HYM. 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. was collected. Furthermore we do not Billick, I. 2001. Density dependence and colony know whether males are produced growth in the ant species Formica neorufibarbis. throughout the year or during restricted journal ofAnimal Ecology70: 895-905. seasons. Bolton,B. 1995.A newgeneralcatalogueoftheantsofthe The increase in queen number may zvorld. Harvard University Press, Cambridge and London. increase the size of the colony, and finally Djieto-Lordon, C. and A. Dejean. 1999. Tropical the number of reproductives. Although in arboreal ant mosaics: innate attraction and im- this study queen number positively affect- printingdeterminenestsiteselectionindominant ed the number of workers and immatures, ants. Behavioral Ecologi/ Sociobiology45: 219-225. the relationship between queen number Elias, M., R. Rogengren, and L. Sundstrom. 2005. naontdpnoseiwtivqe.ueTehnis(aslhsoowsmalthea)t nouthmebrefracwtoarss Syternauosnuocsnoarpulompp.uollBayetdhiaoovmniyoorafanltdhEeucnoirlceoodgloywnioSaoolcdiitoyabinintolaoFgpoyorlmiy5cg7a-: may have operated in determining the 339-349. development of queens and males as Emery, C. 1922. Hymenoptera, Fam. Formicidae, reported for many other genera (for the subfam. Myrmicinae. Pp. 95-206 in: Wytsman, Argentine ant, see Aron et al. 2001). In M. P., Genera insectorum. Fasc. 174B. araclinoides maintaining a large worker Hanstehne,UnL.itDe.dSatnadtesJ.anHd.CKalnoatdz.a.2C00o5m.stCaorcpkenPtuebrliansthsinogf force on one tree itself may be important Associates, Ithaca and London. under certain conditions (e.g., presence of Holldobler, B. and E. O. Wilson. 1990. The ants. competitors) at the cost of producing more Harvard University Press,Cambridge. 314 Journalof Hymenoptera Research Karavaiev, W. 1935. Neue Ameisen aus dem Indo- Majer,J. 1993. Comparison ofthearboreal ant mosaic Australischen Gebiet, nebst Revisioneiniger For- in Ghana, brasil, papua New Guinea and Aus- men. Treubia 15: 57-117. tralia - its structure and influence on arthropod Keller, L. 1995. Social life: the paradox of multiple- diversity. Pp. 115-141 in: LaSalle, J. and I. D. queencolonies. Trendsin Ecologx/andEvolution 10: Gauld eds. Hymenoptera and Biodiversity. CAB 353-360. International, Wallingford. Kenne, M. and A. Dejean. 1999. Spatial distribution, Oster,G. F.and E.O.Wilson. 1978.Casteandecologyin size and density of nests of Myrmicaria opaciven- the social insects. Princeton University Press, tris Emery (Formicidae, Myrmicinae). Insectes Princeton. Sociaux46: 179-185. Savolainen, R., K. Vepsalainen, and R. J. Deslippe. Kenne, M., B. Schatz, R. Feneron, and J. L. Durrand. 1996. Reproductive strategy of the slave ant 2000. Changes in worker polymorphism in Formica podzolica relative to raiding efficiency of Myrmicaria opaciventris Emery (Formicidae, Myr- enslaver species. Insectes Sociaux43: 201-210. micinae). Insectes Sociaux47: 50-55. Sudd, J. H. and N. R. Franks. 1987. The behavioral Kenne,M.,B. Schatz,R. Feneron,and A. Dejean.2001. ecologyofants. Chapman and Hall, New York. Hunting efficacy of workers from incipient Tschinkel, W. R. 1993. Sociomerryand sociogenesis of colonies in the myrmicine ant Myrmicaria opaci- colonies of the fire ants Solenopsis invicta during ventris (Formicidae: Myrmicinae). Sociobiology 37: one annual cycle. Ecological Monographs 63: 121-134. 425-457. 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