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Karasawa S, Beaulieu F, Sasaki T, Bonato L, Hagino Y, Hayashi M

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Karasawa S, Beaulieu F, Sasaki T, Bonato L, Hagino Y, Hayashi M ISSN 0389-1445 EDAPHOLOGIA No.83 July 2008 Bird's nest ferns as reservoirs of soil arthropod biodiversity in a Japanese subtropical rainforest Shigenori Karasawa1'2'*, Frederic Beaulieu3, Takeshi Sasaki4, Lucio Bonato5, Yasunori Hagino6, Masami Hayashi7, Ryousaku Itoh8, Toshio Kishimoto9, Osami Nakamura10, Shiihei Nomura11, Noboru Nunomura12, Hiroshi Sakayori13, Yoshihiro Sawada14, Yasuhiko Suma15, Shingo Tanaka16, Tsutomu Tanabe17, Akio Tanikawa18, Naoki Hijii19 1Iriomote Station, Tropical Biosphere Research Center, University ofthe Ryukyus, Okinawa 907-1541, Japan 2Fukuoka University ofEducation, Fukuoka 811-4192, Japan (Present address) 3Canadian National Collection ofInsects, Arachnids andNematodes, Agriculture andAgri-Food Canada, Ottawa K1A 0C6, Canada 4University Museum, University ofthe Ryukyus, Okinawa 903-0213, Japan 5Department ofBiology, University ofPadova, 1-35131 Padova, Italy 6Natural History Museum and Institute ofChiba, Chiba 260-8682, Japan 7Faculty ofEducation, Saitama University, Saitama 338-8570, Japan 8Showa University, Tokyo 142-8555, Japan 9Japan Wildlife Research Center, Tokyo 110-8676, Japan 10 2507-9 Omaeda, Saitama 369-1246, Japan 11 National Museum ofNature and Science, Tokyo, 169-0073 Japan 12 Toyama Science Museum, Toyama 939-8084, Japan 13 Mitsukaido-Daini Senior High School, Ibaraki 303-0003, Japan 14 Minoh Park Insects Museum, Osaka 562-0002, JAPAN 15 6-7-32 Harutori, Hokkaido 085-0813, Japan 16 5-9-40 Juroku-cho, Fukuoka 819-0041, Japan 17 Faculty ofEducation, Kumamoto University, Kumamoto 860-8555, Japan 18 Laboratory ofBiodiversity Science, School ofAgriculture andLife Sciences, The University ofTokyo, Tokyo 113-8657, Japan 19 Graduate School ofBioagricultural Sciences, Nagoya University, Aichi 464-8601, Japan The Japanese Society of Soil Zoology 0 *±* £ Edaphologia, No. 83: 11-30, July 31, 2008 11 Bird's nest ferns as reservoirs of soil arthropod biodiversity in a Japanese subtropical rainforest Shigenori Karasawa1,2'*, Frederic Beaulieu3, Takeshi Sasaki4, Lucio Bonato5, Yasunori Hagino6, Masami Hayashi7, Ryousaku Itoh8, Toshio Kishimoto9, Osami Nakamura10, Shiihei Nomura11, Noboru Nunomura12, Hiroshi Sakayori13, Yoshihiro Sawada14, Yasuhiko Suma15, Shingo Tanaka16, Tsutomu Tanabe17, Akio Tanikawa18, Naoki Hijii19 1IriomoteStation, Tropical BiosphereResearch Center, University oftheRyukyus, Okinawa 907-1541, Japan 2Fukuoka University ofEducation, Fukuoka 811-4192, Japan (Present address) 3Canadian NationalCollection ofInsects, Arachnids andNematodes, Agriculture andAgri-FoodCanada, Ottawa K1A 0C6, Canada 4UniversityMuseum, University oftheRyukyus, Okinawa 903-0213, Japan 5Department ofBiology, University ofPadova, 1-35131 Padova, Italy 6NaturalHistoryMuseum andInstitute ofChiba, Chiba260-8682, Japan 7Faculty ofEducation, Saitama University, Saitama338-8570, Japan 8ShowaUniversity, Tokyo 142-8555,Japan 9Japan Wildlife Research Center, Tokyo 110-8676,Japan 10 2507-9 Omaeda, Saitama369-1246, Japan 11 National Museum ofNature andScience, Tokyo, 169-0073 Japan 12 Toyama Science Museum, Toyama 939-8084, Japan 13 Mitsukaido-Daini SeniorHighSchool, Ibaraki 303-0003, Japan 14 Minoh ParkInsects Museum, Osaka 562-0002, Japan 15 6-7-32 Harutori, Hokkaido 085-0813, Japan 16 5-9-40 Juroku-cho, Fukuoka 819-0041, Japan 17 Faculty ofEducation, Kumamoto University, Kumamoto 860-8555, Japan 18 Laboratory ofBiodiversity Science,SchoolofAgriculture andLifeSciences, The University ofTokyo, Tokyo 113-8657, Japan 19 Graduate School of Bioagricultural Sciences, Nagoya University, Aichi 464-8601, Japan Received: 16 December; 2007, Accepted: 1 July 2008 Abstract Litter trapped in epiphytic ferns (Asplenium nidus L.) and their rootballs were sampled for arthropods and compared with the forest floor (soil and litter) in a Japanese subtropical forest. A total of ca 40,000 individuals were identified to species, representing 13 arthropod groups and 553 species. The densities (numbers per 100 g dry wt of substrate) of the five most abundant taxa (Oribatida, Mesostigmata, Collembola, Coleoptera, Formicidae) were generally higher in the fern habitats (trapped litter and roots) than in the forest-floor habitats (soil and litter). From the 553 species, 267 were collected from bird's nest ferns and 403 from the forest floor. Rarefaction curves indicate that for a same number of individuals collected, species richness was at least twice higher on the forest floor than in the 12 Shigenori Karasawa et al. ferns for four of the five most diverse groups (Oribatida, Mesostigmata, Coleoptera, Collembola) whereas Araneae had slightly more species in ferns. Multivariate ordination showed that overall species composition clearly differed between fern habitats and the forest-floor habitats. Although species richness in bird's nest ferns may not be as high as for the forest floor, our study supports the hypothesis that these epiphytes are important reservoirs of arthropod biodiversity. Key words: Asplenium nidus;Bird's nest ferns; Species level; Subtropical rainforest; Suspended soil invertebrate communities in tropical and temperate forests. Introduction Because soil-dwelling invertebrates belong to many animal Epiphytes can trap enormous amounts of litter in the groups representing a width of body sizes, feeding and forest canopies. This litter decomposes into humid soil dispersal strategies (Coleman et al, 2004; Bardgett, 2005), ( 'suspended soils' ; Delamare-Debouteville, 1948) which faunal similarities between suspended soils and forest-floor holds abundant and diverse soil-dwelling communities (e.g., habitats may differ among animal groups. Comparative Paoletti et al, 1991; Ellwood and Foster, 2004; Yanoviak studies between suspended soil and ground are often based et al, 2006). Thus, epiphytes not only have significant on identification at higher taxonomic levels (usually order impacts on the recycling of organic matter in the canopy or family; e.g., Delamare-Debouteville, 1948; Paoletti et (Clark et al, 1998; Fonte and Schowalter, 2004), but also al, 1991; Ellwood et al, 2002; Ellwood and Foster, 2004), represent significant components of biodiversity in forest while species-level analyses are mainly limited to a few ecosystems (Basset, 2001; Benzing, 2004). If the faunal studies on mites, collembolans, and ants (e.g. Longino and composition of soil-dwelling invertebrates in suspended soils Nadkarni, 1990; Rodgers and Kitching, 1998; Karasawa and are similar to those in the forest floor (at least in higher taxa), Hijii, 2006a, b, c; Lindo and Winchester, 2006; Beaulieu and their contribution to decomposition processes in arboreal Walter, 2007). Further research targeting invertebrate groups environments may be similar to that in forest floors (Fonte at lower taxonomic levels and contrasting suspended soil with and Schowalter, 2004). Moreover, if suspended soil faunas other forest habitats should clarify the role of suspended soils differ from those in forest-floor habitats at the species level, as biodiversity reservoirs. then epiphytes and suspended soils may enhance and help The aims of this study were to: 1) compare the densities, maintain species diversity in forest ecosystems (Basset et al, species richness, and species composition of soil-dwelling 2003). This would represent a major argument in favor of the arthropods between bird's nest ferns and the forest floor; and 2) protection of old-growth forests, which are characterized by a test whether the arthropod fauna in bird's nest ferns is distinct fully developed epiphytic flora whereas secondary forests can to the one on the forest floor. be almost devoid ofepiphytes and dead organic matter in their canopy (Nadkarni et al, 2004). Materials and Methods Bird's nest ferns (Asplenium nidusL. complex) are widely Study area distributed from tropical to temperate regions (Murakami et The study was carried out in an old-growth, evergreen, al, 1999) and can develop on a variety of host tree species broad-leaved forest dominated by Castanopsis sieboldii (Annaselvam and Parthasarathy, 2001; Karasawa and Hijii, (Makino) Hatusima ex Yamazaki & Mashiba, at the foot 2006b). Because of their large size (fresh weight > 200 kg in of Mt. Nishime, in the northern part of Okinawa Island, Borneo; Ellwood et al, 2002) and their basket-shaped rosette, southwestern Japan (26°49'N, 128°5' E; 250-330 m a.s.l.). the ferns can trap substantial amounts of leaf litter from the The area is characterized by a subtropical climate with canopy. This suspended litter as well as the rootball below mean annual temperature of 23.0 °C, and average annual the rosette have been shown to harbour rich invertebrate precipitation of 2330 mm between 1992.and 2003 (Yona communities (e.g. Walter et al, 1998; Ellwood et al, 2002; Experimental Forest at the University ofthe Ryukyus). Ellwood and Foster, 2004; Karasawa and Hijii, 2006a, b, c). Thus, bird's nest ferns may foster the biomass and diversity of Soil arthropods in bird's nest ferns 13 Table. 1. Characteristics ofhost trees and bird* s nest ferns. Summer Autumn Winter 20 June 01 13 Nov. 01 18 Feb. 03 Host tree Ficus septica Schefflera octophylla Species Dead tree Burm.f. (Lour.) Harms Height (m) 5 8 10 Bird's nest fern Height (m) 1.5 3.0 3.3 Number ofleaves 38 62 27 Amount (g dry wt) Green leaves 221.5 518.9 171.1 Litter 286.7 354.5 132.0 Rootball 515.5 1047.3 796.2 Sampling Subsamples were clumped together for analysis. In total, three We removed a single bird's nest fern (Asplenium nidus) independent samples(trappedlitter+ fern rootball,or 3 forest- from
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