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Polyrhachis Loweryi (Formicinae): a Guest Ant Parasitizing Rhytidoponera Sp

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Polyrhachis Loweryi (Formicinae): a Guest Ant Parasitizing Rhytidoponera Sp Insectes Soc. 50 (2003) 69–76 0020-1812/03/010069-08 Insectes Sociaux © Birkhäuser Verlag, Basel, 2003 Research article Polyrhachis loweryi (Formicinae): A guest ant parasitizing Rhytidoponera sp. (Ponerinae) in Queensland, Australia U. Maschwitz 1,C.Go1,W.H.O.Dorow2,A.Buschinger 3 and R. J. Kohout 4 1 Johann Wolfgang Goethe-Universität, Fachbereich Biologie (Zoologie), AK Ethoökologie, Siesmayerstrasse 70, D-60054 Frankfurt am Main, Germany, e-mail: [email protected] 2 Forschungsinstitut Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany, e-mail: [email protected] 3 Technische Universität Darmstadt, Fachbereich Biologie (Zoologie), Schnittspahnstrasse 3, D-64287 Darmstadt, Germany, e-mail: [email protected] 4 Queensland Museum, PO Box 300, South Brisbane, Queensland 4101, Australia, e-mail: [email protected] Received 6 May 2002; revised 6 September and 9 October 2002; accepted 15 October 2002. Summary. At the type locality near Miles in Queensland, to rear them in neighbouring host nests. The adaptive and Australia a queenright colony of Polyrhachis loweryi was predispositional reasons for the phyletic development of this found (1 dealate queen, 91 workers, males and brood) within parasitic relation remain unclear. a colony of Rhytidoponera sp. (near aciculata), whereas oth- er colonies of this Rhytidoponera species contained only a Key words: Diacamma sp., Formicidae, Polyrhachis low- few workers of P. loweryi. For experiments and behavioural eryi, Rhytidoponera sp., social parasitism observations P. loweryi and its hosts were kept in the labora- tory for some time. The queenright P. loweryi colony con- structed closed nests out of soil and plant fibres, within the host nest, and only containing Polyrhachis specimens. P. low- Introduction eryi workers in part, remained amid the assembly of Rhyti- doponera, from whom they obtained carbohydrate and pro- Social parasites are not uncommon in most major groups of tein food. In addition, Polyrhachis workers regularly left the social insects. In ants, they have evolved in comparatively host nest. They were able to collect liquid carbohydrate food high numbers and in a unique sociobiological diversity. Most and to distribute it among conspecific nest mates via troph- parasitic ants, i.e. inquilines, slave-makers and temporary allaxis. The care of the P. loweryi brood was fully done by parasites, are closely related to their respective host species, P. loweryi itself. However, Rhytidoponera workers were a fact which was termed “Emerys rule” (Le Masne, 1956). observed carrying P. loweryi brood in addition to their own Xenobiotic or guest ants, on the other hand, coexist with host brood. In artificially induced nest relocations, P. loweryi was species belonging to different tribes or even subfamilies. The able to move to a new nest independently from its host ants, recent discovery of a formicine parasite, Polyrhachis lama displaying the typical invitation behaviour, trail laying Kohout, 1994, living nearly like an inquiline within the nests behaviour, and leader independent trail communication of a ponerine, Diacamma sp. in SE-Asia (Maschwitz et al., found in other members of the genus. Rhytidoponera sp., 2000), seemed to indicate an evolutionary pathway to a per- which in addition to inconspicuous trail marking, performs manent, true social parasitism in which a close systematic nest relocation via social carrying, rarely but regularly also relationship was not a necessary predisposition, a parasitism carried P. loweryi to the new nest. Whereas Rhytidoponera not obeying ”Emery’s rule”. The highly sophisticated para- sp. workers were carried in the typical ponerine posture, P. sitic relation suggested a long phyletic development. P. lama loweryi workers were carried by them in the formicine pos- belongs to the viehmeyeri species-group of the subgenus ture. We conclude that P. loweryi is best described as a guest Myrmhopla Forel, 1915 (Kohout, 1994b). The range of P. ant of Rhytidoponera sp.. The species seems to be less well lama apparently extends from Java and Hong Kong to the integrated into the societies of its Rhytidoponera sp. host than Himalayas (Maschwitz et al., 2000; Kohout, 1994a and the closely related P. lama, a social parasite of Diacamma sp. unpublished data). Further nine species of this group are in Java. restricted to Australia and neighbouring islands, i.e. New In both Polyrhachis species workers from queenright Guinea, Aru Islands, Solomon Islands and Seram (Kohout, colonies perhaps emigrate with a few eggs or larvae in order 1990). Collecting data of three Australian species, P. eremi- 70 U. Maschwitz et al. Polyrhachis loweryi a guest ant of Rhytidoponera sp. ta Kohout, 1990, P. rustica Kohout, 1990 and P. loweryi Dissection of ants Kohout, 1990 indicated that they perhaps also were asso- ciated with ponerine ants, though with species of the Dissections of the Polyrhachis loweryi specimens were made to check their reproductive status as described by Buschinger and Alloway genus Rhytidoponera (Kohout, 1990, 1994b). In February (1978) under a dissecting microscope. No efforts were made to identify and March 2000 we searched for these 3 species at their egg-laying specimens of the Rhytidoponera sp. whose queens are mor- former collection sites. One of them, P. loweryi, was found, phologically indistinguishable from the workers. associated with a Rhytidoponera species. This record and the preliminary results of behavioural studies are presented Laboratory culture of Polyrhachis loweryi and Rhytidoponera sp. here. The large queenright mixed colony of P. loweryi which had the original colony composition was kept in a wooden arena (50 ¥ 150 ¥ 10 cm), Material and methods with a plastic box (20 ¥ 20 ¥ 10 cm) inside, as a nest chamber. The nesting ground was covered with plaster of Paris and cardboard (egg- Field collecting containers) was offered as a nest shelter. The smaller colonies were held in arenas (60 ¥ 30 ¥ 10 cm), also provided with nest chambers of the We visited the type localities of Polyrhachis rustica (at Mareeba, same size as above. Inside the arena, water, honey water and protein food (cut cockroaches and Tenebrio larvae) were provided ad libidum in 17°01¢S, 145°24’E and near Collinsville, 20°31¢S, 147°49¢E), of small dishes. Further experimental details are reported in the results Polyrhachis eremita (near Marlborough, 22°43¢S, 149°45¢E) and of section. Polyrhachis loweryi (near Miles, 26°36¢S, 150°10¢E) (see Appendix). Since all three species originally had been found in more or less close association with large Rhytidoponera species, we searched for nests of such species which then were excavated as completely as possible. Results Nests of smaller Rhytidoponera species (belonging to the metallica species-group) also were checked for the presence of Polyrhachis specimens. Nests were dug out and the nest material thrown onto a Characterization of the host ant Rhytidoponera sp. large plastic sheet. The ants were sorted from the soil, transferred (nr. aciculata) into a plastic vessel, and then carefully checked for the presence of a different species. At Mareeba (21 colonies of a large brownish P. loweryi was discovered in colonies of a large Rhytidopon- Rhytidoponera sp. checked), and at Collinsville (15 colonies of a large era species at the type locality, 5 km from Miles in a rela- black and one of a large brownish-black Rhytidoponera sp. checked) tively little disturbed sclerophyll forest. The Rhytidoponera we did not find any Polyrhachis rustica. At the site of P. eremita, near Marlborough, no nests of any large Rhytidoponera sp. were found, workers were 1.2–1.3 cm in length and blackish-brown in probably due to serious deforestation. At Miles, a total of 10 colonies colour, with the gaster slightly lighter. The colonies were eas- of a large Rhytidoponera sp. (belonging to the aciculata species-group, ily recognized, because they had entrance hills of up to H. Reichel, pers. comm.) were dug out in a dry sclerophyll forest. Six 9.5 cm height and 36 cm diameter. The nests could be more of these colonies contained specimens of Polyrhachis loweryi (Table than 65 cm deep (where we had to stop excavation because of 1), including one queenright mixed colony. They were transferred to the laboratory for behavioural studies. In five colonies of another, extremely hard soil). They consisted of a more or less verti- black Rhytidoponera sp. (belonging to the convexa species group, cal entrance tube, widening up to three superimposed floors H. Reichel pers. comm.) no Polyrhachis were detected. Voucher speci- of large flat chambers, up to 30 cm wide, of irregular shape mens have been deposited in the Queensland Museum, Brisbane, and partly ending in short distal tunnels. As the workers Australia and the Forschungsinstitut Senckenberg, Frankfurt am Main, quickly fled into deeper chambers, their actual distribution in Germany. the nest remained unknown. Most ponerine colonies had no brood. In one colony pupae and a few small Rhytidoponera larvae were found. A plethora of “guests” inhabited the Table 1. Census data of Rhytidoponera sp. (nr. aciculata) colonies with and without the parasitic Polyrhachis loweryi Col. # number of Rhytidoponera sp. number of Polyrhachis loweryi adults larvae pupae workers queens males adult females eggs, larvae, pupae 141––6––– – 2 108 – – 7 ––– – 3 167 – – 5 ––– – 446–––––– – 5 238 – – >91 1 + – + 6 132 – – 6 ––– – 7 >240 – – – – – – – 8 ӷ400 – + – ––– – 9 >251 + + >19 – – – – 10 >100 – – – – – – – Insectes Soc. Vol. 50, 2003 Research article 71 Rhytidoponera nests. Great numbers of larvae and adults of a Demographic development of the queenright colony # 5 cockroach species, many woodlice (Nagaroides tumidus after excavation (Trachelipodidae); det. F. Taiti, Firenze), a few specimens of a small reddish spider and a number of crickets and lepis- In the large queenright P. loweryi colony egg laying soon matids crowded all nest chambers and tunnels of Rhytido- commenced as well as larval development, pupation and ponera nests with and without Polyrhachis.
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