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The Influence of Deforestation on the Land Snail Fauna of Kuromatsunai District, Southwestern Hokkaido, Japan

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The Influence of Deforestation on the Land Snail Fauna of Kuromatsunai District, Southwestern Hokkaido, Japan VENUS 77 (1–4): 15–26, 2019 DOI: http://doi.org/10.18941/venus.77.1-4_15Influence of Deforestation on Land Snail Fauna©The Malacological Society of Japan15 The Influence of Deforestation on the Land Snail Fauna of Kuromatsunai District, Southwestern Hokkaido, Japan Yuta Morii1,2* 1Department of Forest Science, Research Faculty of Agriculture, Hokkaido University, Kita-9-Jo, Nishi-9-Chome, Sapporo, Hokkaido 0608589, Japan 2Ecology Group, School of Agriculture and Environment, Massey University, Private Bag 11-222, Palmerston North 4410, New Zealand Abstract: Forest clear-cutting and the subsequent habitat fragmentation causes catastrophic damage to plant and animal communities as well as the entire forest ecosystem. Many studies have suggested that forest disturbance decreases the biomass and species richness in forests. However, the long-term influence of deforestation on local-scale patterns of diversity is poorly understood. I investigated the land snail fauna in the soil of primary and secondary forests and compared the number of individuals and species richness in each. Two primary forests and two secondary forests were surveyed in Kuromatsunai District (Hokkaido, Japan), and they were all dominated by the Japanese beech tree, Fagus crenata. Six soil blocks (50 cm × 50 cm) were sampled from each forest, and all the land snails from each soil block were collected and identified. The number of individuals and species richness were subsequently compared between the primary and secondary forests. A significantly larger number of individuals and significantly greater species richness were recorded in the primary forests (generalized linear mixed models (GLMMs), likelihood ratio tests, P < 0.05). The diversity of the land snail fauna in one of the two primary forests, Utasai Forest, was particularly high, with an average of 239.2 individuals and 7.2 species per soil block. In contrast, only 12.3 individuals and 4.8 species, on average, were detected in the two secondary forests. In addition, the number of individuals of smaller species (2.0 mm or less) was significantly lower in the secondary forests, but that of the larger species (greater than 2.0 mm) was not. I also estimated the age of the two secondary forests using an increment borer and found that both secondary forests were cleared approximately 100–150 years ago. My results imply that the deforestation of more than 100 years ago continues to impact the land snail fauna, thus affecting the soil fauna of the forest. Keywords: Terrestrial molluscs, primary forest, intact forest, clearance, clear-felling, Japanese beech tree, Kuromatsunai Depression Introduction Forest clear-cutting and the subsequent habitat fragmentation is one of the most radical landscape changes in forest ecosystems, and it decreases the biomass and species richness in forests (Turner, 1996; Siira-Pietikainen, 2001; Hylander et al., 2004; Lindo & Visser, 2004; Palviainen et al., 2005; Ewers & Didham, 2006; Chiba et al., 2009). However, the long-term influences of deforestation and local extinction on local-scale patterns of diversity are relatively poorly understood (Taylor et al., 2003; Watters et al., 2005; Ewers & Didham, 2006; Graham et al., 2006; Kappes, 2006; Chiba et al., 2009; Ström et al., 2009), as most studies of historical land use have been conducted less than 100 years after a change to the landscape (Ewers & Didham, 2006). * Corresponding author: [email protected] 16 Y. Morii Several studies suggest that there is a time lag between environmental change and a species-level effect (Hanski & Ovaskainen, 2002; Chiba et al., 2006) and that different species and habitats vary in their responses to habitat destruction and human intervention (Lindo & Visser, 2004; Gotmark et al., 2008). Terrestrial molluscs are particularly useful taxa for investigating spatial patterns of species diversity and population density as well as the influence of historical land use on these factors because of their habitat preferences, low mobility, high population density and high species richness (Chiba et al., 2006; Gotmark et al., 2006; Kappes, 2006). Moreover, land snails are likely able to persist in smaller habitat patches than other taxa due to their small body sizes and restricted mobility (Kerney & Cameron, 1979; Baur & Baur, 1993). Globally, the total area of primary forests continues to decline, so their value is increasing (Gibson et al., 2011; Morales-Hidalgo et al., 2015). Hokkaido Island, Japan, has been undergoing deforestation since 1869, and almost all primary forests around Kuromatsunai District in southwestern Hokkaido were heavily logged until 1923 (Saito, 2012), but some forests remain intact. Kuromatsunai District is well known as the northernmost distribution limit of the Japanese beech tree, Fagus crenata (Saito, 2006, 2015; Matsui et al., 2012; Kitamura et al., 2015), and the primary forests in this area were mainly dominated by this species (Saito, 2006, 2012, 2015). In particular, Utasai Forest is known as the largest patch of primary forest in this area (92 ha) and was therefore designated a natural monument of Japan in 1928 and named “Utasai buna north limit” (Saito, 2006, 2012, 2015). However, the snail fauna of the primary forests in Kuromatsunai District has not been surveyed. I investigated the land snail fauna in the soil of the primary and secondary forests in this region and here discuss the legacy effect of historical forest disturbances. Material and Methods Study sites Two primary forests, Utasai Forest (42.65396°N, 140.32848°E, Alt. 80 m; Site ID: Pri1) and Shiroikawa Forest (42.70482°N, 140.39041°E, Alt. 180 m; Site ID: Pri2), and two secondary forests, Soibetsu Forest (42.68711°N, 140.26756°E, Alt. 50 m; Site ID: Sec1) and Shimo- choposhinai Forest (42.69782°N, 140.35242°E, Alt. 210 m; Site ID: Sec2) were surveyed. All are in Kuromatsunai District (Hokkaido, Japan) and are dominated by F. crenata (Figure 1). The land spaces of two primary forests, Pri1 and Pri2 are 92 ha and 20 ha, respectively (Saito, 2006, 2012, 2015), but there are younger forests adjacent to these forest patches. Other two research sites, Sec1 Fig. 1. Research sites in Kuromatsunai District, Hokkaido, Japan. Two primary forests, (A) Utasai Forest (Site ID: Pri1) and (B) Shiroikawa Forest (Pri2), and two secondary forests, (C) Soibetsu Forest (Sec1) and (D) Shimo-choposhinai Forest (Sec2). Influence of Deforestation on Land Snail Fauna 17 and Sec2 are also one part of larger forest area in Kuromatsunai District. I tried to choose unbiased geologic feature among research sites; the geological base of the Pri1 and Sec1 is igneous rock, Pri2 and Sec2 is sedimentary rock (Geological Survey of Japan & AIST, 2017). Quantitative investigation of the snail fauna Six soil blocks (50 cm × 50 cm), which were separated by at least 5.0 m, were collected from each forest from 5 October to 8 November 2015, and all the land snails in each block were sorted by hand and identified. Both live snails and empty shells still at least partly covered by the periostracum were collected and used for the analyses, and the numbers of individuals and species within each soil block (i.e., densities of individuals (DI) and species (DS)) from the primary and secondary forests were subsequently compared. The differences between the DI and DS of the primary and secondary forests were analysed using generalized linear mixed models (GLMMs) with a Poisson distribution and a log link using R version 3.3.2 software (R Development Core Team, 2016). The DI or DS was the response variable, and the forest type (primary or secondary) and site were the fixed- and random-effect explanatory variables, respectively. The influence of forest type on the DI and DS were statistically examined using likelihood ratio tests (LRTs), in which the deviance of the full model was compared with that of a model lacking the explanatory variable of forest type. The shell diameter of each species was based on the descriptions in Azuma (1995). The micro-snail species (those with shell diameters of 2.0 mm or less based on the descriptions in Azuma (1995)), were highly abundant in terms of the total number of individuals collected in this research, and these species seem to occur more in primary forests than secondary forests (Figure 2). To test this hypothesis, I also analysed the differences in the DI and DS of micro-snails, those with shell diameters of 2.0 mm or less (DI-small and DS-small), and larger snails of greater than 2.0 mm (DI-large and DS-large), between primary and secondary forests (Appendix Table S1). Moreover, I also analysed DI and DI-small without Fig. 2. (A) Proportions of each category of snails based on shell diameter to total number of individuals and species. (B) Proportions of four study sites to total number of individuals for each species. Numerals written with species name are consistent with the species numbers in Table 1. 18 Y. Morii Carychium pessimum, because C. pessimum was much more abundant in one of the two primary forests, Pri1 (68.2% of total individuals collected from Pri1). Age estimation of secondary forests Woody materials were sampled in winter from 20 to 24 February 2016 to estimate the age of the two secondary forests. Age estimation of beech trees in this study basically followed Matsui et al. (2012). The girth of all beech trees was measured within a 30-m × 30-m quadrat in each forest, and core samples were collected from 16 of these beech trees in each forest using an increment borer. The number of annual growth rings was counted under a stereoscopic microscope. Linear regressions between girth and the number of annual growth rings were conducted for each secondary forest and were used to estimate the ages of all other beech trees in the quadrats. I did not conduct this age estimation for two primary forests to avoid an adverse environmental impact to the invaluable forests.
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