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Taxonomy of the genus Longipeditermes Holmgren (Termitidae, Nasutitermitinae) from the Greater Sundas, Southeast Asia
expand article infoSyaukani Syaukani, Graham J Thompson§, Takeshi Yamasaki|, Ahmad Sofiman Othman, Samsul Muarrif, Muhammad Ali Sarong#, Djufri Djufri#, Katsuyuki Eguchi|
‡ Universitas Syiah Kuala, Banda Aceh, Indonesia
§ Western University, London, Ontario, Canada
| Tokyo Metropolitan University, Tokyo, Japan
¶ Universiti Sains Malaysia, Penang, Malaysia
# Universitas Syiah Kuala , Banda Aceh, Indonesia
Open Access

Abstract

More than 200 colonies of the genus Longipeditermes were collected in our field surveys across the Sundaland region of Southeast Asia from 1998 to 2014. Two species, L. kistneri Akhtar & Ahmad and L. logipes Holmgren, are recognized and redescribed with color photographs of the workers and major soldiers. We use variation in characters of soldier caste (head capsules, antennae, and pronotum) and worker caste (antennae and mandibles) to distinguish these two species. Longipeditermes kistneri seems to prefer high-altitude forests (above 1,000 m) and has so far been found exclusively in Java and Sumatra, while L. logipes seems to prefer lowland and swamp forests and is widespread in the Greater Sundas.

Key Words

Morphological characters, Nasutitermitinae, Open-air processional columns termites, species description

Introduction

A termite colony typically consists of a large number of workers and defensive soldiers together with a single king and a queen. This arrangement can, however, vary with seasonal cohorts of reproductive nymphs and dispersive alates (Watson and Gay 1991; Pearce 1997). The castes within termite societies are morphologically and behaviorally specialized to perform particular tasks (Lee and Wood 1971; Grimaldi and Engel 2005) to the point where workers and soldiers are so specialized for non-reproductive helping and defensive tasks that they are considered to be functionally sterile (Roisin 2000). The sexuals, by contrast, are highly specialized for reproduction.

As the largest subfamily among the higher termites (Termitidae), Nasutitermitinae consists of more than 550 species (Emerson 1955), which belong to more than 63 genera (Collins 1989). This subfamily probably originated in the Neotropical region during the Cretaceous period (Emerson1955) and is characterized by a highly specialized defensive caste (Collins 1989) that has a modified rostrum or “nasus” (Deligne et al. 1981; Prestwich 1984). This subfamily was erected by Hare (1937), who emphasized the conspicuous prolongation on the front of the head, which is often accompanied by degeneration of mandibles and the concomitant development of a frontal gland. The development of the nasus is continuously variable among species within the Nasutitermitinae from mild prolongation, as in Syntermes, to pronounced prolongation as in Subulitermes, Convexitermes, and Nasutitermes.

Longipeditermes, in the Nasutitermitinae, is a small genus with only two species so far recorded. These are L. longipes and L. kistneri from the Sundaland area (Gathorne-Hardy 2001; Hoare and Jones 1998). Longipeditermes is, however, well known for its remarkable foraging habit. Workers and soldiers form open-air processional columns on the ground, shrubs, and tree trunks typically inside forests (Gray and Dhanarajan 1974; Hoare and Jones 1998; Miura and Matsumoto 1998; Takematsu et al. 2013; Syaukani et al. 2016). The foraging activities of both species seem to be greatest during the day (Hoare and Jones 1998), as evidenced by workers carrying packed balls of organic matter between their mandibles and returning to the nest.

In the course of our long-term inventory and taxonomic research on termites in Southeast Asia, more than 200 colonies of Longipeditermnes have been sampled from different habitats and altitudes across the Greater Sunda. In this paper, the two species of the genus are redescribed and illustrated using newly obtained nest series. Information on their life history is provided.

Materials and methods

We examined 216 colonies of Longipeditermes from various habitats and altitudes across the Greater Sunda (Table 1; Fig. 1) . The specimens examined are deposited at the Museum Zoologicum Bogoriense (MZB), Cibinong, Indonesia; the Natural History Museum, London (UK); and the Universitas Syiah Kuala (Biology Department), Darussalam, Banda Aceh (Indonesia).

Figure 1. 

Distribution of Longipeditermes longipes (yellow dot) and L. kistneri (red dot) on the Greater Sundas, Southeast Asia.

The head, body (in profile), pronotum, and antenna of the soldier caste (preserved in 70% ethanol) were photographed using a digital microscope (KEYENCE HFVH-8000, Osaka). From these images, multifocused montages were produced using Helicon Focus v. 6.2.2 (Helicon Soft Ltd, Kharkov, Ukraine). Morphological terms and measurement characters follow those of Roonwal and Chhotani (1989), Sands (1998), Tho (1992), and Syaukani et al. (2011). We measured the characters (in millimeters): head capsule length including nasus (HLN), head capsule length excluding nasus (HL), nasus length (NL), nasus index = NL/HL, maximum head width at anterior part (HWA), maximum head width at posterior part (HWP), maximum height of head capsule excluding postmentum (HH), pronotum length (PL), and pronotum width (PW).

Summary of collection sites.

Site Vegetation type Altitude (m)
The Leuser Ecosystem, Sumatra Lowland & subalpine forests Protected & unprotected 50–1400
Aceh Province (outside the Leuser Ecosystem), Sumatra Lowland forest 20–600
Batang Gadis National Park, Sumatra Lowland forest 800–1200
North Sumatra Province (except the Leuser Ecosystem) Lowland & subalpine forests 20–1100
Kerinci Seblat National Park, Sumatra Lowland & subalpine forests 300–1350
West Sumatra Province (outside Kerinci Seblat N. P.), Sumatra Lowland forest 50–400
Jambi Province (outside Kerinci Seblat N.P.), Sumatra Lowland forest 50–600
Bukit Barisan Selatan National Park, Sumatra Lowland forest 50–600
Bengkulu Province (outside Bukit Barisan SelatanNational Park), Sumatra Lowland forest 100–700
Lampung Province (outside Bukit Barisan SelatanNational Park), Sumatra Lowland forest 10–300
Endau Ronpin National Park, Malay Peninsula Lowland forest 150–600
Teluk Bahang National Park, Malay Peninsula Lowland forest 5–200
Bukit Tangkiling Nature Preserve, Borneo Lowland forest 25–170
Pararawen Nature Preserve, Borneo Lowland forest 50–350
Barito Ulu, Borneo Lowland & subalpine forests 900–1200
Gunung Palung National Park, Borneo Lowland forest 50–300
Betung Kerihun National Park, Borneo Lowland forest 400–850
West Kalimantan Province (outside Betung Kerihun NP) Borneo Lowland forest 50–650
East Kalimantan Province (outside Gunung Palung NP) Borneo Lowland forest 150–500
Bukit Suharto, Borneo Lowland forest 40–300
North Kalimantan Province (outside Kayan Mentarang NP), Borneo Lowland & subalpine forests 500–1200
Gunung Halimun National Park. West Java Lowland & subalpine forest 800–1350
Pangandaran Natural Reserve, West Java Lowland 10–40
Bukit Lengkong, West Java Lowland & subalpine forest 800–1250

Results

Taxonomic accounts

Longipeditermes Holmgren, 1913

Soldier

Bimodal in its size distribution. Head capsule pale brown to blackish; antenna much paler than head capsule in coloration, with the basal segments (first and second) generally darker than the remaining segments; pronotum paler than or as pale as head capsule; abdominal tergites pale brown to dark sepia brown; coxae yellowish to pale brown; femora yellow to brown; tibiae pale yellow to yellow. In dorsal view head capsule excluding rostrum pear-shaped to somewhat triangular, weakly constricted behind antennal sockets; its posterior margin weakly to strongly convex; dorsal outline in profile weakly to strongly concave; rostrum excluding the apex somewhat cylindrical rather than conical. Antenna long, with 14 segments; 3rd segment at least twice as long as 4th. Mandible relatively long, with sharp apical processes. Legs very long.

Worker

Monomorphic but showing size variation. Pale brown to blackish. Antenna with 15 segments, with the basal segments darker than the following ones. Left mandible with 3rd marginal tooth weakly to moderately protruding from cutting edge, and 4th partially hidden behind molar prominence.

Remarks

In Longipeditermes, the soldier caste is dimorphic. The caste is subdivided into major and minor soldiers that differ markedly in size (Thapa 1981; Tho 1992; Gathorne-Hardy 2001; Miura and Matsumoto 1998) and slightly in coloration; the former being darker than the latter. The major soldier has more characters useful for species recognition and identification. Two varied species, L. longipes and L. kistneri, were recognized in our collection (Table 1), while no additional species were found by our morphology-based examination. The two species are relatively easily distinguished from each other by a combination of characters given in Table 2. These characters are re-described below in detail.

Summary of morphological characters for the Longipeditermes based on major soldiers and workers.

Species Soldier Worker
Coloration of head capsule Rostrum Anterior margin of pronotum Antennae Left mandible
L. longipes Sepia brown to blackish Apical 2/3 lighter & basal1/3 darker Nearly straight 4th segment longer than 5th 3rd marginal tooth moderately protruding from cutting edge
L. kistneri Pale brown to dark brown Apical 2/3 darker and basal 1/3 lighter Strongly indented in the middle 4th to 6th segments almost equal in length 3rd marginal tooth weakly protruding from cutting edge

Longpeditermes longipes (Haviland, 1898)

Termes longipes Haviland 1898: 439–440. Type locality: Perak, Malay Peninsula. Syntypes examined.

Eutermes longipes Wasmann 1902: 131.

Eutermes (Longipeditermes) longipes: Holmgren 1902: 215–217; John 1925: 406.

Longipeditermes longipes: Snyder 1949: 317; Ahmad 1958: 126; Tho 1992: 180–181: Miura and Matsumoto 1998: 179–189 (ecology); Hoare and Jones 1998: 1357–1366.

Longipeditermes mandibulatus Thapa 1981: 349–352. Sabah, Borneo. Synonymized by Hoare and Jones 1998: 1357–1366.

Materials examined

Sumatra . SYK1998 & 1999-L-1115, 1117, 1120, 1121, 1124, 1130, 1133, 1136, 1138, 1139,1141, 1143, 1147, 3025: soldiers and workers from undisturbed forest, 300–500 m altitude, Ketambe, Southeast Aceh. SYK1998-L-3005, 3010: soldiers and workers from disturbed forest, 450 m altitude, Lokop, East Aceh, Aceh. SYK-L-1148, 3006: soldiers and workers from disturbed forest, 80 m altitude, Soraya, Singkil, Aceh. SYK1998 & 2000-L-1126, 1127, 1131,1199, 1145, 3007, 3008: soldiers and workers from undisturbed forest, 150-350 m altitude, Bukit Lawang, Langkat, North Sumatra. SYK1998-L-3009, 3022, 3024: soldiers and workers from disturbed forest, 200 m altitude, MRT Logging Concession, South Aceh. SYK1999 & 2001-L-1116, 1123, 1125, 1128, 1134, 1135, 1137, 1144, 1146: soldiers and workers from disturbed forest, 50 m altitude, Sekundur, Langkat, North Sumatra. SYK1999-L-1112, 1118, 1129, 1132, 1140, 1142: soldiers and workers from undisturbed forest, 200–400 m altitude, Bengkung, Southeast Aceh. SYK2006-AL-0104: Soldiers and workers from disturbed forest, 50 m altitude, Maestong, Batang Hari, Jambi. SYK2006-KSNP-0011, 0019, 0080, 0091, 0093, 0095, 0096, 0097, 0104, 0206: soldiers and workers from the undisturbed forest, 300 m in altitude, Sungai Manau, Merangin, Jambi. SYK2006-AL-0100, 0101, 0102, 0103: soldiers and workers from disturbed forest, 50 m altitude, Maestong, Batang Hari, Jambi. SYK2007-LP-0019: soldiers and workers from undisturbed forest, 1350 m altitude, Sumber Jaya, Kota Bumi, Lampung. Java. SYK2001-HL-067, 072: soldiers and workers from protected forest, 1,450 m altitude, Halimun NP, West Java. SYK2006-PD-0011, 048: soldiers and workers from protected forest, 10 m altitude, Pangandaran Nature Reserve. West Java. Malay Peninsula. SYK2009-ER-084, 085, 086, 087, 088: soldiers and workers from protected forest, 50 m altitude, Endau Rompin NP. SYK2011-TB-011, 012, 013, 014: soldiers and workers from protected forest, 10 m altitude, Teluk Bahang NP. Borneo. SYK2014-BT-0054: soldiers and workers from protected secondary forest, 35 m altitude, Palangkaraya, Central Kalimantan. SYK-2014-P-0012: soldiers and workers from undisturbed forest, 220 m altitude, North Barito, Central Kalimantan. SYK-2014-P-0024: soldiers and workers from undisturbed forest, 270 m altitude, North Barito, Central Kalimantan.

Soldier

(Figs 2–4, 7, 8, 1012, 15, 16). Bimodal continuum in size. Major soldier (Figs 2–4, 7, 8). Head capsule sepia brown to blackish, in dorsal view somewhat triangular, very weakly constricted behind antennal sockets, with posterior margin roundly convex; dorsal outline (including nasus) in lateral view moderately concave; rostrum feebly bicolored, with apical 2/3 lighter and basal 1/3 darker. Mandible with short to long apical processes in dorsal view. Antenna with 14 segments, much paler than head capsule; 1st antennal segment darker than the following segments, which are uniformly colored; 3rd segment approximately 1.5 times as long as 4th; 5th slightly shorter than 4th; 6th–14th gradually shortening toward the apex. Pronotum in dorsal view paler than head capsule; anterior margin nearly straight; posterior margin weakly indented in the middle. Coxae pale brown to brown; femora pale brown to yellow; tibiae pale yellow. Abdominal tergites pale brown to dark sepia brown. Measurements and index (20 major soldiers from 10 colonies): HLN 2.30–2.71 mm; HL 1.57–1.62 mm; NL 0.72–0.77 mm; NL/HL 0.47–0.51; HWA 0.81–0.98 mm; HWP 1.87–1.92 mm; HH 1.22–1.32 mm; PL 0.37–0.40 mm; PW 0.75–0.78 mm.

Worker

(Figs 5, 6, 9, 13, 14, 17). Monomorphic but showing size variation. Worker (large-sized) (Figs 5, 6, 9). Blackish brown to blackish. Left mandible with apical tooth shorter than 1st marginal tooth; 3rd marginal moderately protruding from cutting edge; 4th completely hidden, scarcely visible behind molar prominence. Right mandible with posterior edge of 2nd marginal tooth nearly straight or weakly concave; the inner layer of molar plate very moderately convex; notch moderately developed. Antenna with 15 segments, whitish yellow; 1st and 2nd segments darker than the following segments; 3rd clearly longer than 4th; 4th longer than 5th, 6th–15th gradually shortening toward the apex.

Distribution

Sumatra, Peninsular Malaysia, Java (new record), and Borneo.

Figures 2–9. 

(SYK2006-KSNP-0011). Longipeditermes longipes (major soldier & major worker). Soldier (2–4, 7, 8) and worker (5, 6, 9). Habitus in profile (2), head in dorsal view (3), head in profile (4), left (5) and right (6) mandibles, pronotum (7), antennae (8, 9). Scale bars: 0.6 mm (2), 0.3 mm (3, 4), 0.1 mm (5– 9), 1.7 mm (6).

Figures 10–17. 

(SYK2006-KSNP-0011). L. longipes (minor soldier & minor worker). Soldier (10–12, 15) and worker (13, 14, 17). Habitus in profile (10), head in dorsal view (11), head in profile (12), left (13) and right (14) mandibles, pronotum (15), antennae (16, 17). Scale bars: 0.6 mm (10), 0.3 mm (11, 12), 0.1 mm (13– 17), 1,7 mm (15).

Longipeditermes kistneri Akhtar & Ahmad, 1985

Longipeditermes kistneri Ahktar and Ahmad 1985: 215–217. Bogor, West Java.

Material examined

SYK1998 & 1999-L-1098, 2001, 3011: soldiers and workers from the undisturbed forest, 1100–1400 m altitude, Kemiri Mountain, Southeast Aceh, Aceh, Sumatra.

Soldier

(Figs 1820, 23, 24, 26–28, 31, 32). Bimodal continuum in size. Major soldier (Figs 1820, 23, 24). Head capsule pale to dark brown, in dorsal view somewhat triangular, very weakly constricted behind antennal sockets, with posterior margin roundly convex; dorsal outline (including nasus) in profile moderately concave; rostrum feebly bicolored, with apical 2/3 darker and basal 1/3 paler. Mandible with short to long apical processes. Antenna with 14 segments, much paler than head capsule; 1st and 2nd antennal segments darker than the following segments, which are uniformly colored; 3rd approximately 1.5 times as long as 4th; 5th slightly shorter than 4th; 6th–14th gradually shortening toward the apex. Pronotum in dorsal view as pale as or paler than head capsule; anterior margin indented in the middle; posterior margin weakly indented in the middle. Coxae and femora yellowish to pale brown; tibiae pale yellow to yellow. Abdominal tergites brown to dark sepia brown. Measurements and index (20 major soldiers from 10 colonies): HLN 2.22–2.43 mm; HL 1.62–1.68 mm; NL 1.02–1.12 mm; NL/HL 0.66–0.73 ; HWA 0.86–0.90 mm; HWP 1.63–1.71 mm; HH 0.98–1.04 mm; PL 0.40–0.44 mm; PW 0.77–0.82 mm.

Worker

(Figs 21, 22, 25, 29, 30, 33). Monomorphic but showing size variation. Worker (large-sized) (Figs 21, 22, 25). Body pale brown to brown. Left mandible with apical tooth shorter than 1st marginal tooth; 3rd marginal weakly protruding from cutting edge; 4th not completely hidden behind molar prominence. Right mandible with posterior edge of 2nd marginal tooth nearly straight; the inner layer of the molar plate very weakly concave; notch moderately to strongly developed. Antenna with 15 segments, with 1st and 2nd segments slightly darker than the following segments; 3rd clearly longer than 4th; 4th–6th nearly equal in length; 7th–15th gradually shortening toward the apex.

Distribution

Sumatra and Java.

Figures 18–25. 

(SYK1999-L-3011). L. kistneri (major soldier and major worker). Soldier (18–20, 23, 25) and worker (21, 22, 25). Habitus in profile (18), head in dorsal view (19), head in profile (20), left (21) and right (22) mandibles, pronotum (23), antennae (24, 25). Scale bars: 0.6 mm (18), 0.3 mm (19, 20), 0.1 mm (21, 22, 24, 25), 1.7 mm (23).

Figures 26–33. 

(SYK1999-L-3011). L. kistneri (minor soldier and minor worker). Soldier (26– 28, 32) and worker (29, 30, 33). Habitus in profile (26), head in dorsal view (27), head in profile (28), left (29) and right (30) mandibles, pronotum (31), antennae (32, 33). Scale bars: 0.6 mm (26), 0.3 mm (27, 28), 0.1 mm (29, 30, 32, 33), 1.7 mm (31).

Discussion

Longipeditermes species are distinguished from those of the other groups of Southeast Asian nasutitermitine termites by forming open-air processional foraging columns (e.g., Hospitalitermes and Lacessititermes) and by having major and minor soldiers that both have long legs and a long rostrum. No information is available for the ratio of the largest and smallest soldiers in typical colonies, but minor soldiers appear to be more numerous (Figs 34, 35). The soldier does not exhibit a clear dimorphism, but a bimodal continuum in size.

Longipeditermes kistneri was only collected from Sumatra during our intensive surveys in the Greater Sundas, although the type locality is Java. Nine colonies were found in protected forests above 1,000 m in altitude. Four nests were located among buttresses and roots of big trees (under the ground covered with decayed leaf litter and small decayed branches). This nesting habit makes it difficult to estimate the sizes of their colonies. Both soldiers and workers forage on the surface of the ground. The size of foraging columns is smaller in L. kistneri than in L. longipes. Their pale- to dark-brown body color and smaller columns make it difficult to find colonies of L. kistneri.

Longipeditermes longipes seems to prefer lowland and swamp forests and is widespread in the Greater Sundas. Nests of L. longipes are usually constructed among buttresses of trees (Fig. 36) and in dead or decayed stumps, with decayed wood debris and leaves (Fig. 37). Several subterranean nests of this species were found in Lambir Hills National Park (Sarawak, Borneo) by Miura and Matsumoto (1998). Underground nests are sometimes made from consisting of soil and litter. This species forages on the ground during the day. However, Hoare and Jones (1998) reported foraging activity at night, during which no food balls were being carried. We also found foraging columns at night; the columns consisted of limited numbers of workers carrying food balls to the nests. In typical processional foraging columns, numerous soldiers guard workers. Foraging ecology and diet of L. longipes have been poorly studied, but we know that this species feeds on leaf litter (Miura and Matsumoto 1998) and in the Pasoh Forest Reserve on the Malay Peninsula it prefers to consume older leaf litter rather than newly fallen leaves (Matsumoto and Abe 1979). We observed foraging columns consuming a mixture of bark, twigs, and decayed wood in forests of Sumatra and the Malay Peninsula.

Figures 34, 35. 

Foraging column of Longipeditermes longipes on the forest floor in Southeast Asia. Workers collecting and transporting food source from a decayed branch (34) and leaf litter (35), soldiers protecting the worker from predators such as Componotus gigas (Latreille).

Figure 36. 

Nest entrances (red arrows) of subterranean nest of Longipeditermes longipes found in SE Asia tropical forests.

Figure 37. 

Above-ground nest of L. longipes constructed in and around a dead tree stump (Shorea sp.). The cavity was fulfill with a mixture of soil, debris and litter.

Conclusion

In general, from the current study, we conclude that only two species of Longipeditermes (L. longipes Holmgren and L. kistneri Akhtar & Ahmad) have so far been collected in the Greater Sundas. Long legs and size dimorphism of soldier caste can help to separate this genus from other nasute termites in Southeast Asia. The condition of rostrum, head capsule, pronotum, and antennae determine the soldier caste, while the condition of antennae and left mandibles of worker caste serve to distinguish these two species. Longipeditermes kistneri seems to prefer high-altitude forests and has been found exclusively from Java and Sumatra, while L. longipes seems to prefer lowland and swamp forests and is widespread in the Greater Sundas.

Acknowledgements

We thank the Termite Research Group (Universitas Syiah Kuala, Indonesia), Sugesti, M. Rapi, Tarmizi, M. Isa, Usman, and Mat Plin (Leuser Development Program, Indonesia), Erwin Widodo (Conservation International), Epal (WWF, Indonesia), Dolly Priatna and Adnun Salampesi (Zoological Society of London), F.X Susilo, I Gede Swibawa, and Rahmat Pranoto (Lampung University), Idris Ghani and Fazli Rahim (UKM, Malaysia), Rosichon Ubaidillah and Wara Asfiya (MZB, Bogor), Teguh Pribadi (PGRI Palangkaraya University, Indonesia), and M. Shahril (USM, Malaysia) for assistance in the field and laboratory. We are grateful to Seiki Yamane (Kagoshima University Museum, Japan) for support, Paul Eggleton and David Jones (Natural History Museum, UK) and staff at the Museum Zoologicum Bogoriense (MZB, Indonesia) for allowing the first author to examine the type material. We thank to staff in the Forestry Department in Indonesia for their help and assistance during field surveys. This work was financially supported by the Leuser Development Program (LDP 1998-2001), Nagao Environmental Foundation (NEF 2006), the Ministry of Research, Technology and Higher Education RG to Syaukani (International Research Collaboration and Scientific Publication 2012–2018), Syiah Kuala University (Professor Candidate Grant 2015, H-index Publication Grant 2016, International Conference Support 2016 (DITJEN RISBANG, the Ministry of Research, Technology and Higher Education, and the Zoology Laboratory Grant 2018). Eguchi’s research activities were partly supported by Asahi Glass Foundation (Leader: Katsuyuki Eguchi; FY2017-FY2020).

References

  • Akhtar MS, Ahmad M (1985) A new nasute termite from Java (Isoptera: Termitidae: Nasutitermitinae). Pakistan Journal of Zoology 17: 215–217.
  • Collins NM (1984) The termites (Isoptera) of the Gunung Mulu National Park, with a key to genera known from Sarawak. Sarawak Museum Journal 30: 65–87.
  • Gray B, Dhanarajan G (1974) Processional trails of the black termite Longipeditermes Longipes (Haviland) (Isoptera: Termitisae). Insectes Sociaux 21: 151–155. https://doi.org/10.1007/BF02222939
  • Gathorne-Hardy F (2001) A review of the South-East Asian Nasutitermitinae (Isoptera: Termitidae), with descriptions of one new genus and a new species and including a key to the genera. Journal of Natural History 35: 1486–1506. https://doi.org/10.1080/002229301317067647
  • Grimaldi D, Engel MS (2005) Evolution of the Insects. Cambridge University Press, New York, 755 pp.
  • Hoare A, Jones DT (1998) Note on the foraging behaviour and taxonomy of the Southeast Asian termite Longipeditermes longipes (Termitidae: Nasutitermitinar). Journal of Natural History 32: 1357–1366. https://doi.org/10.1080/00222939800770681
  • Holmgren N (1912) Termitenstudien. 3. Systematik der Termiten. Die Familien Mastotermitidae. KungligaSvenskaVetensskapakademiens Handlingar 48: 1–166.
  • Holmgren N (1913) Termitenstudien. 4. Versuch einer systematichen Monographie der Termiten der orientalischen Region. KungligaSvenskaVetensskapakademiens Handlingar 50: 1–276.
  • John O (1925) Termiten von Ceylon, der Malayaischen Halbinsel, Sumatra, Java und den Aru- Inseln. Treubia 6: 360–419.
  • Miura T, Matsumoto T (1998) Open-air litter foraging in the nasute termite Longipeditermes longipes (Isoptera: Termitidae). Journal of Insect Behavior 11: 179–189. https://doi.org/10.1023/A:1021039722402
  • Pearce MJ (1997) Termites: Biology and Pest Management. CAB International, Willingford, 172 pp.
  • Roisin Y (2000) Diversity and evolution of caste patterns. In: Abe T, Bignell DE, Higashi M (Eds) Termites: Evolution, Sociality, Symbiosis, Ecology. Kluwer Academic, Dordrecht, 95–120. https://doi.org/10.1007/978-94-017-3223-9_5
  • Roonwal ML, Chhotani OB (1989) The Fauna of India and the Adjacent Countries. Zoological Survey of India, Calcuta, 672 pp.
  • Sands WA (1998) The Identification of Worker Caste of Termite from Soil of Africa and the Middle East. CAB International, Wallingford, 500 pp.
  • Syaukani S, Thompson GJ, Yamane S (2011) Hospitalitermes krishnai, a new nasute Termite (Nasutitermitinae, Termitidae, Isoptera), from southern Sumatra, Indonesia. ZooKeys 148: 161–169. https://doi.org/10.3897/zookeys.148.1768
  • Syaukani S, Thompson GJ, Zettel H, Pribadi T (2016) A new species of open-air processional column termite, Hospitalitermes nigriantennalis sp. n. (Termitidae), from Borneo. Zookeys 554: 27–36. https://doi.org/10.3897/zookeys.554.6306
  • Takematsu Y, Kambara K, Yamaguchi T, Mitsumaki K (2013) Spatial segregation of four coexisting processional termites (Termitidae: Nasutitermitinae) in tropical rainforest. Entomological Science 16: 355–359. https://doi.org/10.1111/ens.12015
  • Thapa RS (1981) Termites of Sabah. Sabah Forest Record, Sandakan 12: 1–374.
  • Tho YP (1992) Termites of Peninsular Malaysia. Malayan Forest Records, Kuala Lumpur 36: 1–224.
  • Watson JAL, Gay FJ (1991) Isoptera (Termites). In: Naumann D, Carne PB, Lawrence JF, Nielsen ES, Spradbery JP, Taylor RW, Whitten MJ, Littlejohn MJ (Eds) The Insects of Australia – A Textbook for Students and Research Workers. Vol. 1. Melbourne University Press, Australia, 330–347.