Biologia 68/2: 314—318, 2013 Section Zoology DOI: 10.2478/s11756-013-0154-x
The diversity of soil mites (Acari: Mesostigmata) in yellow ant (Lasius flavus) nests along a gradient of land use Jacek Kamczyc & Dariusz J. Gwiazdowicz Pozna´ n University of Life Sciences, Department of Forest Protection, Wojska Polskiego 71c, 60-625 Pozna´ n, Poland; e-mail:
[email protected],
[email protected]
Abstract: Mesostigmatid mites communities in yellow ant (Lasius flavus) hills and phoresis of mites on this ant species were analysed in the Wielkopolska Region, Central Poland. Samples were collected from ant nests located along a gradient of four different types of land use: forest, ecotone, meadow and garden. In total, 132 mites were collected in ant nests among which 26 species were identified. The highest total abundance of mites was observed in the ecotone. Moreover, 14 mite specimens were found on L. flavus workers bodies. These are the first records of phoresis of mesostigmatid mites on this ant species. Key words: Lasius flavus; mites; Mesostigmata; ant nests; land use; forest; ecotone; meadow; garden
Introduction Symbiosis is generally defined as the relationships between organisms without any implication of reciprocal positive or negative influences and commonly occurs with impressive diversity in ants (Witte et al. 2008). Organisms that live in close relationships with ants are referred to as myrmecophiles. This kind of symbiotic interactions between ants and other groups of animals range from mutualism to predation and parasitism and additionally is common in Diptera, Hymenoptera, Coleoptera and Hemiptera, Thysanoptera as well as Strepsiptera (Lachaud & Perez-Lachaud 2009). Myrmecophiles were classified by Wasmann (1890) into four main groups: organisms that are fed and cared as guests of the colony, small and active organisms which feed on dead ants, ant‘s enemies and endo- and ectoparasites. In and around ant nests, various arthropods, including up to 150 families of 15 orders, were considered as myrmecophiluos animals (Ito & Takaku 1994). They comprise various arachnids, e.g., Pseudoscorpionidae, Araneidae, Opiliones and Acarina (Wi´sniewski 1965). Mites are the most abundant arthropod group in ant nests. Similar to other myrmecophiles, mites are ectoparasites or live commensally in ant colonies: they are scavengers, predators, ectoparasites or phoretic on ant bodies (Kranz 1978). Several records of symbiosis of mites with ants have been documented by, e.g., Berlese (1904), Wasmann (1897, 1899/1990, 1902), Donisthorp (1907a, b, 1909), Vitzthum (1918) and Samšiňák (1957, 1962) mainly as single cases. Mites associated with certain ant species, e.g., only Formica rufa L., 1761 and/or F. polyctena F¨ orster, 1850 are well documented by Wi´sniewski (1965, 1966, 1980, 1984). Additionally,
c 2013 Institute of Zoology, Slovak Academy of Sciences
Kielczewski & Wisniewski (1966, 1971) investigated phoresy of mites on various ant species. However, studies on phoretic mites by Kielczewski & Wi´sniewski (1962, 1966, 1971, 1973), Kielczewski et al. (1970), Bloszyk & Olszanowski (1986), Ito & Takaku (1994) and Gwiazdowicz (2001) were restricted only to forest or particular ant species. We studied the species diversity of mesostigmatid mites associated with the yellow ant Lasius flavus (F., 1782) along a gradient of different land use. This ant predominantly lives below ground, feeding on root aphids and their honeydew (Dauber et al. 2006). The aim of the study was to compare the species diversity of mesostigmatid mites in yellow ant nests in forest, ecotone, meadow and garden. In addition, we aimed to find out if mites use L. flavus for transportation and which mite species are phoretic. Due to differences in vegetation surrounding ant hills we expected differences in abundance and species diversity of mites. Material and methods Sampling This study was conducted in the Wielkopolska Region, central Poland (51◦ 41 N, 16◦ 58 E). The climate of the area is continental with annual rainfall of 600 mm and mean annual temperature of 8 ◦C. Four sites were chosen to represent different land use/land cover types with a gradient of management intensity: forest, forest ecotone zone (both were treated as woody vegetation), meadow and garden. Ant nests were identified by careful searching the plots for visual signs and by following the workers. Ant nests were sampled on a single occasion in August 2007. Totally, 12 samples (4 study sites × 3 ant nets ×1 sample), with the size of 125 cm3 were taken from the upper 0–5 cm layer of
Soil mites in yellow ant nests
315
Table 1. List of mesostigmatid mites from ants nests in various microhabitats. Results are shown as dominance D (%). Microhabitat Species
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26
Dendrolaelaps latior (Leitner, 1949) Discourella baloghi Hirschmann & Zirngiebl-Nicol, 1969 Discourella modesta (Leonardi, 1899) Hypoaspis aculeifer (Canestrini, 1883) Hypoaspis brevipilis Hirschmann, Bernhard, Greim, G¨ otz, 1969 Hypoaspis vacua (Michael, 1891) Leptogamasus suecicus (Tr¨ ag˚ ardh, 1936) Macrocheles ancyleus Karauss, 1970 Pachylaelaps furcifer Oudemans, 1903 Pachylaelaps magnus Halbert, 1915 Pachylaelaps siculus Berlese, 1921 Pachyseius humeralis Berlese, 1910 Paragamasus vagabundus (Karg, 1968) Parasitus consanguineus Oudemans & Voigts, 1904 Parasitus fimetorum (Berlese, 1904) Pergamasus brevicornis Berlese, 1903 Pergamasus septentrionalis (Oudemans, 1902) Rhodacarus mandibularis Berlese, 1921 Trachyuropoda formicaria (Lubbock, 1881) Trichouropoda ovalis (C.L. Koch, 1839) Trichouropoda sp. Uropodina sp. Veigaia nemorensis (C.L. Koch, 1839) Vulgarogamasus kraepelini (Berlese, 1905) Zercon peltatus peltatus C.L. Koch 1836 Zercon triangularis C.L. Koch 1836 Total
soil/nest in a 25 cm2 layer. The nest samples were transported to the laboratory, the mites were extracted with Berlese funnels (chambers were enclosed to prevent mites escape) and counted. The samples were heated from above with 40 W bulbs and the extraction lasted 7 days. Additionally, for phoresis investigations samples were collected from the surrounding of ant nests. In total, 12 samples of 100 ant workers were manually collected from each colony. The ants were collected into plastic taps filled with 70% ethanol, transported to the laboratory and checked for phoretic mites on their bodies. Mesostigmatid mites were stored in 70% ethanol, mounted in permanent (using Hoyer’s medium) and semipermanent (using lactic acid) slides, counted, and identified using the latest taxonomical literature. Data analysis Diversity for each sample was measured using the Shannon’s diversity index (H = − (pi ln pi ), where pi = proportional abundance of the i-th species. Eveness index was calculated as E = H / ln S, where S = species richness. Species richness was examined by counting the number of species in each sample. Eveness, Shannon index and species richness were statistically tested with a Kruskall-Wallis H test, since variances were not homogenous and data were not normally distributed (Shapiro-Wilk normality test). The level of significance for all statistical tests was α = 0.05. Comparisons of mesostigmatid mites communities between ant nests were made using Euclidean distance. The index was used to create resemblance matrices of pair-wise comparisons among all the wood decay classes. The resemblance matrices were then analysed in a hierarchical cluster analysis, employing an unweighted pair-group method with arithmetic averaging (UPGMA).
Forest
Ecotone
Meadow
Garden
– – – 2.17 – 4.35 – – 4.35 – – – – – – 2.17 2.17 – 56.52 19.57 – – 4.35 – – 4.35
2.63 – – 5.26 5.26 13.16 – – 2.63 – – 5.26 2.63 – – – – 10.53 28.95 5.26 5.26 10.53 – – 2.63 –
– 3.33 – 10.00 – – 6.67 3.33 – – 3.33 6.67 – 3.33 – – – 36.67 – 16.67 – 3.33 3.33 3.33 – –
– – 5.56 11.11 – 11.11 – 11.11 16.67 5,56 – – – – 11.11 – 11.11 – 11.11 – – 5.56 – – – –
100
100
100
100
Results Totally 132 mites were collected, of which 26 species were identified (Table 1). In general, the total number of species (alpha diversity) was diverse between microhabitats (various land uses) and ranged from 9 species in the forest to 13 species in the ecotone. The mean number of species did not significantly differ statistically between habitats with different land use (microhabitats) (P = 0.96), however, the number of species tended to be highest in the forest (5.3 ± 1.20). Furthermore, the mean abundance of mesostigmatid mites did not significantly differ between study sites (P = 0.47), although the highest abundance increased from 6.0 ± 1.53 in the garden to 15.3 ± 6.17 in the forest. The diversity did not significantly differ between study plots located in sites with different land use and ranged from 1.19 ± 0.08 in meadow to 1.39 ± 0.05 in the ecotone (Table 2). On the other hand, the evenness index (E ) was generally highest in the garden (0.98 ± 0.07) and the lowest in the meadow 0.79 ± 0.10. The similarity relationships between the species composition of the mesostigmatid mites fauna in ant nests in different land uses/habitats is demonstrated by a cluster analysis of the species identity index (Euclidean distance) (Fig. 1). The dendrogram shows that the microhabitats were divided into two main clusters: communities from forest and ecotone as well as meadow and garden. Generally, the most numerous species were Trachyuropoda formicaria (39 ind.), Trichouropoda ovalis
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J. Kamczyc & D.J. Gwiazdowicz
Table 2. Diversity of mesostigmatid mites in Lasius flavus ant nest. Results are shown as a mean ± SEM.
Total number of species Mean number of species Total abundance Mean abundance Shannon (H) Eveness (E) Explanations:
∗
Forest
Ecotone
Meadow
Garden
Significance∗
9 5.3 ± 1.20 46 15.3 ± 6.17 1.32 ± 0.02 0.86 ± 0.12
13 5.0 ± 1.16 38 12.6 ± 2.33 1.39 ± 0.05 0.91 ± 0.06
12 4.6 ± 2.03 30 10.0 ± 4.73 1.19 ± 0.08 0.79 ± 0.10
10 4.3 ± 0.89 18 6.0 ± 1.53 1.37 ± 0.03 0.98 ± 0.07
– NS – NS NS NS
Based on Kruskal-Wallis H test; NS – not significant (P > 0.05).
Table 3. Total abundance and dominance of mesostigmatid mites collected from Lasius flavus workers. Species not recorded in the ant nest are marked by bold. Species 1 2 3 4 5 6 7 8 9 10 11
Abundance (ind.)
Dominance (%)
Discourella modesta (Leonardi, 1899) Eviphis ostrinus C.L. Koch, 1836 Hypoaspis aculeifer (Canestrini, 1883) Hypoaspis vacua (Michael, 1891) Nenteria sp. Pachyseius humeralis Berlese, 1910 Parasitus fimetorum (Berlese 1904) Pergamasus norvegicus (Berlese, 1906) Trachyuropoda formicaria (Lubbock, 1881) Trichouropoda sp. Uropodina sp.
1 1 1 3 1 1 1 1 2 1 1
7.14 7.14 7.14 21.43 7.14 7.14 7.14 7.14 14.29 7.14 7.14
Total
14
100.00
Fig. 1. Cluster analysis of quantitative community data in ant nests based on different land use (UPGMA method, Euclidean distance).
(16 ind.) and Rhodacarus mandibularis (15 ind.). In the forest and ecotone, Trachyuropoda formicaria dominated in the community with the abundance of 26 and 11 individuals, respectively. Rhodacarus mandibularis was the most numerous species in the meadow. The abundance of mites in the garden was similar with the highest abundance of Pachylaelaps furcifer (3 ind.). As a result of phoresis investigation, 14 mites classified to 11 species occurred on Lasius flavus workers. Generally, they occurred in low abundance (mainly 1 individual per 100 workers). We have recorded three specimens of Hypoaspis vacua and two of Trachyuropoda formicaria (Table 3). Moreover, the investigation revealed that Eviphis ostrinus, Nenteria sp. and Pergamasus norvegicus occurred exclusively on ants workers. Discussion The main goal of this study was to determine differences in mite diversity, mite community structure and
the phoresis of mites on Lasius flavus ants in four types of land use. We expected that differences between study sites/land cover types would be reflected in dissimilarity in the mean number of species, mean and total abundance. Although the direct effects of land use/land cover types on either of these parameters were not statistically significant, we observed a tendency of decrease of the mean and total abundance and mean number of mite species with the gradient of land use/land cover types. Our results are partly in accordance with those of Minor & Ciancialo (2007) who studied soil microarthropods in a similar gradient of land use/land cover type from agricultural sites to forests in the order: “corn fields – willow plantation – herbaceous old/shrubby fields – forests”. Generally, they stated that the land use type influenced the diversity of soil microarthropods, but there is a difference between Oribatida and Mesostigmata. This may explain only the tendency we have recorded in this study on mesostigmatid mites. Initially, we assumed that the species diversity of mesostigmatid mites in ant nests would be depended on the aboveground environment and we would find different mite communities in ant nests located in various sites. Our hypothesis was supported by Koehler (1999) who stated that the vegetation layer greatly influenced soil fauna abundance and diversity. Moreover, the aboveground flora affects the development of soil fauna indirectly, by providing favorable microclimate or by influencing their food (Mueller 1990). Surprisingly, the results of clustering analysis revealed that the mite communities were grouped into two groups: communities of ant nests in agricultural sites (meadow and garden) and those with the “forest” vegetation, e.g.,
Soil mites in yellow ant nests ecotone and forest site. Similarly, Maraun et al. (2003), who studied Oribatida and Collembola, indicated that disturbances strongly affect the community structure of soil microarthropods and their diversity was generally reduced due to disturbances. Our study also indicated that the ant nests in forest and ecotone sites were generally colonized by (1) large predatory species common in forest litter, e.g., Pergamasus brevicornis, Pergamasus septentrionalis and Veigaia nemorensis (Karg, 1993) and by (2) species recorded mainly in ant nests, e.g., Trachyuropoda formicaria (Wisniewski & Hirschmann 1993). Moreover, we found 26 mite species in ant nests and only four of them had been earlier recorded form Lasius flavus nests, i.e., Hypoaspis vacua, Pergamasus septentrionalis, Trachyuropoda formicaria and Zercon peltatus peltatus (Wi´sniewski 1983, unpublished; Skorupski 2001; Skorupski & Gwiazdowicz 1996, 2002). Actually, the checklist of mesostigmatid mites associated with L. flavus was enlarged and consists of 55 species. Lower numbers of mite species were recorded form Lasius fuliginosus (Latreille, 1798) – 43 species, Lasius niger (L., 1758) – 44 species, Lasius umbratus (Nylander, 1846) – 24 species and Lasius alienus (Foerster, 1850) – only one species (Gwiazdowicz 2008). Although our investigation did not confirm the hypothesis that the highest diversity and total number of mite species in ant nests occur in ecotones, we observed a tendency. This can be explained by the theory of “positive effect” of boundary zone on diversity and abundance. The theory predicts that the boundary zone (ecotone sites in our study) is characterized by the highest species diversity and abundance/density (Odum 1982). This was confirmed, e.g., by Seniczak et al. (1996) who indicated the highest diversity of Oribatida, Gamasida and Actinetida in ecotone zones between meadows and forest islands. We have also investigated phoresis of mites on L. flavus workers. Our results support previous records of phoresis of mites on insects (Wisniewski & Hirschmann 1992; Bajerlein & Bloszyk 2004; Gwiazdowicz et al. 2011), however, phoresis of mites on L. flavus workers is unique. Generally, we recorded 11 mite species occurring on ant workers, however, except Hypoaspis vacua and Trachyuropoda formicaria, they occurred as single individuals. In conclusions, the species diversity of mesostigmatid mites in nests of the ant L. flavus can be associated with the gradient of land use, the ant nests in disturbed areas are characterized by lower mites diversity. Moreover, our results revealed that the ecotone is characterized by the highest diversity of mites in L. flavus nests. Finally we concluded that some mesostigmatid mites use L. flavus workers for transportation, but this phenomenon has been observed in a small scale.
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