ASIAN MYRMECOLOGY Volume 1, 81 – 99, 2007 ISSN 1985-1944 © S.K.A.ROBSON & R.J. KOHOUT

A review of the nesting habits and socioecology of the genus Fr. Smith

SIMON K.A. ROBSON1* & RUDOLF J. KOHOUT2

1School of Marine & Tropical Biology, James Cook University Queensland 4811, Australia *Corresponding author’s email : [email protected]

2Biodiversity Program, Queensland Museum, PO Box 3300, South Brisbane Queensland 4101, Australia

Abstract. We provide a summary of the nesting and socioecological habits of 197 species of Polyrhachis . Nesting habits range from subterranean to terrestrial, lignicolous, lithocolous and arboreal with varying levels of consistency at the subgeneric level. Species are predominantly either subterranean or terrestrial nesting within the subgenera Campomyrma, Chariomyrma and Hagiomyrma; mainly arboreal nesting within Aulacomyrma, Cyrtomyrma, Hedomyrma, Hemioptica, Myrmatopa and Myrmothrinax; lignicolous within Hedomyrma, and variable within Myrma, Myrmhopla and Polyrhachis. The majority of species within a particular subgenus demonstrate a consistent nesting type but a few species can be highly variable, demonstrating both subterranean and arboreal habits. The inclusion of larval silk in the nests is correlated almost entirely with arboreal nesting, but the presence of spider silk in the nests of at least four species suggests caution when inferring silk origin. Pupal cocoons are present in all but Cyrtomyrma and Myrmatopa. Polydomy and polygyny are known from a number of species and may be more common in arboreal nesting species, but limited data sets prevent definitive interpretations at this time.

Keywords: Polyrhachis, Formicidae, nesting ecology, socioecology, mating systems

INTRODUCTION Studies of the socioecology of ants have provided an important framework for the testing The Old-World ant genus Polyrhachis Fr. Smith and development of numerous evolutionary includes approximately 500 described species theories (Hölldobler & Wilson 1990) and with organised into 12 subgenera, and is the subject of improved phylogenetic knowledge ants are extensive and ongoing taxonomic revisions becoming even more significant model systems to (Bolton 1973; Dorow 1995; Kohout 2006b). The explore the evolution of a variety of ecological, taxonomic diversity of the genus is matched with social. and behavioural characters (Brady et al. an equally diverse socioecology. Nesting habits, 2006; Moreau et al. 2006). The ant genus for example, range from subterranean and Polyrhachis is ideally suited to such approaches terrestrial, to lignicolous, lithocolous and arboreal due its highly diverse socioecological habits, but (Robson & Kohout 2005). Nest walls can include unfortunately much of the information on these both larval and spider silk (Robson 2004), and nests attributes of the genus is located in equally can even be found in such unusual habitats as scattered sources. There exists a variety of intertidal mangrove mud (Nielsen 1997) or within ecological or behavioural studies summarising the colonies of other ant species (Maschwitz et particular biogeographical subgroups or ecological al. 2003). communities (e.g. Lieke et al. 1998; Kohout 1999) 82 Nesting habits of Polyrhachis and the socioecology of a number of species has stems, bamboo internodes, the base of ferns in been studied in detail (e.g. Ofer 1970; Yamauchi trees, cavities under bark, or within myrmecophyte 1987; van Zweden et al. 2007), but much valuable plants), lithocolous (on or within rock crevices) or information remains embedded within the primary arboreal (among leaves or twigs, either against a taxonomic literature. In combination with ongoing single surface such as a trunk or between adjacent taxonomic revisions, this makes it difficult to surfaces such as leaves) sites. In a single case, determine the socioecological state of individual ‘laboratory’ was used to describe the nesting species and subgenera, and the potential location of a captive but originally subterranean significance of new descriptions to our colony. understanding of the evolution of these traits as a whole. The determination of the number of colonies and queens per nest was made with reference to The goal of this paper is to review the entire published literature and our own field studies. literature on Polyrhachis in order to consolidate Polydomy was inferred if either workers were our knowledge of the nesting habits and observed moving between nests or the transfer socioecology of the genus. We hope this of workers between nests did not invoke information and the conclusions we draw from it aggressive interactions. Polygyny was inferred if will provide a solid basis for ongoing more than one dealate queen was observed in a socioecological and phylogenetic studies, and single nest. In a single case this interpretation prompt fellow researchers to further the collection was confirmed by molecular studies (van Zweden of such information. et al. 2007).

MATERIALS AND METHODS Additional nest material was classified as either silk (typically flat sheets thought to arise from Information on the nesting and socioecological larval silk glands), carton (fine plant material), characteristics of Polyrhachis was derived from spider silk (silk known to be removed from spider both the primary literature and our own webs or egg sacks) or none (where the absence of unpublished field observations and museum either silk, carton or spider silk was confirmed). All inspections. In order to optimise the information types were described if more than one type of presented in the face of ongoing taxonomic nesting material was found in a single nest. revisions, records are provided only for described species. Further information will be provided on The presence or absence of pupal cocoons is additional species once they are described. noted, based on existing literature or direct observations of field and laboratory colonies. The taxonomic classification, spelling of species epithets and authors’ names follows that The source of the information typically established by Bolton (1995a) with additions from indicates the earliest published reference to the Kohout (2006b, 2006a). Informal species groups character state of a particular species. We have within Myrma follow those proposed by Bolton deliberately restricted the sources to those that (1973) and Kohout (2006a), while species groups specifically refer to an actual identified species, within Myrmhopla and Polyrhachis follow those and give precedence to published records over of Dorow (1995). our own unpublished observations. The following abbreviations are used in addition to literature Individual nests were described as being citations where required, and refer to reliable located in either subterranean (in the soil with or sources such as field notes, laboratory without any cover such as rocks or fallen logs), observations or information accompanying terrestrial (on the ground beneath logs and stones, museum specimen label data: BBL (Bede B. in or between grass stems), lignicolous (within the Lowery,unpublished), JRF (John R. Fellowes, cavities of living or dead plants, including hollow unpublished), MCZC (Museum of Comparative Zoology, Harvard University, Cambridge, Mass., S. Robson & R. J. Kohout 83

USA), NMNH (National Museum of Natural of P. schoopae moving into an abandoned History, DC, USA), SKR (Simon K.A. Robson, Polyrhachis turneri Forel nest. Pupal cocoons are unpublished), RJK (Rudolf J. Kohout, present in the eight Chariomyrma species for unpublished). In all cases, the absence of which this trait has been described. information means that a particular character state has been neither explicitly described in the Within the subgenus Cyrtomyrma, all species literature, nor determined through our own form arboreal nests, all nests include larval silk unpublished studies. (though some such as Polyrhachis australis Mayr and Polyrhachis pilosa Donisthorpe are known RESULTS to also include spider silk), and all larval appear to pupate without cocoons. Both monodomous and Nesting and socioecological data were determined polydomous nests, and single vs. multiple queen from 197 of the approximately 500 described colonies, have been described. species of Polyrhachis. Representative species were included from all 12 subgenera, but the extent Within the subgenus Hagiomyrma, the to which each subgenus is sampled is variable majority of species are subterranean or terrestrial, (Table 1). Records are particularly sparse from the with a few lignicolous species. There is no evidence smaller subgenera (Aulacomyrma, Hemioptica), for the incorporation of silk into their nest perhaps due to their limited distribution and structures, and all larvae pupate in cocoons. relative rarity. Nothing is known of the queen number or colony structure. Within the subgenus Aulacomyrma, all three species for which nesting habits have been Within the subgenus Hedomyrma, the majority described are lignicolous. The nests of P. dohrni of species are lignicolous, with a few species also Forel were lined with silk and cocoons have been nesting in the ground or inhabiting nests confirmed in two species. constructed of spider silk on the side of rock walls. Colonies are monodomous in the single species Within the subgenus Campomyrma, the for which information is available, P. turneri. The majority of species are subterranean-nesting, presence of silk is variable and all larvae pupate in though lignicolous and arboreal nesting are known cocoons. in two and one species respectively. Silk has been described in the field nest of only a single species, Within the subgenus Hemioptica, all species Polyrhachis phryne Forel, though a captive colony are arboreal nesting, incorporate silk into their of Polyrhachis hexacantha Erichson lined their nests, and appear to produce pupal cocoons. There test-tube nest with silk. Pupal cocoons have been is support for monodomous nesting habits in a described in 12 species. Nothing is known of the single species, Polyrhachis bugnioni Forel. queen number or colony structure of any species in this genus. Nesting and socioecological habits within the subgenus Myrma are extremely variable, and there Within the subgenus Chariomyrma, the appears to be a loose correlation between the majority of species are subterranean-nesting but various species groups and their habits. All species a few species have been described with within the alexisi and monista species groups, for lignicolous nests and with one exception silk has example, are arboreal nesting, while nesting habits never been found as part of their nests. The single range from terrestrial to lignicolous and arboreal exceptional species, Polyrhachis schoopae Forel, within the relucens species group. Nests can has been found in a silk nests on the sides of rock include either carton, silk or neither, though the faces, but based on the type of nests and the presence of pupal cocoons appears consistent identity of other Polyrhachis species at the same within the subgenus. Both monodomous and locality we believe this unusual example is a case 84 Nesting habits of Polyrhachis polydomous nests, and single vs. multiple queen DISCUSSION colonies, have been described. The diversity of nesting habits described within Within the subgenus Myrmatopa, all species Polyrhachis ants is in accordance with previous have arboreal nests, and with the exception of one descriptions of the nesting habits of the genus as variable species, Polyrhachis schang Forel, all a whole (Dorow 1995), but a number of previously species can include silk in their nests. The larvae undetected patterns are now evident. Arboreal of this subgenus appear to pupate without nesting appears to be universal in species of the cocoons. subgenera Aulacomyrma, Cyrtomyrma and Myrmatopa and almost so in Hedomyrma and Nesting and socioecological habits within the Myrmothrinax, where a number of species have subgenus Myrmhopla are also extremely variable. adopted lignicolous nesting habits. Species of Nest habits range from subterranean to lignicolous Campomyrma, Chariomyrma and Hagiomyrma and arboreal, and at least one species (Polyrhachis are predominantly subterranean-nesting, with a few lacteipennis Fr. Smith) appears capable of nesting species in each subgenus utilising alternative nest in all three localities. The incorporation of silk (and sites. Within the subgenus Campomyrma at least occasionally carton) appears to occur in all arboreal one species, P. halidayi, can be found in arboreal species. Both monodomous and polydomous nests and a number of undescribed species are nests, and single vs. multiple queen colonies, have known to be lignicolous in the Australian tropics been described, and all larvae appear to pupate in (C. Burwell, personal communication, 2006). cocoons. The viehmeyeri species group stands Similarly in Chariomyrma, a few species occupy out as an almost entirely parasitic group, with no lignicolous or lithocolous nest sites. Myrma and evidence for the incorporation of silk into their (or Myrmhopla stand out among the Polyrhachis their hosts) nest. subgenera for the diversity of nesting habits, ranging from subterranean to arboreal and in the Within the subgenus Myrmothrinax, the case of Myrmhopla, even as parasites within the majority of species are arboreal nesting, with two nests of other ant species. This diversity is perhaps species, Polyrhachis eudora Fr. Smith and not surprising given that Myrmhopla is most likely Polyrhachis ternatae Karawajew, forming polyphyletic (Dorow 1995). The subgenus lignicolous nests. At least two species are known Polyrhachis is predominantly an arboreal-nesting to form polydomous colonies, and silk and pupal group, but at least one member of the bihamata cocoons are in present in all nests described. species group (P. bihamata Drury), and the single member of the lamellidens-group (P. lamellidens Nesting and socioecological habits within the Fr. Smith) can nest in the soil (Table 1). subgenus Polyrhachis are variable, including examples of subterranean, terrestrial, lignicolous Information on the presence of silk in the nests and arboreal nesting. Nesting habits of of Polyrhachis is less detailed than knowledge of Polyrhachis lamellidens Fr. Smith appear highly their nest locations, but a number of general variable, including all but lithocolous habits. patterns can be discerned. The incorporation of Examples of polydomous and both monogynous silk into the nests is restricted almost entirely to and polygynous colonies are known; silk is arboreal-nesting species, though it has been incorporated into the nest of arboreal nesting recorded occasionally in lignicolous species. Silk species in which nest-wall structure was described, has never been reported in species of the two but appears to be absent in the case of the ground- predominantly ground-nesting subgenera, nesting P. olybria Forel. All larvae appear to pupate Chariomyrma or Hagiomyrma, and within those in cocoons. subgenera containing a diversity of nesting types it is restricted almost entirely to lignicolous or arboreal-nesting species. The presence of silk in a S. Robson & R. J. Kohout 85 laboratory colony of the normally soil-nesting P. Wilson 1977). Similarly, knowledge of the presence (Campomyrma) hexacantha and “woven of polydomous or polygynous nests is extremely material”in the soil-nesting P. (Myrma) esarata limited and it is not yet possible to draw ecological Bolton suggest that silk production may be conclusions. Arboreality seems to be the most facultative or even maintained in a number of common nesting habit of those species described subterranean species. as being either polydomous or polygynous (Table 1) but this may reflect the ease with which arboreal Despite the consistent pattern of nest habits nests can be sampled, rather than an ecological in some subgenera, caution still needs to be correlate of arboreality per se. exercised when reviewing published descriptions of nest habits. Donisthorpe (1941c, 1941a, 1943), In conclusion, the ant genus Polyrhachis for example, appears to have used “carton” and demonstrates a significant diversity in nesting “silk” almost synonymously when describing habits but much information remains to be Polyrhachis nests and so it is not always clear if gathered. The recent descriptions of inter-tidal “carton” implies the absence of silk, as it is nesting habits (Nielsen 1997) and the occupation typically interpreted to mean at present. The of nests on rock faces constructed entirely of location of described nest material should also be spider silk (Robson 2004) suggest that unique noted, as carton can also be present at the habits remain to be discovered. Polyrhachis ants entrance of a number of lignicolous and comprise a significant component of the Southeast subterranean-nesting species (Table 1). The partial Asian ant fauna (Bolton 1995b) and play an inclusion of spider silk in P. (Cyrtomyrma) important role in a variety of environmental studies australis, P. (Cyrtomyrma) pilosa and P. (Myrma) in this region (Yamane et al. 1996; Brühl et al. 1998; laboriosa Fr. Smith and the construction of silk Maryati Mohamed 1998; Hashimoto et al. 2000), nests made entirely of spider silk in P. yet we have knowledge of the nesting habits of (Hedomyrma) turneri also raise the possibility that less than half of the described species. Future “silk” nests need not always imply the use of larval studies that report on the presence and type of silk only. We suspect that most published silk, the presence or absence of larval cocoons, descriptions of silk nests imply a larval origin, as and the number of queens in sampled colonies, the fluffy spider-silk nests of P. turneri are quite would be extremely informative. different from the larval-derived flat sheets of silk found in the nests of most weaving Polyrhachis ACKNOWLEDGEMENTS (Robson 2004), but field observers should be aware that multiple sources of silk exist. We greatly appreciate the many individuals, institutions and funding agencies that have Many important socioecological aspects of the supported our work. In particular, we acknowledge genus are currently poorly known. The absence Prof. Seiki Yamane, Prof. Maryati Mohamed and of pupal cocoons appears limited to species in the fellow members of ANeT, the Australian Research subgenus Cyrtomyrma and Myrmatopa (Table 1; Council (for support to SKAR), the Australian Kohout 1999), but cocoon status in the majority Biological Resource Study (to RJK), an Ernst Mayr of species remains to be described. This is Fellowship (to RJK), James Cook University and unfortunate as the loss of cocoons has been the Queensland Museum. correlated with the evolution of nest-weaving within ants as a whole (Hölldobler & Wilson 1983). Species that produce the most extensive silk nests, such as Oecophylla smaragdina (Fabricius), pupate without cocoons, while species such as Dendromyrmex chartifex (Fr. Smith), considered to represent the lowest grade of nest weaving, continue to produce cocoons (Hölldobler & 86 Nesting habits of Polyrhachis

BBL Polyrhachis

subterranean Kohout (2000) subterranean Kohout subterranean lignicolous present RJK lignicolous present

Forel, 1902 subterranean present RJK Santschi, 1920 Erichson, 1842 Erichson, subterranean BBL André, 1896 subterranean BBL Forel, 1915 Clark, 1930 Clark, lignicolous Kohout (2000) Wheeler, 1911 1911 Wheeler, Emery, 1921 Emery, Wheeler, 1916 subterranean Wheeler (1910) Fr. Smith, 1858 lignicolous none Kohout (2000) Emery, 1887 Emery, subterranean Kohout (2000) Kohout, 2007 lignicolous present Kohout press) (in Emery, 1889 Emery, arboreal (1903) Bingham Mayr, 1862 1862 Mayr, subterrranean entrance carton SKR Emery, 1897Emery, subterranean Kohout (2000) Mayr, 1866 Mayr, subterranean Wheeler (1915a) Santschi, 1920 Santschi, 1920 subterranean silk Brown, pers.comm. cited in Mayr, 1876 Mayr, subterranean (1915a) Wheeler Mayr, 1876 Mayr, subterranean RJK Forel,1907 subterranean Kohout (1990) Forel, 1901 1901 Forel, lignicolous silk Kohout press) (in Clark, 1930 subterranean RJK Kohout, 2007 Kohout, lignicolous present Kohout press) (in : Nesting and social habits of the ant genus Forel, 1907 Forel, sidnica semipolita schwiedlandi prometheus phryne patiens ops micans macropa maculata insularis hexacantha hirsuta gravis fuscipes flavibasis creusa creusa femorata orokana wardi dohrni ulacomyrma Table 1 Table

subterranean subterranean present present RJK RJK (1967) Hung

subterranean present RJK

subterranean subterranean none present SKR RJK

halidayi lignicolous / subterranean present JRF laboratory silk present RJK

lignicolous present RJK subterranean (1915a) Wheeler present RJK Campomyrma subterranean present RJK

Subgenus Nest Multiple Multiple Additional Pupal Source Source Pupal Additional Multiple Multiple Nest Subgenus Species A location nests/colony? material queens/colony? cocoons nest S. Robson & R. J. Kohout 87 Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple Mayr, 1876 1876 Mayr, subterranean (2000) Kohout Mayr, 1876 1876 Mayr, subterranean (1990) Taylor & Kohout Kohout, 2006 Kohout, arboreal silk absent Kohout (2006b) Kohout, 2006 2006 Kohout, arboreal silk absent (2006b) Kohout Kohout, Kohout, 2006 arboreal silk absent Kohout (2006b) Emery, 1897 1897 Emery, subterranean (1996) & Anderson Clay Forel, 1902 Forel, lithocolous Kohout (1999) Forel, 1895 subterranean Kohout (1990) Forel, 1915 Forel, 1902 Forel, 1902 subterranean Kohout (1988b) Kohout, 2006 arboreal silk absent RJK Mayr, 1870 arboreal silk Turner (1897) Forel, 1895 subterranean BBL Forel, 1915 1915 Forel, Emery, 1897 1897 Emery, lignicolous RJK Lowne, 1865 1865 Lowne, subterranean (1990) Kohout Emery, 1887 Emery, arboreal silk Donisthorpe (1938b) Forel, 1895 subterranean BBL Forel, 1902 Forel, 1902 subterranean lignicolous / Kohout (1988a) Forel, 1902 Forel, 1902 lignicolous present RJK Donisthorpe, 1943 1943 Donisthorpe, lignicolous Donisthorpe(1943) Mayr, Mayr, 1876 subterranean RJK Forel, 1901 subterranean Kohout (2000) Emery, 1895 Emery, subterranean lignicolous / Kohout (1988a) Forel, 1879 1879 Forel, lignicolous / subterranean (1988a) Kohout Species Location Nests/C decumbens debilis brevinoda sokolova vermiculosa abbreviata aporema australis schoopae senilis punctiventris nitens obscura lownei limbata lata heinlethii hookeri constricta cyrus gab templi aurea Subgenus Nest Mul

arboreal no silk absent SKR arboreal arboreal arboreal yes yes silk silk (1915b) absent (1994) Wheeler Hinman spider silk & Ebert (1994) Dwyer subterranean present SKR Cyrtomyrma present RJK present RJK

terrestrial / subterranean (2000) Kohout present RJK present RJK subterranean lignicolous none none present RJK present SKR present RJK subterranean present RJK Chariomyrma 88 Nesting habits of Polyrhachis Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple Roger, 1863 Roger, 1863 subterranean BBL Donisthorpe, 1944 1944 Donisthorpe, lignicolous present (1990) Kohout & Taylor Karawajew, Karawajew, 1927 subterranean/often in Mayr, 1876 1876 Mayr, / lignicolous terrestrial (1905) Froggatt Fr. Smith, 1858 Fr.Smith, 1858 arboreal carton Karawajew (1928) Kohout, 2006 arboreal silk absent RJK Kohout 2006 arboreal silk absent Kohout (2006b) Forel, 1902 1902 Forel, lithocolous carton Robson (2004) (Latreille, 1802) 1802) (Latreille, arboreal silk Turner (1897) Wheeler, 1911 Forel, 1916 Forel, subterranean Kohout (1988c) Donisthorpe, 1938 Donisthorpe, arboreal silk Robson Kohout (2005) & Forel, 1915 Forel, 1915 arboreal silk (1915a) Wheeler Forel, 1886 Forel, 1886 subterranean Kohout (1988c) Kohout, 2006 arboreal silk absent RJK Forel, 1897 subterranean present Kohout (2000) Forel, 1902 1902 Forel, subterranean present Kohout (1988c) Kohout, 2006 arboreal yes silk absent Kohout (2006b) (Fabricius, 1775) subterranean present RJK Fr. Smith, 1860 1860 Smith, Fr. subterranean Donisthorpe (1943) Kohout, 2006 Kohout, 2006 arboreal silk absent Kohout (2006b) Fr. Smith, 1863 1863 Fr. Smith, lignicolous present Kohout (1988c) Donisthorpe, 1938 1938 Donisthorpe, arboreal silk (1990) Kohout & Taylor Forel, 1902 Forel, subterranean present Kohout (1988c) Species Location Nests/C delecta ammon ammonoeides laevissima mackayi monteithi rastellata kyawthani robsoni dorsena angusta crawleyi denticulata lignicolous RJK RJK lignicolous lachesis lydiae metella paxilla schenkii semiaurata thusnelda agiomyrma Subgenus Nest Mul H

pilosa arboreal arboreal arboreal arboreal silk silk silk (1929) absent SKR Karawajew absent SKR spider silk Dwyer Ebert & (1994)

arboreal carton / silk Mann (1919)

cocoons RJK arboreal arboreal yes silk no (2006b) Kohout silk absent SKR yorkana arboreal arboreal yes yes yes silk silk vanet Zweden(2007) al. Heinze & Hölldobler (1993)

stumps oldtree rotting present Kohout (1999)

subterranean present RJK

semiobscura terrestrial (1922a) Wheeler present RJK SKR no lignicolous present carton S. Robson & R. J. Kohout 89

RJK RJK

Cocoons

Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple

lignicolous

Emery, 1897 Emery, Forel, 1893 1893 Forel, lignicolous present SKR Forel, 1902 Forel, 1902 lignicolous Mann, 1919 subterranean Kohout (1988c) Mann (1919) Forel, 1907 Forel, lignicolous (1988c) Kohout Forel, 1915 1915 Forel, lignicolous Kohout (1988c) Forel, 1908 1908 Forel, arboreal no silk present Norden & Krombein Emery, 1895 Emery, arboreal silk Kohout (1988c)

Mayr, 1876 1876 Mayr, lignicolous (1897) Turner Forel, 1902 Forel,1902 subterranean present RJK Forel, 1915 1915 Forel, Fr. Smith, 1860 1860 Smith, Fr. lignicolous (1990) Taylor Kohout& p Fr. Smith, 1860 1860 Fr. Smith, arboreal (1871) Smith Dorrow & Kohout, 1995 1995 & Kohout, Dorrow arboreal silk present Kohout (1995) & Dorow Roger, 1862 1862 Roger, Forel, 1895 1895 Forel, lithocolous silk spider (2004) Robson Mayr, 1876 Mayr, subterranean present RJK Mayr, 1876 Mayr, lignicolous (1897) Turner Mann, 1919 1919 Mann, lignicolous / subterranean (1919) Mann Roger, 1862 1862 Roger, arboreal silk (1892) Wroughton Forel, 1902 1902 Forel, lignicolous no present Kohout (1988c) Forel, 1902 lignicolous present RJK rou Billberg, 1820 Billberg, g - i Species Location Nests/C boltoni bugnioni scissa annae argentosa atropos clio cupreata daemeli erato fervens malaensis melpomene mjobergi ornata rufifemur terpsichore turneri tubifera yrma edomyrma emioptica Kohout (1995) & in Dorow cited

lignicolous lignicolous lignicolous lignicolous none present carton SKR RJK (2005) Kohout & present Robson SKR lignicolous lignicolous lignicolous lignicolous present RJK no (2005) Kohout present & SKR Robson

lignicolous lignicolous present RJK present RJK lignicolous lithocolous no silk spider present SKR present RJK trapezoidea H H M

alexis Subgenus Nest Mul 90 Nesting habits of Polyrhachis

Wheeler (1922b) (1922b) Wheeler Wheeler (1922b) (1922b) Wheeler Cocoons

Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple arboreal arboreal , cited (1892) Emery

André, 1889 1889 André, lignicolous (1973) Bolton Chapman, 1963 1963 Chapman, lignicolous (1963) Chapman

Gerstäcker, 1859 1859 Gerstäcker, subterranean silk

Kohout, 2006 2006 Kohout, present RJK Santschi, 1910 arboreal Bolton (1973) Fr. Smith, 1858 1858 Smith, Fr. arboreal silk (1973) Bolton Emery, 1892 Emery, Stitz, 1911 1911 Stitz, / terrestrial lignicolous RJK Emery, 1925 1925 Emery, lignicolous carton (1944), Eidmann (Fabricius, 1782) 1782) (Fabricius, lignicolous none (1973) Bolton André, 1889 1889 André, arboreal (1973) Bolton Santschi, 1910 1910 Santschi, arboreal silk (1973) Bolton Bolton, 1973 1973 Bolton, arboreal 1858 Smith, Fr. subterranean (1973) Bolton material' like 'paper group group Bolton, 1973 1973 Bolton, subterranean (1973) Bolton Forel, 1901 1901 Forel, stumps / tree lignicolous (1901) Forel Fr. Smith, 1857 1857 Smith, Fr. lignicolous (1919) Wheeler Donisthorpe, 1942 1942 Donisthorpe, lignicolous (1942) Donisthorpe group (1973) (1973) arboreal carton in cited (1916), Forel Bolton Bolton Mayr, 1902 1902 Mayr, arboreal silk (1922a) Wheeler group group group André, 1890 1890 André, arboreal (1973) Bolton chapmani parabiotica arboreal present Wheeler (1922a) (1922a) Wheeler present latharis conops spinifera decemdentata esarata fissa arboreal latispina militaris rufipalpis schistacea hosei laboriosa concava Species Location Nests/C arabiotica-

present RJK Bolton in cited (1973) curta in Bolton gagates (1994) subterranean monista- p arboreal arboreal arboreal terrestrial / subterranean material' 'woven cited (1924), Arnold (1973) silk silk spider no silk (1976) Lévieux (1932) Collart Dejean & Lenir (1922a) Wheeler continua- inermis- monista

arboreal terrestrial silk (1928) Karawajew MCZC alluaudi alluaudi militaris- vindex in Bolton (1973) Subgenus Nest Mul S. Robson & R. J. Kohout 91

Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple terrestrial Hung (1962) (1962) Hung terrestrial Fr. Smith, 1859 Fr. Smith, lignicolous Kohout (1989) Emery, 1900 1900 Emery, lignicolous yes present RJK Roger, 1863b 1863b Roger, lignicolous / arboreal silk (1988c) Kohout Mayr, 1872 Mayr, / terrestrial lignicolous RJK Mayr, 1879 1879 Mayr, lignicolous MCZ Mayr, 1862 Mayr, lignicolous silk in Bolton Arnold (1924), cited

Bolton, 1973 Bolton, arboreal carton (1973) Bolton Walker, 1859 1859 Walker, lignicolous yes no none et (1998)al. Liefke

(Latreille, 1802) 1802) (Latreille, lignicolous Mann (1919) Roger, 1863 subterranean yes yes none et Liefke (1998)al. Emery, 1897 Emery, lignicolous RJK (Guérin-Méneville, 1831) 1831) (Guérin-Méneville, terrestrial (1943) Donisthorpe Stitz, 1911 Stitz, lignicolous RJK Forel, 1914 1914 Forel, arboreal silk in Bolton Arnold cited (1924), Fr. Smith, 1858 1858 Smith, Fr. subterranean (1973) Bolton Forel, 1910 1910 Forel, / subterranean lignicolous yes silk Mann (1919) Fr. Smith, 1861 Fr. Smith, present Donisthorpe (1941b) Santschi, 1910 1910 Santschi, arboreal silk (1973) Bolton Kohout, 1989 lignicolous/subterranean Kohout (1989) Forel, 1916 Forel, lignicolous silk (1967) in Hung Forel cited (1916) group group group group Species Location Nests/C viscosa cubaensis transiens weissi otleti sericata semitestacea salomo rufofemorata proxima nigropilosa obliqua litigiosa

foreli illaudata zopyra andromache opyra- (1973)

viscosa- arnoldi (1973)

revoili-

pubescens relucens lignicolous no no present SKR terrestrial (1962) Hung

lignicolous yes no none present RJK et (1998)al. Liefke

terrestrial lignicolous / lignicolous terrestrial none (2000) none Kohout present SKR Robson & Kohout (2005) lignicolous / none carton present RJK Robson (2005) & Kohout

relucens- z Subgenus Nest Mul 92 Nesting habits of Polyrhachis

argentea

Karawajew, 1906 Karawajew, Donisthorpe, 1937 alexandri taurus Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple

Emery, 1898 Emery, arboreal yes silk (1990) Kohout Taylor & Fr. Smith, 1860 Fr. Smith, arboreal silk present SKR Fr. Smith, 1863 1863 Fr. Smith, arboreal (1871) Smith Emery, 1900 arboreal carton/silk Karawajew (1928) Fr. Smith, 1871 1871 Fr. Smith, arboreal carton Donisthorpe as (1943) listed

1902 Forel, arboreal silk (1990) & Maschwitz Dorow Mann, 1919 arboreal silk Mann (1919) Emery, 1896 Emery, arboreal none (1928) Karawajew Forel, 1879 Forel, 1879 arboreal carton/silk/none (1928) Karawajew Forel, 1910 Forel, 1910 arboreal silk Robson & Kohout (2005) Fr. Smith, 1858 Fr. Smith, 1858 arboreal carton (1928) Karawajew Fr. Smith, 1863 Smith, Fr. absent RJK Fr. Smith 1858 Fr.Smith arboreal yes no silk et (1998)al. Liefke (Le Guillou, 1842) 1842) Guillou, (Le arboreal silk (1928) Karawajew Forel, 1915 Forel, Emery, 1887 Emery, arboreal / silk carton (1906) Karawajew as listed Forel, 1915 Mann, 1919 Mann, arboreal yes carton/silk Mann (1919) -group -group -group ulysses ulysses subtridens solmsi solmsi bicolor lombokensis osae schang saevissima tibialis tibialis wheeleri dolomedes lignicolous Bingham (1903) (1903) Bingham arachne hodgsoni lignicolous armata Species Location Nests/C yrmatopa yrmhopla arboreal absent SKR

arboreal silk present SKR

arboreal no silk/carton absent SKR / lignicolous arboreal arboreal silk silk RJK Mann as (1919) listed bicolor arboreal silk 1862 Mayr, Robson & Kohout (2005) alphea derecyna arachne armata lignicolous arboreal arboreal lignicolous yes no none silk (1903) silk Bingham present RJK et (1998)al. Liefke (1990) & Maschwitz Dorow Robson & Kohout (2005)

M M Subgenus Nest Mul

S. Robson & R. J. Kohout 93

Karawajew, 1927 Karawajew, mystica Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material

tiple Multiple

lignicolous yes no none present Schellerich-Kaadenet al. (1997)

Forel, 1910 1910 Forel, present RJK

Emery, 1897 1897 Emery, present RJK

Karaw.,1927 Karaw.,1927 arboreal silk (1928) Karawajew

Emery, 1887 Emery, lignicolous silk (1995) Dorow

Karawajew, 1927 arboreal silk Karawajew (1928)

Dorow, 1996 Dorow,

Fr. Smith, 1858 1858 Fr. Smith, arboreal carton silk / (1928) Karawajew

Fr. Smith, 1858 1858 Fr. Smith, arboreal yes silk (1995) Dorow

-group

-group

Forel, 1893 1893 Forel, arboreal silk (1929). Karawajew

Forel, 1892 1892 Forel, arboreal silk (1910) Wheeler in (1894), cited Forel

Wheeler, 1919 1919 Wheeler, arboreal yes silk (1995) Dorow

Fr. Smith, 1858 1858 Smith, Fr. arboreal yes no silk al. et (1998) Liefke

Fr. Smith, 1858 1858 Smith, Fr. lignicolous present RJK

Karawajew, 1926 Karawajew, arboreal silk Karawajew (1928)

Fr. Smith, 1857 1857 Smith, Fr. arboreal present RJK

Chapman, 1963 1963 Chapman, lignicolous (1963) Chapman

Fr. Smith, 1857 1857 Fr. Smith, arboreal silk (1910) Wheeler in cited (1894), Forel

-group

-group -group

Species Location Nests/C

tubifex

schellerichae schellerichae

muelleri -group

hector

abdominalis

rufipes

furcata

etheli etheli

flavoflagellata

lacteipennis

dives

daphne

jerdonii

cryptoceroides

bicolour nigripes bicolour

bicolour concolor bicolour

arboreal yes no silk al. et (1998) Liefke

arboreal silk present Dorow& Maschwitz(1992)

lignicolous present RJK

hector

present RJK

furcata

flavoflagellata

subterranean silk (1962) Hung

subterranean/ arboreal silk Wheeler& Mann (1916)

subterranean/ lignicolous / arboreal silk present (1970) Ofer

arboreal silk present SKR

terrestrial silk (1928) Karawajew

arboreal yes yes silk al. et (1987) Yamauchi

arboreal carton (1988c) Kohout

dives

daphne

lignicolous / arboreal silk (1928) Karawajew

arboreal silk Karawajew (1928)listed as

Subgenus Nest Mul cryptoceroides

bicoloratrocastanea 94 Nesting habits of Polyrhachis

Donisthorpe 1941 Donisthorpe 1941 arcuspinosa present RJK Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple Forel, 1901 Forel, arboreal yes silk SKR Donisthorpe, 1937 1937 Donisthorpe, lignicolous arboreal/ Donisthorpe (1937)

Fr. Smith, 1859 Fr. Smith, arboreal silk (1990) Kohout Taylor & Clark, 1930 1930 Clark, arboreal silk (1990) Taylor & Kohout (Latreille, 1802) 1802) (Latreille, arboreal carton/silk (1928) Karawajew 1927Karawajew, arboreal carton/silk (1928) Karawajew Fr. Smith, 1863 1863 Fr.Smith, arboreal (1871) Smith Karawajew, 1927 Karawajew, arboreal silk (1995) Dorow Fr. Smith, 1860 Fr. Smith, arboreal (1871) Smith Emery, 1887 arboreal silk Karawajew (1928) Emery, 1897Emery, arboreal carton Donisthorpe as listed (1941c) F. Smith F.1863 Smith arboreal carton Donisthorpe (1943) Menozzi, 1926 arboreal NMNH Forel, 1911 arboreal silk Robson & Kohout (2005) Kohout, 1990 Kohout, 1990 subterranean Kohout (1990) Menozzi,1932 arboreal Menozzi (1932) Kohout, 1990 subterranean Kohout (1990) -group -group -group -group Kohout, 1994 1994 Kohout, subterranean (2000) al. et Maschwitz -group Species Location Nests/C lama rustica loweryi sexpinosa mitrata mucronata paromala ochracea rugifrons follicula keratifera aureovestita esuriens esuriens calypso glabrinota reclinata bismarckensis expinosa Subgenus Nest Mul

subterranean no none (2003) al. et Maschwitz viehmeyeri arboreal lignicolous arboreal no silk present SKR silk (1999) Kohout Robson & Kohout (2005)

arboreal no no silk present SKR ochracea lignicolous lignicolous lignicolous no yes / carton present SKR Robson (2005) & Kohout (1990) Kohout Taylor & arboreal silk Robson & Kohout (2005)

present RJK bubastes / lignicolous arboreal lignicolous no no silk present carton SKR Robson (2005) & Kohout present RJK

mucronata s S. Robson & R. J. Kohout 95 Cocoons Additional Pupal Source olony? Queens/Colony? Nest Material tiple Multiple

Emery, 1895 1895 Emery, arboreal yes / silk carton Kohout (1999)

Donisthorpe, 1937 arboreal Donisthorpe (1937)

Forel, 1886 arboreal silk Robson & Kohout (2005) Fr. Smith, 1874 1874 Smith, Fr. / lignicolous subterranean terrestrial / (1967) Yanoin Hung (1911) cited Mayr, 1862 1862 Mayr, arboreal / silk carton (1928) Karawajew Emery, 1900 1900 Emery, arboreal no silk (1988b) Kohout Forel, 1915 Donisthorpe, 1948 1948 Donisthorpe, arboreal silk Robson & Kohout (2005) (Drury, 1773) (Drury, 1773) arboreal silk (1903) Bingham Fr. Smith, 1859 arboreal Wilson (1959) x Donisthorpe, 1938 1938 Donisthorpe, arboreal silk Donisthorpe(1938a) Karawajew, 1933 Karawajew, lignicolous (1933) Karawajew Crawley, 1915 arboreal yes silk Kohout (1999) Fr. Smith, 1957 Smith, Fr. Roger, 1863 1863 Roger, arboreal / silk carton (1928) Karawajew Emery, 1887 arboreal silk SKR Forel, 1912 1912 Forel, terrestrial yes no none et (1998)al. Liefke F. Smith, 1860 1860 Smith, F. lignicolous inquirenda (1871) Smith (species -group Kohout, 1988 1988 Kohout, lignicolous yes silk present Kohout (1988b) Fr. Smith, 1857 1857 Fr. Smith, arboreal (1871) Smith -group Species Location Nests/C thrinax lamellidens ternatae textor saigonensis ypsilon queenslandica taylori frauenfeldi eudora olybria abnormis cheesmanae durvillei delicata bihamata erosispina bellicosa yrmothrina olyrhachis Subgenus Nest Mul

lamellidens

arboreal silk present RJK

Kohout see (1998)) arboreal silk present SKR arboreal silk (2005) Kohout present RJK

M P bihamata subterranean SKR arboreal arboreal no carton / present silk SKR Kohout (1988b)

96 Nesting habits of Polyrhachis

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Received: 20 November 2006; accepted: 19 February 2007; published: August 2007

ASIAN MYRMECOLOGY Published by the Institute for Tropical Biology & Conservation, Universiti Malaysia Sabah, Malaysia on behalf of ANeT — the International Network for the Study of Asian Ants