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Dendrobaena Octaedra Changes the Oribatid Community and Microarthropod Abundances in a Pine Forest

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Dendrobaena Octaedra Changes the Oribatid Community and Microarthropod Abundances in a Pine Forest Soil Biology & Biochemistry 32 (2000) 1671±1681 www.elsevier.com/locate/soilbio Introduction of the epigeic earthworm Dendrobaena octaedra changes the oribatid community and microarthropod abundances in a pine forest M.A. McLean*, D. Parkinson Department of Biological Sciences, University of Calgary, Calgary, AB, Canada T2N 1N4 Accepted 5 April 2000 Abstract The eects of the activities of the epigeic earthworm Dendrobaena octaedra on the oribatid community and microarthropod abundances were studied in a 90-year old pine forest over 2 years. Oribatids were extracted from the L and FH layers and the Ah and Bm horizons at 1 and 2 years and data were analyzed using principal component analysis (PCA). High worm biomass correlated positively with oribatid species richness and diversity in the L layer. In the FH layer, worm biomass accounted for 83% of the variation in the oribatid community data and correlated negatively with oribatid species richness. High worm biomass correlated with decreases in the abundances of 18 oribatid species, and the total abundances of adult and juvenile oribatids, astigmatids, mesostigmatids, Actinedida and Arthropleona in the FH layer. These eects were attributed to the changes in the physical structure of the organic layers of the soil. In the Ah and Bm horizons the C±N ratio accounted for 72± 97% of the variation in the oribatid species and microarthropod group data. The abundances of O. nova, other Oppioidea, several Brachychthoniidae, C. cuspidatus and adult (in the Ah horizon only) and juvenile oribatids, and Arthropleona were positively correlated with the C±N ratio. This re¯ected the mixing of less decomposed organic matter into the lower horizons by D. octaedra. 7 2000 Elsevier Science Ltd. All rights reserved. Keywords: Dendrobaena octaedra; Earthworm invasion; Oribatid community; Microarthropods 1. Introduction have been invoked to explain these eects have included: (1) alteration in the physical structure of the soil (Marinissen and Bok, 1988; Hamilton and Sillman, There is con¯icting evidence of the eects of earth- 1989; Loranger et al., 1998; McLean and Parkinson, worms on soil microarthropods (arthropods between 1998; Maraun et al., 1999); (2) alteration of the chemi- 200 mm and 2 mm, including mites and Collembola). cal or physical characteristics of organic matter (OM) Increased microarthropod abundance and diversity and its eects on the soil microbes (Yeates, 1981; (Marinissen and Bok, 1988; Loranger et al., 1998), Hamilton and Sillman, 1989; Loranger et al., 1998; decreased abundance (Dash et al., 1980; Yeates, 1981) Maraun et al., 1999); (3) competition for food (Brown, and mixed eects (Yeates, 1981; Hamilton and Sill- 1995); and (4) predation (Dash et al., 1980). Some of man, 1989; McLean and Parkinson, 1998; Maraun et the discrepancies between these studies are undoubt- al., 1999) have all been reported. Mechanisms which edly due to the diering eects of earthworms of dierent ecological strategy on soil physical structure and OM dynamics. Feeding of anecic earthworms (lar- * Corresponding author. Louis Calder Center, Fordham Univer- ger litter feeding species with permanent vertical bur- sity, 53 Whippoorwill Road, Armonk, NY 10504, USA. Tel.: +1- 914-273-3078 X 18; fax: +1-914-273-2167. rows) increases the organic matter content and E-mail address: [email protected] (M.A. McLean). porosity of mull soils, and therefore might be expected 0038-0717/00/$ - see front matter 7 2000 Elsevier Science Ltd. All rights reserved. PII: S0038-0717(00)00083-3 1672 M.A. McLean, D. Parkinson / Soil Biology & Biochemistry 32 (2000) 1671±1681 to improve the physical and chemical characteristics of of the forest where earthworms had already been the soil for microarthropods. However, feeding by epi- established for a few years. geic earthworms (smaller litter feeding species con®ned In September 1994 and September 1995 the plots to the organic and upper mineral layers) mixes mineral were sampled for microarthropod abundances and for material into the organic layers, and might be expected assessment of worm abundance and biomass. to reduce physical and chemical soil qualities for microarthropods in the organic layers of the soil. 2.3. Earthworm abundance and biomass Given the paucity of data on the eects of earth- worms on soil microarthropods and the diculties of At each sampling time one core 10.5 cm diameter arriving at any conclusions based on soils in which was taken from each plot and the earthworms present earthworms have previously been active, we were for- were heat extracted (Kempson et al., 1963) and tunate to be able to study the recent invasion of the counted as small (<10 mm long) immature, large epigeic earthworm Dendrobaena octaedra into lodge- immature, mature and aclitellate adults. Oven dry pole pine forest in SW Alberta, Canada. We used two weights of each of the worm size classes were used to approaches: short-term (6 months) laboratory studies obtain estimates of worm biomass at each of the (McLean and Parkinson, 1997a, 1998), and longer sampling times. Mean biomass of a mature worm was term (2 years) ®eld studies (McLean and Parkinson, 27 mg DW. 1997b, 2000, the present study). Under conditions of optimum moisture and temperature in mesocosms 2.4. Microarthropod abundances (intact soil cores 30 cm diameter  25 cm high), the ac- tivities of D. octaedra increased oribatid diversity and At each of the sampling times one core 5.5 cm diam- abundances (McLean and Parkinson, 1998). This was eter was taken from each plot to assess microarthro- attributed to an increase in spatial heterogeneity pod abundances. Cores were separated into L and FH through the addition of casts to the organic materials layers and into Ah and Bm horizons where possible already present. However, since organic layers in the and the microarthropods were heat extracted using a mesocosms with the highest worm numbers were com- high gradient extractor (Merchant and Crossley, 1970) pletely homogenized at the end of 6 months, we hy- from each layer and horizon. Microarthropods were pothesized that the longer term (2 years) eects of D. preserved in 70% ethanol and identi®ed: adult oribatid octaedra would be decreased oribatid diversity and to species where possible; juvenile oribatids to genus microarthropod abundance. where possible; other mites and Collembola to subor- der. Due to the heterogeneity of the soil and the pre- sence of rocks, some samples did not include the Bm horizon. Due to worm activities, an A horizon devel- 2. Materials and methods h oped in some plots but not in others, and was there- fore not sampled in all cases. 2.1. Site description Oribatid community parameters (species richness (S ), dominance (d ), diversity (1/D )) were calculated The site of this experiment was a 90-year old lodge- from the abundance data for all horizons in all plots. pole pine (Pinus contorta var. latifolia Engelm.) forest in the Kananaskis Valley of SW Alberta, Canada. For 2.5. Statistical analysis a more detailed description see McLean and Parkinson (1997b). At the time the plots were set up, earthworms were already invading the forest and therefore some of the 2.2. Experimental design ``control'' plots contained earthworms (McLean and Parkinson, 1997b) so ®nal earthworm biomass was Five pairs of plots 1 m  2 m were set up in August included in the analysis. Data were analyzed using 1993 in a part of the lodgepole pine forest which sur- ordination which allows the simultaneous analysis of veys had shown to be free from earthworms. Within the whole community. Principal Components Analysis each pair of plots, two treatments (control without (PCA), a linear and indirect ordination method was earthworms and treatment with worms) were randomly chosen since (i) the range of sample scores was low, assigned. The epigeic earthworm Dendrobaena octaedra making a linear method preferable to a unimodal (Savigny) was added to the worm plots at numbers method, (ii) indirect ordination methods allow the dis- equivalent to its 1993 ®eld density of 250 immatures covery of the largest variation in the species data with- and 70 matures m2, with a total biomass of 3.3 g d out being constrained by possibly irrelevant wt m2. The earthworms used were heat extracted environmental variables, and (iii) analysis can be fol- (Kempson et al., 1963) from pine forest ¯oor in a part lowed by correlation of the extracted axes with en- M.A. McLean, D. Parkinson / Soil Biology & Biochemistry 32 (2000) 1671±1681 1673 Table 1 Mean (standard error) oribatid species richness, dominance (d ), diversity (1/D ) and number of adult oribatids identi®ed to species in the L and FH layers and the Ah and Bm horizons n 20, 22, 13, 17, respectively) per 5.5 cm diameter core over all plots and sampling times LFHAh Bm Richness 1.6 (1.5) 12.9 (5.8) 5.6 (3.4) 4.1 (3.1) d 0.62 (0.40) 0.42 (0.14) 0.67 (0.23) 0.60 (0.27) 1/D 1.79 (3.71) 5.00 (2.62) 2.85 (1.91) 3.41 (2.64) Number of Individuals 3 (3) 136 (122) 51 (56) 17 (17) vironmental variables to discover which, if any, of the 3. Results supplied environmental variables (in this case, ®nal worm biomass, organic matter content (OM), moisture 3.1. Earthworm numbers and biomass content, pH, C±N ratio) account for a signi®cant pro- portion of the variation in the species data (ter Braak, Earthworm numbers ranged from 0 to 3349 individ- 1995). Since the conditions in each soil layer/horizon uals m2, with a mean of 854 individuals m2. Earth- were dierent, the analysis was conducted on each worm biomass ranged from 0 to 39.9 g DW m2, with layer/horizon separately.
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