Table Of ContentHYM. RES.
J.
Vol. 15(2), 2006, pp. 307-316
Nesting Biology of a Tropical Myrmicine Ant, Myrmicaria arachnoides
(Formicidae), in West Java, Indonesia
Bakhtiar Effendi Yahya and Seiki Yamane
(BEY, SKY) Department of Earth and Environmental Sciences, Faculty of Science,
Kagoshima University, Korimoto-1, Kagoshima, 890-0065 Japan,
BEY email: [email protected], SKY email: [email protected]
(BEY) Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Kota Kinabalu,
Sabah, Malaysia
—
Abstract. Nesting biology of a myrmicine ant, Myrmicaria arachnoides, was studied on Java
island, Indonesia. The colonies were polydomous and polygynous. Nests ofa colony were located
close to each other on different leaves of a tree. The number of dealated queens was positively
correlated with the numbers of nests, adult workers and immatures, but not with the numbers of
males and new queens present. Adaptive aspects of nesting site, polygyny and polydomy are
discussed.
The genus Myrmicaria is widely distrib- and polygynous colonies but on vegetation
uted in the Old World tropics, i.e., South- (Karawajew 1935, this study, Bakhtiar and
east Asia, South Asia and tropical Africa. Yamane unpubl.). This may allow them to
Emery (1922) sorted Myrmicaria species develop relatively large colonies similar to
into two principal groups based on mor- Polyrhachis species nesting in similar situa-
phology, i.e., the M. brunnea group and M. tions (R. Kohout pers. com.).
arachnoides group, and briefly mentioned Research on colony growth in eusocial
that species of the former construct huge insects has focused on the relationship
nests underground while those of the latter between colony size and productivity. At
construct carton nests on trees. Our ob- each growth period, colonies make invest-
servations have confirmed this for most of ment decisions about whether to produce
the Southeast Asian forms (also see Kar- more workers and grow larger, or rather to
avaiev 1935). However, there are few invest that energy in reproductive output
detailed studies on the nesting habits of (Billick 2001). Because colony size is often
Myrmicaria. In Cameroon, Africa, one the most important factor determining
species, M. opaciventris Emery, belonging reproductive output (Odum and Pontin
to the M. brunnea group, has been in- 1961, Michener 1964, Fowler 1986, Tschin-
tensively studied for its ecology/bionomics kel 1993, Savolainen et al. 1996), maximiz-
including nesting habits (Kenne and De- ing the long-term size of the colony is an
jean 1999, Kenne et al. 2000, 2001). This important component of colony fitness
species has polydomous and polygynous (Oster and Wilson 1978). Colony size is
colonies in the soil. Interconnected nests known to be related with queen number
are built and trenches and tunnels are and also to affect caste/sex composition in
constructed as underground trails connect- ant nests. In this respect nesting behaviour
ing nests. Species of the M. arachnoides of the M. arachnoides group is oi special
group also often construct polydomous interest.
308 Journalof Hymenoptera Research
Little has been studied on the colony RESULTS
—
structure of the Southeast Asian Myrmi- Nesting site and structure. In Salak and
caria arachnoides F. Smith (but see Kara- Salak-Halimun Corridor, colonies were
vtaaixeovno1m9y35)o.f ODruireintnagl oMuyrrmisctaurdiya wone otbh-e afonudnbduisnhepsl,anwthaitlieonianrethaes wboittahniscpaalrsgeatrrdeeens
tained relatively good samples of M. they were found in a forested area with
anersaicah.noiWdees croelpoonritesheorne Jtahvea niessltaindn,g Isnitdeos-, trehleatuinvedleyrshiidgeh torfeesl.eaNveesstsofwevraerilooucsatepdlaonnt
colony size, and reproductive output of species at 0.5-1.5 m above the ground in
these colonies. the former two sites (Fig 1 a,b), but in the
MATERIALS AND METHODS botanical garden they were positioned
m
higher at around 3-4 above the ground.
The taxonomy of Myrmicaria is still In the case of polydomous colonies com-
unresolved, particularly the status of in- ponent nests were generally constructed
fraspecific forms of some species. Myrmi- separately on different leaves of the same
caria arachnoides was originally described plant, but in one case, two nests were built
from Borneo and consists of several 'sub- narrowly connected on one and the same
sspoemcieesw'ou(Blodltobne 1g9o9o5)d, sopfecwiehsi.chThaet floearsmt leaNfe(stBsOGw2e5r-e2m&ad-4e) o(fApcparetnodni-lxik1e).material
studied here is in coloration mostsimilar to (probably chewed plant tissues), flattened
'M. arachnoides arachnoides'.
domes in shape, and greyish brown in
Samples were collected from three dis- colour. Various sizes of nests were built on
turbed sites in West Java in September
42H60a'0l4:iEm,fuoon5t60oCfmoG)r.riS(adBloOarGk3--,SiS1tie0te1&(62138)9,'6SS,a41l5a0'kS6-, nt(hAuepmpeuernnoddueisrxsci1dh).eamWboifetrhvsianrfiotorhueasdnueslsittzs,esatnhoedfreilmewamevare-es
B10o6go3r7'EB,ota7n1i0caml)G(aBrOdGe2n4,- 2S5ite&3(26()6,36a'nSd, gtaulrelse;riessomfoer otfhethmeosevewmoeunltds boef uwsoerdkerass
106 48'E, 220 m) (BOG 38). The distance (Fig. 1 b,c).
between each plant where these colonies The number of nests per colony varied
were found was approx. 5-50 m in Site 1, from 1 to as many as 12; nests of a colony
and 5-10 m in Site 2. These habitats have were usually built close to each other on
been infringed by plantation or agricultur- one plant (approx. 15—-30 cm apart).
al activities or surrounded by residential Colony composition. Dealated queens (Q)
areas. Nests constructed on a same plant were considered to be foundresses or those
were thought to constitute a single colony thathavejoined later (simplycalled 'queens'
as the ants use same foraging trails. Nests hereafter) andwerefound inallcoloniesand
of each colony from different plants were all nests except in BOG18-3 and BOG25-11.
collected intact and put into plastic bags The two queenless nests had immatures,
separately. In total, seven colonies of suggesting the transportation of them from
different sizes were collected. other nests where a laying queen(s) existed.
Nests were measured fortheir maximum The number of queens differed from colony
width and length, and then dissected tocolonyandnesttonest(Fig. 2). Numberof
carefully. Workers, reproductives (dealat- queens is highly correlated with the number
ed queens, young winged queens and of nests (Fig. 3a). The highest mean number
males), immatures (eggs, larvae and pu- of queens per nest, 11.67, was found in a 3-
pae) were counted and preserved in 80% nest colony (BOG18) (range: 0-22) (Fig. 2).
alcohol. Pupae were sorted into sexes and However,BOG38whichalsohad3nestshad
castes as much as possible. the smallestnumber ofqueens ineach ofthe
Volume 15, Number2, 2006 309
Fig. la-d. a, Nests constructed on undersideofleaves,b,Neston the underside a leaf, c,Structureofa nest;
top ofenvelop removed to show the interior, d, Compartments within nest; outerwalls removed.
nests (mean=1.33 with a range of 1 to 2). constitute relatively lower ratios, while
BOG24 (1-nest colony) and BOG3 (2-nest other life forms (stages) were much fewer.
colony) contained relatively small numbers Most of the nests had queen(s), workers
(1-3) of queens per nest. In larger colonies, and immatures, and there was no striking
BOG10 (4-nest colony) and BOG25 (12-nest specialization for a certain nest(s) in a sin-
colony),themeannumberofqueenspernest gle colony with respect to worker/repro-
was larger and relatively stable (4.00 ± 2.16 ductive production. However, BOG18-3,
& 4.00 ± 3.34 BOG38-1, BOG26-3 and BOG10-2 con-
respectively).
Within each colony, workers represent tained higher percentages of workers com-
the greatest number among inhabitants pared with other nests composing these
(approx. 40%) except in BOG 25 and colonies (Appendix 1).
BOG26 (approx. 30.3%), followed by eggs In the single 1-nest colony (BOG24), all
(approx. 20-40%) and larvae (approx. 10- theadults wereworkers (approx. 40% ofall
30%). Worker pupae and male adults the inhabitants) except for three queens
310 Journalof Hymenoptera Research
BOG10
Nests/colony
Fig. 2. Number ofqueens in each nest and colony.
that were possibly egg layers. There were all colonies), and winged adult queens in
numerous eggs (approx. 45%), while larvae colony BOG26 (Appendix 1). As in smaller
and pupae were relatively few, being colonies mentioned earlier, workers again
approx. 10% and 5%, respectively. Further- constitute between 30 and 40% of all
more, all the pupae were workers, all this inhabitants. Following the workers, eggs
suggesting that the colony was in its and larvae also occupied large proportions
ergonomic stage (reproduction and dis- except in BOG18-3, just as in the smaller
persal are not its immediate concern) (cf. colonies. In BOG18-3, more adult males
Oster and Wilson 1978). were observed. Worker pupae were ob-
In the single 2-nest colony (BOG3) all life served to constitute approximately 5% in
stages were present except the pupae of the colonies BOG38 and BOG26.
new queens (Appendix 1). Males had In the 12-nest colony (BOG25; Appen-
started to be produced, representing ap- dix 1), winged queens were seen only in
proximately 5% of the total adults for the two nests in small numbers, while males
colony (Appendix 1). Male pupae also were distributed more evenly; the pupae of
existed in BOG3-1 (Appendix 1). Adult new queens were absent in this colony. For
workers, eggs and larvae each had similar the whole colony, workers, eggs and larvae
percentages for the whole colony and also had approximately the same numbers
for each component nest. (around 30%) while approximately 10%
In the three 3-nest and 4-nest colonies were worker pupae. —
(BOG18, 26 and 38), all life stages were Reproductive output. There were strong
presentexceptthe pupae ofnew queens (in relationships between the number of
Volume 15, Number 2, 2006 311
12000-
312 Journalof Hymenoptera Research
Table 1. Egg:worker (E:W) ratio.
Volume 15, Number 2, 2006 313
ortrees with a group ofworkers. However, reproductives. Furthermore we cannot
we do not have any evidence supporting know how many reproductives have been
this view. produced in a colony at the time of
—
Coloni/ size and reproductive production collection because dispersed individuals
Smaller colonies (eg. BOG24) obviously do not leave any indication of their pre-
had lower numbers of inhabitants (Appen- vious presence in the colony unlike the
dix 1). These may have been at stages just case for social vespids where reproductive
after the colony foundation (cf. Sudd and production can be measured rather pre-
Franks 1987). Data for other colonies show cisely by observing pupal remnants.
that as a colony grows in terms of the
number of nests per colony, the number of ACKNOWLEDGMENTS
colony members increases dramatically
(Appendix 1), as suggested by Oster and We would like to express our appreciation to the
Wilson (1978). In this subsequent stage, Indonesian Institute ofSciences (LIPI) for grantingus
aprnodfitisnfarraestrmuacitnulrey sruec-ihnvaessttehde inneswto(rSkuedrds Wthee paelsromiwsosuilondtloikceatroryexopurtesthsisousrurgvreeyatinapWpreesctiaJtaivoan.
and Franks 1987). Mtor.ouRrorseiscehaornchUhcaoiudnitlelrapharatsndinhiIsndsotnaefsfifa,ropmarMtiucsuleaurlmy
At some critical size, a colony begins to Zoologicum Bogoriense (Cibinong); and also Mr.
produce sexual offspring in order to realize Akhmad Rizali for giving us an immense amount of
its inclusive fitness. In our case even the 2- help during the survey. Our sincere thanks are
mneasltes.coBluotnythhearde waalsreandoyrpelraotdiuoncebdetswoemeen eMxutseenduemd tfoorDrg.iviRnugdyusKovhaoluutabloef tihnefoQrmuaeteinosnlanodn
Polyrhachis nesting habits.
the colony size and number of males
present. For example, in the 12-nest colony LITERATURE CITED
(BOG25), the number of males was rela-
tively small as compared with BOG38 and Aron, S., L. Keller, and L. Passera. 2001. Role of
BOG10 with fewer nests. Several factors resource availability on sex, caste and reproduc-
may be responsible for this. Some males tive allocation ratios in the Argentine ant Eine-
might have already left the nest when it pithema humile. journal of Animal Ecology 70:
831-839.
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Volume 15, Number 2, 2006 315
Appendix 1. Composition of life forms (stages) in colonies and nests.
316 Journalof Hymenoptera Research
Appendix 1. Continued.
Nest
