. https://doi.org/10.15298/rusentj.20.2.03
  • Ibanez S, Manneville O, Miquel C, Taberlet P, Valentini A, Aubert S, Coissac E, Colace M, Duparc Q, Lavorel S, Moretti M (2013) Plant functional traits reveal the relative contribution of habitat and food preferences to the diet of grasshoppers. Oecologia 173: 1459–1470. https://doi.org/10.1007/s00442-013-2738-0
  • Ingrisch S (1998) Monograph of the Oriental Agraeciini (Insecta, Ensifera, Tettigoniidae): Taxonomic revision, phylogeny, biogeography, stridulation and development. Courier Forschungsinstitut Senckenberg 206: 1–391.
  • Ingrisch S (2011) New taxa of Mirolliini from South East Asia and evidence for an abdominal gland in male Phaneropterinae (Orthoptera: Tettigoniidae). Zootaxa 2943: 1–44. https://doi.org/10.11646/zootaxa.2943.1.1
  • Ingrisch S (2018) New taxa and records of Gryllacrididae (Orthoptera, Stenopelmatoidea) from South East Asia and New Guinea with a key to the genera. Zootaxa 4510: 1–278. https://doi.org/10.11646/zootaxa.4510.1.1
  • Ingrisch S (2020) New subgenera and species of Agraeciini (Orthoptera, Tettigoniidae, Conocephalinae) from South Asia found in historical insect collections. Evolutionary Systematics 4: 119–132. https://doi.org/10.3897/evolsyst.4.60525
  • Karny HH (1923) On Malaysian katydids (Gryllacridae and Tettigoniidae) from the Raffles Museum, Singapore. Journal of the Malaysian Branch of the Royal Asiatic Society 1: 117–193.
  • Karny HH (1925a) List of some katydids (Tettigoniidae) in the Sarawak Museum. The Sarawak Museum Journal 3: 35–53.
  • Karny HH (1925b) On some cricket-locusts (Gryllacrididae) from Mt. Dulit and Mt. Murud, Sarawak. The Sarawak Museum Journal 3: 54–62.
  • Karny HH (1926) On Malaysian katydids. Journal of the Federated Malay States Museums 13(2–3): 69–157.
  • Kevan DKME, Jin X (1993) Remarks on the tribe Phlugidini Eichler and recognition of new taxa from the Indo-Malayan region and East Africa (Grylloptera: Tettigonioidea: Meconematidae). Invertebrate Taxonomy 7: 1589–1610. https://doi.org/10.1071/IT9931589
  • Liana A (1999) The type material of Pseudophyllidae (Orthoptera) in the Museum and Institute of Zoology PAS, Warsaw. Bulletin of the Museum and Institute of Zoology PAS 2: 45–62.
  • Ocampo-Peñuela N, Garcia-Ulloa J, Kornecki I, Philipson CD, Ghazoul J (2020) Impacts of four decades of forest loss on vertebrate functional habitat on Borneo. Frontiers in Forests and Global Change 3: 1–13. https://doi.org/10.3389/ffgc.2020.00053
  • Skejo J, Pushkar TI, Kasalo N, Pavlović M, Deranja M, Adžić K, Tan MK, Rebrina F, Muhammad AA, Abdullah NA, Japir R, Chung AYC, Tumbrink J (2022) Spiky pygmy devils: revision of the genus Discotettix (Orthoptera: Tetrigidae) and synonymy of Discotettiginae with Scelimeninae. Zootaxa 5217: 1–64. https://doi.org/10.11646/zootaxa.5217.1.1
  • Song H, Béthoux O, Shin S, Donath A, Letsch H, Liu S, McKenna DD, Meng G, Misof B, Podsiadlowski L, Zhou X, Wipfler B, Simon S (2020) Phylogenomic analysis sheds light on the evolutionary pathways towards acoustic communication in Orthoptera. Nature Communications 4939: 1–16. https://doi.org/10.1038/s41467-020-18739-4
  • Tan MK (2014) An annotated checklist of the bush katydids (Orthoptera: Phaneropteridae: Phaneropterinae) from Singapore, including an illustrated key to species. Zootaxa 3884: 573–593. https://doi.org/10.11646/zootaxa.3884.6.6
  • Tan MK, Duncan J, Wahab RA, Lee CY, Japir R, Chung AYC, Baroga-Barbecho JB, Yap SA, Montealegre-Z F (2023) The calling songs of some katydids (Orthoptera, Tettigonioidea) from the tropical forests of Southeast Asia. Journal of Orthoptera Research 32: 1–24. https://doi.org/10.3897/jor.32.84563
  • Tan MK, Grumo KC, Gono AA, Bahoy DCM, Rivera RR, Nuñeza OM, Japir R, Chung AYC (2024b) New species of Itara (Orthoptera: Grylloidea: Itarinae) and descriptions of calling songs from Mindanao and Sabah. Zootaxa 5424: 61–79. https://doi.org/10.11646/zootaxa.5424.1.3
  • Tan MK, Ingrisch S, Wahab RA, Japir R, Chung AYC (2020) Ultrasonic bioacoustics and stridulum morphology reveal cryptic species among Lipotactes big-eyed katydids (Orthoptera: Tettigoniidae: Lipotactinae) from Borneo. Systematics and Biodiversity 18: 510–524. https://doi.org/10.1080/14772000.2020.1769223
  • Tan MK, Japir R, Chung AYC (2019) Uncovering the Grylloidea and Tettigonioidea (Orthoptera: Ensifera) in the Forest Research Center (Sepilok) Entomological Collection. Zootaxa 4701: 301–349. https://doi.org/10.11646/zootaxa.4701.4.1
  • Tan MK, Japir R, Chung AYC (2024c) An account on some katydids of the tribes Agraeciini (Conocephalinae) and Meconematini (Meconematinae) from the highlands of western Sabah. Zootaxa 5419: 584–600. https://doi.org/10.11646/zootaxa.5419.4.6
  • Tan MK, Japir R, Chung AYC (2024e) New species of the elusive crickets from the genus Pendleburyella Chopard, 1969 (Gryllidae, Pentacentrinae) from Sabah, Borneo. Zootaxa 5397: 264–272. https://doi.org/10.11646/zootaxa.5397.2.7
  • Tan MK, Japir R, Chung AYC, Robillard T (2022) A review of the genus Monseremus Ingrisch, 2018 (Orthoptera, Stenopelmatoidea, Gryllacrididae) from Borneo. Zootaxa: 5165: 107–114. https://doi.org/10.11646/zootaxa.5165.1.5
  • Tan MK, Kamaruddin KN (2014) Orthoptera of Fraser’s Hill, Peninsular Malaysia. Lee Kong Chian Natural History Museum, National University of Singapore, Singapore, 88 pp.
  • Tan MK, Liu CX, Artchawakom T (2015) Taxonomic review of Tapiena (Orthoptera: Phaneropteridae: Phaneropterinae), with key to species and new species from Thailand. Zootaxa 920: 40–50. https://doi.org/10.11646/zootaxa.3920.1.2
  • Tan MK, Liu CX, Ingrisch S, Japir R, Chung AYC (2024a) An account on the Phaneropterinae (Tettigonioidea: Tettigoniidae) from the highlands of western Sabah. Zootaxa 5443: 451–494. https://doi.org/10.11646/zootaxa.5443.4.1
  • Tan MK, Muhammad AA, Abdullah NA, Japir R, Chung AYC (2024d) A taxonomic review of Odontogryllodes Chopard, 1969 (Gryllidae: Landrevinae). Zootaxa 5410: 267–279. https://doi.org/10.11646/zootaxa.5410.2.7
  • Tan MK, Wahab RA (2017) New taxa and notes on crickets of the Subfamily Landrevinae (Orthoptera: Gryllidae) from Brunei Darussalam, Borneo. Zootaxa 4365: 440–454. https://doi.org/10.11646/zootaxa.4365.4.4
  • 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: 119–142. https://doi.org/10.3897/jor.27.24152
  • Tan MK, Wahab RHA, Japir R, Chung AY, Robillard T (2021) Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species. European Journal of Taxonomy 761: 1–75. https://doi.org/10.5852/ejt.2021.761.1449
  • Tumbrinck J (2014) Taxonomic revision of the Cladonotinae (Orthoptera: Tetrigidae) from the islands of South-East Asia and from Australia, with general remarks to the classification and morphology of the Tetrigidae and descriptions of new genera and species from New Guinea and New Caledonia. In: Telnov D (Ed.) Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea 2, 350–396.
  • Van der Ent A, Repin R, Sugau J, Wong KM (2015) Plant diversity and ecology of ultramafic outcrops in Sabah (Malaysia). Australian Journal of Botany 63: 204–215. https://doi.org/10.1071/BT14214
  • Willemse CJM (1921) Bijdrage tot de kennis der Orthoptera s. s. van den nederlandsch indischen Archipel en omliggende gebieden. Zoologische Mededeelingen 6: 1–44.
  • Willemse CJM (1930) Fauna Sumatrensis (Bijdrage Nr. 62) Preliminary revision of the Acrididae (Orthoptera). Tijdschrift voor Entomologie 73: 1–210.
  • Willemse CJM (1935) Description of some new Traulia sp., with key to the known species (Orthopt. Acrididae). Entomologische Berichten 9: 137–143.
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