Indian Journal of Entomology Review Article (2020) DoI No.:

NESTING HABITAT AND COMB GEOMETRY OF STINGLESS BEE TETRAGONULA BENGALENSIS SMITH IN WEST BENGAL

Gautam Kunal*, RaKesh Das, nsneha latha, amit layeK anD shantanu Jha

Department of Agricultural Entomology, Bidhan Chandra Krishi Viswavidyalaya, Mohanpur, Nadia, West Bengal

*Email: gautamkunalbg@gmail.com (corresponding author)

ABSTRACT

A detailed survey was carried out at Purulia and Bankura districts of West Bengal to study the nesting habitat and comb geometry of stingless bee, Tetragonula bengalensis. A total of 55 colonies were found to nest in different habitats out of which 31 colonies were studied in both the districts, among them 11 colonies were found in tree nesting habitat and 20 found in wall nesting habitat. A colony nested in wall cavity in Bankura district was found to build their nest in a nest of swarmed wasp. The tree and wall nested colonies having the mean height of 179.57 cm, 173.25 cm and 99.71 cm, 79.15 cm in two respective districts. The mean dimensions of cavity and entrance tube nested in tree cavities of both the districts were 60.57 x 22.29 cm; 4.21 x 2.90 mm and 35 x 17.75 cm; 3.75 x 3.5 mm whereas for wall nested colonies 21.21 x 30.00 cm; 3.90 x 3.71 mm and 20.87 x 28.69 cm; 3.98 x 3.60 mm respectively. The brood cells, honey and pollen pots were brown to dark brown in colour, oval in shape and arranged in clusters. The mean dimensions of brood cells, pollen and honey pots in tree nested combs of both the districts were 2.26 x 1.66 mm, 7.42 x 5.18 mm, 7.60 x 4.77 mm and 2.26 x 2.00 mm, 7.44 x 5.16 mm, 7.80 x 4.99 mm while for wall nested combs 2.28 x 1.57 mm, 7.44 x 4.99 mm, 7.57 x 4.71 mm and 2.13 x 1.77 mm, 7.30 x 4.85 mm, 7.77 x 4.65 mm in length and width, respectively.

Key words: Tetragonula bengalensis, nesting habitat, comb geometry, tree nesting, wall nesting, nest morphometrics, cavity and entrance of nests, colour, shape, arrangement

20049-- Gautam Kunal

Stingless bees are the smallest of the honeyproducing bees, belonging to the subfamily Meliponinae consisting of eight general 5 subgenera and > 500 species (Wille, 1983). It has been known in tropical areas for over 65 million years which is quite longer than Apis bees (Camargo and Pedro, 1992; Michener, 2000). Despite of diverse flora and fauna of India till now only six species of stingless bees (Hymenoptera: Apidae: Meliponini), belonging to three genera are known from India (Sakagami, 1978; Rasmussen, 2008; Rasmussen and Cameron, 2007; 2010): these are- Tetragonula Bengalensis (Cameron), T. aff. laeviceps (Smith), T. ruficornis (Smith), Lepidotrigonaarcifera (Cockerell), Lisotrigonacacciae (Nurse) and L. mohandasi Jobiraj and Narendran. Recently, T. gressitti is has been observed at Arunachal Pradesh (Rathor et al., 2013).

Stingless bees are opportunistic in nest site selection often utilizing anthropogenic cavity and are very less prone to disease and the sting of these bees are highly reduced which make them suitable as a pollinator in protected condition.In both natural and agricultural environments bees act as key pollinators throughout the world. But in many crops due to mismatch of

size of flowers with that of honey bees (Apis spp.) those cannot extent pollination services to the crop. In those cases alongside with other small sized flower visitors, stingless bees can be right options (Heard, 1999). This is more so, when buzz pollination remains as a requirement. As stingless bees are prevalent in tree cavities, wall crevices, under the soil or other undisturbed places, their domestication is not an easy task. But because of their low swarming tendency and high adaptability they can be domesticated. Therefore, domestication and practicing with the stingless bees needs to have a sound knowledge about their natural habitat, structure of nest, size of colony and positions of the cells inside the colony. Hence, the present study on the habitat nesting and comb geometry of stingless bee T. bengalensis in West Bengal.

MATERIALS AND METHODS

For the study of habitat and comb geometry of T. bengalensis, a survey was conducted in two districts of West Bengal namely, Bankura and Purulia. The locations of surveyed areas of Bankura district are villages Kulmura (23017’7.29”N, 8705’43.92”E) and Palla (23019’9.84”N, 86053’14.6”E); and in Purulia district,

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2 Indian Journal of Entomology Review Article 2020

the villages Kenda (23012’15.1”N,86029’59.3”E), Ramnandapally (23°32’35.8”N, 86°40’22.9”E), Bundla (23°33’27.1”N, 86°41’08.2”E) and Ramchandrapur (23°34’57.8”N, 86°48’24.1”E).

A random survey was done to locate the natural nesting habitat of stingless bees. The colonies from walls and tree cavities were harvested by a method descried herein. A rectangular wooden box of dimension 15x7x8’’ were made. The upper opening of box is provided with a wooden slab of thickness 0.5 inch, adjusted in such a way that it can be easily pulled out and pushed in to open and close the box. Inside the sliding wooden slab a transparent plastic sheet was provided to take the inside observation. Generally, the colony should be harvested in early morning or in late evening to avoid swarming. A simple plastic bottle wrapped with cotton cloth were inserted into the entrance of bee’s colony. The wall or tress branches were gently tapped around the entrance with the help of a stone or stick to transfer the bees inside the bottle. The bottle was then capped and kept under shady place. About 0.25 m2 area around the entrance of bee’s colony on wall and about 0.25 m vertical portion in the tree branches were removed carefully. Small portions of pollen were carefully kept inside the wooden box near the entrance (0.70 cm dia). Few broods (deep yellow coloured and light yellow coloured) were kept next to the pollen pots. Few undamaged honey pots were kept next to the broods. The entrance were gently rubbed with the propolis of harvested colony. The bottle was then fitted to the entrance of the box and taped from bottom end to shift the collected bees into the box. The box was then covered and kept near the colony for one hour to settle the bees. The box was then shifted to the Commercial Apiculture Unit of BCKV, Mohanpur, Nadia for further studies.

To study the nesting habitat, observations onvarious parametersviz., nesting place, number of colonies nested in each place, height from the ground level (cm), cavity dimension i.e., length and width (cm) and length and width of entrance tube (mm)had been taken with simple meter scale from their respective natural habitat. The comb geometry/ internal structure of the nest was studied after removing and opening the colonies from their respective habitats (wall crevices and tree cavities) and parameters observed viz., dimensions of brood cells, pollen and honey pots by using Vernier Calliper; and shape, colour and location of different cells inside the comb were recorded by keen observation through naked eyes.

RESULTS AND DISCUSSION

Nesting habitatsA total of 55 colonies of T. bengalensis nesting in

tree-cavities (Fig. 1a), concrete wall (Fig. 1b), mud wall (Fig. 1c), iron pipe (Fig. 1d) and wash basin (Fig. 1e) were reported. Smith (1954) found T. gribodei Margretti nesting in cavities of tree trunk and branches, similar to the present findings. Similar observations were made by Drumond et al. (1995) who had collected nests of Plebiapoecilochora Moure and Camargo from hollows in house walls and earth banks. Heard (1988) reported Trigona carbonifera Smith to form nests inside hollows in trees or wooden pillars of houses. Pooly and Michener (1969) found T. gribodei nests in cavity of tree trunk and branches. Sakagamiet al.(1983) reported that Tetragonulalaevicepes nested in tree cavities. A colony nested in wall cavity in Bankura district was found to build their nest in a nest of swarmed wasp (Fig. 1f). Rasmussen (2004) also reported that Trigona cilipes establishes its nest within the nest of the social paper wasp Epiponatatua. The present findings added new information of T. Bengalensis forming nests in the others. Darchen (1971) reported such case in T. oyuni Darchen and Sakagami et al. (1989) in T. moorei which nested within the nests of ants, whereas Smith (1945) reported T. denoiti to nest inside the termite mound.

Geometry of comb External: Out of 55 a total of 31 colonies were

selected from their natural habitat to study their nesting habitat and internal geometry in two districts of West Bengal (Table 1). Of these, 14 were reported from Purulia and rest were from Bankura. Out of 14 from Purulia district 7 were found nesting in tree cavities at the mean height of 179.57 cm from the ground level (Fig. 2a) and 7 were found to be nesting in wall cavities at the mean height of 99.71 cm while at Bankura a total of 17 colonies were studied out of which 4 colonies were observed in tree cavities nesting at mean height of 173.25 cm from the ground surface while the remaining 13 were observed nesting in the wall cavities whose mean height was 79.15 cm from the ground surface (Fig. 2b).

In earlier study Danaraddi (2007) reported that the T. iridipennis was found to be nesting in tree cavity and wall cavity at Dharwad. Roopa (2002) and Gajanan et al. (2005) found the similar nesting behaviour of T. iridipennis in Bangalore and Muthuraman (2006) in Tamil Nadu. It was observed that T. iridipennis mostly preferred wall cavities for their nesting site than tree

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Nesting habitat and comb geometry of stingless bee Tetragonula bengalensis smith in West Bengal 3 Gautam Kunal et al.

a b

c d.

e f

Fig. 1: Habitats of T. bengalensis in nature- a. Trees cavity; b. Concrete wall; c. Mud wall; d. Iron pipe; e. Wash basin; f. In swarmed wasp net

Fig. 1: Habitats of T. bengalensis in nature- a. Trees cavity; b. Concrete wall; c. Mud wall; d. Iron pipe; e. Wash basin; f. In swarmed wasp net

cavity or other places. During present study, it was observed that T. bengalensis inhabited mostly in dead and dry date palm trees and at one place a maximum of five colonies could be found out from a single wall. Starr and Sakagami (1987) reported that aggregation of colonies of T. fuscobalteata and T. sapiens Cockerell were found in bamboo cavities. The reasons for aggregation of nests in a site might be due to availability of nesting sites for long duration and short swarming distance.

Colonies nested in tree cavities at Purulia had the

dimensions ranging from 48-70 cm length with a mean of 60.57 cm and the width of the cavity ranged from 18-27 cm with a mean of 22.29 cm whereas the dimension of tree nested colonies in Bankura ranged from 11-25 cm in length with mean of 35 cm and the mean width of 17.75 cm ranging from 11-25 cm. The wall nested colonies in Purulia had the cavity dimension ranging from 17-25.5 cm (Fig. 2c) in length with a mean of 21.21 cm and the width of the cavity ranged from 22- 35 cm with a mean of 30 cm whereas the cavity dimension of colonies nested in walls of Bankura was more or less similar to Purulia which ranged from 17.8-

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Nesting habitat and comb geometry of stingless bee Tetragonula bengalensis smith in West Bengal 4 Gautam Kunal et al.

Nesting place Cavity dimensionsNo. of colony

Height from the ground (cm)

Length (cm) Width (cm) Entrance tube width (mm)

Entrance tube length (mm)

Purulia, West-BengalTree cavity 07 179.57 ± 25.91

(145-215)60.57 ± 8.28

(48-70)22.29 ± 3.25

(18-27)4.21 ± 1.07

(3-6)2.90 ± 0.70

(2-4)Wall cavity 07 99.71 ± 74.22

(32-215)21.21 ± 3.39

(17-25.5)30.00 ± 4.24

(22-35)3.90 ± 1.42

(2-6)3.71 ± 0.86

(3-5)Bankura, West-Bengal

Tree cavity 04 173.25 ± 20.55 (145-192)

35 ± 6.06 (11-25)

17.75 ± 6.40 (11-25)

3.75 ± 0.96 (3-5)

3.5 ± 0.41 (3-4)

Wall cavity 13 79.15 ± 59.14 (33-235)

20.87 ± 2.94 (17.8-25.5)

28.69 ± 4.91 (20-35)

3.98 ± 1.30 (2-6)

3.60 ± 0.87 (2.7-5)

ParameterTree Cavity Wall Cavity

Brood cells Pollen pots Honey Pots Brood cells Pollen pots Honey PotsPurulia, West Bengal Length (mm) 2.26 ± 0.58

(1.34-3.2)7.42 ± 1.04

(5.8-9.2)7.60 ± 0.61 (6.42-8.93)

2.28 ± 0.65 (1.18-3.2)

7.44 ± 1.14 (5.9-9.8)

7.57 ± 0.70 (6.4-8.90)

Width (mm) 1.66 ± 0.37 (0.98-2.5)

5.18 ± 0.88 (3.16-6.36)

4.77 ± 0.57 (3.81-5.9)

1.57 ± 0.42 (0.98-2.5)

4.99 ± 0.96 (3.22-6.33)

4.71 ± 0.69 (3.65-5.94)

Shape Oval Oval Oval Oval Oval OvalColour Brownish Dark brown Dark brown Brownish Dark brown Dark brown

Location Centre Periphery Inner Periphery Centre Periphery Inner PeripheryBankura, WLength (mm) 2.26 ± 0.37

(1.63-2.8)7.44 ± 1.06

(5.85-9)7.80 ± 1.04 (5.64-9.28)

2.13 ± 0.44 (1.37-2.8)

7.30 ± 1.08 (5.3-9)

7.77 ± 0.84 (6-9.30)

Width (mm) 2.00 ± 0.45 (1-2.6)

5.16 ± 1.23 (3.-6.8)

4.99 ± 0.83 (3.8-6.2)

1.77 ± 0.3 (1-2.5)

4.85 ± 0.96 (3-6.89)

4.65 ± 0.69 (3.5-6.2)

Shape Oval Oval Oval Oval Oval OvalColour Brownish Dark brown Dark brown Brownish Dark brown Dark brown

Location Centre Periphery Inner Periphery Centre Periphery Inner Periphery

Table 1. Nesting habitats and comb geometry of nests of T. bengalensis

25.5 cm in length with a mean of 20.87 cm and the width of the cavity had a mean measurement of 28.69 cm which ranged from 20-35 cm. The present findings agree with those of Roopa (2002) who reported a nest width ranging from 210.00- 375.00 mm in Bangalore. Similar results had also been documented by Gajanan et al. (2005).

In some colonies nested in wall cavities, a distinct entrance tube was absent (Fig. 2d) similar result was found by Drumond et al. (1995) who reported that the nest entrance of P. poecilochora had small and circular hole, surrounded by dark resin and with no outer tube. The remaining colonies had a distinct entrance tube. The entrance tube made in the tree cavities in Purulia ranged from 2- 4 mm in length with a mean of 2.90 mm and the width ranged from 3- 6 mm with a mean of 4.21 mm (Fig. 2e) whereas the entrance tube of wall nested colonies ranged from 3- 5 mm and 2- 6 mm in length

and width (Fig. 2f), respectively with a mean length of 3.71 mm and mean width of 3.90 mm. In Bankura the tree nested colonies had a mean entrance tube length of 3.5 mm ranged from 3- 4 mm and the width of entrance tube had a mean of 3.98 mm ranged from 3- 5 mm. On the contrary the entrance made in wall cavities was found to range from 2.7- 5 mm in length and 2- 6 mm in width with a mean of 3.60 mm and 3.98 mm in length and width, respectively.

The present findings contrast with those of Pooley and Michener (1969) who reported that the mean length of entrance tube in T. gribodei was 6.25 mm. The results of Roopa (2002), Gajanan et al. (2005) and Danaraddi et al. (2009) also about the length of entrance tube does not go in the lines of the present finding. Wille (1983) reported that the entrance tube of T. pectaralis might be as long as one m and continued inside the nest cavity along the inner wall of the entrance cavity.

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Nesting habitat and comb geometry of stingless bee Tetragonula bengalensis smith in West Bengal 5 Gautam Kunal et al.

a b

c d

e f

Fig. 2: External comb geometry of T. bengalensis- a. Date tree; b. In wall cavity; c. Dimension of colony in wall cavity; d. No distinct entrance tube in wall nest; e. Distinct entrance tube in Tree nest; f. Length of entracne tube in wall cavity

Fig. 2: External comb geometry of T. bengalensis- a. Date tree; b. In wall cavity; c. Dimension of colony in wall cavity; d. No distinct entrance tube in wall nest; e. Distinct entrance tube in Tree nest; f. Length of entracne tube in wall cavity

Melo (1996) reported that the nest entrance of Melipona capixaba was quite variable, ranging from simple to very elaborate structure.

Internal: All the colonies consisted of brood portion and food storage portion separately (Table 1). Within the food storage part, the honey and pollens were stored in separate pots, but sometimes these pots were often intermixed (Fig. 3a). The present findings are in agreement with those of Dollin (1996) who reported that the honey pots tend to be in the outer parts of nest but often the cluster would have contained both honey

and pollen pots. Such pots are filled with a mixture of wax, resin/ propolis and mud called “cerumen”. The cells inside the comb were interconnected by thin thread like structure made up of cerumen (Fig. 3b). Pooley and Michener (1969) also found that the cells of comb were connected to each other by small pillars or connectives of soft cerumen. Such little grains like cells, were surrounded at their inner and outer edges by the honey and pollen pots. Those were located at the centre of the comb and placed in a cluster that provided sufficient room for easy movement of the bees. The present findings derive support from Roopa

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Nesting habitat and comb geometry of stingless bee Tetragonula bengalensis smith in West Bengal 6 Gautam Kunal et al.

a b

c d

e f Fig. 3: Internal comb geometry of T. bengalensis- a Intermixed (Brood, pollen and honey pots); b Thread like cerumen; c. Brood pots; d. Honey pots; e. Unsealed honey pots; f. Pollen pots

Fig. 3: Internal comb geometry of T. bengalensis- a Intermixed (Brood, pollen and honey pots); b Thread like cerumen; c. Brood pots; d. Honey pots;

e. Unsealed

(2002) who found that the storage pots or food pots were present either above, below or at both sides of the brood area and the whole cluster looked like a densely packed bunch of grapes. In another study, Danaraddi et al. (2007) reported that the brood cells appeared like Jowar grains and were arranged in a network of narrow vertical pillars with horizontal connectives.

The oval shaped brood cells containing eggs and larvae were brownish whereas the pupae cells showed little creamy white in colour and were found to be smaller than the food pots (Fig. 3c). Similar

observations had been made by Gajanan et al. (2005). The brood cells of tree nested colonies measured from 1.34-3.2 mm in length and 0.98- 2.5 mm in width with a mean of 2.26 and 1.66 mm, respectively at Purulia; while at Bankura it ranged from 1.63- 2.8 mm in length and 1- 2.6 mm in width with a mean of 2.26 and 2.00 mm, respectively. The wall nested colonies of Purulia had mean brood cell length of 2.28 mm ranging from 1.18- 3.2 mm and mean width of 1.57 mm ranging from 0.98- 2.5 mm; whereas, colonies at Bankura had mean brood cell length of 2.13 mm ranging from 1.37- 2.08 mm and mean width of 1.77 mm ranging from 1- 2.5

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Nesting habitat and comb geometry of stingless bee Tetragonula bengalensis smith in West Bengal 7 Gautam Kunal et al.

mm. These observations corroborate with those of Roopa (2002).

The honey pots were oval-shaped, dark brown (Fig. 3d) and slightly larger than the pollen pots. Those were mounted at the colony’s inner periphery and were found tooften intermix with pollen pots. In sealed conditions, several pots were found although many were not sealed (Fig. 3e). Generally, bees sealed their honey pots after ripening of stored honey. The honey pots made in the tree cavities in Purulia ranged from 6.42- 8.93 mm in length with a mean of 7.60 mm and the width ranged from 3.81- 5.9 mm with a mean of 4.77 mm whereas for wall nested colonies it ranged from 6.4- 8.90 mm and 3.65- 5.94 mm in length and width, respectively with a mean length of 7.57 mm and mean width of 4.71 mm. Similar results were also found by Roopa (2002). In Bankura the tree nested colonies had a mean length of honey pots was 7.80 mm ranging from 5.64- 9.28 mm and mean width of 4.99 mm ranging from 3.8- 6.2 mm. On the contrary, the honey pots made in wall nested colonies were found to range from 6- 9.30 mm in length and 3.5- 6.2 mm in width with a mean of 7.77 mm and 4.65 mm in length and width, respectively. Likepollen pots, honey pots were also sealed after ripening of honey.

The pollen pots were also oval in shape and dark brown, like honey pots (Fig. 3f). Many of the pots were closed because they were completely filled with pollen, although few were left open to store the incoming pollen loads of foraging bees. In both the colonies, pollen containing pots were found more than the honey pots, suggesting the abundance of pollen plants surrounding the nest habitat. The pollen pots were usually observed at the periphery of the colony surrounding the honey pots. The length and width of pollenpots found in tree nested colonies in Purulia varied from 5.8 - 9.2 mm and 3.16 - 6.36 mm with a mean of 7.42 mm and 5.18 mm, respectively; whereas in wall nested colonies length and width varied from 5.9- 9.8 mm and 3.22- 6.33 mm with mean of 7.44 mm and 4.99 mm, respectively. The pollen pots in Bankura were somewhat smaller as compared to Purulia which was 7.30 mm which ranged from 5.9- 9 mm in length and the width ranged from 3- 6.89 mm with a mean of 4.85 mm. These observations corroborate with those of Roopa (2002). The stored pollens were found to be little wet with sour taste. Similar observations were made by Dollin (1996) who reported that the cluster of the food pots resembled bunch of grapes and contained honey and pollen pots.

Thus, the study of nesting habitats of T. bengalensis reveal that the height of the colonies from the ground level was found to be more in tree nesting colony than that of wall nesting. But the colony found in both tree and wall cavities at Purulia were found to be located at greater height from the ground level than that of Bankura. The colonies found in trees in both the districts having more cavity dimensions than that of wall nested in both length and width whereas the cavity dimensions of tree and wall nesting colonies at Purulia were observed to be more than that of Bankura. The length and width of entrance tubes of the colonies found in tree and wall cavities were approximately similar in both the places. The dimensions of brood cells were found to be least in size in the colony whereas the length and width of honey pots were observed to be more than that of pollen pots in both types of colonies found at both the districts. The brood cells were oval, creamy/brownish and found at the centre of the colony, whereas the pollen pots and honey pots were oval, dark brown in colour and were located at the periphery and inner periphery of the colony, respectively.

ACKNOWLEDGEMENTS

The authors acknowledge the AICRP on Honey Bees and Pollinators, BCKV Centre for providingfinancial and technical facilities. Authors thankthe local people of the survey area for providing necessary support.

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(Manuscript Received: February, 2020; Revised: April, 2020; Accepted: April, 2020; Online Published: May, 2020)

Online published (Preview) in www.entosocindia.org Ref. No. 20049

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