Journal of Earth Science and Engineering 3 (2013) 723-729

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The Effect of Selective Logging on the Nest Density, Foraging Range and Colony Size on the Ground-Dwelling Ant Aphaenogaster Swammerdami Forel, 1886 in Kirindy ForestMadagascar
Ferdinand Tornyie1, Stefan Jahnel2 and Eliezer Bortei Borketey-La3
1. Council for Scientific and Industrial Research-Institute of Industrial Research, Accra 00233, Ghana, 2. Department of Behavioural Biology, University of Vienna, Vienna 11010, Austria 3. Department of Crop/Soil Science Education, University of Education, Winneba 00233, Ghana Received: September 25, 2013 / Accepted: October 20, 2013 / Published: November 25, 2013. Abstract: Studying the ecology of ants can be a powerful tool for conservation. While the effect of logging is mainly investigated by the comparison of species richness and composition, the impact on individual species are often neglected. This study investigated the effect of selective logging on the nest density, foraging range and colony size on the ground-dwelling ant Aphaenogaster swammerdami in Kirindy forestMadagascar. This ant is a common ground-dwelling species in Kirindy, a western dry deciduous forest of Madagascar. Sampling was done in two sites of the forest: One part that was selectively lodged and another that have not been logged. Here we show that selective logging led to a decrease in colony size and density, while the foraging range seemed to be unaffected. Higher desiccation stress and lower food availability in the logged forest are most likely to be responsible for these results. Key words: Ground-dwelling, selective logging, colony size, nest density, species richness.

1. Introduction
The family of ants include only about 1% of all insect species. Nevertheless they are one of the most dominant groups in terrestrial habitats both in number and ecological importance. Six of the worldwide 16 subfamilies have been recorded to date in Madagascar including 4 endemic genera [1, 2]. Despite their huge diversity, importance in ecosystem functioning and high levels of species turnover, invertebrates are often unjustifiably excluded from inventories of natural areas [3, 4]. Ants, in particular ground-dwelling ones, are a powerful tool for conservation, because they
Corresponding author: Ferdinand Tornyie, MPhil, research fields: Insect ecology, Pollination biology, integrated pest management, apiculture and meliponiculture, E-mail: ferditor@yahoo.com.

show a high level of species turnover and therefore explain fine-scale patterns of diversity [1]. They provide important information for: (1) detecting invasive species, (2) detecting trends among threatened species, (3) detecting trends among keystone species, (4) evaluating land management actions and (5) assessing long-term ecosystem changes [1, 5]. Selective logging is a common form of forest exploitation in tropical regions [6]. It can affect the invertebrate community of the forest floor. Downed wood and slash from logging operations may increase the food availability, create predation refugia, nest sites and mesic microhabitats for a variety of litter taxa like ants [7, 8]. Former studies in Kirindy forest suggest that selective logging has minor effects on the

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The Effect of Selective Logging on the Nest Density, Foraging Range and Colony Size on the Ground-Dwelling Ant Aphaenogaster Swammerdami Forel, 1886 in Kirindy Forest-Madagascar

litter invertebrate community as a whole. No significant difference of species richness and ant species composition has been found between logged and unlogged forest [8, 9]. However, little is known about the effects of logging on individual species. In Kirindy forest a total of 86 ant species are recorded. One of the most common species is Aphaenogaster swammerdami. It is a ground-dwelling ant, which lives in conspicuous nests, easily recognisable by the excavated soil around the nest entrance (mount), reaching a diameter of up to more than 1 m. The workers are approximately 1.2 cm long and forage solitary. Foraging ranges longer than 15 m have been observed and they are known to be expert thieves. Besides that A. swammerdami has been shown to play a crucial role in the dispersal of seeds of Commiphora guillaumini [2, 10]. In our study, we focus on the effect of selective logging on the nest density, foraging range and colony size of A. swammerdami in Kirindy forest.

colonies and found out that they are most active during the morning (5: 30 a.m.-8: 30 a.m.) and the evening (3: 00 p.m.-6: 00 p.m.). 2.3 Measurement of Foraging Range For each site we chose 15 active nests randomly (30 for the two sites together) and these nests were used in measurement of foraging range. Each nest was observed for at least 1 h. To determine how far the workers go away from their nest to collect food, we offered them food (pieces of bread) at different distances from the nest and in different directions. Once the food was taken to the corresponding nest, the distance was increased. This was done for at least 10 ants until the maximum feeding distance from the nest was reached. 2.4 Estimation of Colony Size We measured the diameter of the mostly circular mounts. For noncircular mounts we measured the maximum and minimum diameter and recorded the average. Former studies [13] found out that the mount diameter is correlated to the colony size and therefore measuring this provides a simple method for estimating the colony size. 2.5 Determination of Nest Density Since the grid cells of the different sites are not of the same size, we choose 20 sample areas with a size of 15 m 15 m. We got the impression that nests occur more frequently on the trails. For this reason we wanted to make sure that the same length of trail is included in each sample area. Sample points were chosen randomly. From these points we measured 15 m along the northern and western trail for each sample area. Within these 15 m 15 m squares active nests were counted and the mount diameters were recorded (Fig. 1). 2.6 Data Analysis Microsoft Excel was used for data entry, management and graphical output. Minitab (version

2. Materials and Methods

2.1 Study Site The study was carried out in Kirindy forest, a Western dry deciduous forest located approximately at 40 km ENE of Morondava of western Madagascar (2004 S, 4439 E; 30 m elevation). The area covers about 10,000 ha of which a part was selectively logged until 1991 [11]. Several parts of the forest were divided into grid systems for the purpose of research. For the unlogged site the eastern part of the grid (site) N5 was chosen, whereas grid (site) CS5 served as the study site of the logged forest (extensively logged until 1980) [12]. Kirindy forest faces highly seasonal climate changes with the wet season usually running from November to April. Our study was carried out at the end of the dry season (November). 2.2 Preliminary Studies To determine the time of day most ideal for sampling, we observed the activities of different

The Effect of Selective Logging on the Nest Density, Foraging Range and Colony Size on the Ground-Dwelling Ant Aphaenogaster Swammerdami Forel, 1886 in Kirindy Forest-Madagascar Upper part of nest Entrance of nest

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3.2 Foraging Range Considering the 30 (15 for each side) nests used for the foraging range measurement, the foraging range in most cases was not equal in all directions. This was influenced by some factors such as adjacent nests and presence of fallen dead trees on the forest floor. The foraging ranges extend from approximately 600 cm to 1,800 cm in both sites (CS5: min = 610 cm, max = 1,770 cm; N5: min = 550 cm, max = 1,750 cm) with a mean distance to the nest of about 9.5 m. Although N5 (903.67 cm [SE 72.60]) showed a slightly higher mean of foraging range than CS5 (1,029.67 cm [SE 100.31]), the difference was not significant (U-Test, df = 19, P = 0.6334) (Fig. 3). 3.3 Colony Size Among all the 64 nests the mount diameter ranged of 60-150 cm. The mean mount diameter of the logged and the unlogged site differ significantly with the unlogged part showing a mean mount diameter of 110.93 cm (SE 5.36) and the logged part a mean of 87.53 (SE 4.83) (U-Test, df = 19, P = 0.0037) (Fig. 4). 3.4 Relationship between Foraging Range and Mount Diameter A correlation between foraging range and the mount diameter could be observed in both sites, with a strong correlation (R2 = 0.8987, P < 0.001) in the unlogged forest and a weak correlation (R2 = 0.3154, P = 0.029) in the logged forest (Fig. 5).

Fig. 1 Schematic representation of the nest mount. Top: side view, bottom: view from above.

16.0) was used to determine statistical differences, associations and relationships.

3. Results
Sixty four (64) active nests were recorded for the two sites together (grid CS5: 22, grid N5: 42), however, 30 (15 for each site) of them were used in the measurement of foraging range. 3.1 Density In total, an area of 9,000 m2 (4,500 m2 for each site) was covered for the 2 sites. Besides the 64 active nests recorded, we also observed a much higher number of inactive nests, which were partially used by other animals (e.g., spiders, lizards). The number of nests per study area (15 15 m) varied little (CS5: min 0, max 3; N5: min 0, max 4). On average there were 1.1 (SE 0.03) nests/study area (N = 20) in CS5 and 2.1 (SE 0.26) nests/study area (N = 20) in N5 (Fig. 2). A comparison between these two sites revealed a significant difference (U-Test, df = 19, P = 0.0106).
2.5 Nests/15x15m 2 1.5 1 0.5 0
Logged (CS5) Unlogged (N5)

Site 1
Fig. 2 Comparison of nest density in the logged and unlogged forest.

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The Effect of Selective Logging on the Nest Density, Foraging Range and Colony Size on the Ground-Dwelling Ant Aphaenogaster Swammerdami Forel, 1886 in Kirindy Forest-Madagascar
1200 Logged (CS5) Unlogged (N5) 1000 Foraging range (cm)

800

600

400

200

0 1 Site

Fig. 3

Comparison of mean foraging ranges in the logged and unlogged forest.

140 Mount diameter (cm) 120 100 80 60 40 20 0

Logged (CS5) Unlogged (N5)

1 Site
Fig. 4 Comparison of mean mount diameter in the logged and unlogged forest.

Unlogged
Foraging range (cm)

Logged
2000 1500 1000 500 0 0 50 100 R2 = 0.3154 150

Foraging range (cm)

2000 1500 1000 500 R2 = 0.8987 0 0 50 100 150 200 Mount diam eter (cm )

Mount diam eter (cm )

Fig. 5

(a) (b) Correlation between foraging range and the mount diameter in the unlogged and logged forest.

4. Discussion
4.1 Nest Density Our study has shown that there is a significant difference in the density of nests between the logged and the unlogged forest. Density in the unlogged

forest was almost twice as high as in the logged forest. This can be due to both direct and indirect effects of logging. As the opening of the canopy increases and the vegetation structure and understory plant richness are reduced, more sunlight and wind reaches the soil. This can lead to increasing desiccation, higher soil

The Effect of Selective Logging on the Nest Density, Foraging Range and Colony Size on the Ground-Dwelling Ant Aphaenogaster Swammerdami Forel, 1886 in Kirindy Forest-Madagascar

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temperatures, decreasing relative humidity and litter moisture. Indirectly, logging can eliminate other invertebrates, which provide a reliable source for food and moisture [14, 15]. Bhning-Gaese [10] showed that colonies of A. swammerdani are significantly associated with trees of C. guillaumini, suggesting that the arils might be an important food source for this ant species. Therefore it is also likely that the decreased density of C. guillaumini in the logged forest is responsible for the lower nest density of A. swammerdani. 4.2 Foraging Range There foraging range between the logged and unlogged forest did not differ. One could argue that nest density will be a limiting factor for the foraging range, because of increased competition for resources [16]. Since we found significant differences (U-Test, df = 19, P = 0.0106) in density between the two types of forest, we suggest that the critical threshold for the capacity of nests per area has not been reached. Covich [17] also recorded that refuge-inhabiting animals can increase their foraging area without simultaneously increasing predation risk and travel costs by using additional refuges scattered throughout the home range. Since these ants are grand dwelling, refuging and foraging in leave litters, they probably had enough cover away from predators in both the logged and unlogged forest. This probably gave them the advantage to forage any distance away from their nest without predators being a limiting factor. On the other hand, it is likely that the nest density of both areas is not high enough to have any influence on the foraging range, thus the effect of logging has no influence on the foraging range of this ant species. 4.3 Colony Size The mean mount diameter of nests in the unlogged forest is slightly higher (significant) than in the logged forest. Since mount diameter has been shown to correlate to the colony size [13], we assume that the

mean colony size is bigger in the unlogged forest. When food abundance, microenvironment and competition change, ant colonies may respond by altering home range size, the frequency of foraging trips and the duration of foraging trips [18]. Possible explanation for that could be a higher availability of food in the unlogged forest [19] thus the large colonies found here. In the logged forest on the other hand, a dryer and hotter microclimate could have limited the population size of the ants [20, 21]. 4.4 Relationship between Foraging Range and Mount Size Body size is often positively correlated with ecologically related traits such as fecundity, survival, resource requirements and home range size [22, 23]. A positive correlation between these two parameters was shown at both sites. This is not surprising, since larger colonies require more space to forage and also have the advantage of effective food source communication [13, 24]. McGlynn [19] also found a strong correlation between home range and colony size, and suggested that home range was independent of habitat quality. However, there is a stronger correlation in the unlogged site than in the logged one. In the unlogged site, one can easily predict the foraging range just by measuring the mount size. This is quite different in the logged site, where this relation seems to be disturbed. Factors like, general lower food availability or higher desiccation stress alone cannot explain this deviation, because it is a weak positive correlation. It is more likely that selective logging caused a disturbance in the uniform distribution of food sources and microclimate (e.g., patches of opened canopies), leading to a different environmental stress [21, 25] caused this deviation foraging range in the logged forest.

5. Conclusions
Our results suggest that logging heavily affects the nest density and colony size of A. swammerdani. The

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logged forest shows a decrease in both the density of nests and the colony size probably due to lower food availability and higher desiccation stress. The foraging range, however, is not affected. An explanation for this could be that the density of nests is not high enough to have an influence on the foraging range. Also there was enough leave litter for refuge for the ants against predators during foraging. The fact that the correlation between colony size and foraging range is weaker in the logged forest suggests that the composition of food sources and the microclimate are not uniform. It should be mentioned that the recorded densities of nests is relative, because the area of trails in the study areas do not represent the total trail proportion of the whole forest (nests seem to occur more likely on trails).

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Acknowledgments
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We want to extend our sincere gratitude to Tropical Biology Association for sponsoring our stay in Madagascar for a field course and to the British Society of Ecology for their financial support. Special thanks go to all the resource persons on the field course, especially Henry and Melanie for their assistance in the analyses of our data.

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