a captive study on activity-rest, nesting and aggression behavior of an indian ant species,...

8/11/2019 A Captive Study on Activity-Rest, Nesting and Aggression Behavior of an Indian Ant Species, Polyrhachis Lacteipennis (Smith 1858) (Hymenoptera Formicidae

1/8

R E S E A R C H A R T I C L E

A Captive Study on Activity-Rest, Nesting and AggressionBehavior of an Indian Ant Species, Polyrhachis lacteipennis

(Smith 1858) (Hymenoptera: Formicidae: Formicinae)

Ranajit Karmakar Sarmistha Banik

Tanima Biswas Ratanlal Brahmachary

Chitta Ranjan Sahu

Received: 4 May 2011 / Revised: 28 June 2012/ Accepted: 17 July 2012 / Published online: 4 August 2012

Zoological Society, Kolkata, India 2012

Abstract In the present study, the effect of captivity on

the behavioral patterns like activity-rest time budget,nesting, aggression etc. in a group ofPolyrhachis lectei-

pennishas been documented. The ants were collected from

the natural population and maintained in the laboratory

keeping them either in a single formicarium or prepared

formicaria interconnected with tubes to observe the

behavior. About 8 types of behavioral pattern of ants were

noticed. In case of necrophoresis (carrying of dead nest-

mates), the space inside the formicarium was not found to

be sufficient to dispose off the dead nest-mates and there-

fore, all the time the workers carried the dead nest-mates

haphazardly possibly in search of suitable disposal area.

When the worker ants with dead nest-mates were allowed

to leave the formicarium, they left, disposed and came back

to the formicarium. With regard to the nesting behavior, it

was observed that the silk secreting property was not the

only larval character, the adult workers, on the other hand,

could also secrete some amount of silk. When the naturalhibiscus leaf-nest dried up in the formicarium, the workers

along with queens left the natural leaf-nest and make tube-

nest in the formicarium. The workers carried all the larvae

and pupae to the newly forming tube-nest. It was also

observed that sometimes workers took the animal food

inside the tube-nest and sometimes not. They were very

aggressive towards the intruder ants but when fellow

members, separated for several months were re-introduced,

the residents did show almost no aggression. The maxi-

mum number (n = 10) of worker ants were observed as

active during the months of May and June. Afterwards, the

number gradually declined to become lowest (n = 3) in the

month of January. Among the different behavior patterns

studied herein, most of them were found to be innate or

hard-wired, did not require the presence of stimuli like

large sized natural colony, queen, egg, larvae, pupae etc.

inside the formicarium.

Keywords Polyrhachis Captivity Activity-rest

Nesting Necrophoresis

Introduction

Socio-biological works provide important information

about various evolutionary theories and that has been

summed up inAnt Bible (Holldobler and Wilson1990). In

this regard the ants have attracted attention since ancient

times, the study provides useful data for behavioral

observations or bio-geographical subgroup study (Kohout

1990; Leifke et al. 1998). Various authors (Hickling and

Brown 2000; Holldobler and Wilson 2009; Jones et al.

2004; Quinet et al. 2005; Roces 1996) have studied

R. Karmakar (&)

Department of Zoology, Hooghly Mohsin College,

Chinsurah, Hooghly, WB, India

e-mail: [email protected]

S. Banik (&)

Department of Zoology, Bethune College,

Kolkata 700 006, India

e-mail: [email protected]

T. Biswas C. R. Sahu

Department of Zoology, University of Kalyani,

Nadia, Kalyani 741235, WB, India

R. Brahmachary

21B Moti Jheel, Kolkata 700 074, India

1 3

Proc Zool Soc (July-Dec 2012) 65(2):7178

DOI 10.1007/s12595-012-0040-5T

HEZOOLO

GICALSO

CIE

TY

KOLKATA


2/8

different sp. of ants for their interesting behavior. It has

been reported that workers ofCamponotus sp. reveal only

one of the two circadian activity patterns (diurnal or noc-

turnal) suggesting the fact that every worker is hard-

wired for either one or the other (Sharma et al. 2004a,b)

which has later been recognized as an ethological identi-

fier. According to Bingham (1903) two marked worker ants

of Phedole yensis have been found to return to the nestafter detecting food source and they have led the advance

parties for collecting food from that source. Different types

of such behavior is also noticed in Polyrhachis species

including recruitment, aggression, nesting etc. have been

studied in natural population (Robson 2004). In India

*600 species of ants are available (Narendra and Sunil

Kumar2006) and more to be discovered but this rich field

of ant ethology lays unexplored by Indian biologists

excepting the brilliant series of ethological works on

Diacamma vagans (Bhattacharyya1943,1945).

With a view to these facts the present study is designed

with an approach to get in depth insight on the instinctivebehavior of an ant, Polyrhachis lacteipennis during cap-

tivity. The objective is to study (i) necrophoretic behavior

(ii) activity-rest time plan (iii) nest building capacity and

(iv) the aggression behavior particularly during captive

condition, in a population consisting of eggs larvae and

queen etc. Such study will help to generate data on ants

behavior that may further strengthen the existing knowl-

edge about the behavior of the ant and in particular sp.

studied in general.

Materials and Methods

Description

The ant species P. lacteipennis in the present study

belongs to the subfamily Formicinae of the family

Formicidae. It is black in color and can be easily iden-

tified by three pair of spines. The largest pair located in

the petiole takes the shape of abdomen i.e. the spines

curve to the shape of abdomen. Another pair has been

found in the propodeum. The tips of the spines curve

outward. These are moderate sized spines. The last pair of

short and thick spines is divergent, located in pronotum.

The petiolar spines look like horn of a bull and hence

they are commonly termed as Bullhorn ant (Narendra and

Sunil Kumar 2006). The head and the mesosoma are

finely punctured and the surface is granular. The abdomen

is opaque and non-granular as compared to the head and

mesosoma. The body size varies from 5.0 mm to 6.0 mm.

The ant species was identified by Zoological Survey of

India, Alipore, Kolkata.

Preparation of Formicarium

The formicarium was a box made up of transparent Perspex

sheet with a lid of similar material at the top measuring

about 25.5 9 20.0 9 11.0 cm3. There was a circular

opening of about 6.0 cm diameter at the centre of the lid

and the opening was blocked by nylon mesh so that ants

could not come out of the formicarium but there was free

flow of air. The bottom of the formicarium was provided

with a layer of sand which was covered by soil collectedfrom the same locality.

Another formicarium of different pattern was also pre-

pared with same measurement as previous one. It was four

chambered and were interconnected by transparent Perspex

tubes of about 11.5 cm long. For four chambers there were

four such tubes (Fig. 1). The tubes were fitted in such a

way that ants of one formicarium could go to another

through these transparent tubes with ease. The tube mouth

could be blocked by rubber cork at will at both the ends in

each tube. The chambers were numbered I, II, III and IV.

Collection of Ants

From a natural population, the ants (P. lacteipennis) were

collected from a suburban area located 10 km away from

Calcutta, India. In a very close vicinity, the nests were

observed in the hibiscus (Hibiscus rosasinensis), sugarcane

(Saccharum officianarum) plants and also in guava (Psid-

ium guava) trees in the same area. The ants were collected

either with nest or without nest.

Fig. 1 A photograph of four formicaria made up of perspex sheet and

interconnected by perspex material transparent tube

72 Proc Zool Soc (July-Dec 2012) 65(2):7178

1 3


3/8

Laboratory Maintenance

The nest was brought to the laboratory and subsequently

kept in the formicarium and the ants collected without nest

were kept in the formicarium marked IV only to observe

typical movement. They were given sugar and honey

(soaked in cotton), boiled fish and egg and butter at dif-

ferent time as food. Guava or hibiscus leaves were alsoprovided to them in the formicarium. Dead ants of other

species (Camponotus sp.) and live caterpillar found in

marigold flower (monarch butterfly) were also made

available to them. They were regularly given food and

water ad libitum. The nest was sheltered with workers,

larvae, queens etc.

Experimental Procedure

The behavioral study of the ant sp. was conducted within

the single formicarium, formicarium with chamber as well

as in the tube itself for several days to months in a year.The number of sample varied in different experiments. The

experiment was conducted three times to observe the

behavioral pattern of the sp.

Statistical Analysis

All results are presented as mean SD of three replicate

measurements. Statistical analysis of the data was per-

formed by the Students t test using one-way ANOVA

and Tukey test. The data was considered significant either

at p \0.05 or p \ 0.01 and p \0.0001.

Results and Discussion

Activities-1

When the ants (n = 50) were put in a single formicarium it

was seen that they move haphazardly and do not show any

typical notable movement.

When the ants (n = 50) were put in other formicarium

with tube attachment for about 11 days they showed typi-

cal behavior of movement in a regular fashion. Though

sufficient quantities of food were there still death of the

ants occurred in the formicarium and the size of the captive

colony became smaller. Side by side as the number of dead

nest-mates increased, the number of workers with dead

nest-mates also increased. In this situation they always

moved haphazardly within the formicarium. When the

workers were allowed to escape from the formicarium with

dead nest-mates, they quickly came out with the dead nest-

mates and went some 50100 cm away from the formica-

rium and were lost in garbage. Repeated observation

showed that the ants returned to the formicarium without

the dead nest-mates after about 1025 min. The distance

covered by the ants is shown in Table 1. It is seen from the

table that groupB was more efficient in carrying the dead

nest mates, which covered maximum distance. Further,

group B when compared with group A, shows a significant

difference in covering travelling area. It is also evident

from the table that though maximum (102 cm) and theminimum distance (56 cm) was covered by the ants of

group A, group B showed its efficiency in covering mean

distance.

Constant checking of behavior of ants in the formica-

rium, it was seen after 10 days that the ants still remained

under the dried up leaf of guava which was given in the

formicarium. Next 15 days onwards, there were only 710

ants inside and were busy in preparing structure by gluing

the sand particles in the guava leaf (Fig. 2).

Table 1 Distance travelled by the resident worker ants carrying deadnest-mates from the formicarium

No. of ants

(n = 24)

Size of the animal

(mm)

Distance range

(cm)

Mean SD

Gr. A (n = 8) 5.05.5 56102 76.5 16.16*

Gr. B (n = 8) 5.56.0 62101 81.3 11.43*

Gr. C (n = 8) 5.56.5 59100 80.5 14.18

Data represents mean SD of the three replicates

* Significant at p \ 0.05 (ttest)

Fig. 2 A dried up guava leaf from which the mingled sand particles

are hanging. These sand particles are glued together by the adults only

in the formicarium

Proc Zool Soc (July-Dec 2012) 65(2):7178 73

1 3


4/8

Activities-2

When Polyrhachis lacteipennis were collected along with

the guava leaf-nest, it was found that almost every night

they were busy moving inside the formicarium. The nest

collected was in forming stage and there was no egg, larva

or pupa inside but there were only workers. All the workers

(n = 25) were found to be of similar size. No queen(female) caste was recognized inside the nest as the queen

is larger in size as compared to the workers. After about

one and half day, most of the workers were found active in

carrying dead nest-mates as was seen in previous case. The

number of ants carrying the corpse on different days is

given below (Table 2).

It is clearly evident that from day 1 through day 11, the

number of dead nest-mate carrying ants were increasing

gradually. Since the number of dead mates were increasing

day by day the ants were very much active in taking the

corpse outside showing efficient activity of the workers

removing the corpse. When the data were plotted as daysvs number of ants it shows a significant (p\ 0.001) rela-

tionship (R2 = 0.528) (see Plate 1).

Activities-3

The activity and rest time for the ants throughout the

months for 9 months (May 2009Jan 2010) study showed

an interesting result. It was observed that in the month of

June, the maximum number of ants individuals were busy

or highly active outside the tube nest (shown graphically in

Plate2) in foraging or carrying the dead nest mates fol-lowed by May and the lowest number was shown in the

month of January. When the data was considered for the

total activity in days it was seen that in the month of July,

17 (Table3) days were considered as active days. When

the data was analyzed through one way ANOVA (see

Table4) it shows a significant result (p\ 0.0001).

Tukey test also reveals a significant (p\ 0.0001) vari-

ation when the data were compared between months: Jan

versus June, Jan versus May, and Jan versus July. Partic-

ularly, month of May and June when compared with Oct.

Table 2 Number of ants involved in carrying corpse in different days

Days 1 2 3 4 5 6 7 8 9 10 11

No. of ants with

corpse*

4 2 2 4 5 5 5 5 6 5 5

* Group strength (n)

Table 3 Showing number of active days in a month for the ant sp.

(n = 50)

Month May

09

Jun

09

Jul

09

Aug

09

Sep

09

Oct

09

Nov

09

Dec

09

Jan

10

Number ofactive days

8 6 17 9 8 6 6 8 10

Table 4 ANOVA table of the data represented in Plate 2

Source DF Sum of squares Mean squares F Pr[F

Month 8 404.738 50.592 18.626 \0.0001

Error 69 187.416 2.716

Total 77 592.154Plate 1 Relationship between activities on days and the number of

ants

Plate 2 Graphical representation of the activity of the number of ants

with different months of the year (200910)


1 3


5/8

and with Dec. they also showed a significant (p\ 0.0001)

result.

Activities-4

The ants of this species always preferred sugar and honey

to the other food items. When the dead ants of other

species were provided to them, they quickly took theminside. When live caterpillar was given, it was killed and

taken inside the nest. In another experiment, a similar

pattern of behavior was also observed (*40 ants). After

20 days, the leaves of the nest were found dried and curled

up. There were larvae inside the nest. In next 30 days, no

food was given except water and no larva was found live

inside except workers of similar size. No queen was

ascertained inside the nest in formicarium according to the

size. The worker ants, foraged within the formicarium,

were of at least of two sizes. After 30 days, food was

again given to them. The number of individuals reduced to

about 25 members. After 75 days from the start ofexperiment, the leaves of the nest inside the formicarium

further dried up. A paper box of about 7.5 9 5.0 9

5.0 cm of size was placed inside the formicarium. After

7 days, all the worker ants were found to migrate from

dried up leaf nest to the paper box. The lid of the paper

box was not properly fitted and the gap was filled up by

the ants with earthen clod, twig etc. in a similar fashion as

found in the natural nest management. The death toll

increased day by day and the last two live individuals

were observed up to 190 days.

When workers were found foraging, their activity and

body posture were normal but with any disturbance on the

floor or in the formicarium, they became stalled first but

the antennae were moving slowly in the air. Not only that,

the body posture was also changed. The gaster ofOeco-

phylla smaragdina becomes raised when they apprehend

any danger (Holldobler and Wilson1990) but somewhat a

different type of behaviour was observed in the present

study with the apprehension of danger. The gaster became

lowered i.e. they bent their gaster under the body towards

the mouth and the dorsal side of gaster almost touched the

ground. They stayed in that posture for a short while and

after that they fled, entered into the nest or performed

normal activity depending on the degree of danger they

apprehended.

Activities-5

Nests of P. lacteipennis were collected from same natural

population in the month of July 2009 and brought to the

laboratory on the same day and kept in the formicarium no.

I. The ants in formicarium stayed inside the leaf nest. After

about 1 h, few of them came out and started taking food.

After 4 days, one end of the interconnecting transparent

tube between formicarium I and IV (towards the formica-

rium I) was opened up and the other interconnected tube

opening between formicarium I and II remained as it was.

Nothing happened on that day but on the next day, a few

ants were noticed within the transparent tube (810 indi-

viduals). Some other worker ants (23) were also found at

the rim of the transparent tube towards the formicarium I(Fig.3). On 6th day, some ants (56) were found carrying

woodchip, soil particles, dried up leaf materials etc. and

two other ants were found carrying large sized larvae

(Fig.3) to the newly made tube-nest. Another group was

found transporting the larvae into the forming tube-nest; in

the transparent tube. It was noticed that the adult workers

also have the ability to secrete some amount of nest making

silk and while checking for suitability of a future nest, the

workers secrete some amount of silk at the rim site for the

foundation of the nest. But the major work was found to be

performed by large sized larvae. The worker ants held the

larvae from the above towards the forward half of the body.The larva spun its head with repeated forward and back-

ward movement. Gently held larva was then made to touch

the rim surface once and after that the head was retracted.

The phenomenon occurred cyclically for 45 min for that

day before noon. The same activity was repeated on the

next day for the specified period of time. It was observed

utmost two larvae were used for the job and sometimes

there was only one. The nest making activity was observed

for three consecutive days and they were found busy during

first half of the day in silk secretion. Duty like carrying nest

Fig. 3 One Polyrhachis lacteipennis is seen with a larva and other

two are with earthen clod at the future nest entrance of the tube-nest

Proc Zool Soc (July-Dec 2012) 65(2):7178 75

1 3


6/8

materials was generally performed all along the day.

Finally, it took 3 days to almost complete the tube-nest

and in those 3 days, the worker ants transferred all the

larvae, pupae from the drying leaf-nest to the new tube-

nest (probably there were no egg inside the old nest and

that might be due to the fact that queen(s) stopped laying

eggs after removal of leaf nest from the natural host plant).

In those three days time, the tube-nest entrance was notfinished and that duty was still on up to the 4th day. They

made the tube-nest entrance with three almost circular

passages for their entry and exit. It was also found that the

tube-nest entrance was so meticulously made that any

worker with dead nest-mate could not easily enter the nest.

At three occasions, the worker ants with dead nest-mate

tried to enter (inadvertently!) the new tube-nest but failed.

On 21st day, it was observed that there was connection

between the old leaf nest and new tube-nest as workers

were always found running between the two. They were

supposed to doing the transfer of remaining eggs, larvae

and pupae left over there after settling down in the newtube-nest. They were found to maintain the communica-

tion for*30 days from the start of captive study within the

transparent tube. Afterwards we did not notice any such

communication between the two nests. After opening up

the leaf-nest we did not find any remnants of egg, larva etc.

Meanwhile, the corks in the connecting tube between

formicaria I and II were now removed and subsequently the

passage from the formicarium I to the formicarium II was

now opened up. After 3 h, three ants were seen in the

formicarium II. On 5th day some more ants were noticed in

the formicarium II and some of them with dead nest-mates.

On that day, the connection between formicaria II and III

was also opened up. The ants now started to use all the

three formicaria as their foraging ground. In the formica-

rium III more ants were found with dead nest-mates. In

following days the number of dead carrying ants increased

in the formicarium III as compared to the formicarium I

and II.

The ants of formicarium IV were allowed to mix with

their colony members after 8 months. The eight-month

separation between the two groups did not elicit any

noticeable hostility. The residents touched the intruder co-

specifics with the antennae and after sometime the resi-

dents went away.

Activities-6

A totally different scenario was recorded when a Leptog-

enyssp. (live) was introduced into the formicarium I to test

the aggression behaviour ofP. lacteipennis. Immediately

after the intruder ant was recognized, the residents chased

it. The intruder anyway ran away to safer place. The

intruder hid itself sometimes under water pot or under old

natural dried-up nest or under pebbles but 2 days later the

residents finally traced it out and killed it.

In one months time during JulyAugust 2009, the

queen laid a new batch of eggs inside the tube nest . When

larvae were emerged they were observed cling to the upper

lateral wall. Newly laid eggs were not found on the floor of

tube-nest. Almost every week a new batch of eggs clinging

to the upper lateral wall of the tube-nest was observed. Thephenomenon continued till the end of Sep. to early Oct. but

the frequency of laying eggs was slowed down in the

month of September. Some winged and large sized castes

were observed inside. The winged castes were slender and

the larger ones were probably the queens. The number of

large sized castes was more than one inside and they were

never seen outside the new tube-nest. Within 45 days

larvae emerged from eggs and sometimes, the large-sized

larvae (last instar) were used for nest management.

The tube-nest inside the formicarium I was made by

pure larval silk and a thin inner sheet of silk was clearly

seen from outside of the tube-nest.

Activities-7

When they were provided with dead ants of other species

(Camponotus compressusand caterpillar found in marigold

flower) in the formicarium, the Polyrhachis behaved dif-

ferently towards the animal food at different time. Some-

times they took the animal food inside the tube-nest but

sometimes they did not. When there was no egg, larva etc.

inside the nest, the worker ants either took the whole ani-

mal food or part of it inside. But a different scenario

altogether was noticed when there was egg, larva inside the

tube-nest. During that time they never took or allowed to

take the dead animal inside the tube-nest. It was tested

that particular behavior by placing a dead ant ofC. com-

pressusat the entry point of tube-nest for six times and on

each occasion the worker ants inside the tube-nest threw

it away after varying time intervals at different occasions.

Activities-8

The existence of common nest weaving behaviour as found

in O. smaragdina was also discovered in the Asiatic spe-

cies Polyrhachis dives by Jacobson and Wasmann (1905).

The detailed study on the behavior was made by Holldobler

and Wilson (1983). In nature, the ants construct nest among

the leaves and twigs of a wide variety of bushes and trees.

Most of the nests are built in two opposing leaves. Poly-

rhachis ants have never been observed to make chains of

their own bodies or to line up in rows in the manner routine

for Oecophylla (Holldobler and Wilson 1990). Occasion-

ally a single Polyrhachis worker pulls and slightly bends

the tip or edge of a leaf, but ordinarily the leaves are left in


1 3


7/8


8/8

are also found in captivity and that common behavior could

not be altered in captivity. The ants under observation also

developed home-instinct behavior as recorded their return

to the formicarium after disposing off the dead nest-mates.

The present study gives an idea that all the common ant

behavior like nesting, foraging, activity-rest budget pattern,

necrophoresis etc. are innate or hard-wired and did not

change even in captivity. From another experiment, it canbe stated that even a single ant of C. compressus under

observation exhibited the innate behavior of dead nest-

mate removal in a formicarium. This investigation warrants

further study with regard to make solid comment on the

innate or hard-wired behavior of the P. lacteipennis ant

species.

References

Bhattacharyya, G.C. 1943. Reproduction and caste determination inaggressive red ants Oecophylla smaragdina. Transactions of the

Bose Research Institute CalcuttaXVI: 137156.

Bhattacharyya, G.C. 1945/2004. Life history of the wood ant,

Diacamma vagans (trans: Sen, S.). Science and Culture 70:

232235 (in Bengali).

Bingham, C.T. 1903. The Fauna of British India including Ceylone

and Burma. Hymenoptera, ants and cuckoo-wasps. Vol. II.

Taylor and Francis, London.

Hickling, R., and R.L. Brown. 2000. Analysis of acoustic commu-

nication by ants. Journal of the Acoustical Society of America

108: 19201929.

Holldobler, B., and E.O. Wilson. 1983. The evolution of communal

nest-weaving in ants. American Scientist71: 490499.

Holldobler, B., and E.O. Wilson. 1990. The ants. USA: Harvard

University Press.Holldobler, B., and E.O. Wilson. 2009. The superorganism: The

beauty, elegance, and strangeness of insect societies, 5. New

York: W. W. Norton.

Jacobson, E., and E. Wasmann. 1905. Beobachtungen ueber Poly-

rhachis dives auf Java, die ihre Larven Zum Spinnen der Nester

benutzt. Notes from the Leyden Museum 31: 221253.

Jones, T.H., D.A. Clark, A.A. Edwards, D.W. Davidson, T.F. Spande,

and Roy R. Snelling. 2004. The chemistry of exploding ants,

Camponotus spp. (Cylindricus complex). Journal of Chemical

Ecology 30: 14791492.

Kohout, R.J. 1990. A review of the Polyrhachis viehmeyeri, species

group (Hymenoptera, Formicidae, Formicinae). Memoirs of the

Queensland Museum28: 499508.

Leifke, C., W.H.O. Dorow, B. Holldobler, and U. Maschwitz. 1998.

Nesting and food resources of syntopic species of the ant genus

Polyrhachis (Hymenoptera, Formiciade) in west-Malaysia. In-

sectes Sociaux 45: 411425.

Narendra, A., and M. Sunil Kumar. 2006. On a trail with ants: A

handbook of the ants of peninsular India. Bangalore, India:

Tholasi Publication.

Olfer, J. 1970.Polyrhachis simplex, the weaver ant of Israel. Insectes

Sociaux 17: 4952.

Quinet, Y., N. Tekule, and J.C. de Biseau. 2005. Behavioural

interactions between Crematogaster brevispinosa rochai Forel

(Hymenoptera: Formicidae) and two nasutitermes species (Isop-

tera: Termitidae). Journal of Insect Behavior18: 117.

Robson, S.K.A. 2004. Comparative nesting biology of two species of

Australian lithocolous ants: Polyrhachis (Hedomyrma) turneri

Forel and P. (Hagiomyrma) thusnelda Forel (Hymenoptera:

Formicidae: Formicinae). Australian Journal of Entomology 43:

59.

Robson, S.K.A., and R. Kohout. 2007. A review of the nest habits and

sociobiology of the ant genus Polyrhachis Fr. Smith. Asian

Myrmecology 1: 8199.

Robson, S.K.A., and R. Kohout. 2008. Nest construction in arboreal

ant Polyrhachis tubifex Karavaiev. Asian Myrmecology 2:

121123.

Roces, F., and B. Holldobler. 1996. Use of stridulation in foraging

leaf-cutting ants: Mechanical support during cutting or short-

range recruitment signal? Behavioral Ecology and Sociobiology

39: 293.

Sharma, V., S. Lone, A. Goel, and M. Chandrasekhar. 2004a.

Circadian consequences of social organization in the ant species,

Camponotus compressus. Naturwissenschaften 91: 386390.

Sharma, V., S. Lone, D. Mathew, A. Goel, and M. Chandrasekhar.

2004b. Possible evidence for shift work schedules in the media

workers of the ant species. Chronobiology International 21:

297308.

Yamauchi, K., Y. Ito, K. Kinomura, and H. Takamine. 1987.

Polycyclic colonies of the weaver ant Polyrhachis dives.Kontyu

Tokyo55: 410420.


1 3

a captive study on activity-rest, nesting and aggression behavior of an indian ant species,...

Documents