Journal of Threatened Taxa | www.threatenedtaxa.org | 26 October 2018 | 10(11): 12582–12588

 

 

New species of termite Pericapritermes travancorensis sp. nov. (Isoptera: Termitidae: Termitinae) from India

 

Jobin Mathew 1 & Chinnu Ipe 2

 

1,2 Department of Zoology, CMS College, Kottayam, Kerala 686001, India

1 jobin@cmscollege.ac.in (corresponding author), 2 chinnurupen@gmail.com

 

 

 

Abstract: A new species of Pericapritermes, P. travancorensis sp. nov. (Isoptera, Termitidae, Termitinae), is described from Kerala, India, based on the characters of the king, queen, imago, soldiers, and workers.  Morphological measurements of the king, queen, imago, soldiers, and workers are given with suitable illustrations.  List of species of the genus Pericapritermes from the Oriental region is provided.

 

Keywords: Isoptera, Kottayam District, new species, Pericapritermes, Termitidae.

 

 

 

 

doi: https://doi.org/10.11609/jott.3389.10.11.12582-12588  |  ZooBank: urn:lsid:zoobank.org:pub:20ABAF80-205F-4DF8-ABC7-06D387C9CE33

 

Editor: R. Sundararaj, Institute of Wood Science & Technology, Bengaluru, India.      Date of publication: 26 October 2018 (online & print)

 

Manuscript details: Ms # 3389 | Received 02 March 2017 | Final received 28 August 2018 | Finally accepted 24 September 2018

 

Citation: Mathew, J. & C. Ipe (2018). New species of termite Pericapritermes travancorensis sp. nov. (Isoptera: Termitidae: Termitinae) from India. Journal of Threatened Taxa 10(11): 12582–12588; https://doi.org/10.11609/jott.3389.10.11.12582-12588

 

Copyright: © Mathew & Ipe 2018. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use of this article in any medium, reproduction and distribution by providing adequate credit to the authors and the source of publication.

 

Funding: None.

 

Competing interests: The authors declare no competing interests.

 

Acknowledgements: We thank Edwin Jose, T.R. Sreeraj, Nebin Ninan, and Shanavas Moitheen for their help in termite survey and sample collection.  The authors are obliged to the officials of Zoological Survey of India (ZSI), Kolkata, for facilities and encouragement.

 

 

 

 

India is a region rich in termite diversity.  Roonwal & Chhotani (1989) listed and comprehensively described 337 species of termites under 59 genera from the Indian sub-region (comprising of India, Pakistan, Nepal, Bhutan, Bangladesh, Myanmar, and Sri Lanka).  Krishna et al. (2013) listed 290 species of termites belonging to six families and 54 genera from India.  Mathew (2004) reported 56 species under 23 genera belonging to four families of termites from Kerala.  The current termite fauna of Kerala can be estimated to be 66 species belonging to three families and 29 genera (Amina et al. 2016).

Silvestri (1914) described the genus Pericapritermes based on Pericapritermes urgens.  The genus has been reported from the Ethiopian region by Snyder (1949) and Emerson (1955).  The studies of Krishna (1968) transferred many species of the genus Capritermes with Pericapritermes, collected from Indo-Malayan region and New Guinea.  Presently, Pericapritermes is known from the Oriental, Ethiopian, Palearctic, and Papuan regions (Thomas et al. 2008); Krishna et al. (2013) listed 40 species under the genus Pericapritermes.  The oriental region contains 24 species in which six species are known from India (Table 3).

The soldiers of Pericapritermes are the most advanced in the Capritermes group in that their mandibles are robust and strong.  The left is twisted and arched in the middle, with its apex blunt, and the right is flat, straight, and blade-like; in defence, they are locked together and released with a loud click, flipping the soldier several inches through the air (Thomas et al. 2008).  The head is flat, the forehead is not steep, and the frontal gland is much reduced.  The labrum of the soldier is straight at the anterior margin and with small anterio-lateral points.  Imago has a large, oval fontanelle (Roonwal & Chhotani 1989).  The apical tooth is shorter than the fused first plus second marginal tooth, the posterior margin of the fused first plus second marginal tooth is elongated and sinuate, and the third marginal tooth is prominent (Roonwal & Chhotani 1989).  The members of this genus are soil feeders.  Soil-feeding termites constitute 38.3% of Termitidae species, which dominate several subfamilies (Jones & Eggleton 2011).  Pericapritermes build subterranean diffuse gallery systems that consist of clusters of small cells connected with tunnels, with the cells usually adjacent to underground hard materials.

 

Material and Methods

Specimens were collected from CMS College Campus, Kottayam District of Kerala State, southern India, which is situated between 9.5960N 76.5200E.  The area is characterized by humid tropical climate with a mean annual rainfall around 3,600mm, and temperature ranging from 20–37 0C.  CMS College Campus contains 35 acres of protected land.  The tropical climate of this region supports rich biodiversity.

 

Collection and Identification of Termites

All the termites encountered in the colony were collected using an aspirator and preserved in 80% alcohol.  Measurements and photographs were taken using Labomed Luxeo 4D binocular microscope with attached camera and PixelPro software at magnification 8X–35X.  Morphological terminology, measurements, and indices for describing soldiers, workers, and imago follow Roonwal & Chhotani (1989), and Sands (1998).  Important measurements and indices used in the study were total body length with wing, total body length without wing, length of head to lateral base of mandibles, maximum width of head with eye, maximum diameter of compound eye, maximum diameter of lateral ocellus, minimum eye-ocellus distance, maximum length of labrum, maximum width of labrum, length of left mandible, length of right mandible, maximum length of Pronotum, maximum width of Pronotum, minimum length of postmentum, maximum width of postmentum, minimum width of postmentum, minimum length of hindwing without scale, minimum length of forewing without scale, head width/head length index, and mandible length/head length index.  The population study was done using direct count method. 

 

Museum details

The holotype and paratypes are preserved in 80% alcohol and deposited in the Zoology Museum, Department of Zoology, CMS College, Kottayam, Kerala, India.

 

Pericapritermes travancorensis sp. nov.

(Image 1)

urn:lsid:zoobank.org:act:E49C1081-D644-4E51-877A-E147BD91B7C1

Material examined

Holotype: CMSZMAI-101, Soldier, 08.xi.2015, CMS College Campus, Chungam, Kottayam, Kerala, India, 9.5960N & 76.5200E, 2.97m, coll. Jobin Mathew. 

Paratypes: CMSZMAI-102, Soldier; CMSZMAI-103, Imago; CMSZMAI-104, King; CMSZMAI-105, Queen; CMSZMAI-106, Worker; 08.xi.2015, CMS College Campus, Chungam, Kottayam, Kerala, India, 9.5960N & 76.5200E, 2.97m, coll. Jobin Mathew. 

 

Diagnosis

Five species of Pericapritermes were known from India. Diagnostic characters of the soldiers from India with their distribution are given in the Table 4. Pericapritermes found in the Indian subcontinent classified as large, medium and smaller species. P. dunensis was the only species coming under the medium group (Roonwal & Chhotani, 1989)  P. travancorensis sp. nov. is a medium sized termite showed affinity with P. dunensis. P. dunensis was described from Dehra Dun, India; and has later been reported from Meghalaya, Arunachal Pradesh, Uttar Pradesh West Bengal and Bhutan (Roonwal, M.L. & Sen-Sarma, 1960; Roonwal & Chhotani, 1989). Imago of P. dunensis have body length with wings 14.10-15.10 mm, body length without wings 7.40-8.00 mm and length to the base of mandibles 0.90-1.00mm. Whereas In P. travancorensis sp. nov. body length with wings 10.77-11.23, body length without wings 5.58 – 6.45 mm and length to base of mandibles 0.78-0.82. P. travancorensis sp. nov. can be easily distinguished from P. Dunensis by larger soldier and mandible of the soldier. In this species right mandible is equal or slightly longer than left (Tables 1  & 3). The imago of the P. travancorensis sp. nov. smaller than the P. Dunensis, fontanelle large and sub-squarish.  Moreover P. dunensis confined to the northeast of India.

 

 

 

 

Table 1. Table showing the various measurements of CMSZMAI-101 holotype of P. travancorensis sp. nov. (in mm)

 

Characters

Soldier

Total body Length (Without wings)

7.10

Total Head Length

4.32

Head Width

1.40

Head Length (Without Mandibles)

2.63

Left Mandible Length

1.73

Right Mandible Length

1.74

Labrum Length

0.24

Labrum Width

0.29

Pronotum Length

0.35

Pronotum Width

0.82

Postmentum Length

1.16

Postmentum Width Maximum

0.42

Postmentum Width Minimum

0.28

Head W/Head L

0.53

Mandible L /Head L

0.66

 

 

 

Table 2. Table showing the various measurements of  CMSZMAI-102, 10 Soldiers; CMSZMAI-103, 4 Imagos; CMSZMAI-104, 1 King; CMSZMAI-105, 1 Queen; CMSZMAI-106  soldier, 10 workers of P. travancorensis sp. nov. (in mm).  Measurements are based on 1 king,

1 queen, 4 imagos, 10 soldiers, and 10 workers.

 

  Characters

King

Queen

Imago

Soldier

Worker

Total body length (with wings)

-

_

10.77–11.23

_

_

Total body length (without wings)

6.36

17.82

5.58–6.45

6.29–7.30

4.48–5.33

Total head Length

1.27

1.28

1.26–1.34

4.18–4.43

1.26–1.36

Head width

1.23

1.22

1.18–1.23

1.35–1.51

0.93–1.06

Head length (without mandibles)

0.82

0.79

0.78–0.82

2.49–2.73

0.74–0.83

Left mandible length

_

_

_

1.65–1.83

_

Right mandible length

_

_

_

1.71–1.87

_

Maximum diameter of eye

0.25

0.24

0.24–0.30

 

_

Eye ocilla distance

0.13

0.11

0.11–0.13

 

 

Ocellar diameter

0.11

0.12

0.11–0.13

 

 

Labrum length

0.30

0.31

0.29–0.31

0.22–0.26

0.34–0.36

Labrum width

0.29

0.32

0.29–0.32

0.27–0.31

0.41–0.43

Pronotum length

0.64

0.63

0.61–0.68

0.29–0.39

0.19–0.24

Pronotum width

1.12

1.16

1.04–1.16

0.74–0.92

0.53–0.62

Postmentum length

0.27

0.26

0.26–0.30

1.14–1.19

0.27–0.33

Postmentum width (maximum)

0.32

0.35

0.34–0.39

0.40–0.45

0.25–0.30

Postmentum width (minimum)

0.34

0.33

0.32–0.36

0.26–0.30

_

Head width / Head length

1.5

1.54

1.46–1.54

0.55–0.59

1.24–1.35

Mandible length /Head length

_

_

_

0.65–0.73

_

Fore wing length

_

_

8.47–9.10

_

_

Hind wing length

_

_

7.75–8.46

_

_

 

 

 

Description

Holotype: Soldier

Head-capsule yellowish-brown, uniformly coloured, antennae yellowish darker anteriorly, labrum and pronotum pale with yellowish tinge, left mandible blackish, right mandible reddish-brown, legs and abdomen pale yellowish; head and body sparsely hairy; total body length 7.10 mm; head-capsule sub-rectangular, sides sub-straight (length to base of mandibles 2.63 mm, width 1.4 mm, index width/length 0.56 mm); in profile frons sloping in front and shallowly depressed medially, median suture present, not extending up to fontanelle, fontanelle minute, circular, situated anteriorly, occipito-fontanelle distance 1.87 mm; antennae 14-segmented, segment 4 smallest; labrum fleshy, anterior margin depressed medially, antero-lateral corners produced into small, pointed tips; mandibles strongly asymmetrical,  longer than half the length of head-capsule, left mandible strongly twisted at middle, with a spoon like tip, right blade-like tip pointed and weakly out curved apically, longer than left mandible (length: left 1.73 mm, right 1.74 mm, index left mandible length/head-length 0.66mm) (Fig. 1D); postmentum long club-shaped, weakly narrowed at waist (length 1.16 mm, max. width 0.42 mm, width at waist 0.28 mm) Pronotum saddle shaped, anterior margin convex, posterior margin with a weak median notch (length 0.35 mm, width 0.82  mm) (Figs. 6–12, Table 1).

 

Colony

The termite colony was found in porous laterite soil.  The colony consists of clusters of small cells connected with tunnels, with the cells usually adjacent to small crevices of the laterite stones.  The colony occupied an area of 3697cm3 and was located between the depth of 3.6–9.6 cm from the surface.  The royal chamber is an enlarged cell with smooth, almost polished, and quite clean inner surface.  It contains the royal pair and a number of soldiers and workers.  Maximum number of workers and soldiers were observed below 3.6cm.  Foraging workers were noticed 1.5m away from the colony.  The numbers of individuals of different castes of the colony were as follows: king - 1, queen - 1, imago - 5, workers - 1265, soldiers - 119, immature workers – 1153, and immature soldiers - 86.

 

Paratype: Imago

Head dark brown, postclypeus, labrum, antennae and legs brownish-yellow, pronotum dark brown paler anteriorly, abdomen dark brown above and yellowish below; head and body with a coat of fine short hairs and several long hairs; total body-length with wings 10.77–11.23 mm and without wings 5.58–6.45 mm; head-capsule subcircular (length to base of mandibles 0.78–0.82 mm, width with eyes 1.18–1.23 mm); fontanelle golden yellow, slightly depressed around fontanelle; epicranial suture indistinct; eyes round, strongly projecting, nearly equidistant from antennae and ocelli, (max. diameter 0.24-0.30 mm); ocelli oval, 0.11–0.13 mm long; eye ocillar distance 0.11–0.13 mm; postclypeus swollen, hairy; antennae with 15 segments; segment 3 shortest; 4 longer than 5 and 5 subequal to 3; mandibles with a prominent apical teeth and a pair of teeth basaly; pronotum flat, length 0.61–0.68 mm, width 1.04–1.16 mm, anteriorly weakly notched medially and posteriorly slightly emarginated; postmentum subsquarish length 0.26–0.30 mm, width 0.34–0.39 mm; wings brownish, membrane with distinct cubitus, media, and radius veins, in forewing and hindwing media arising from radius inside wing-membrane near to the wing scale, forewing-length 8.47–9.10 mm, hindwing-length 7.75–8.46 mm; body dimensions and colouration of king similar to imago but with slightly enlarged paler abdomen;total body length of queen 17.82 mm, abdomen whitish with yellowish tinge nearer to the sclerite (Figs. 1–5, Table 2).

 

Paratype: Soldier

Colour and characters same as holotype; total body length 6.29–7.30 mm; head-capsule sub-rectangular, sides sub-straight (length to base of mandibles 2.49–2.73 mm, width 1.35–1.51 mm, index width/length 0.55–0.59 mm); in profile frons sloping in front and shallowly depressed medially, median suture present, not extending up to fontanelle, fontanelle minute, circular, situated anteriorly, occipito-fontanelle distance 1.80–1.99 mm; antennae 14-segmented, segment 4 smallest; labrum fleshy, anterior margin depressed medially, antero-lateral corners produced into small, pointed tips; mandibles strongly asymmetrical,  longer than half the length of head-capsule, left mandible strongly twisted at middle, with a spoon like tip, right blade-like tip pointed and weakly out curved apically, longer than left mandible (length: left 1.65–1.83 mm, right 1.71–1.87 mm, index left mandible length/head-length 0.65–0.73 mm) (Fig. 1D); postmentum long club-shaped, weakly narrowed at waist (length 1.14–1.19 mm, max. width 0.40–0.45 mm, width at waist 0.26–0.30 mm) Pronotum saddle shaped, anterior margin convex, posterior margin with a weak median notch (length 0.29–0.39 mm, width 0.74–0.92 mm) (Figs. 6-12, Table 2).

 

Paratype: Worker

Head-capsule pale yellowish, antennae yellowish paler basely, pronotum and legs creamish, abdomen pale, intestinal contents clearly visible; head sparsely and body moderately hairy; total body length 4.48–5.33 mm; head capsule sub-circular (length to base of mandibles 0.74–0.83 mm, max. width 0.93–1.06 mm); fontanelle indistinct; antennae with 14 segments; segment 4 shortest; post clypeus swollen, length half of width; Pronotum saddle-shaped; length 0.19–0.24 mm, width 0.53–0.62 mm; anterior margin rounded, posterior margin with a median notch (Table 2).

 

Etymology

The name travancorensis was taken from the name of the area from where the specimens were collected.  The specimens were collected from the CMS College campus, a 200 year old College.  CMS College was the first college in the Kingdom of Travancore.

 

Distribution

Pericapritermes travancorensis sp. nov. is currently known from the CMS College campus, Kottayam, Kerala, India.

 

 

 

 

Table 3. Described Pericapritermes species, type localities and distribution from oriental region

 

 

Species

Type locality

Distribution

1

P. assamensis (Mathur & Thapa, 1965)

West Bengal

India

2

P. brachygnathus (John, 1925)

Sumatra, Indonesia

Malaysia, Indonesia

3

P. buitenzorgi (Holmgren, 1914)

Java, Indonesia

Malaysia, Indonesia

4

P. ceylonicus (Holmgren, 1911)

Peradeniya, Sri Lanka 

Sri Lanka 

5

P. dolichocephalus (John, 1925)

Selangor, Malaysia

Malaysia, Indonesia

6

P. dunensis (Roonwal & Sen-Sarma, 1960)

Dehradun, Uttarakhand

India

7

P. durga (Roonwal & Chhotani, 1962)

Cherrapunji, Meghalaya

India

8

P. fuscotibialis (Light, 1931)

Hong Kong, China

China

9

P. gutianensis Li & Ma, 1983

Fujian, China

China

10

P. hepuensis Gao & Yang, 1990

Guangxi, China

China

11

P. latignathus (Holmgren, 1914)

Tjibodas, Java

Bangladesh, Indonesia, Thailand, Vietnam, China, Malaysia

12

P. Modiglianii (Silvestri, 1922)

Sumatra, Indonesia

Indonesia

13

P. mohri (Kemner, 1934)

Buitenzorg, Indonesia

Indonesia, Malaysia

14

P. nitobei (Shiraki, 1909)

Maruyama, Taiwan

Sumatra, Indonesia

15

P. paetensis (Oshima, 1920)

Luzon, Philippines

Philippines

16

P. parvus Bourguignon & Roisin, 2008

Irian Jaya, Indonesia

Indonesia

17

P. perparvus (Holmgren, 1911)

Sri Lanka

Sri Lanka

18

P. planiusculus Ping & Xu, 1988

Guizhou, China

China

19

P. semarangi (Holmgren, 1913)

Semarang, Java

Bangladesh,Thailand, Malaysia, Indonesia, China

20

P. speciosus (Haviland, 1898)

Borneo Malaysia

Indonesia, Malaysia

21

P. tetraphilus (Silvestri, 1922)

Rangamati, Bangladesh

India, Bangladesh, China, Burma

22

P. topslipensis Thakur, 1976

Coimbatore, Tamil Nadu

India

23

P. wuzhishanensis (Li, 1982)

HainanIsland, China

China

24

P. travancorensis sp. nov.

Kerala, India

India

 

 

Table 4. Diagnostic characters of soldiers of the different species of Pericapritermes from India with their distribution

 

 

Name

Characters

Distribution

1

P. assamensis

Head capsule yellowish-brown, darker anteriorly, head length without mandibles 2.09-2.28 mm, head-width 1.19–1.27 mm, left mandible length 1.19–1.27 mm, pronotum width 0.72–0.91 mm.

Assam, West Bengal

2

P. dunensis

Head capsule yellowish-brown to brown, head length without mandibles 2.30–2.50 mm, head-width 1.30–1.45 mm, left mandible length 1.43–1.70 mm, pronotum width 0.80–0.88 mm.

Meghalaya, Arunachal Pradesh, Uttar Pradesh

West Bengal

3

P. durga

Head capsule yellow to pale brown, head length without mandibles 2.45–3.05 mm, head width 1.45–2.00 mm, left mandible length 1.45–2.00 mm.

Meghalaya, Arunachal Pradesh Manipur

4

P. tetraphilus

Head capsule yellow to reddish-yellow, head length without mandibles 2.45–3.06 mm, head width 1.48–1.70 mm, left mandibles length 1.60–1.84 mm, pronotum width 0.90-1.13 mm.

Assam, West Bengal, Madhya Pradesh

5

P. topslipensis

Head capsule pale yellow to deep straw yellow, head length without mandibles 2.5–2.75 mm, head width 1.32–1.41 mm, left Mandibles length 1.45–1.48 mm, pronotum width 0.85–0.90 mm.

Tamil Nadu, Karnataka

6

P. travancorensis sp. nov.

Head capsule yellowish-brown, head length without mandibles 2.39–2.73mm, head width 1.35–1.51 mm, left mandible length 1.65–1.83 mm, pronotum width 0.74–0.92 mm.

Kerala (Kottayam District)

 

 

 

Key to the soldiers of the Indian species of pericapritermes

 

1   Smaller species: Head-length with mandibles less than 4.0mm …………………………............................................................. 2

-    Head length without mandibles 2.09–2.28mm, head width 1.19–1.27 mm, mandible length 1.32–     1.37 mm  ..... P. assamensis

 

2   Medium sized species: Generally smaller in size. Head-length with mandibles 4.00–4.50 mm ........................................ 3

-    Head length without mandibles 2.30–2.50 mm, head width 1.30–1.45 mm, Left mandible length 1.43–1.70 mm……

     .................................................................................................................................................................................. P. dunensis

-    Head-length with mandibles 4.18–4.43 mm, head length without mandibles 2.49–2.73 mm, head width 1.35-1.51 mm, left

     mandible length 1.65–1.83 mm …..................................................................................…………….... P. travancorensis sp. nov.

 

3   Generally larger in size. Head-length with mandibles 4.5–5.0 mm, head length without mandibles 2.45–3.05 mm, head

     width 1.45–2.00 mm, left mandible length 1.45–2.00 mm.  Anterio-lateral corners of labrum small …………………………….… 1

-    Body weakly hairy, with fewer longer hairs ………………………………………………........................................................................... 4

-    Head-capsule generally larger, head length without mandibles 2.45–3.06 mm head width 1.48–1.70 mm.  Left mandible-

     length 1.60–1.84 mm, Antennae with segment 4 subequal to 2  .............................................................................  P.  tetraphilus

-    Head-capsule generally smaller, head length without mandibles 2.5–2.75 mm, width 1.32–1.41 mm left mandible length

     1.45–1.48 mm Antennae with segment 4 shorter than 2 .................................................................................... P. topslipensis

 

4   Body densely hairy with many long hairs, Labrum with anterior margin convex and antero-lateral corners minute. Segment

     4 of antennae shorter than 2. Pronotum not emarginate anteriorly .......................................................................... .P. durga

 

 

References

 

Amina, P., K. Rajmohana, K.V. Bhavana & P.P. Rabeeha (2016). New records of Termite species from Kerala (Isoptera: Termitidae). Journal of Threatened Taxa 8(11): 9334–9338; https://doi.org/10.11609/jott.3067.8.11.9334-9338

Chhotani, O.B. (1987). Distribution and Zoogeography of Oriental Termitidae (Isoptera). Beiträge zur Entomologie 37(2): 407–420.

Emerson, A.E. (1955). Geographical origins and dispersions of termite genera. Fieldiana: Zoology 37: 465–521.

Jones, D.T. & P. Eggleton (2011). Global biogeography of termites: a compilation of sources, pp. 477–498. In: Bignell, D.E., Roisin Y. & Lo N. (eds.). Biology of Termites: A Modern Synthesis. Springer, Netherlands, 576pp.

Krishna, K. (1968). Phylogeny and generic reclassification of the Capritermes complex (Isoptera, Termitidae, Termitinae). Bulletin of the American Museum of Natural History 138(5): 294-304.

Krishna, K., D.A. Grimaldi, V. Krishna & M.S. Engel (2013). Treatise on the Isoptera of the world. Bulletin of the American Museum of Natural History B377(6): 2282–2310; https://digitallibrary.amnh.org/handle/2246/6430

Mathew, G. (2004). Biodiversity Documentation for Kerala, Part 7: Insects. Kerala Forest Research Institute KFRI Handbook No. 17, 284pp.

Roonwal, M.L. & O.B. Chhotani (1989). The Fauna of India and the Adjacent Countries: Isoptera (Termites) - Vol. 1. Zoological Survey of India, Calcutta, viii+672pp.

Roonwal, M.L. & P.K. Sen-Sarma (1960). Contributions to the Systematics of Oriental Termites. ICAR, New Delhi, 407pp.

Sands, W.A. (1998). The Identification of Worker Castes of Termite Genera from Soils of Africa and the Middle East. CAB International, Wallingford, 500pp.

Silvestri, F. (1914). Contribuzione alla conoscenza dei Termitidi e Termitofili dell’Africa occidentale. I. Termitidi. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scoula Superiore d’Agricoltura, Portici 9: 1–146.

Snyder, T.E. (1949). Catalog of the termites (Isoptera) of the world. Smithsonian Miscellaneous Collections 112: 1–490.

Thomas, B., L. Maurice & R. Yves (2008). Revision of the Termitinae with snapping soldiers (Isoptera: Termitidae) from New Guinea. Zootaxa 1769: 1–34.