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Deforestation Reduces Nest Mound Size and Decreases the Production of Sexual Offspring in the Wood Ant Formica Aquilonia

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Deforestation Reduces Nest Mound Size and Decreases the Production of Sexual Offspring in the Wood Ant Formica Aquilonia Ann. Zool. Fennici 42: 259–267 ISSN 0003-455X Helsinki 28 June 2005 © Finnish Zoological and Botanical Publishing Board 2005 Deforestation reduces nest mound size and decreases the production of sexual offspring in the wood ant Formica aquilonia Jouni Sorvari1* & Harri Hakkarainen2 1) Department of Biological Sciences, 8888 University Drive, Simon Fraser University, Burnaby, BC V5A 1S6, Canada; present address: Section of Ecology, Department of Biology, FI-20014 University of Turku, Finland (*corresponding author’s e-mail: jouni.sorvari@utu.fi) 2) Section of Ecology, Department of Biology, FI-20014 University of Turku, Finland Received 8 Oct. 2004, revised version received 10 Dec. 2004, accepted 23 Dec. 2004 Sorvari, J. & Hakkarainen, H. 2005: Deforestation reduces nest mound size and decreases the pro- duction of sexual offspring in the wood ant Formica aquilonia. — Ann. Zool. Fennici 42: 259–267. Little is known about how anthropogenic changes in nature affect reproduction in social insects such as ants. We studied the effects of clear-cutting on the production of sexual offspring and on the mound size in Formica aquilonia, using neighbouring uncut forests as controls. Nest mounds were smaller in clear-cuts, apparently because they tended to be new bud nests. The basal area of nest mounds decreased towards forest edges. The production of sexual offspring increased with nest mound size. Clear-cutting reduced the production of sexual offspring also when the effect of nest mound size had been factored out. Our results show that anthropogenic changes do have the potential to modify the production of sexual offspring in social insect. Introduction Many social insects, such as termites, wasps, bees and ants, live in colonies and exploit local The current rate of forest loss and the fragmen- resources. Among them, ants and termites are tation of natural habitats are considered to be excellent study organisms for studies on the a potential threat to the persistence of animal effects of habitat structure change, because their populations (Barbault & Sastrapadja 1995, colonies are perennial and due to wingless work- Fahrig 1999). The main forest renewal method ers they are using resources on a very local is clear-cutting where all or most of the trees scale. Workers can be either females or males are harvested from the renewal area. In Fin- in termites, but in Hymenoptera all workers are land clear-cuts are normally planted with single females (Wilson 1971). Whether a female devel- tree species monoculture and are re-harvested ops into a worker or a queen is determined in approximately at 100-year intervals (Virkkala Hymenoptera by environmental factors such as & Toivonen 1999). This kind of effective forest quality or quantity of food (Wilson 1971, Oster harvesting is likely to have reflections in animal & Wilson 1978, Brian 1983, Bourke & Franks populations through food shortage (Zanette et al. 1995), by worker behaviour (Brian 1973), or by 2000) and changes in species composition (Punt- genetic factors (Volny & Gordon 2002). Work- tila et al. 1991, Esseen et al. 1997). ers care for queens and offspring, so a sufficient 260 Sorvari & Hakkarainen • ANN. ZOOL. FENNICI Vol. 42 number of workers are needed before sexual off- specialized for producing mainly queens or spring (i.e. reproductive offspring) can be reared males and the average sex ratio across colonies (Oster & Wilson 1978, Hölldobler & Wilson is usually female biased (Pamilo & Rosengren 1990, Bourke & Franks 1995). Virtually noth- 1983, Rosengren & Pamilo 1986). New sexual ing is known about the effects of anthropogenic offspring are raised mainly from late April to late changes on sexual and worker offspring produc- May, before worker progeny. Winged males and tion in social insects. Especially in the boreal gynes (young queens) can be observed on the region forest management may have a major nest mound or in the vicinity of nests from late impact on the ant fauna dependent on mature May to early June when mating takes place. F. old-growth forests. rufa group wood ants rear sexual offspring early Wood ants of the Formica rufa group are in the spring by using food resources collected dominant ants in boreal coniferous forests. They the previous season and stored by young workers have ecological effects that extend over several called repletes (Rosengren et al. 1987, Gösswald trophic levels, and hence the study of the asso- 1989). Sexual offspring are reared with lipids ciation between man-made habitat alteration and and glandular nutrients stored by repletes in the breeding success of colonies is important. In fat-bodies. The utilization of these lipid reserves terms of resource and interference competition may initiate metabolic heating of a nest (Martin and predation, wood ants affect arthropod species 1980), which also seems to be needed for sexual composition (Adlung 1966, Niemelä et al. 1992, offspring production (Rosengren et al. 1987). Punttila et al. 2004) and thus, also the breeding Clear-cutting likely reduces the food success of some insectivorous birds (Aho et al. resources of wood ants, because the majority 1999). Further, they affect the distribution and of their food comprises honeydew, aphids and abundance of other ant species by their competi- other arthropods collected from trees (Rosen- tive superiority (Savolainen & Vepsäläinen 1988, gren & Sundström 1987, 1991, Whittaker 1991, 1989, Savolainen et al. 1989). Nest mounds are Punttila et al. 2004). Lenoir (2003) found that shared with numerous other arthropod species, when excluded experimentally from trees, wood so-called myrmecophiles (Päivinen et al. 2004) ants search for new trees further away instead as well as earthworms (Laakso & Setälä 1997). of increasing predation on the forest floor. In F. Wood ants spread seeds of many plants (Gorb aquilonia, new colonies are frequently estab- & Gorb 1995, 1999, Gorb et al. 2000) and influ- lished by budding in clear-cuts, which alters the ence the growth of trees due to aphid tending age and size distribution of nests (Rosengren & (Rosengren & Sundström 1991). Pamilo 1978, Rosengren et al. 1979). An abrupt As in other ants, colony kin structure varies change in habitat structure in conjunction with among wood ant species. A colony can have decreased nest sizes may alter the probability single egg-laying queen (monogyny) or multi- of sexual offspring production. We suggest that ple queens (polygyny). In the latter case colo- an increase in colony size increases the colony’s nies can also comprise multiple nests (poly- potential of producing sexual offspring and that domy, Rosengren & Pamilo 1983, Hölldobler & deforestation has a negative effect on sexual Wilson 1990, Bourke & Franks 1995). The most offspring production by reducing nest sizes and common wood ant in boreal forests is the highly food resources. Several studies have found that polygynous and polydomous Formica aquilonia production of sexual offspring increases with (Collingwood 1979, Rosengren et al. 1979). A an increase of colony size (Oster & Wilson large nest mound may contain a million workers 1978, Nonacs 1986, Tschinkel 1993, Deslippe & and hundreds of queens (Pamilo 1982, Rosen- Savolainen 1995, Wagner & Gordon 1999), but gren et al. 1987). to our knowledge there are no published studies All colonies of F. aquilonia do not normally on the effects of clear-cutting on the production produce sexual offspring every year, possibly of sexual offspring in ants of the Formica rufa due to the age of colony and/or the availability group. We studied this by measuring nest mounds of resources (Pamilo & Rosengren 1983, Rosen- and collecting information on the production of gren & Pamilo 1986). Colonies are normally sexual offspring in polydomous F. aquilonia in ANN. ZOOL. FENNICI Vol. 42 • Deforestation and production of sexual offspring in ant 261 clear-cuts and forest interiors. Our goal in this worker ants in other Formica species (Deslippe study was to examine whether the size distribu- & Savolainen 1994, Liautard et al. 2003). In this tions of nest mounds differ between clear-cuts study, the basal area of the nest mound was used and forests, and whether nest mound size, habitat as a rough estimate of worker population. The type and year-to-year variation affects the pro- basal was measured from the part of the mound duction of sexual offspring. not covered by plants. In our study species, the shape of the nest mound base is normally circu- lar, hence we used a common equation for cir- Materials and methods cular area (pr2) to calculate the basal area of the nest mound. We also measured the distance from Study area each nest to the nearest forest edge. The study was conducted in the coniferous boreal zone in central Finland, near the town of Statistical analyses Jyväskylä (62°15´N, 25°45´E) in 2000–2001. We used two types of study areas of different We used generalized linear model (Genmod) for habitat quality, forest interior areas (n = 13) and comparing nest mound sizes, habitat types and clear-cuts (n = 18). Forest interior areas and year as fixed factors; and colony (nested within clear-cuts were chosen from the same region and habitat type) as a repeated subject and normal were initially similar Vaccinum myrtillus growth error term. The relationship between the nest mature forests dominated by Norway spruce mound size and the distance to the forest edge (Picea abies) located 3.8 ± 0.3 km (mean ± S.E.) in both habitat types was also analysed with a apart. Clear-cuts varied from 0.3 to 4.0 ha (mean generalized linear model (Genmod), with the ± S.E. = 1.5 ± 0.2 ha). Nine of the clear-cuts distance to the forest edge as a quantitative vari- were cleared in the winter 1998–1999 and the able, and the study area as a repeated subject and remaining nine in the winter 1999–2000. The normal error term.
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