Biologia 65/1: 122—127, 2010 Section Zoology DOI: 10.2478/s11756-009-0222-4

Chemical properties of forest soils as affected by nests of ruginodis (Formicidae)

Adam Véle1,2,JanFrouz3,4, Jaroslav Holuša1,5 &JiříKalčík3

1Forestry and Game Management Research Institute Office Frýdek-Místek, Nádražní 2811,CZ-73802 Frýdek-Místek, Czech Republic; e-mail: [email protected] 2Department of Ecology and Environmental Sciences, Faculty of Science, Palacky University in Olomouc, tř. Svobody 26, CZ-77146 Olomouc, Czech Republic 3Institute of Soil Biology, Biology Centre AS CR, Na Sádkách 7,CZ-37005 České Budějovice, Czech Republic 4Institute for Environmental Studies, Faculty of Science, Charles University, Benátska 2,CZ-12801 Praha, Czech Republic 5Department of Forest Protection, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 1176,CZ-16521 Praha 6–Suchdol, Czech Republic

Abstract: Chemical properties (total and available P concentration; oxidizable C concentration; available K, Na, and Ca concentration; and pH) were quantified for 33 nests of the Myrmica ruginodis and in surrounding soil in young spruce forest stands. All properties, except total P, were significantly higher in the nests than in the surrounding soil. Total P was not higher in nests than in surrounding soil across all nests because nests had higher total P than surrounding soil if the soil contained low concentrations of total P but nests had lower total P than surrounding soil if the soil contained high concentrations of total P. The effect of nests on total P in the surrounding soil corresponded with effects of nests on oxidizable carbon (an indicator of organic matter) in the surrounding soil (concentrations of oxidizable carbon and total P were closely correlated). Available P concentrations were much higher in nests than in surrounding soil. Overall, the results indicated that two main processes explain the chemical changes of soil in the ant nests: (i) mixing due to excavation of deeper soil layers and (ii) deposition of excreta and food residues. The effect of soil mixing (whereby transport mineral soil from deeper layers to layers near the surface) is more pronounced in soils with high organic content near the surface because mixing increases the proportion of mineral soil in the nest while decreasing the proportion of organic matter and the concentration of total P.

Key words: ant nests; Cox ; Myrmica; phosphorus; soil chemical properties; spruce forest

Introduction Frouz et al. 2003; Holec & Frouz 2006). Transfer of ma- terial from deeper soil layers and the mixing of this Nest-building ants extensively modify the physical and material with surface soils can change soil texture (Fol- chemical properties of soil (Dlusskij 1967; Petal 1978, garait 1998). 1991; Lobry de Bruyn & Conacher 1990; Lobry de These changes to soil caused by ants affect many Bruyn 1999; Nkem et al. 2000), and create specific and other organisms. Ant nests contain a greater biomass patchy environments for other soil biota and plants of microfauna and microflora than surrounding soil (Petal 1978, 1991; Wang et al. 1995; Laakso & Setala (Dauber & Wolters 2000; Folgarait et al. 2002; Holec 1997; Wagner et al. 1997; Dauber et al. 2006). The effect & Frouz 2006). Plant rhizomes are thinner and have of ants on soil structure and nutrient enrichment, how- longer internodes and more branches in the nests of the ever, is highly variable (Dlusskij 1967; Petal 1978, 1991; ant Lasius flavus (F., 1782) than in the surrounding Lobry de Bruyn & Conacher 1990; Lobry de Bruyn soils (Dostál et al. 2005). Ants also can increase the 1999). How ants affect soil properties is determined by colonization of roots by arbuscular mycorrhizal fungi many factors, including -specific factors such as (Dauber et al. 2008). Soil affected by ants has higher nutritional demand, feeding behavior, nesting strategy, productivity than unaffected soil (Folgarait et al. 2002), mound size, and colony age (Petal 1978, 1980; Dauber and ants can change the course of vegetation succes- & Wolters 2000; Nkem et al. 2000; Wagner et al. 2004). sion (Jonkman 1978). Even abandoned nests can pro- Moreover, the effect of ants on soil also varies according vide habitat for shrubs (DeSlippe & Savolainen 1995). to the properties of the surrounding soil (Dlusskij 1967; Although much information is available concern- Czerwinsky et al. 1971; Jakubczyk et al. 1972; Frouz et ing how soil properties are affected by ants that build al. 2003) and may vary along successional gradients and aboveground nests (Dlusskij 1967; Dauber & Wolters with landscape position (Whitford & DiMarco 1995; 2000; Frouz et al. 2003, 2005), little information is avail-

c 2009 Institute of Zoology, Slovak Academy of Sciences Chemical properties of soils affected by Myrmica ruginodis 123 able on how soil properties are changed by subterranean passed through a 2-mm sieve. The concentration of total ant nests. In this study, we focused on the effect of P(Ptot) was determined after P was mineralised with per- the ant Myrmica ruginodis Nylander, 1846 on soils of chloric acid (Sommers & Nelson 1972). Available P (Pav ) a spruce forest of various ages after clear cutting. Myr- in filtered extracts was measured by the cation exchange mica ruginodis is a common species in European forests resin method (Macháček 1986). Water-soluble P (Pwat )was measured in a filtered water extract (sample:water 1:100). and builds shallow subterranean nests (Seifert 1996). In P content in mineralized soil and extracts was measured particular, we wanted to learn how the changes in soil according to Murphy & Riely (1962) as modified by Wa- properties caused by ants correspond with the chem- tanabe & Olsen (1965). Concentration of total oxidizable istry of the surrounding soil, and how the changes in C(Cox ) was determined using wet combustion (oxidation soil properties are affected by nest size and stand age with chromic acid in excess of sulfuric acid) (Jackson 1958). (time since the last clear cut). Concentrations of available Ca, Na, and K were determined using ion sensitive electrodes in water extracts in 1% cit- ric acid solution (sample:liquid ratio 1:5; shacked 1 h) ac- Methods cording K¨onig (1923) as modified by Kubíková (1970). The pH was measured either in water or in a KCl solution (1:5 The study was done in one large Norway spruce, Picea abies sample liquid ratio). The electrical conductivity (EC) was (L.) Karts., forest complex located on Černá Studnice Hill measured in filtrate of 1:5 suspension in H2O. ◦   ◦   (869 m a.s.l., 50 42 29.57 N; 15 12 31.72 E) near Jablonec The dependence between proportions of nest and be- nad Nisou (Czech Republic) at 620–760 m altitude. The tween nest volume and age of forest stand were determined ◦ area has a mean annual temperature of 7 C and an annual by correlation. Individual chemical properties in nests and precipitation of 1000 mm (Culek 1996). The soil was an in surrounding soil were compared with a paired t-test. The oligotrophic clayey sandy cambisol ranging in depth from relationship between chemical properties of the nests vs. the 30 to 50 cm. The original beech woods had been replaced surrounding soils was determined by correlation. Statistical by spruce monocultures at the end of eighteenth century evaluations used the SPSS 10.0 program package. (Culek 1996) so that now spruce is the predominant species in this rugged upland. Results Myrmica ruginodis nests were sampled in 15 young sec- ondary spruce (Picea abies) forest stands during spring and summer months of 2005. We separated these forest stands We removed 33 nests from 10 forest stands; in five into three age categories: 0–2, 3–5, and 8–19 years old. Each stands, nests were either absent or inaccessible. The category was represented by five replicate stands. These average density of these nests was 1,777 per hectare forests stands were about 1 ha wide. In each forest stand, (4.8 per plot). Minimum of nests were found in younger we randomly marked three squares; each square was 3 × 3 2 forest stands. Excavated nests were typically circular m, giving an area of 27 m for each stand. Using shovels, and only slightly longer than wide; the average length we attempted to excavate all the M. ruginodis nests in each (±SD) was 16.5 ± 12.9 cm and the average width was square but stones and stumps sometimes prevented nest re- ± moval. During excavation, the nest’s longest horizontal di- 13.4 10.9 cm. Length and width were closely corre- mension and the dimension perpendicular to it were mea- lated (r = 0.938, P < 0.05). There was no correlation sured and are referred to as length and width. The depth of between the nest horizontal dimensions and nest depth; the nest (the deepest part with chambers) was also recorded. average nest depth was 7.8 ± 3.5 cm. Average nest vol- The nest volume was determined as the volume of roll. Four ume was 2.1 ± 2.5 dm3, and the nest volume was more 3 evenly spaced subsamples of soil (ca 100 cm each) were closely correlated with both horizontal dimensions (r = also collected from the soil surrounding (within 1 m) each 0.838 and 0.849 for length and width) than with depth nest; the subsamples were combined to form one sample of (r = 0.424), although all of these correlations were sig- surrounding soil per nest. nificant (P < 0.05). Nest volume was not correlated The excavated nest material and soil samples were placed in plastic bags and transported to the laboratory with stand age. The number of ants in a nest was cor- where samples were frozen at –20 ◦C for 24 h. Ants were hand related (r = 0.56, P < 0.01) with nest volume (ant num- 3 sorted out of the nest material, and the nest material (minus ber = 72.43V + 88.51, where V is nest volume in dm ). ants) and surrounding soil samples were air-dried and then On average, each nest contained 248 ± 263 workers.

Table 1. Concentration (mean and SD) of chemicals in Myrmica ruginodis nests and in surrounding soil; P values indicate significant differences based on paired t-tests (n = 33).

Chemicals Nest Soil P

pH 3.8 (0.36) 3.4 (0.16) >0.001 −1 Ptot (mg kg ) 943.6 (265.2) 855.6 (217.0) 0.194 −1 Pav (mg kg ) 112.3 (65.8) 29.6 (16.1) >0.001 −1 Pwat (mg kg ) 88.3 (57.0) 27.0 (14.6) >0.001 Cox (%) 19.2 (7.6) 13.9 (5.1) 0.006 Na (mg kg−1) 114.4 (98.4) 57.6 (39.0) 0.007 K(mgkg−1) 414.5 (279.2) 142.1 (104.7) >0.001 Ca (mg kg−1) 130.5 (148.6) 24.9 (17.8) >0.001 EC (µScm−1) 392.6 (192.6) 223.3 (75.9) >0.001 124 A. Véle et al.

1000 centrations were more than 3-fold higher in the nest y = -1.2014x + 1115.9 800 than in the surrounding soil. The proportion of Pav rel- R2 = 0.4985 600 ative to Ptot was higher in nest than in surrounding soil. 400 The difference between nest Ptot and soil Ptot was nega- 200 tively correlated with soil Ptot, such that the difference 0

increase in nest was positive in soils with low Ptot (higher values in the -200 tot tot

P nest than in the soil) and negative in soils with high P -400 -600 (higher values in the soil than in the nest) (Fig. 1, Ta- 400 600 800 1000 1200 ble 2). This pattern corresponded with organic matter properties as indicated by Cox. In the surrounding soil, P in soil tot Ptot was correlated with Cox (Table 3). As was true for changes in nest Ptot relative to soil Ptot, the difference 30 y = -1.311x + 23.552 between nest Cox and soil Cox was negatively correlated 25 2 R = 0.4514 with soil Cox, such that the difference was positive in 20 15 soils with low Cox (higher values in the nest than in the 10 soil) and negative in soils with high Cox (higher values 5 in the soil than in the nest) (Fig. 1, Table 2). Conse- 0 quently, Ptot increased in the nests built in mineral soils increase in nest -5

ox with low Cox and decreased in highly organic soils with

C -10 -15 high Cox concentration (Fig. 1, Table 2). 0 102030 Changes in nest pH relative to soil pH were nega- tively correlated with soil pH and positively correlated Cox in soil with soil Cox (Table 2); in other words, increases in nest pH were more pronounced in soil with low pH and high 1000 y = -42.492x + 680.36 2 organic matter content. Highly organic soils are likely 800 R = 0.3506 to be more acidic than mineral soils because soil Cox 600 and soil pH were negatively correlated (Table 3). 400 Changes in the chemical properties in the nests 200 were not correlated with nest size. Nest EC and nest 0

increase nest in -200 available K were positively correlated with stand age tot

P -400 (Table 2). Also, soil EC was positively correlated with -600 stand age (Table 3). 0102030 Discussion Cox in soil

Fig. 1. Correlation between Ptot increase in nests and Ptot in In Lesica (1998), the volume of Myrmica nests ranged soil (top), Cox increase in nests and Cox in soil (middle), Ptot from 0.1 to 15.5 dm3, with an average of 1.1 dm3.In increase in nests and Cox in soil (bottom). this study, the average nest volume was 2.1 ± 2.5 dm3, and nest volume and dimensions were not correlated with stand age. This indicates that all of the stands Concentrations of all of the measured soil chemi- in this study, regardless of age (2–19 years), had the cals, except (Ptot), were higher in the nest than in the same properties for ants. Previous research has demon- surrounding soil (Table 1). In contrast to Ptot,Pav con- strated that ant abundance per nest decreases when re-

Table 2. Correlation coefficients (R2) between properties of forest soil and difference between ant nest and surrounding soil for chemical properties (n = 33). Only significant (P < 0.05) correlation coefficients are given. The first column contains the full name and units for individual properties.

Difference between ant nest and soil properties Forest soil properties pH Ptot Pav Pwat Cox Na K Ca EC

Stand age (years) 0.350 0.448 pH –0.494 0.410 0.406 0.520 −1 Ptot (mg kg ) –0.706 –0.455 –0.449 –0.561 −1 Pav (mg kg ) 0.433 −1 Pwat (mg kg ) 0.494 –0.350 Cox (%) 0.456 –0.592 –0.466 –0.498 –0.672 Na (mg kg−1) –0.682 –0.478 –0.456 –0.598 –0.438 –0.607 –0.490 K(mgkg−1) –0.587 –0.359 –0.366 –0.403 –0.477 Ca (mg kg−1) EC (µScm−1) –0.672 –0.438 –0.381 –0.490 Chemical properties of soils affected by Myrmica ruginodis 125

Table 3. Correlation coefficient (R2) among forest soil properties (soil unaffected by ants; n = 33, P < 0.05).

Difference between ant nest and soil properties Forest soil properties EC Ptot Pav Pwat Cox Na K Ca

Stand age (years) 0.472 pH –0.365 –0.582 EC (µScm−1) 0.889 0.607 0.508 0.766 0.448 0.551 −1 Ptot (mg kg ) 0.486 0.521 0.755 0.508 0.524 −1 Pav (mg kg ) 0.938 0.756 −1 Pwat (mg kg ) 0.736 Cox (%) 0.434 Na (mg kg−1) 0.841 K(mgkg−1) Ca (mg kg−1) sources are insufficient (Sorvari & Hakkarainen 2007). ganic content. Because pH is negatively correlated with We found that the number of ants in a nest correlates organic matter content, mixing in organic soils tends with nest volume. This finding agrees with the results to reduce nest Cox but increase nest pH. No correla- of Seifert (1991), who studied aboveground rather than tion was found between change in nest chemical prop- belowground nests. erties and nest size, although Nkem et al. (2000) noted Concentrations of Cox and most other chemicals a negative relationship between mound size and Ca con- were higher in the nests than in the surrounding soil, centration. This may be because nests of Myrmica are which is in agreement with several authors (Czerwinsky too short lived (Seifert 1996) to produce an increase et. al. 1971; Jakubczyk et al. 1972; Petal 1980; Frouz et in nutrient accumulation with nest age as observed by al. 2003) who observed higher concentrations of avail- Zacharov et al. (1980) for some Formica nests. Gen- able K, Ca, Na, and P and increased pH in nests of erally, two main processes seem to be responsible for Formica, Lasius,andMyrmica spp. An increase in P the changes in chemical properties of ant nests: soil concentration, as well as K concentration (King 1977), mixing during nest building and accumulation of ant was also described for other ant species (Dlusskyj 1967; excreta and food residues in the nest (Kilpelainen et Petal 1978; Pokarzhevskij 1981; Zacharov et al. 1981; al. 2007). Of the chemical properties measured in this Culver & Beattie 1983; Lobry de Bruyn 1999; Frouz study, the concentrations of Ptot and Cox were most af- et al. 2003, 2005). In the current study, however, Ptot fected by soil mixing. Second group of processes is con- did not differ between the nest and surrounding soil. nected with accumulation of excreta and food residues This result reflects the negative relationship between in the nest (Frouz et al. 1997; Lobry de Bruyn 1999). Ptot and organic matter. We expect that a decrease in Our study documents a positive effect of M. ruginodis P concentration in the nest is due to soil mixing (to on soil chemical properties in their nests. The el- transport of mineral soil from deeper layers to shallower ements that were in higher concentration in the nests layers). Because Ptot is mainly concentrated in organic than in the surrounding are also important nutrients for matter, such mixing reduces not only organic matter plants (Procházka et al. 1998). Similar results have been (Cox) but also Ptot concentration, and as indicated by reported for other ant species (Folgarait 1998; Cammer- Frouz et al. (2005), the effect of soil mixing is more aat & Risch 2008), and research has documented that pronounced when soil layers near the surface soil have these changes benefit plants and soil (Dauber a high organic matter content. &Wolters 2000; Folgarait et al. 2002; Dostál et al. 2005; In contrast to total P concentration, the concentra- Holec & Frouz 2006). The effect of ant nests is especially tions of available forms of P were greater in the nests important in spruce monocultures, where soils have raw than in the surrounding soil. This is in agreement with humus accumulation, low pH, and poor nutrient dy- Frouz & Kalčík (1995) and Frouz et al. (2003, 2005), namics (Podrázský & Remeš 2005; Pavlů et al. 2007). who observed that ant nests caused much higher in- Ants affect the rate of decomposition, which affects soil creases in available forms of P than in total P. De- fertility (Troeh & Thompson 2005; Cammeraat & Risch position of food residues and excreta in the nest may 2008). Because Myrmica ants improve the soil and be- increase the ratio of Pav relative to Ptot. When the top cause young forest stands in secondary spruce forests layers of the soil are largely mineral (with low organic are important environments for Myrmica ants (Véle et matter content), soil mixing by ants will decrease to- al., in litt.), such young forest stands should be encour- tal P only to a small degree because such soils already aged. contain low levels of total P. In this case, deposition of feed and excreta may even cause an increase in P Acknowledgements concentration (Fig. 1). 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Received March 6, 2009 Accepted September 15, 2009