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J. Env. Bio-Sci., 2018: Vol. 32 (1):115-120 ISSN 0973-6913 (Print), ISSN 0976-3384 (On Line)

DESCRIPTION OF THE NEST AND THE IMMATURES OF MEGACHILE STUDIOSA


BINGHAM (HYMENOPTERA, APOIDEA, MEGACHILIDAE)
Damayanti Patel, *Abhishek Rajpurohit, Imran, Habib Pathan and Rajiv K. Gupta
Department of Zoology, Jai Narain Vyas University, Jodhpur-342001, Rajasthan, India.
*Department of Zoology and Environmental Sciences,
Lachoo Memorial College of Science and Technology (Autonomous), Jodhpur, Rajasthan, India.
(Corresponding author email: abhiraj_rajpurohit@yahoo.com)

Received: 19-05-2018 Accepted: 29-05-2018


Three nests of Megachile studiosa Bingham (Hymenoptera, Apoidea, Megachilidae) were collected from a natural crevice made
amidst stones in a wall in the Department of Zoology, JNV University, Jodhpur, during rainy seasons of year 2017. They had leaf
cutting coating outside their nest chambers. Each nest consisted of 3 chambers, making a count of total 09 chambers. They
contain larvae, pupae and, two chambers were occupied by pupae of cleptoparasites. Later these also emerged to adults and
were identified as those of Brachymeria spp (Hymenoptera: Chalcididae). The paper describes the nest, immature stages and
developmental consequences of the broods collected out of the nests of M. studiosa. A pair of adult bees which were reared from
the nest shall be deposited at the national museum of Zoological Survey of India, Kolkata as Lactotypes.
Key words: Megachile studiosa, Brachymeria sp., Lactotypes

Bingham (1898a) described Megachile studiosa from Shimla MATERIALS AND METHODS
(H.P.) later, Gupta (1993) Gupta and Yadav (2001), Gupta et
Nesting site: The nesting site was located at Zoology
al. (2003), recorded it from Nagaur and Sikar in Rajasthan and
Department, J.N.V. University, New campus, Jodhpur. The nest
Deesa in Gujarat respectively. Present paper describes nesting
was found in natural crevices formed between stones at the
biology of this leaf-cutting bee from Jodhpur city of Western elevation of the departmental building. The height of nest was
Rajasthan, India. The nesting biology studied and presented about 162 cm from ground level. Temperature was recorded
in this paper has reared two females and three males from the as 32oC during excavation of the nest. The nest elevated angle
nests described in following pages. The male recorded hitherto of about 60o-65o.The direction of nest opening was facings
constitutes the very first record for M. studiosa. The pair reared towards southeast. No direct sunlight rays fell on the nest but
out of its nest in laboratory has been designated as Lactotypes entrances received light that was an indication that bee activity
that will be placed at the Zoological Survey of India, Kolkata. initiated with sunrise. The activity of this bee was kept under
This is very first collection record of M. studiosa from Jodhpur. observations from 9.30 AM to 04:30 PM.
The bees identified belong to Megachile studiosa with sub OBSERVATIONS
generic placement under Eutricharaea Thomson. Bees
Three nests of M. studiosa were observed on July 27th of year
incoming subgenus Eutricharaea are well known for use of
2017.
leaf cutting to line their nests such as recently observed in
case of M. gathela Cameron (Gupta and Yadav, 2001). The The bees were busy constructing their nests in the cervices
Megachile contains six common, mostly wide- ranging species and they continued to work for around three weeks. Their
that nest in a wide variety of substrates and are among our activities were closely observed and information were noted
best-studied leafcutter bees, especially M. brevis (Michener, down .The bee was seen to enter the nest at 08:10 a.m., with
1953b; Pengelly, 1955). Megachile texana nests in the soil leaf cuttings that was gathered in their mandibles and front
and under rocks. Hicks (1926) described nest-digging behaviour two pairs of legs. After 3 minutes, it again came out of the
and nest structure; Krombein (1953) described two nests and nest for second trip. Thereafter 2-3 weeks later, bee was
later (1970) described nest-searching behaviour and Pengelly observed at guard chamber opening with no forage activities
(1955) extensively described nest structure, behaviour of that was the sign of completion of nests. Later on 22-8-2017
females, development of immature. nests were excavated out of crevices and the collected material
NAAS Rating (2017)-4.43
PATEL, RAJPUROHIT, PATHAN AND GUPTA (116)

was brought to the laboratory along with the trapped adult Nesting sites of M. studiosa are mainly found in walls of picked
bees. stone and sometime hallow cervices at two stones and any
other crevice. In summary, almost any short of small crevice
The nests excavated from wall were slightly variable in their
may be used for nesting by this bee. When there are several
pattern of linear cell arrangement. It was probably due to
nests, of course, the first constructed nest is the one farthest
availability of vacant space between the masonry stones. M.
from the entrance into crevice while the last constructed is the
studiosa is known to have cutting edges in between their
one closest to the entrance. Mainly 3-4 nests were found at
innermost space of teeth in tetra-dentate mandibles. Males
one place and each nest consisted of 3-4 cells. Another aspect
were never observed sharing the work of nest construction. It
about nest shape was that as per available space of various
was only female that continued the efforts for approximately
crevices between stones. Usual length of crevice occupied by
10-12 days. Normally one cell was constructed in a single
the nest was measured from 6 cm to 10 cm. At one place,
day as in the case of M. brevis and M. gathela (Michener,
nests were constructed in a continuous one by one condition,
1953b),
i.e. one built after other and nearby other, no overlapping of
Structure of the nest: Overall nine chambers were collected nests were found in any place. The direction of their opening
out of three nests. The outermost chamber, termed as guard was found towards east or southeast. Nest was situated at an
cell, was found occupied by the owner female and often by its average height of 1.5 m from the ground. Tables 1 to 3
companion male. Plenty of provision deposit was observed in summarize the various characters and dimensions of the
the guard chamber and it was plugged from behind so that collected nests.
remaining posterior part of the nest had no internal connection
Contents of the nest: The content used in the construction of
with the guard cell. The outermost layer of chambers had small
a nest chamber was slightly mud, often mixed with coarse
leaf pieces of Verbicinia encicloides and some of the Tephrosia
and fine pebbles. M. studiosa larvae produce variable amounts
hamictoni. Gupta (1993) recorded that female collected nectar
of silk during the feeding period of the last instar to separate
and pollens from two referred species of Angiosperms(Fig.1-
the fecal pellets from the food. The female deposited leaves
3).
and petals fillers mixed with mud and mandibular secretions
The pattern of arrangement of leaf cutting in the cell was different in spiral pattern arrangement. The sizes of the leaves used in
at place such as the small leaf cutting at base of cell had a different parts of cell cap construction may vary. Cap and plug
broad concavity at center. Normally 2-4 leaf pieces were portion were mainly constructed by round cutting of leaves.
arranged at the base, having a deep concavity at centre. The
One larva, two pupa and three mature adult bees were collected
cylinder part of the chamber contained leaf pieces normally 8
from 9 chambers/cells from a nest until 5 September 2017.
to 14 in numbers. They were arranged in semi lunar whorl
The cocoons of Megachilids were rigid and composed of
patterns and sometimes loosely fitted superficially in some
several layers. In addition, many live parasite larvae and pupae
chambers. The cap part of chamber had 3-4 leaves pieces
were also collected from several cells.
which were arranged in a concavo-convex pattern like the basal
part of the chamber. Identical arrangement on each cell wall Immature stages: Description of first/second instar
and base and apices were observed in every nest. larva: Fig. 4-6 show a second instar larva of M. sttudiosa.
Total body length was 4.0 mm with maximum width of head
The chamber walls (of the cylinder) consisted of flat leaf cutting
1.0 mm and median length of head was 1.01mm.
slightly bent to accommodate lateral pieces. Almost similar
Head: Shape of head was almost triangular with broadly bulging
arrangements of leaf cutting were recorded in all nest chambers.
ocular lobes. The dorsal surface with a few bristles; lateral
However cap portion of chamber had less number of leaves
lobes emarginated concealing a narrow invagination at center;
cutting so that emerging adult had to make lesser efforts in
mandible slightly elongated, bidentate, massive, half dentate
excavating out from cap of the nest chamber and it was
margin pigmented; mandibular lower tooth slightly exceeding
observed latter also (Batra, 1976).
upper in length, dentate margin smooth; maxillae elongated,
massive; sub basal line pigmented; labio-maxillary region
(117) DESCRIPTION OF THE NEST AND THE IMMATURES OF MEGACHILE STUDIOSA

Table 1: Observations on different Nest Characters - Nest No. 1

S.N o. N es t cha ra cters C ha m ber - 1 C ha m ber - 2 C ha m ber - 3

1. O u ter w id th o f t h e n es t cell (cm) 0 .4 2 0 .4 6 0 .4 4

2. In n er d iam eter o f ch am b ers (cm) 0 .5 1 0 .5 3 0 .5 0

3. Wid th o f i n n er cell u l ar s ep t a (cm) 0 .11 0 .1 3 0 .1 2

4. In n er len g th o fcel l (cm) 0 .6 9 0 .7 1 0 .6 7

5. O u t er l en g t h o f cell (cm ) 0 .7 4 0 .7 6 0 .7 3

6. L arv a(L ), Pu p a(P), Paras it e larv a(PL ) P P PL

Table 2: Observations on different Nest Characters - Nest No. 2


S.No. Nest characters Chamber - 1 Chamber - 2 Chamber - 3

1. Outer width of the nest cell (cm) 0.45 0.41 0.43

2. Inner diameter of chambers (cm) 0.55 0.58 0.57

3. Width of inner cellular septa (cm) 0.10 0.13 0.11

4. Inner length of cell (cm) 0.68 0.70 0.71

5. Outer length of cell (cm) 0.72 0.74 0.74

6. Larva(L), Pupa(P), Parasitelarva(PL) P P P&PL

Table 3: Observations on different Nest Characters - Averages of nest dimensions


S. N o. N e st ch ar act ers N e st-1 N e st- 2 N est- 3

1. N e st angle f r om hor izonta l 65° 64° 65°

2. Total length of tube up to the end ( cm ) 3 2.8 3. 1

3. I nner dia m ete r of tube ( cm ) 0. 62 0.63 0.61

4. N umber of ce ll in eac h ne st 3 3 3

5. O uter w idth of the ne st ce ll ( cm ) 0.4 0.38 0.40

6. Aver age w idth of inner ce llular se pta ( cm ) 0.1 0.09 0.11

7. Ave r age inner dia m ete r of c ha mber ( c m) 0. 51 0.55 0.52

8. Aver a ge inne r le ngth of cham be r ( cm ) 1.1 1.25 1.09

9. Aver a ge oute r le ngth of cham be r ( cm ) 1.3 1.4 1.25

10. D istance of ne st entr ance f r om w all ( c m ) 1.2 2.1 3. 2

11. Ave r a ge w e ight of eac h cha m ber ( gr a m) 0.1018 0.1280 0.1251

12. N est height f r om gr ound level ( cm ) 162 162 162


PATEL, R AJPUROHIT, PATHAN AND GUPTA (118)

slightly concealing the ventral margin of mandible. Maxillae with cell cap. Normally nest construction lasts for 2-3 weeks.
slightly elongated, cardo and stipes smooth without setae; Many species nest in the thatched roofs, wooden walls, on
maxillary palpi prominent; labium divided into a prementum rocks, stone cervices and smaller aggregation in the ground.
and postmentum; hypopharynx finely bilobed, salivary slit On the basis of their nesting activities, megachilids can be
transverse. Antennal papillae longer than median width and divided into three groups: lodger, burrowing and mason bees.
arise from broad and strong produced protuberances; frons Most species are lodger bees and females make nest in pre-
with median setae. Ocular lobes are strongly protuberances. existing cavities. They are known to construct nest in the
passages of xylophagous beetles, in old nests of bees and
Body: Body with dense setae all over the dorsal surface;
wasps in wood and in the soil, in hollow dry stems and
Intersegmental lines easily defined and dorsal tubercles feebly
branches, in the galls of Cynipoids, and they readily accept
markable in between thoracic segments 2 and 3, thoracic
artificial nesting sites.
segments 3 and 1st abdominal segment, abdominal segments
1 and 2, 2 and 3, 3 and 4. Dorsal tubercles are reduced in Megachile studiosa is a lodger bee that uses leaves and petals
between segment 4 and 5 and after 5 segment no markable of plants to construct its nest chambers. It belong to subgenus
tubercles found. Spiracles in each segment found; last anal Eutricharaea. The characters of difference in nest as well as
segment with anus transverse slit at the posterior-dorsal in the larva are clear, compared to the pre described Megachile
surface(Fig. 7-12). cephalotes which belong to subgenus Callomegachile. These
differences can establish the phylogenic relationships between
Description of pupa (male): Total body length was 8 mm;
two subgeneric categories by adopting a multi-dimensional
maximum width of head was 2.5 mm, median length of head
approach.
was 3 mm.
The fine lining secreted by Dufour's gland (Cane and Michener,
Head: Antennal scape without median tubercle, instead finely
1983) helped in protection. The problem of parasitic infestation
striated; vertex dorso-laterally with dense setae; frons without
is a serious matter for survival rate of species. The periodicities
a pair of low tubercles in front of the summit of each eye; all
of development of bee larvae and those of cleptoparasites were
ocelli prominent and bulging; clypeal apically margin at centre
noted as significant regulatory aspect controlling the population
deeply invaginated, baso-centrally with a projecting edge;
of bees and should be further studied. The huge infestation by
vestibule quite long.

Thorax: Lateral angles of pronotum finely produced to acute


spine just before tegulae, posterior-lateral lobes not produced;
scutum with two median sized patches of erect bristles
localized to either side of mid-line, non-tubercles; scutellum
broadly rounded posteriorly; metanotum with a fine conical
projection at centre; wing without any median tubercle; coxae
with a distinctly projecting spine; trochanters.

Abdomen: On terga 3-6 basal margin of apical rim with a row


of erect prominent bristles.

Material examined : First and second instar larva and pupae.

RESULTS AND DISCUSSION

The female of Megachilidae constructs typical nests. She builds Fig. 1-3: Nest structure of Megachile studiosa.
and provisions her own nest. The chambers are built [1- Three chambers of single nest,
successively one by one; she fills them with provision for their 2- Third chamber of nest, 3- Cell cap.]
offspring's and lays one egg in each cell. The cells are closed
(119) DESCRIPTION OF THE NEST AND THE IMMATURES OF MEGACHILE STUDIOSA

Fig. 4-6: Second instar Larva [4- Body, lateral view, 5-


Head, front view, 6- Head, lateral view.]
Abbreviation used- Lab- Labium,
Tub- Tubercles, Man- Mandible
Fig. 9

Fig. 7 Fig. 10

Fig. 8 Fig.11
PATEL, RAJPUROHIT, PATHAN AND GUPTA (120)

2. Batra, S. W. T. (1977a). Bees of India (Apoidea), their behaviour,


management and a key to the genera. Orient. Insec., 11(3/4): 289-
324.
3. Bingham, C. T. (1898a). On some species of Indian Hymenoptera.
Jour. Bomb. Nat. Hist. Soc. 12: 115-130.
4. Cane, J. H. and Michener, C.D. (1983). Chemistry and function of
mandibular gland products of bees of the genus Exoneura
(Hymenoptera:Anthophoridae). Jour. Chem. Ecol. 9(12):1525-
1531.
5. Gupta, R. K. (1993). Taxonomic studies on the Megachilidae of
northwestern India. Scientific Publishers (India). (Reprint 1999). v
+1-294 p.
6. Gupta, R. K. and Yadav, S. ( 2001). Apoidean species composition
Fig. 12: on Crotalaria juncea L., Cajanus cajan (L.), Helianthus annus L.,
and Brassica compestris L. var sarson Prain in eastern Rajasthan,
micro-hymenopterous wasps restricted the adult emergence India (Hymenoptera). Opuscula Zoological Fluminensia 198: 1-
of bees in fields. 10.
7. Gupta, R. K., Naval, R.K. and Charan, S.K. (2003). Nesting biology
of the solitary bee, Megachile (Pseudomegachile) creusa Bingham
ACKNOWLEDGEMENTS (Hymenoptera: Megachilidae). Mellifera 3(5): 11-18 (Turkish) & 43-
50 (English).
The authors are thankful to Dr. L.S. Rajpurohit, Head, 8. Hicks, C.H. (1926). Nesting habits and parasites of certain bees of
Department of Zoology, Jai Narayan Vyas University, Jodhpur Boulder County, Colorado. Univ. Colo. Stud. 35:217-252.
(for providing the necessary research and laboratory facilities) 9. Krombein, K.V. (1953). A note on the nesting habits of Megachile

and all seniors colleagues of wild bee laboratory, J.N.V. texana Cresson (Hymenoptera: Megachilidae). Proc. Entomol. Soc.
Wash. 55:84-85.
University, Jodhpur.
10. Michener, C. D. (1953a). Comparative morphological and systematic
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1. Batra, S. W. T. (1976). Nests of Ceratina, Pithitis and Braunsapis 11. Michener, C.D. (1953b). The biology of a leafcutter bee (Megachile
from India (Hymenoptera: Anthophoridae). Orient. Insec. 10: 1-9. brevis) and its associates. Univ. Kans. Sci. Bull. 35: 1659-1748
12. Pengelly, D.H.(1955).The biology of bees of the genus Megachile
with special reference to their importance in alfalfa seed production
in southern Ontario. Ph.D. diss., Cornell Univ., 269 pp.

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