nesting and colony structure in the giant forest ant, camponotus gigas

nesting and colony structure in the giant forest ant, camponotus gigas nesting and colony structure in the giant forest ant, camponotus gigas

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Orr et al.: Camponotus <strong>gigas</strong> <strong>colony</strong> <strong>structure</strong><br />

AREA AND METHODS<br />

The study was carried out <strong>in</strong> primary lowl<strong>and</strong> mixed dipterocarp <strong>forest</strong> near <strong>the</strong> Kuala<br />

Belalong Field Studies Centre, Temburong District, Brunei. Nests were located by present<strong>in</strong>g<br />

forag<strong>in</strong>g <strong>ant</strong>s with a piece of tuna, <strong>and</strong> follow<strong>in</strong>g <strong>the</strong>m back to <strong>the</strong>ir nests. Although C.<br />

<strong>gigas</strong> is known to be polydomous, a previous study (Orr & Charles, 1994) suggested that<br />

most nests (85%) <strong>in</strong> this area, which occurred at a density of 9-15 per hectare, were associated<br />

with a dist<strong>in</strong>ct <strong>colony</strong> as <strong>the</strong>re was little exchange of marked <strong>ant</strong>s between different nests.<br />

At <strong>the</strong> nest chosen for study, diel activity patterns were determ<strong>in</strong>ed by cont<strong>in</strong>uously monitor<strong>in</strong>g<br />

all <strong>ant</strong>s enter<strong>in</strong>g <strong>and</strong> leav<strong>in</strong>g <strong>the</strong> nest, <strong>and</strong> <strong>the</strong> extent of <strong>the</strong> terrestrial forag<strong>in</strong>g territory was<br />

established by follow<strong>in</strong>g marked <strong>ant</strong>s on <strong>the</strong>ir forag<strong>in</strong>g excursions. Subsamples of 40 <strong>and</strong> 75<br />

<strong>ant</strong>s were marked ei<strong>the</strong>r yellow or white dur<strong>in</strong>g <strong>the</strong> day, <strong>and</strong> night observation periods<br />

respectively. On 25/8/94 at 0800 h, a time at which activity was m<strong>in</strong>imal <strong>and</strong> most <strong>colony</strong><br />

members were expected to be <strong>in</strong>side <strong>the</strong> nest, all entrances were sealed <strong>and</strong> approximately<br />

11 of ethyl acetate was <strong>in</strong>troduced <strong>in</strong>to <strong>the</strong> nest. At 1400 h <strong>the</strong> nest was opened by saw<strong>in</strong>g<br />

<strong>the</strong> log <strong>in</strong>to sections approximately 30cm <strong>in</strong> length us<strong>in</strong>g a cha<strong>in</strong>saw. Any <strong>ant</strong>s which had<br />

not succumbed fully to <strong>the</strong> fumes of <strong>the</strong> ethyl acetate were sprayed with a pyrethr<strong>in</strong> based<br />

aerosol <strong>and</strong> <strong>the</strong> section with its contents was sealed <strong>in</strong> a large plastic bag. These were<br />

subsequently opened <strong>in</strong> <strong>the</strong> laboratory on a plastic sheet <strong>and</strong> all contents removed <strong>and</strong> placed<br />

<strong>in</strong> alcohol.<br />

RESUL TS AND DISCUSSION<br />

Activity patterns. - Activity patterns were similar to those reported by Orr & Charles<br />

(1994) for o<strong>the</strong>r nests <strong>in</strong> <strong>the</strong> area. Up to 144 foragers emerged from <strong>the</strong> nest <strong>in</strong> <strong>the</strong> even<strong>in</strong>g,<br />

versus only 29 dur<strong>in</strong>g <strong>the</strong> day. Temporal specialization <strong>in</strong> forag<strong>in</strong>g was also noted i.e. daymarked<br />

<strong>ant</strong>s were subsequently observed only forag<strong>in</strong>g by day <strong>and</strong> night-marked <strong>ant</strong>s were<br />

observed only forag<strong>in</strong>g by night. On 3 Jul. 1994 heavy ra<strong>in</strong> at 1900 h completely <strong>in</strong>hibited<br />

activity. Thirty-one marked <strong>ant</strong>s which had left <strong>the</strong> nest earlier <strong>in</strong> <strong>the</strong> even<strong>in</strong>g did not return<br />

until <strong>the</strong> follow<strong>in</strong>g even<strong>in</strong>g, but were not observed forag<strong>in</strong>g by day. It is probable that many<br />

sheltered under leaves on <strong>the</strong> <strong>forest</strong> floor, as five were located do<strong>in</strong>g this, but at least <strong>in</strong> two<br />

<strong>in</strong>stances, <strong>the</strong>y were observed enter<strong>in</strong>g a second, nearby small nest at <strong>the</strong> base of a large tree.<br />

This nest probably did not conta<strong>in</strong> a queenright <strong>colony</strong>, <strong>and</strong> was apparently a satellite of <strong>the</strong><br />

nest be<strong>in</strong>g studied s<strong>in</strong>ce <strong>ant</strong>s marked at <strong>the</strong> ma<strong>in</strong> nest were observed to emerge from it, <strong>and</strong><br />

it was completely unoccupied three weeks after <strong>the</strong> ma<strong>in</strong> nest was destroyed. Such satellite<br />

nests were not detected <strong>in</strong> an earlier study (Orr & Charles, 1994) <strong>and</strong> might arise temporarily<br />

to exploit a food source associated with <strong>the</strong> crown of a particular tree, while m<strong>in</strong>imiz<strong>in</strong>g<br />

travel over <strong>the</strong> ground at night which might expose <strong>the</strong>m to predation risks. As previously<br />

found by Orr & Charles (1994) only a small m<strong>in</strong>ority (6%) of <strong>ant</strong>s return<strong>in</strong>g were carry<strong>in</strong>g<br />

food, mostly fragments of diverse fungal material.<br />

Nest location <strong>and</strong> <strong>structure</strong>. - The nest (Fig. 1) was located <strong>in</strong> a fallen log 7m <strong>in</strong> length,<br />

<strong>and</strong> vary<strong>in</strong>g <strong>in</strong> diameter between 25 <strong>and</strong> 30cm. It lay at an <strong>in</strong>cl<strong>in</strong>ation of 15° with <strong>the</strong> nest<br />

entrance at its lower end, thus ensur<strong>in</strong>g that water could not enter <strong>the</strong> cavity. The lower part<br />

of <strong>the</strong> log was almost completely hollow with a large open<strong>in</strong>g to <strong>the</strong> exterior on its ventral<br />

surface. This had been almost completely covered by a wall of mud 2-3cm <strong>in</strong> thickness with<br />

two entrance holes (6cm X 2.4cm <strong>and</strong> 2.8cm X 2.2cm respectively) on ei<strong>the</strong>r side of<strong>the</strong> log.<br />

The ma<strong>in</strong> cavity of <strong>the</strong> nest, which extended back for some l.4m, was deeply grooved with<br />

considerable qu<strong>ant</strong>ities of decay<strong>in</strong>g wood fragments <strong>and</strong> some mud. Lead<strong>in</strong>g off this cavity<br />

248


THE RAFFLES BULLETIN OF ZOOLOGY 1996 44(1)<br />

ma<strong>in</strong><br />

chamber<br />

brood<br />

chamber<br />

Fig. 1. Sectional view of <strong>the</strong> nest (rotated up through 30°), show<strong>in</strong>g <strong>the</strong> ma<strong>in</strong> chamber, 'brood' chamber,<br />

o<strong>the</strong>r passages. The entrances <strong>in</strong>dicated are located <strong>in</strong> a wall of mud seal<strong>in</strong>g most of <strong>the</strong> hollow end<br />

of <strong>the</strong> log.<br />

from its <strong>in</strong>nermost recesses were two narrow tunnels, one exp<strong>and</strong><strong>in</strong>g almost at once <strong>in</strong>to<br />

what is presumed to be <strong>the</strong> brood chamber, judg<strong>in</strong>g from <strong>the</strong> number of eggs found <strong>the</strong>re,<br />

(approximately 13cm X 4cm X 2.5cm), <strong>the</strong> o<strong>the</strong>r cont<strong>in</strong>u<strong>in</strong>g <strong>in</strong>to <strong>the</strong> heart of <strong>the</strong> log for at<br />

least ano<strong>the</strong>r metre <strong>and</strong> end<strong>in</strong>g bl<strong>in</strong>dly without widen<strong>in</strong>g. There was no evidence of fungal<br />

culture with<strong>in</strong> <strong>the</strong> nest, hence fungus ga<strong>the</strong>red is probably used only for immediate food<br />

requirements.<br />

Colony composition. - Although no queen was recovered, <strong>the</strong> presence of eggs <strong>and</strong> young<br />

larvae <strong>in</strong> what was apparently a brood chamber suggests that it was a queenright <strong>colony</strong> <strong>and</strong><br />

<strong>the</strong> queen had ei<strong>the</strong>r escaped or had been destroyed by <strong>the</strong> cha<strong>in</strong> saw as <strong>the</strong> log was opened.<br />

A total of 834 <strong>ant</strong>s were recovered from <strong>the</strong> nest, <strong>in</strong>clud<strong>in</strong>g 45 majors (5.4%). S<strong>in</strong>ce this<br />

count was made when most foragers had probably returned to <strong>the</strong> nest this is a good estimate<br />

of <strong>colony</strong> size except<strong>in</strong>g a few escapees <strong>and</strong> <strong>in</strong>dividuals which may have been stationed at<br />

a nearby satellite nest. Based on observed activity at this latter nest it is thought that it probably<br />

conta<strong>in</strong>ed fewer than 100 <strong>ant</strong>s.<br />

As <strong>in</strong>dicated by Fig. 2 <strong>the</strong> size frequency distribution of workers was bimodal, reflect<strong>in</strong>g<br />

<strong>the</strong> existence of two subcastes of workers (i.e. majors <strong>and</strong> m<strong>in</strong>ors). However <strong>the</strong> size frequency<br />

II..<br />

o<br />

c::<br />

W<br />

!Xl<br />

::E<br />

:J<br />

Z<br />

1.6<br />

HEAD CAPSULE WIDTH (MM)<br />

Fig. 2. Size frequency distribution of major (right) <strong>and</strong> m<strong>in</strong>or (left) workers based on head capsule<br />

width.<br />

249


THE RAFFLES BULLETIN OF ZOOLOGY 1996 44(1)<br />

LITERA TURE CITED<br />

Chung, A.Y.C. & M. Mohamed, 1993. The organization <strong>and</strong> some ecological aspects of <strong>the</strong> Gi<strong>ant</strong><br />

Forest Ant, Camponotus <strong>gigas</strong>. Sabah Soc. J., 10(1): 41-55.<br />

Dartigues, D. & L. Passera, 1979. Polymorphisme larvaire et chronologie de l'apparition des castes<br />

femelles chez Camponotusaethiops Latrielle (Hymenoptera, Formicidae). Bull. Soc. Zool. Fr., 104(2):<br />

197-207.<br />

Holldobler, B. & E.O. Wilson, 1990. The <strong>ant</strong>s. Spr<strong>in</strong>ger-Verlag, Berl<strong>in</strong>, Heidelberg. xii + 732 pp.<br />

Orr, A.G. & 1.K. Charles, 1994. Forag<strong>in</strong>g <strong>in</strong> <strong>the</strong> <strong>gi<strong>ant</strong></strong> <strong>forest</strong> <strong>ant</strong>, Camponotus <strong>gigas</strong> (Smith)<br />

(Hymenoptera: Formicidae): evidence for temporal <strong>and</strong> spatial specialization <strong>in</strong> forag<strong>in</strong>g activity. J.<br />

Nat. Hist., 28(4): 861-872.<br />

Porter, S.D. & W.R. Tsch<strong>in</strong>kel, 1985. Fire-<strong>ant</strong> polymorphism: <strong>the</strong> ergonomics of brood production.<br />

Behav. Ecol. Sociobiol. , 16(4): 323-336.<br />

Tho, Y. P. , 1981. The Gi<strong>ant</strong> Forest Ant. Nat. Malay. , 6(4): 32-35.<br />

Wilson, E.O., 1978. Division oflabour <strong>in</strong> flIe <strong>ant</strong>s based on physical castes (Hymenoptera: Formicidae).<br />

J. Kansas ent. Soc., 51(4): 615-636.<br />

Wilson, E.O., 1985. The sociogenesis of <strong>in</strong>sect colonies. Science, 228: 1489-1495.<br />

251<br />

Received 27 Apr 1995<br />

Accepted 21 Nov 1995

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