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Corresponding author: Ming Kai Tan ( orthoptera.mingkai@gmail.com ) Academic editor: Juliana Chamorro-Rengifo
© 2018 Ming Kai Tan, Rodzay bin Haji Abdul Wahab.
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:
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
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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 Belalong Field Studies Centre (KBFSC) within the Ulu Temburong National Park. We present here an annotated and illustrated checklist of Orthoptera 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 monophyletic 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. Despite 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.
checklist, diversity, natural history, Southeast Asia, species richness, taxonomy
Orthoptera is an order of insects consisting of grasshoppers (suborder Caelifera), and crickets and katydids (suborder Ensifera). These are among the most diverse (around 28,000 species worldwide) and common terrestrial macro-invertebrates (
Southeast Asia is made up of numerous biodiversity hotspots (
On the island of Borneo, pockets of forests remained unstudied for orthopterans. This includes Brunei Darussalam, particularly in the Temburong District (Fig.
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 published separately in taxonomic journals (
Surveys were conducted by the first author (MKT) in the primary lowland and ridge dipterocarp forests (Fig.
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.
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 accomplished using Adobe Photoshop CC 2014. Scales were given with the images. Acoustic analysis was done using the open source R package WarbleR (
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 identities of the orthopteran specimens were verified by orthopterists: Andrej V. Gorochov (for Grylloidea), Sigfrid Ingrisch (for Tettigonioidea), 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).
Classification and nomenclature of species were based on the Orthoptera Species File (OSF) Online Version 5.0/5.0 (
Overall, 72 species representing nine families were collected between 2016 and 2017 (Table
A summary of the coverage of orthopteran superfamilies and families that were represented in the collection in KBFSC.
Ranking | Nomenclature | Species Richness | |
---|---|---|---|
1. | Order | Orthoptera | |
1.1. | Suborder | Caelifera | |
1.1.1. | Superfamily | Acridoidea | 5 |
1.1.1.1. | Family | Acrididae | |
1.1.2. | Superfamily | Tetrigoidea | 13 |
1.1.2.1. | Family | Tetrigidae | |
1.1.3. | Superfamily | Trigonopterygoidea | 1 |
1.1.3.1. | Family | Trigonopterygidae | |
1.2. | Suborder | Ensifera | |
1.2.1. | Superfamily | Grylloidea | 21 |
1.2.1.1. | Family | Gryllidae | |
1.2.1.2. | Family | Mogoplistidae | |
1.2.2. | Superfamily | Gryllotalpoidea | 1 |
1.2.2.1. | Family | Gryllotalpidae | |
1.2.3. | Superfamily | Rhaphidophoroidea | 1 |
1.2.3.1. | Family | Rhaphidophoridae | |
1.2.4. | Superfamily | Stenopelmatoidea | 5 |
1.2.4.1. | Family | Gryllacrididae | |
1.2.5. | Superfamily | Tettigonioidea | 25 |
1.2.5.1. | Family | Tettigoniidae |
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
We tentatively identified this species to be near to P. borneensis 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. Interestingly, 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
A female specimen was identified using the generic key by
This species was found near the forest edge. It was identified based on the description of the color in
The two Traulia species were found co-occurring in similar habitat, although this species can also be found in more forested areas of the Canopy Walk. This species was identified based on the description of the color in
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
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
We found this pygmy grasshopper on dead logs and tree trunks along Ashton Trail. We used the key by
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 (
A single specimen (a female) collected from the boardwalk 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.
Unlike most ground-dwelling pygmy grasshoppers, this species was found on foliage of understory plants. This species was identified by J. Tumbrinck.
These pygmy grasshoppers were collected near the sandy bank of the Sungai Belalong at KBFSC. We compared our specimens with type images from OSF (
A single female specimen was found attracted to light. We compared our specimen with type images from OSF (
This pygmy grasshopper was found near small drains near the forest edge at KBFSC. The species was identified by J. Tumbrinck.
This pygmy grasshopper was also found near small drains near the forest edge at KBFSC. The species was identified by J. Tumbrinck.
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 specimen with type images from OSF (
We compared our single female specimen with type images from OSF (
This pygmy grasshopper was found near small drains near the forest edge at KBFSC, probably because they are hydrophilic. The species was identified by J. Tumbrinck.
These crickets are sometimes encountered among vegetation in the understory of the dipterocarp forest. The nymphs can sometimes be found on the forest floor. We compared our specimens with images from
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 (
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 frequency = 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 Belalong resemble the images and description by
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.
This rare and beautiful black cricket was encountered only once on the forest floor of the dipterocarp forest. A.V. Gorochov assisted with the identification of this species and this could represent an undescribed species (see
This species is one of the two species of the cosmopolitan genus Velarifictorus Randell, 1964 (Modicogryllini) found in Kuala Belalong. Male genitalia resemble that of the species illustrated in
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
Prior to the description, we could only tentatively identify the male specimen to be close to D. luzonensis (
One of the two Landrevinae newly described from the collection; this cricket can be commonly heard after dusk in the dipterocarp forest, often while hiding among dead leaves hanging on branches. They were also observed to feed on dead leaves. Refer to
One of the two Landrevinae newly described from the collection; this cricket was found among branches of trees in the dipterocarp forest. Refer to
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 (
Our female specimen may belong to Sonotrella (Megatrella) 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.
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 (
14. Amusurgus or Metiochodes sp.?
Fig.
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.
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.
The male can be distinguished from the other Trigonidiinae above by the developed stridulatory apparatus, and foretib>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.
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 (
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 images from
The males produce trilling calling songs after dusk, often 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 (
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: 1mm.
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 (
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 species was used for identification but this genus is likely to be paraphyletic and taxonomic work is needed (
We recorded the calling songs heard in the evening sporadically across the forest floor, even though we did not manage 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.
This group is speciose and can be hard to identify for Southeast Asian representatives (A.V. Gorochov, in litt.).
Our single specimen (a male) resembles the drawing of the habitus of A. genualis (Walker, 1869) from Borneo. The abdominal apex (particularly the medial apical sclerotized processes of ninth abdominal tergite) also resembles that in
We tentatively identified our female specimen to be close to C. superba (Brunner von Wattenwyl, 1888) using an old key (
This is among the most common Stenopelmatoidea that was found in the understory of the Kuala Belalong forest. Unfortunately, we could only tentatively identify our specimens to this genus using an old key (
Gryllacrididae: A.Asarcogryllacris (Pseudolarnaca) cf. genualis male; B. Face; C. Dorsal view of head and pronotum; D, E. Abdominal apex; F.Capnogryllacris (Capnogryllacris) cf. superba; G. Melaneremus (?) sp. mating pair; H. Male adult; I. Male stridulatory file; J. Male subgenital plate. Scale bars: 1mm.
This stunningly red raspy cricket was found among foliage 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 similar and was described from Sarawak close to Brunei Darussalam.
We tentatively identified our specimens to this genus using an old key (
This species was found on the foliage of understory plants in the dipterocarp forest. We used key to genera of Agraeciini by
This katydid is not uncommon in the understory of Kuala Belalong forest. However, curiously, we have yet to encounter the males. Unfortunately, the male of this species has also not been described. Our females correspond to the description by
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 Kästner, 1932 from northern Sumatra (
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 (
This species was encountered on foliage of the dipterocarp forest understory at night. Refer to
This species was encountered on foliage of the dipterocarp forest understory at night. Refer to
We tentatively identified this species to be close to either N. longipennis Jin, 1992 or N. curvata Jin, 1992. This species was found fairly frequently on foliage of the dipterocarp forest understory at night.
This species was quite frequently encountered on foliage of the dipterocarp forest understory at night. The peculiar shape of the abdominal apex led us to consider this to be a completely different genus. Refer to
Refer to
This large katydid can be heard calling at night among shrubby bushes. The calling song resembles the train call (see
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 Baryprostha from
This species was found attracted to light, thus appeared around the station. We compared our male specimens with images of type specimens from OSF (
Only one male specimen was collected during light trapping. We used the key to genera of Mirolliini by
One male specimen was collected during light trapping. Upon freezing the katydid, a horsehair worm (Nematomorpha) emerged from the abdominal apex. This parasite is not uncommon 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 abdominal tergite with two apical lobes, area around left stridulatory file strongly swollen (
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
This species was found attracted to light and we postulate 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 (
We compared our male specimen with the original description (
We tentatively identified the species to be close to P. marginata Brunner von Wattenwyl, 1878. One female was collected 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. marginata 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.
We tentatively identified this species to be similar to R. profunda Liu & Kang, 2007. We compared our single specimen (a male) with the descriptions of all species of Rectimarginalis Liu & Kang, 2007 (see
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
This species is among the more common species of orthopterans attracted to light. They are, however, not encountered during opportunistic surveys in the understory, which is indicative of this being a canopy dweller. We used original descriptions (
Our male specimen, collected during light trapping, resembles Tapiena (
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 costal margin of the tegmina distinguish our specimen from allied genera from the region (such as Sathrophyllia Stål, 1874 and Tegra Walker, 1870) (
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 epiphylls that were found commonly on the leaves of the trees in the forest around KBFSC (notice the leaf of Fig.
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.
The authors are grateful to Andrej V. Gorochov (for Grylloidea), Sigfrid Ingrisch (for Tettigonioidea), Tony Robillard (for Eneopterinae and Phalangopsidae), Luc Willemse (for Acridoidea), Josip Skejo (for Tetrigoidea), Josef Tumbrinck (for Tetrigoidea), 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]). The authors are grateful to the Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam, for allowing us to work in Kuala Belalong Field Studies Centre, and to the Biodiversity and Research Innovation Centre (BioRIC), Ministry of Primary Resources and Tourism, Brunei Darussalam for the issuance of our export permits (BioRIC/HoB/TAD/51-73, 51-80, and 51-90). The work of MKT was supported by the Lady Yuen Peng McNeice Graduate Fellowship of the National University of Singapore.