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1 This article was published in an Elsevier journal. The attached copy is furnished to the author for non-commercial research and education use, including for instruction at the author s institution, sharing with colleagues and providing to institution administration. Other uses, including reproduction and distribution, or selling or licensing copies, or posting to personal, institutional or third party websites are prohibited. In most cases authors are permitted to post their version of the article (e.g. in Word or Tex form) to their personal website or institutional repository. Authors requiring further information regarding Elsevier s archiving and manuscript policies are encouraged to visit:
2 Soil Biology & Biochemistry 39 (2007) Oribatid mite communities and foliar litter decomposition in canopy suspended soils and forest floor habitats of western redcedar forests, Vancouver Island, Canada Zoë Lindo, Neville N. Winchester Department of Biology, University of Victoria, P.O. Box 3020, Station CSC, Victoria, BC, Canada V8W 3N5 Received 20 April 2007; received in revised form 20 June 2007; accepted 22 June 2007 Available online 16 July 2007 Abstract Litter decomposition and changes in oribatid mite community composition were studied for 2 years in litterbags collected from arboreal organic matter accumulations (canopy suspended soils) and forest floors associated with western redcedar trees on Vancouver Island, British Columbia. We tested the hypotheses that lower rates of mass loss, higher nutrient levels, and different patterns of oribatid mite richness and abundance in decomposing western redcedar litter would be observed in litterbags associated with canopy suspended soils compared to forest floors. Decomposition, measured by mass loss of cedar litter in litterbags, was not significantly different in canopy and forest floor habitats, although reduced in the canopy. Abundance and richness of oribatid mites inhabiting litterbags were significantly greater on the forest floor compared to the canopy suspended soils. Canopy suspended soils had higher levels of total nitrogen, available phosphorus and potassium than the forest floor, but moisture content was significantly lower in the suspended soils. Higher nutrient levels in the canopy system are attributed to differences in coarse woody debris input (but not foliar litter), combined with reduced nutrient uptake by roots and lower mobilisation rates of nutrients by detritivorous and fungivorous microarthropods. Moisture limitation in the canopy system possibly contributed to lower mass loss in litterbags, and lower abundance and richness of oribatid mites in litterbags placed on canopy suspended soils. Patterns of oribatid mite community composition were related to mite communities associated with the underlying substrate (forest floor or canopy suspended soil) which act as source pools for individuals colonising litterbags. Successional and seasonal trends in oribatid mite communities were confounded by moisture limitation at 24 months, particularly within the canopy habitat. r 2007 Elsevier Ltd. All rights reserved. Keywords: Canopy; Decomposition; Litterbag; Oribatid mites; Suspended soil; Temperate rainforest; Western redcedar 1. Introduction Decomposition is an important process in the functioning of forest ecosystems (Swift et al., 1979), releasing nutrients bound in organic matter and enabling plant nutrient uptake (Aber and Melillo, 1980). Rates of decomposition are often correlated with climate (temperate and precipitation) (Trofymow et al., 2002), the quality of the decomposing material (Preston et al., 2000; Trofymow et al., 2002), and the composition of the decomposer community (Moore et al., 1988). Soil microarthropods Corresponding author. Tel.: ; fax: address: zlindo@uvic.ca (Z. Lindo). (mites (Acari) and springtails (Collembola)) contribute to decomposition and nutrient cycling processes (Seastedt, 1984), though their participation is mostly indirect via their interactions with the microbial community (Moore et al., 1988). The abundance, composition and activity of the decomposer community are also directly affected by available litter quality, and thus interactions among climate, litter quality and the decomposer community are important regulator of decomposition and nutrient release. Decomposition processes have primarily been studied in belowground habitats such as forest floor systems, but decomposition starts in the canopy with senescence of litter, litterfall, and litter deposition to the forest floor (Hempfling et al., 1991). Additionally, in structurally /$ - see front matter r 2007 Elsevier Ltd. All rights reserved. doi: /j.soilbio
3 2958 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) complex forest ecosystems like natural temperate and tropical rainforests, litter is also deposited and retained within the canopy (Nadkarni and Matelson, 1991; Lindo and Winchester, 2007a). In ancient temperate rainforest ecosystems, where individual trees range from years old, litter deposition within the canopy, combined with dead organic matter derived from epiphytic plants and bryophytes, has decomposed to form canopy suspended soils (Wallwork, 1976; Enloe et al., 2006). In a tropical cloud forest, Vance and Nadkarni (1990) found canopy suspended soils were more acidic than forest floor organic horizons, but concentrations of carbon, other nutrients, and microbial biomass were similar to those for the forest floor. Organic suspended soils within canopy systems experience greater temperature extremes and more frequent wet dry cycles than the forest floor (Bohlman et al., 1995), which can further alter decomposition rates in canopy systems (Nadkarni and Matelson, 1991) compared with ground habitats. In addition to being a repository for nutrients, canopy suspended soils are an important habitat for many invertebrate fauna, in particular, oribatid mites (Behan- Pelletier and Walter, 2000; Lindo and Winchester, 2006). Oribatid mites inhabiting canopy suspended soils generally are lower in diversity and abundance than those of forest floor habitats, and differ in species composition where canopy specificity is approximately 30% (Behan-Pelletier and Walter, 2000; Lindo and Winchester, 2006). Oribatid mites are often the most dominant fauna represented in litter decomposition studies (Gonza lez and Seastedt, 2000) and the diversity and composition of the faunal community colonising litterbags is usually determined by local habitat attributes and site factors such as elevation (Walter, 1985). Differences in microclimate between canopy and forest floor soils may also contribute to the observed high complementarity (low similarity) of oribatid mite species colonising litterbags in canopy versus forest floor habitats (Fagan et al., 2006). Decomposition in canopy ecosystems is expected to differ from that of forest floor ecosystems due to biotic factors such as differences in diversity of oribatid mite and other microarthropod assemblages, and abiotic parameters (e.g. temperature, moisture) that operate at microscale levels. To date, mass loss from decomposing litter in a canopy ecosystem has not been studied in temperate forests, and in this study we quantified: (1) decomposition in canopy (suspended soils) and ground (forest floor organic layer) habitats based on mass loss of foliar western redcedar (Thuja plicata D. Don) litter in litterbags over two years in a coastal temperate forest; (2) oribatid mite abundance and species richness, and described the oribatid mite community which colonizes litterbags placed in canopy and forest floor habitats over two years; and (3) nutrient levels (total N, P, K, and ph) in canopy suspended soil and forest floor habitats. We addressed the following three hypotheses: (1) decomposition will be lower in the canopy due to lower richness and abundance of the oribatid mite fauna, and lower moisture content of suspended soils compared to that of forest floor habitats (Nadkarni and Matelson, 1991); (2a) abundance of oribatid mites would be lower in canopy versus forest floor litterbags since colonisation abundances are expected to reflect abundances in canopy and forest floor habitats (Behan-Pelletier and Walter, 2000), and (2b) the oribatid mite community composition would differ between canopy and forest floor litterbags because the composition of source pools available for colonisation (suspended soil and forest floor communities) have been shown to be different (Lindo and Winchester, 2006); and (3) nutrient levels would be similar in canopy suspended soils due to similarities in microbial biomass (Vance and Nadkarni, 1990). 2. Materials and methods 2.1. Site description and experimental design The study site was an ancient ( years old) temperate coniferous forest in the Walbran Valley located on the southwest coast of Vancouver Island, British Columbia, Canada ( N, W) (Lindo and Winchester, 2006). Mean annual temperature is 9 1C (70.6), and mean annual precipitation is 2990 mm ( e.html [cited 9 April 2007]). The experimental site is 1 ha, facing southeast at 200 m elevation. Western hemlock (Tsuga heterophylla (Rafn.) Sarg.), Sitka spruce (Picea sitchensis (Bong) Carr.), silver fir (Abies amabilis (Dougl.) Forb.), and western redcedar (Thuja plicata D. Don) are dominant conifers in this rainforest. The soils of the Walbran Valley are deep, organic, overlying shallow mineral, unconsolidated or fractured bedrock. Moisture levels are high throughout the year, though saturation levels vary greatly depending on the slope. The forest floor organic layer consists of 10 cm or more LFH horizon derived from leaf and needle litter, twigs, branches and mosses. Well-decomposed woody material from dead trees is also abundant in these soils. Soils within the study area conform to the Folisol great group under the Canadian System of Soil Classification (Soil Classification Working Group, 1998). The western redcedar litter used for decomposition in litterbags was collected from the study site one month prior to the commencement of the experiment. Senesced western redcedar (cedar) foliage (scale leaves) was stripped from branch tips and brought to the laboratory. The litter was mixed thoroughly, dried in a 70 1C oven for 24 h, removed and left at room temperature for 2 3 h before filling. Five grams of cedar foliage were added to each of 144 nylon mesh (1 mm) 15 7 cm litterbags. The initial moisture content of the litter (expressed as percent dry weight (d.w.)) used in each litterbag was 3.5% based on further drying of foliage (% moisture ¼ (wet weight (g) d.w. (g)/d.w. (g)) 100)).
4 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) Litterbags were placed on forest floor and canopy suspended soil habitats associated with six western redcedar trees on 10 September Three replicate litterbags were placed 1.5 m from the base of each tree at the four cardinal directions on the forest floor. Single rope climbing methods were used to access the suspended soils and fix litterbags in the high canopy of the same six trees. Suspended soils in western redcedar form only where trunk reiterations, limb junctions and other structural formations allow for the accumulation of litter, thus suspended soils occur in discrete patches. Three replicate litterbags were placed on the top of the four largest suspended soil patches (average surface area of 0.85 m 2, min. ¼ 0.30 m 2, max. ¼ 2.76 m 2 ) at a mean height of 35 m from the ground (min. ¼ 21 m, max. ¼ 45 m) (Lindo and Winchester, 2006). One litterbag from each sample point was collected after 6, 12 and 24 months in the field ((4 forest floor+4 suspended soil) 6 trees ¼ 48 litterbags per collection date; 144 litterbags total). Litterbags were placed in separate plastic bags during collection and transported to the laboratory for microarthropod extraction. The outside litter accumulation was brushed off and litterbags were placed on modified Berlese-type extractors over 75% EtOH for 48 h. Following microarthropod extraction, litterbags were placed in a 70 1C drying oven for 24 h. Litterbags were then cut open and the contents removed and weighted. There were no ingrown roots in the litterbags, but any moss propagules were carefully removed prior to weighing the sample. Litter decomposition was estimated as percent mass loss from decomposing foliage, calculated as the difference in 70 1C oven dry weight before and after 6, 12 and 24 months (% mass loss ¼ (g d.w. cedar litter before g d.w. cedar litter after /g d.w. cedar litter before ) 100)). At the time of litterbag placement, core samples of canopy and ground habitats (PVC corers 160 series, cm diameter) were collected from each sample point ((4 forest floor+4 suspended soil) 6 trees ¼ 48 core samples) for evaluation of canopy suspended soil and forest floor nutrient levels and moisture content. The average depth of core samples collected for both canopy suspended soils and the forest floor organic horizon was 6.2 (72.4) cm, and no mineral soil was included for these analyses. The ph of the samples was measured in water with a glass electrode ph meter. Distilled water was added to field-moist organic soils to create a slurry of 1:10 soil (d.w.) to deionized water (2 g d.w. in 20 ml water). Slurry ph was measured after 1 h with a glass electrode digital ph metre. Total N concentrations were determined by the Kjeldahl procedure. Air-dried subsamples were ground to pass through a 40 mesh (0.5 mm) sieve in a Wiley mill and further dried at 80 1C for 24 h. Samples (1 g d.w.) were digested in 5 ml solution (sulphuric acid mercuric sulphate potassium sulphate solution with K 2 SO 4, H 2 SO 4 ) by heating the mixtures for 2.5 h at 380 1C. The final dilution was to 75 ml with colourimetric determination of NH 4 + by the Indophenol blue method. Total available phosphorus and potassium were extracted from 5 g d.w. using 50 ml modified Kelowna extractant (Ashworth and Mrazek, 1995) and analysed by emission spectroscopy using inductively coupled plasma (ICP). Moisture content of the canopy and ground substrates were determined at the time of litterbag placement and measured repeatedly at 12- and 24-month collection dates. Moisture content was estimated gravimetrically from soil cores by the difference in wet weight (w.t.) and dry weight (d.w.) following desiccation at 70 1C for 24 h (% moisture ¼ (w.t. (g) d.w. (g)/d.w. (g)) 100)) Specimen identification and data analyses Extracted microarthropods were sorted into major taxonomic groups: mites (Acari), springtails (Collembola) and other microarthropods, which included pseudoscorpions (Pseudoscorpiones), beetles (Coleoptera), millipedes and centipedes (Myriapoda) and spiders (Araneae). The Acari were further identified to suborder (Mesostigmata, Prostigmata, Astigmata and Oribatida) and all adult oribatid mites were identified to species. Summary results for abundances of major microarthropod groups other than oribatid mites are available in Appendix A. Oribatid mite abundance is expressed as # individuals g 1 d.w. litter. Representative oribatid mite specimens were slide mounted using Hoyer s medium and a reference collection is deposited at the Canadian National Collection of Insects and Arachnids, Ottawa, Canada. We used repeated measures ANOVA to test for the effect of time and habitat (forest floor versus canopy suspended soils) on decomposition in litterbags as measured by mass loss of needle litter, oribatid mite abundance and species richness in litterbags, and moisture content of the underlying substrate. Pearson s correlation was used to relate mass loss to oribatid mite abundance and species richness for all litterbags, and for litterbags within each collection time. A one-way analysis of variance (ANOVA) was used to test for differences in soil nutrients and ph between canopy and forest floor organic soil samples. These analyses were performed using Statistica 7.0 (StatSoft, 2004) with a significance level of a ¼ Community composition of oribatid mites inhabiting canopy and forest floor litterbags during the three sampling periods was analysed using standardized abundances (# ind. g 1 d.w.) of all species. We compared a community compositional similarity matrix based on Bray Curtis similarity of square root transformed oribatid mite species for each habitat at each sampling period in Primer 5 (Primer-E Ltd., 2001). Final assessment of significance of the random occurrence of a priori main factor effects (habitat, collection time) was based on analysis of similarities (ANOSIM) with 10,000 randomized permutations. Oribatid mite community compositional data were further analysed using principle components analysis (PCA) in Statistica 7.0. The 20 most abundant species over all sampling periods were used to create PC factors
5 2960 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) which were analysed with ANOVA for main effects of habitat and sampling period. Factor coordinates based on species correlations were used to interpret the PCA. 3. Results There was a significant effect of time on the mass loss of needle litter in litterbags placed in the canopy and on the forest floor over two years (Table 1). Decomposition, as measured by mass loss of needle litter, although not significantly so, was greater on the forest floor than in the canopy during all collection time periods (Fig. 1), and increased significantly over time in both habitats (Table 1) with a total cumulative mass loss of 51.5% (78.1 SD) in the canopy suspended soil and 54.2% (76.5 SD) on the forest floor over two years. The average standardized oribatid mite abundance (# ind. g 1 d.w. litter) at all collection dates was significantly greater in litterbags on the forest floor compared to litterbags in the canopy (Fig. 2). Time of collection had a significant effect on oribatid mite abundance with the highest abundances observed in litterbags collected after 12 months (Table 1). A total of 62 species of oribatid mites were identified from 3443 adult specimens collected from the 144 litterbags (Appendix B). Oribatid mite species richness was significantly greater in litterbags on the forest floor compared with those of litterbags from canopy suspended soils at all collection times, and highest in the forest floor habitat after 12 months (Table 1, Fig. 2). Correlation patterns between oribatid mite abundances and species richness showed a significant positive correlation, overall and within each litterbag collection time (overall: r 2 ¼ 0.835, Po0.001). Mass loss was positively correlated with oribatid mite abundance in litterbags from both canopy and forest floor materials during the highest abundance collection period (12 months) (canopy: r 2 ¼ 0.446, P ¼ 0.033; forest floor: r 2 ¼ 0.559, P ¼ 0.006), but not at 6 or 24 months. Litterbag placement habitat (canopy versus forest floor) had a significant effect on the oribatid mite community composition as did litterbag collection time based on the analysis of Bray Curtis percent community similarity (habitat: r ¼ 0.838, Po0.001; time: r ¼ 0.455, Po0.001). The 20 most abundant species showed similar results in the principle components analysis with six principle components (PC) having eigenvalues 41 and the first 5 components accounted for 72.1% of the cumulative variation in the data set. PC 1 was significantly related to litterbag habitat (F 1,32 ¼ 16.36, Po0.001) and litterbag collection time (F 2,32 ¼ 16.36, Po0.001) (Fig. 3). PC 2 was also significantly related to the litterbag collection date (F 2,32 ¼ 3.61, P ¼ 0.038) (Fig. 3). The species drivers of PC 1 based on the species correlations with the principle component coordinates were Dentachipteria sp., Liacarus sp. nr. bidentatus, andliochthonius sp.3; species driving PC 2 were Scheloribates (Scheloribates) sp., Eupterotegaeus rhamphosus, andramusella (Ramusella) sp. (Fig. 4). Litter mass loss (%) Canopy Forest floor Time in field (months) Fig. 1. Decomposition (% mass loss) of 5 g western redcedar litter in litterbags placed on the surface of canopy suspended soils and on the forest floor associated with western redcedar trees in the Walbran Valley, Vancouver Island, Canada after 6, 12 and 24 months. Table 1 Results of repeated measures ANOVA for decomposition (mass loss) of needle litter, and abundance and species richness of oribatid mites in litterbags collected from canopy and forest floor habitats at 6, 12, and 24 months, and soil moisture content of canopy and forest floor substrate underlying litterbags at 0, 12 and 24 months (September 2004, September 2005, September 2006) Variable Source of variation (df) SS F P Mass loss Repeated measures Habitat (1,10) Time (2,20) o0.001 Time Habitat (2,20) Abundance Repeated measures Habitat (1,10) Time (2,20) o0.001 Time Habitat (2,20) o0.001 Richness Repeated measures Habitat (1,10) o0.001 Time (2,20) o0.001 Time Habitat (2,20) o0.001 Moisture Repeated measures Habitat (1,10) o0.001 Time (2,20) o0.001 Time Habitat (2,20)
6 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) Canopy # ind. g -1 d.w. litter Forest floor 0 6 months 12 months 24 months Litterbag collection time 15 Canopy # species per litterbag 10 5 Forest floor 0 6 months 12 months 24 months Litterbag collection time 350 Moisture content (%) Canopy Forest floor 0 Sept Sept Sept Sampling date Fig. 2. (A) Average oribatid mite abundances (# ind. g 1 d.w.) in litterbags, (B) average oribatid mite species richness (per litterbag collected) in litterbags placed on the surface of canopy suspended soils and on the forest floor associated with western redcedar trees in the Walbran Valley, Vancouver Island, Canada after 6, 12 and 24 months, and (C) moisture content (%) of canopy suspended soil and forest floor substrate at the time of litterbag placement (September 2004) and during litterbag collection at 12 and 24 months. * denotes significance of Po0.05, ** Po0.001 between canopy and forest floor at time of litterbag collection. Error bars are standard deviations. The ph values of the canopy and forest floor environments were not significantly different, though the nutrient content (N, P, K) within canopy suspended soils was significantly greater than the forest floor (Table 2). The moisture contents were consistently significantly wetter on the forest floor compared to those of the canopy (Fig. 2). There was a significant effect of collection time and interaction effect on substrate moisture content as September 2006 was shown to be a significantly drier year than September 2004, or September 2005 (Table 1). 4. Discussion Decomposition of litter in forest ecosystems is influenced, primarily by climate conditions (macro and microclimate), and secondarily by litter quality which incorporates a suite of substrate characteristics such as nutrient (particularly N) concentration, organic composition and physical properties (Preston et al., 2000). The reduction in mass loss of western redcedar litter in the canopy versus the litterbags on the forest floor is possibly
7 2962 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) related to abiotic conditions at the microscale level, such as moisture availability, because litter quality was standardized within the litterbags. Canopy suspended soils in temperate forests have low bulk density and are susceptible to desiccation, particularly in surface layers, and in soils of exposed areas within the tree crown (e.g. on a limb versus formation within a crotch) (Enloe et al., 2006). These drier conditions compared to the forest floor may impede microbial activity, slowing decomposition in the canopy (Salamanca et al., 2003). Moisture limitation as a mechanism for reduced decomposition in canopy habitats was supported in this study by the observed difference in moisture content between the canopy suspended soil and forest floor habitats. PC factor PC factor 1 Canopy 6 mo. Forest floor 6 mo. Canopy 12 mo. Forest floor 12 mo. Canopy 24 mo. Forest floor 24 mo. Fig. 3. Plot of the 1st and 2nd principle components with average (7SE) case scores for litterbag samples collected from canopy suspended soil and forest floor habitats at 6, 12, and 24 months. Litterfall is a major process for nutrient input into soil systems (Hempfling et al., 1991). Sources of nutrient input to canopy suspended soils include autochthonous sources such as senescent bryophytes, needle litterfall, herbaceous epiphyte litter and roots, leachate from live foliage, bark decomposition, and animal inputs (fecal pellets and animal death), as well as allochthonous sources such as atmospheric wet and dry deposition or gaseous inputs (Nadkarni and Matelson, 1991), however, the canopy is less likely to input as much woody debris as the forest floor. Woody debris has lower concentrations of total N, P and K and thus the lower concentrations of total N, P and K in forest floor may be due to greater inputs of lower quality detritus. However, the main constituents of litter input to the canopy suspended soils are western redcedar litter and bryophytes (Lindo and Winchester, 2007a), which also typically have low nutrient levels, and high concentrations of polyphenols, waxes and non-polar compounds (Preston et al., 2000; Turetsky, 2003). Uptake of nitrogen from soil by bryophytes may be lower than for vascular plants because bryophytes are efficient in assimilating nitrogen from atmospheric sources and recycling nitrogen from older segments of the plant into new growth (Turetsky, 2003), thus differences in nutrient levels in the canopy are most possibly related to a reduction in nutrient uptake by roots of vascular plants. Nutrient levels and decomposition may also be related to abundances of detritivorous and fungivorous microarthropods (Persson, 1989; Joo et al., 2006), although correlations of oribatid mite abundance and richness with decomposition in this study were only observable when 1.0 Schel. E.rham T.vel R.ram PC factor 2: 13.80% K.lut Arch.1 Pilo. Denta. Allo.2 L.ros Cera.1H.gib Lioc.2 L.bid O.nova P.pelt Lioc.1 P.hyst T.tect Lioc PC factor 1: 34.21% Fig. 4. Plot of the 1st and 2nd principal components with % explained variation for adult oribatid mite assemblages (top 20 species based on standardized abundances) collected from litterbags in canopy and ground habitats after 6, 12 and 24 months. Species abbreviations are for: Palaeacarus hystricinus, Liochthonius sp.1, Liochthonius sp.2, Liochthonius sp.3, Archiphthiracarus sp.1, Platynothrus peltifer, Tryhypochthonius tectorum, Hermannia gibba, Eupterotegaeus rhamphosus, Liacarus sp. nr. bidentatus, Liacarus sp. nr. rostratus, Ceratoppia sp.1, Kodiakella lutea, Tectocepheus velatus, Oppiella nova, Ramusella (Ramusella) sp., Allosuctobelba sp.2, Dentachipteria sp., Scheloribates (Scheloribates) sp., Pilogalumna sp.
8 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) Table 2 Soil nutrient analysis (mean7sd) for canopy suspended soils and forest floor habitat associated with western redcedar trees in the Walbran Valley on Vancouver Island, Canada (collected September 2004) Canopy Forest floor F 1,10 P Total Kjeldahl nitrogen (mg g 1 ) 9.99 (3.4) 4.42 (1.3) Total available phosphorus (mg g 1 ) 0.85 (0.20) 0.39 (0.18) Total potassium (mg g 1 ) 6.62 (1.77) 2.39 (8.94) o0.001 ph 3.55 (0.26) 3.69 (0.12) high abundance levels were observed in litterbags. In general, investigations of the role of microarthropods in soil processes have shown that soil fauna accelerate decomposition, contributing an average of 23% to litter decomposition when measured as mass loss (Seastedt, 1984).HigherlevelsofNH 4 - N, NO 3 -N, and PO 4 -P have been measured in soils with microarthropods than in defaunated soils (Heneghan and Bolger, 1998; Bardgett and Chan, 1999), and also, rates of net N mineralisation have been shown to increase by 10 49% (average 30%) in the presence of microarthropods (Persson, 1989), suggesting that soil fauna enhance nutrient mobilisation. The role of microarthropods in decomposition and nutrient cycling processes is primarily indirect through their interactions with the microbial community, and, any direct contribution to these processes may not be measurable, or difficult to separate from indirect effects (Moore et al., 1988). Studies exploring the contribution of microarthropods to soil processes have found no measurable results (Faber and Verhoef, 1991; Ca rcamo et al., 2001). In addition, soil physical and chemical properties may confound the measurement of microarthropod participation in the litter decay process (Brussaard et al., 1995), and environmental variables, specifically moisture, have a stronger effect on decomposition rates than the presence of microarthropods (Douce and Crossley, 1982). Oribatid mite abundance and species richness were lower in canopy litterbags compared to litterbags on the forest floor, which is consistent with our hypothesis of litterbag oribatid mite assemblages reflecting soil source pool assemblages (Lindo and Winchester, 2006). Furthermore, oribatid mite abundance in canopy suspended soils has been linked to the moisture content and depth of the suspended soil (Lindo and Winchester, 2007b) and Prinzing (2001) suggested that oribatid mites utilise vertical stratification and compensatory redistribution to maintain viable moisture requirements in arboreal systems. Lower oribatid mite abundance and species richness in canopy litterbags may also be a result of moisture limitation compared to litterbags on forest floor, as litterbags on the surface of canopy suspended soils may not provide adequate resistance to desiccation for oribatid mites. Low habitat heterogeneity in cedar litter, lower moisture content of litterbag contents and smaller colonising source pool of oribatid mites in the canopy are the parsimonious explanations for observed richness and abundance patterns in canopy litterbags. Canopy and forest floor litterbag assemblages of oribatid mites were shown to be different (high complementarity) and reflect differences in forest floor and suspended soil assemblages, which colonise litterbags (Fagan et al., 2006). Addison et al. (2003) found similar patterns in Collembola assemblages in forest floor litterbags over a 4-year period, showing that source communities influence colonising assemblages of litterbags. Our ANOSIM and PCA results support the observed difference between canopy and forest floor litterbag community composition. Species on the positive side of PC 1 of the PCA ordination plot are typically canopy species (e.g. Kodiakella lutea, Eupterotegaeus rhamphosus), while species on the negative of PC 1 are mainly forest floor dwellers (e.g. Dentachipteria sp., Liacarus sp. nr. bidentatus). Oribatid mites are generally considered late colonisers of decomposing conifer litter (Hasegawa, 1997; Osler et al., 2004), and densities of oribatid mites in litterbag studies often increase over the first 12 months (Hasegawa, 1997; Fagan et al., 2006). Plant litter quality is a major driver of decomposer invertebrate community (Wardle et al., 2006) and microarthropod communities may respond to structural components of litterbags such as root biomass (Lindo and Visser, 2003), moisture holding, and overall complexity of the litter environment (Hansen, 2000). Successional changes in oribatid mite communities during litter decomposition in the field are often superseded or confounded by seasonal affects on species composition (Osler et al., 2004; Fagan et al., 2006). Seasonality is likely to be a contributing factor in abundance differences between 6 (January 2005) and 12 months (September 2005), although at 24 months (September 2006) conditions were very dry in both canopy and forest floor habitats, and microarthropod abundance, richness and composition patterns were most likely to reflect this. We are not able to discern whether changes in oribatid mite community composition over time were successional, seasonal or driven by abiotic differences at sampling periods, but we suggest a combination of these factors contributed to the observed community patterns in oribatid mites colonising litterbags over 24 months. Acknowledgements The authors gratefully acknowledge Kevin Jordan (arbor nautaccess@hotmail.com) for his contribution in climbing
9 2964 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) methodology, canopy sampling, and general field expertise, Drs. Tony Trofymow and Suzanne Visser for helpful comments on the manuscript, and Drs. Richard Ring and Valerie Behan-Pelletier for inspiration and continued support. This research was supported by National Science and Engineering Research Council of Canada (ZL and NNW). Appendix A Average abundances (# ind. g 1 d.w.) (7SD) of major microarthropod groups in litterbags placed on the surface of canopy suspended soils and forest floor habitats associated with western redcedar trees in the Walbran Valley, Vancouver Island, Canada after 6, 12 and 24 months. Different superscript letter denotes significant (Po0.05) difference between canopy and forest floor habitats based on one-way ANOVA performed separately for each sample period. 6 months 12 months 24 months Canopy Ground Canopy Ground Canopy Ground Prostigmata 0.72 (1.5) 0.46 (0.6) 1.06 (1.1) b 9.20 (4.9) a 0.49 (1.0) b 3.44 (3.0) a Astigmata 0.00 (0.0) b 2.68 (2.6) a 1.87 (3.6) b (26.0) a 0.00 (0.0) 0.00 (0.0) Mesostigmata 3.98 (4.6) b (11.9) a 0.00 (0.0) b 6.77 (6.3) a 0.00 (0.0) b 8.89 (18.3) a Collembola 8.32 (6.9) (11.8) 5.73 (8.3) b (22.9) a 0.05 (0.2) 2.44 (10.1) Other microarthropods 0.27 (0.3) b 2.09 (1.9) a 0.52 (0.6) b 6.67 (5.2) a 0.04 (0.1) b 1.03 (1.4) a Total microarthropods (16.7) b (24.2) a (38.7) b (86.3) a 2.36 (5.1) b (41.9) a Appendix B Oribatid mites collected from litterbags placed in canopy suspended soil and forest floor habitats associated with six western redcedar trees in the Walbran Valley on Vancouver Island, Canada following 6, 12 and 24 months. Species 6 months 12 months 24 months Canopy Ground Canopy Ground Canopy Ground Palaeacarus hystricinus (Tra ga rdh, 1932) Eniochthonius minutissimus (Berlese, 1904) Pterochthonius angelus (Berlese, 1910) Liochthonius sp Liochthonius sp Liochthonius sp Liochthonius sp Archiphthiracarus sp Archiphthiracarus sp Maerkelotritia sp. nr. alaskensis (Hammer, 1967) Epilohmannia sp Camisia foveolata (Hammer, 1955) Platynothrus sp. nr. peltifer (C.L. Koch, 1839) Trhypochthonius tectorum (Berlese, 1896) Malaconothrus sp Nanhermannia elegantula (Berlese, 1913) Nanhermannia sp. nr. comitalis (Berlese, 1916) Hermannia gibba (C.L. Koch, 1840) Hermanniella sp. nr. occidentalis (Ewing, 1918) Damaeidae sp. nr. Epidamaeus Belba (Protobelba) californica (Banks, 1904) Eupterotegaeus rhamphosus (Higgins and Woolley, 1963) Eupterotegaeus sp. nr. rostratus (Higgins and Woolley, 1963) Sphodrocepheus anthelionus (Woolley and Higgins, 1968)
10 Z. Lindo, N.N. Winchester / Soil Biology & Biochemistry 39 (2007) Eueremaeus marshalli (Behan-Pelletier, 1993) Megeremaeus montanus (Higgins and Woolley, 1965) Peltenuiala pacifica (Norton, 1983) Liacarus sp. nr. bidentatus (Ewing, 1918) Liacarus sp. nr. robustus (Gervais, 1844) Liacarus sp Furcoribula sp Ceratoppia sp Ceratoppia sp Ceratoppia sp Gustavia sp Kodiakella lutea (Hammer, 1967) Tectocepheus velatus (Michael, 1880) Medioppia sp Multioppia sp Oppiella nova (Oudemans, 1902) Oppiella sp Ramusella (Ramusella) sp Quadroppia quadricarinata (Michael, 1885) Suctobelbella sp Suctobelbella sp Suctobelbella sp Suctobelbella sp Allosuctobelba sp. nr. gigantea (Hammer, 1955) Allosuctobelba sp Autogneta sp. nr. longilamellata (Michael, 1885) Achipteria sp. nr. curta (Aoki, 1970) Dentachipteria sp Eupelops sp Scheloribates (Scheloribates) sp Oribatula sp Phauloppia sp Melanozetes crossleyi (Behan-Pelletier, 2000) Sphaerozetes winchesteri (Behan-Pelletier, 2000) Mycobates corticeus (Behan-Pelletier, 2001) Pilogalumna sp Sum Richness References Aber, J.D., Melillo, J.M., Litter decomposition: measuring relative contributions of organic matter and nitrogen to forest soils. Canadian Journal of Botany 58, Addison, J.A., Trofymow, J.A., Marshall, V.G., Abundance, species diversity, and community structure of Collembola in successional coastal temperate forests on Vancouver Island, Canada. Applied Soil Ecology 24, Ashworth, J., Mrazek, K., Modified Kelowna test for available phosphorus and potassium in soil. Communications in Soil Science and Plant Analysis 26, Bardgett, R.D., Chan, K.F., Experimental evidence that soil fauna enhance nutrient mineralization and plant nutrient uptake in montane grassland ecosystems. Soil Biology & Biochemistry 31, Behan-Pelletier, V.M., Walter, D.E., Biodiversity of oribatid mites (Acari: Oribatida) in tree canopies and litter. In: Coleman, D.C., Hendrix, P.F. (Eds.), Invertebrates as Webmasters in Ecosystems. CAB International, Wallingford, pp Bohlman, S.A., Matelson, T.J., Nadkarni, N.M., Moisture and temperature patterns of canopy humus and forest floor soil of a montane cloud forest, Costa Rica. Biotropica 27, Brussaard, L., Noordhuis, R., Geurs, M., Bouwman, L.A., Nitrogen mineralization in soil in microcosms with and without bacteriovorous nematodes and nematophagous mites. Acta Zoologica Fennici 196, Cárcamo, H., Prescott, C.E., Chanway, C.P., Abe, T.A., Do soil fauna increase rates of litter breakdown and nitrogen release in forests of British Columbia, Canada? Canadian Journal of Forest Research 31, Douce, G.K., Crossley Jr., D.A., The effect of soil fauna on litter mass loss and nutrient loss dynamics in Arctic tundra at Barrow, Alaska. Ecology 63, Enloe, H.A., Graham, R.C., Sillett, S.C., Arboreal histols in oldgrowth redwood forest canopies, Northern California. Soil Science Society of America Journal 70, Faber, J.H., Verhoef, H.A., Functional differences between closelyrelated soil arthropods with respect to decomposition processes in the
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