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 the trunk of a host tree in summer (20 June 2001), floor litter subsamples + 3 soil subsamples) were taken from autumn (13 November 2001), and winter (18 February 2003). each habitat. All sampling was conducted between 09:00 and Each fern was divided into living fern leaves, the litter trapped 17:00 in sunny weather. in the basket-shaped rosette, and the rootball, consisting of live fern roots and humus (made of dead roots and decomposed Extraction of arthropods litter). The litter and roots were placed in individual plastic We extracted arthropods from all samples using Tullgren bags at the sampling site. The ferns were collected at 1.5-3.3 funnels; samples were placed in the dark for 24 h to avoid m high, and the number of leaves and total dry weight of the a sharp rise in temperature, which could kill arthropods ferns ranged from 27 to 62 and ca. 1-2 kg, respectively (Table intolerant of desiccation, and then under 40-W electric 1). Characteristics of the bird's nest ferns (host trees, spatial bulbs for 72 h. The adults of Araneae, Pseudoscorpiones, distribution, size parameters, litter accumulation, etc.) in this Mesostigmata, Oribatida (excluding Astigmata), Isopoda, forest were described in details elsewhere (Karasawa and Diplopoda, Pauropoda, Chilopoda (Geophilomorpha only), Hijii, 2006b). The fern size is positively correlated with the Protura, Collembola, Diplura, Hemiptera, and Coleoptera amount of accumulated litter and microarthropod abundances were identified to the lowest taxonomic level possible (usually (Karasawa and Hijii, 2006a, b). The ferns sampled in this morphospecies; Appendix 1). After extraction, we dried the study were of average size or slightly larger than average remaining substrate of the samples and the fern live leaves (therefore minimizing bias in microarthropod abundances). in an oven (70°C, 72 h) and weighed the dry matter on a Moreover, our previous studies showed that in this forest the microbalance to determine the invertebrate density per 100 g size and height of the fern, the tree species, and whether the dry wt ofsubstrate. tree is alive or dead, did not affect the species composition of oribatid communities in the ferns (Karasawa, 2006; Karasawa Analyses and Hijii, 2006a). Species richness was compared between bird's nest In each season (6 July 2001, 22 November 2001, fern and the forest-floor habitats for all groups and the five 27 February 2003), we haphazardly collected three litter most diverse groups using rarefaction curves showing the subsamples of 25 cm x 25 cm within 3 m from the base of cumulative number of species as a function of the number each host tree and then three (mineral) soil subsamples (25 cm of individuals collected (= sampling effort). The abundance x 25 cm x depth 5 cm) beneath the litter samples. of individuals was used as the sampling effort instead of the 14 Shigenori Karasawa et al.
number of samples in order to compensate for variation in the forest floor, densities were much higher in the litter than sample volumes within and among habitat. The curves were in the soil for all groups (Fig. 1). Density levels for the five computed using the software Estimates (Colwell, 2005) and most abundant groups (Oribatida, Mesostigmata, Collembola, each represents the average curve of 100 randomizations Coleoptera, Formicidae) do not show any clear trend: the ofthe order in which samples (fern's trapped litter or roots, highest and lowest densities generally do not correspond to forest-floor soilor litter) were added. the same season between habitats (ferns and forest floor) Faunal similarity between the bird's nest ferns and the nor across taxonomic groups. One possible exception are forest floor was expressed using a species turnover index, Mesostigmata, which show highest densities inwinter for both S0rensen's distances (Kimoto and Takeda, 1989): ferns and forest floor. S0rensen's distance = l-[2a / (b + c)] where band c denote the total number ofspecies present Species richness andcomposition in each of the two communities (or samples), respectively, and We identified a total of553 species (40,119 individuals) a denotes the number ofspecies present inboth communities. in 13 arthropod groups collected from the ferns and the forest This value has an upper limit of1(no species are present in floor in the three seasons. From the total, 136 species (25%) both communities) and a lower limit of0 (all species in both were represented by asingle individual, and 267 species (48%) communities are the same). The turnover index was calculated were collected from only one sample. Almost two-thirds of among samples for within habitat (fern vs. fern; and forest species (64%) could not be assigned to named species. Table floor vs. forest floor) and between habitats (fern vs. forest 2 shows the number ofspecies collected from the fern and the floor), for all groups and for the five most diverse groups. forest floor habitats for each group. To visualize the dissimilarities in species composition Acari (Oribatida and Mesostigmata) represents the most among subsamples (n = 12), atwo-dimensional ordination was diverse group in bird's nest ferns (46% ofall species) and conducted using semi-strong hybrid multi-dimensional scaling forest floor (54%) (Table 2). Coleoptera were second with with the software PATN (Belbin, 1995), based on all arthropod 25% and 16% of species in ferns and forest floor, followed by groups. In order to minimize bias due to differences in sample Collembola (13% and 14%, respectively) and Araneae (10% volumes between habitats, abundance data was transformed and 5%, respectively). The eight remaining groups represented into presence-absence data (l's and 0's) prior to analysis. less than 5% and 12% of species in ferns and forest floor, The ordination was based on Bray-Curtis distances between respectively (Table 2). Although differences in sample volumes' samples; however, because of the data transformation, do not allow rigorous comparison, the number ofspecies for distances were equivalent to Sorensen's distances. Analysis each group was generally higher inforest floor than bird's nest ofSimilarity (ANOSIM) was conducted using PATN to test ferns, except for Araneae and Coleoptera which had slightly whether dissimilarity in species composition between ferns more species in ferns. Rarefaction curves.indicate that for a and forest floor habitats was significant. same number of individuals collected, species richness was at least twice higher on the forest floor than in the ferns for
Results all groups except Araneae which had slightly higher species richness in ferns after 40 individuals were collected (Fig. 2). Densities and numerical dominance ofarthropod groups Differences in species richness between the litter and roots A total of 70,707 invertebrates were collected from the ofthe ferns are minimal, except for Araneae which has three bird's nest ferns (51,999 individuals) and the forest floor times more species in the litter. Forest-floor litter and soil (18,708 individuals) in the three seasons. Ofall invertebrates also have similar species richness, although Oribatida and collected from the ferns, Acari were the most abundant (32,346 Coleoptera show substantially more species in soil than litter individuals), followed by Collembola (8970). Similarly, these (Table 2). two groups were also dominant in the forest floor (87% ofall Ordination of the samples shows that the fern habitats invertebrates). (litter and roots) were inhabited by an arthropod fauna distinct For the ferns, the densities were generally similar to the one on the forest floor (Fig. 3). This distinction was between litter and roots (except for Formicidae in the litter significant (ANOSIM, P = 0.006). It also indicates that the which have low densities in summer and winter), whereas for fauna extracted from the roots of a given fern was more Soil arthropods in bird's nest ferns 15
bo o o
Summer Autumn Winter ^: litter in BNF •: rootball ofBNF •: FF litter •: FF soil
Fig. 1. Seasonal variation in the densities of thefive most abundant groups in bird's nest ferns (trapped litter and rootball) and theforest floor (litterandsoil), a: Oribatida, b: Mesostigmata, c: Collembola, d: Coleoptera, and e: Formicidae. BNF: bird's nestfems, and FF: forest floor. 16 Shigenori Karasawa et al.
Table. 2. Number ofspecies in 13 invertebrate groups collected from bird's nest fems and the forest floor
Bird's nest fem Forest floor Class Order (or Suborder) Total Litter Rootball Total Litter Soil Total Arachnida Araneae 27 8 27 13 12 19 40
Pseudscorpiones - - - 3 3 5 5 Acari (Mesostigmata) 34 28 43 46 58 71 94 Acari (Oribatida) 53 68 81 88 135 147 177 Crustacea Isopoda 2 2 2 3 4 5 5 Diplopoda 4 4 4 6 12 13 14 Pauropoda Tetramerocerata 1 1 1 3 3 4 4 Chilopoda Geophilomorpha 2 2 2 3 4 5 5 Insecta Protura - - _ 1 5 5 5 Collembola 30 26 36 37 49 56 74
Diplura - - - - 4 4 4 Hemiptera 4 3 4 2 5 6 9 Coleoptera 42 48 67 27 58 63 117
Total 199 190 267 232 352 403 553
All taxa Oribatida 450 180 Mesostigmata
400 160 350 I 140 70 300- 120 o 60 250 100 - 50 |H 200 80 40 150 60 - ) i 30 100 - 40 20 - t 11 50 20 10 - 0 0 0 500 1000 1500 2000 2500 O Bird's nest ferns Coleoptera 80 - Collembola Araneae • Forest floor 70 i 35 - 70 - il i 60 30 - 60 - < i 50 - 25 - 50 - 40 20 - 40 ( H1 il 30 - 30 • 15 - I 11 11' j 20 - /, 20 10 - * { 10 - 10 - 5 - 0 - ,ifSl 0 0 - 0 500 1000 1500 2000 2500 0 2000 4000 6000 8000 0 25 50 75 100 Cumulative no. individuals Cumulative no. individuals Cumulative no. individuals
Fl8' T;pR0wa5°LCUrVeSine yDYo confidence intervals°/SPeideSare°fua11shown.invertebrates and the five most diverse taxa collected from bird's nest fems and the forest floor Soil arthropods in bird's nest ferns 17
1.5
b 1.0 • b a ♦ • a O Litter in bird's nest ferns 0.5 a a ° ♦ V CM c V Rootball ofbird's nest ferns " ♦ c • b • Forest floor litter b v -0.5 o ♦ Forest floor soil
c -1.0 V c stress = 0.2289 o
-1.5 I i 1 1 i -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5
Axis 1
Fig.3. Semi-strong hybrid multi-dimensional scaling ordination of arthropod communities in samples from thelitter and roots of bird's nest fems and forest-floor litter and soil, a: summer, b: autumn, c: winter.
similar to the fauna extracted from the litter trapped in the s nest ferns were not found on the forest floor; and about a same fern than to the fauna from other ferns (however, this third ofspecies ofOribatida (37%) and others groups (Others, similarity may also be due to a seasonal effect, but see below). 31%) collected from ferns were not found on the forest floor. In concordance with the ordination, the species turnover Considering species collected on the forest floor, between index (S0rensen's distance) was higher for bird's nest fern 65% (for Oribatida) and 81% (for Others) ofspecies were not vs. forest floor paired comparisons (index averages vary found in ferns. between 0.66 to 0.87 across groups) than for within habitat Species turnover index based on samples (Sorensen comparisons (index for bird's nest fern vs. bird's nest fern: distance) also show differences among taxa. Species turnover 0.40-0.69; for forest floor vs. forest floor: 0.37-0.64) for each between fern and forest-floor samples was higher for group (Fig. 4). Species turnover was higher for fern vs. fern Coleoptera, Araneae and Others than for microarthropods than forest floor vs. forest floor comparisons for the five most (Oribatida, Mesostigmata, and Collembola) (Fig. 4). However, diverse groups, indicating that species composition was more turnover within habitat was also high for Araneae (0.69 and heterogeneous among fern than among forest-floor samples. 0.64) and somewhat high for Coleoptera (0.63 and 0.56). Overall analysis of complementarity (species turnover) Species turnover among ferns was lowest for Collembola shows that for each group relatively few species are found (0.41) and Others (0.40), indicating that different ferns had a in both the ferns and the forest floor (Fig. 5). Among the five more homogenous fauna for these groups than for other taxa. most diverse.groups, the proportion of the number of species Species turnover among forest-floor samples was lower for shared between habitats range from 11% (for Coleoptera) microarthropods (Mesostigmata, 0.39; Oribatida, 0.43; and to 29% (for Oribatida) of all species collected. Considering Collembola, 0.37). species collected in bird's nest ferns, a high proportion of Species turnover values for fern vs. forestfloor samples Coleoptera (81%) and Araneae (78%) species were apparently taken in different seasons are quite similar to values for fern absent from the forest floor; about half of species of vs. forest floor taken in the same date, for each group (Fig. 4), Mesostigmata (53%) and Collembola (50%) collected in bird* suggesting that there was little seasonal effect on the species 18 Shigenori Karasawa et al.
O §
CO a
Q CO
All Ori Meso Coleo Coll Ara Others • BNF vs BNF • FF vs FF U BNF vs FF (different season) II BNF vs FF (same season)
Fig. 4. Sarensen's distances between pairs ofbird's nest fern vs. bird's nest fern (BNF vs. BNF; n= 3), forest floor vs. forest floor (FF vs. FF; n = 3), bird's nestfern vs. forest floor from different seasons (BNF vs. FF; n = 6), and bird's nest fern vs. forest floor from the same season (BNF vs. FF; n= 3),for the five most diverse groups. Standard deviations are shown. All: all taxa, Ori: Oribatida, Meso: Mesostigmata, Coleo: Coleoptera, Coll:Collembola, Ara: Araneae, andOthers: other taxa.
27867 13284 2124 2128 7599
An Ori Meso Coleo Coll Ara Others •In forest floor only HIn both habitats Din bird's nest ferns only Fig. 5. Relative proportion ofspecies found inbird's nest ferns only, inthe forest floor only, and in both habitats. The number ofspecies is shown inside thebars. The total number of(identified) individuals represented ineach habitat is indicated above each bar(inferns / in forest floor). All: alltaxa, Ori: Oribatida, Meso: Mesostigmata, Coleo: Coleoptera, Coll: Collembola, Ara: Araneae, and Others: other taxa. Soil arthropods in bird's nest ferns 19 composition. by a similar fauna to the one in litter trapped inside the Because of the low level of replication, it is difficult to ferns' crown. However, our data is confounded by the fact draw conclusions on the habitat preferences of individual that trapped litter and roots were taken from the same ferns. species.However,our data suggests that many speciesmay be Although some species showed high abundances in both restricted to either bird's nest fern or the forest-floor habitats: fern habitats, others were much better represented in either, from species collected in at least two samples (out of three particularly the rootball (Karasawa and Hijii, 2006a, c). The samples), 46 were restricted to bird's nest fern habitats (litter high densities and species numbers observed in fern rootballs and/or roots) and 123 were restricted to forest-floor habitats indicate that they represent a suitable habitat for a diversity (litter and/or soil). From these 46 species apparently specific of arthropods. Live and humified rhizoids provide a food to bird's nest ferns, Coleoptera contributed the most with 15 source and/or a habitat for herbivores (e.g. the ortheziid scale species, followed by Oribatida (9 species), Collembola (8 insect Nipponorthezia ardisiaewas abundant in rootballs) and species), Mesostigmata (6 species), Araneae (6 species) and numerous saprophages, fungivores and predators, some of other groups (1 Hemiptera and 1 Diplopoda spp.). From the which were several-fold more abundant in rootballs than in 123 species apparently specific to the forest floor, Oribatida litter. In Malaysia, rootballs of bird's nest ferns were used as contributed the most (39 species), followed by Mesostigmata nesting sites by ants and termites (Ellwood et al, 2002). (28 species), Collembola (20 species), Coleoptera (17 Our results suggest that macroarthropods, especially species),Araneae (3 species) and other groups (16 species: 2 Coleoptera, are more habitat-specific than microarthropods, Pseudoscorpiones, 3 Isopoda, 4 Diplopoda, 1 Pauropoda, 2 with 89% of beetle species restricted to either the ferns (54 Chilopoda, 1 Protura, 2 Diplura and 1 Hemiptera). species) or the forest floor (51 species). Spiders also had a high percentage of species found in a single habitat (85%) and
Discussion most of these species were collected from bird's nest ferns (21 Over 500 species in 13 arthropod groups were identified species). Whether macroarthropod taxa divide habitats more from bird's nest ferns and the forest floor in this subtropical finely (despite their larger size) than mites and springtails will forest. Given the low number of samples taken and the requirefurther comparativestudies. A few studieshave shown number of taxa not identified to species level, the actual associations of macroarthropods with epiphytes and canopy species richness of the soil-dwelling fauna in this forest soil. Many ants and beetles were common in suspended may be well over 553. Moreover, the high percentage of soil but rare or absent from the forest floor in Costa Rica unidentified species highlights the need for further taxonomy and Australia, respectively (Longino and Nadkarni, 1990; in Japan, especially for groups in soil habitats. Soil (including Rodgers, 1999). In Panama and Brazil, some spiders were suspended soil) represents one of the most understudied found to be associated with particular bromeliads and orchids ecological systems although its contribution to local and (Stuntz et al, 2001; Romero and Vasconcellos-Neto, 2005). global biodiversity is unquestionably very high (Andre et al, Karasawa and Hijii (2006c) reported that the presence 2002). of bird's nest ferns could not dramatically raise the species This study shows that bird's nest ferns represent a richness of oribatid mites in the whole forest, because the significant habitat for arthropods in the studied forest: the oribatid fauna in the ferns was similar to the fauna in the densities of the five abundant groups were similar to or forest-floor litter and soil. Bird's nest ferns may nevertheless higher than the forest floor, and the number of species was contribute to local mite diversity because several oribatid remarkably high (267 species) and of similar magnitude to species appeared to be restricted to or to show clear the species richness on the forest floor (403 species). Our data preferences for bird's nest ferns over other habitats (forest also suggests that numerous arthropod species (belonging floor, bark, branches and leaves) (Karasawa and Hijii, 2006c). mainly to beetles, mites, springtails and spiders, although Research elsewhere showed relatively high habitat specificity several groups, e.g. ants, were not identified to species) favour of canopy microarthropods. In Canadian and South American bird's nest fern habitats over the forest floor. Thus, bird's nest rainforests, 34-51%.oforibatid species found in suspended soil ferns may contribute significantly to the local species richness and epiphytes were absent from the ground (Behan-Pelletier offorest arthropods. and Walter, 2000; Lindo and Winchester, 2006). Three out of The rootball of bird's nest ferns seem to be inhabited six dominant species of arboreal springtails were completely 20 Shigenori Karasawa et al.
or almost restricted to canopy litter in a cedar plantation in among bird's nest fern than among ground samples observed Japan (Yoshida and Hijii, 2005), and about half of the 24 in this study. springtail species found in bird's nest ferns in an Australian rainforest showed a preference for the canopy (Rodgers and Acknowledgements Kitching, 1998). Preliminary work on mesostigmatic mites We thank Mr. Kenshi Gotoh (University ofthe Ryukyus), indicate thatthere may be few arboreal specialists in Western Mr. Nobuto Shimada (Tokyo University) and Mr. Issei Canada, whereas inAustralian rainforests most suspended soil Kawashima (Okinawa) for supporting the field work; Dr. dwellers may berestricted toabove-ground habitats (Walter et Jun-ichi Aoki (Kanagawa Prefectural Museum of Natural al ,1998; Beaulieu andWalter, 2007). History) and Mr. Norihide Ohkubo (Mie Plant Protection Several factors, however, may have obscured Office) for valuable advice on oribatid mites; Dr. Pawel patterns of habitat specificity in our study. First, the higher Jatoszynski (Poland) for identifying the family Scydmaenidae; complementarity observed in beetles and spiders may in part Dr. Gen Takaku (Hokkaido University), Dr. Matthew Shaw be due to the higher proportion of beetles (62% of species) (Queensland Museum, Australia) and Dr. Jenny Beard (AQIS, and spiders (63%) collected in a single sample, compared Australia) foradvice onmesostigmatic mite identification; Dr. to microarthropods (42%). Thus, increasing sampling effort Valerie Behan-Pelletier (Agriculture and Agri-Food Canada) might show that many species only found in ferns so far are for comments on a previous version of this manuscript. This infact generalists, frequent in both suspended soil and on the study was supported in part by a Grant-in-Aid for Scientific ground. Second, sampling ofother canopy habitats may reveal Research from the Japanese Ministry of Education, Culture, that some species are primarily bark orfoliage dwellers. Third, Sports, Science and Technology (No. 17380091). it is possible that many arboreal specialists occur mainly in the mid- and/or upper canopy (Rodgers and Kitching, 1998), which was not considered in the present study, although the height of the fern (0.1-10 m) had no apparent effect on the t 907-1541 tttm±m 870, mm&i?±¥WLii¥U oribatid fauna in this forest (Karasawa and Hijii, 2006a). T 811-4192 £ft7|T/*^;£ifc!gri-l (5itt0f)), Frederic Last, suspended soil may act as a source habitat for arboreal Beaulieu (Canadian National Collection of Insects, Arachnids species while many individuals may nevertheless visit the and Nematodes, Agriculture and Agri-Food Canada, Ottawa forest floor, which may act as a sink habitat. Similarly, ground kiaoc6), fe^m^ mm±^mm t 903-0213 mm 'specialists' may breed principally in the forest floor though W^-f-Jgl), Lucio Bonato (Department ofBiology, University also wandering opportunistically into suspended soil patches of Padova 1-35131 Padova), $tff«I9 (^m^AL^^WMi near the ground. These stray individuals can impede analyses T 260-8682 cfffeEWItPrr 955-2), #jE§£ (*0*^ife of habitat distribution based on presence-absence data or low W^SK t 338-8570 3 V>fciflf#BT*A#255)f « replication. &ft (H8«;fc3Mi*W t 142-8555 iSJIIEJ»<7)& 1-5-8), The forest floor is inhabited bya relatively more diverse ^*¥6R (Smmmfflft^y?- t 110-8676 &JtCETS fauna than is bird's nest ferns. This difference is not dueto a 3-10-10), **tfcH (t 369-1246 *«*/h«ffl 2507-9). sampling bias orhigher arthropod densities on the ground (in »ttfl¥ (B£#*«»« t 169-0073 #1&KSAW this study, ferns had higher densities): for a same number of 3-23-1), m$& (•OjT&ft^ift*;'*- t 939-8084 individuals collected, over twice more species were collected •ujiiiB+ffw 1-8-30. ««k (%®m$.*Bmm~^ on the ground than in bird's nest ferns. Higher diversity on ^$ t 303-0003 «*ff*»ii«*Hr 3549-4), ?fffl* the ground may be due to its more favorable conditions. The ft (*EJff#*ffi&HfiA«| t 562-0002 «ffi7tT»ffi& forest floor may provide a more stable microclimate and a • 1-18). mmttB (t 085-0813 illK?tT#»6-7-32), ffl* more predictable food supply for soil-dwelling arthropods, *ff (t 819-0041 SK&AWT 5-9-40), BSltl 0»*** reducing chances of local extinctions (Hijii, 1989; Bowlman ifcW^glS t 860-8555 J»#fljJi« 2-40-1). m\W% (M etal, 1995). It is also physically morecontinuous thanabove- ' £*¥*¥IK«¥£frW¥ffi%# t 113-8657 JmEM ground habitats, favoring patch colonization (Rodgers and 1-1-1), W#a:« (££S;fc¥£fra*ffljfe«. T464-8601 Kitching, 1998; Proctor et al, 2002). These factors could ^SMrff^ttE^^fflr) also explain the higher heterogeneity in species composition Soil arthropods in bird's nest fems 21
Autonoma Mexico, Sociedad Latinoamericana de Acarologia. MJtkS# £ 45 It &* * * - 7 * U (Asplenium nidus)
predatory mites: diversity and distribution of the genus London. Athiasella (Mesostigmata: Ologamasidae) in forest floor and Hijii, N., 1989. Arthropods communities in a Japanese cedar suspended soils in Australian rainforests. In: Acarologia XL- (Cryptomeria japonica D. Don) plantation: abundance, Proceedings ofthe International Congress (eds. J.B. Morales- biomass and some properties. Ecological Research, 4:
Malacara, V. Behan-Pelletier, E. Ueckermann, T.M. Perez, 243-260. E. G. Estrada-Venegas, and M. Badii), pp 129-135. Instituto Karasawa. S., 2006. Diversity of oribatid-mite communities de Biologia, Facultad de Ciencias, Universidad Nacional (Acari: Oribatida) in suspended soils. Edaphologia, 79: 22 Shigenori Karasawa et al.
27-40. (In Japanese with English abstract) and forest floor ofa Venezuelan cloud forest. Journal of Karasawa, S. and Hijii, N„ 2006a. Effects of distribution and TropicalEcology, 7: 373-383. structural traits ofbird' s nest ferns (Asplenium nidus) on Proctor, H. C, Montgomery, K. M., Rosen, K. E. and Kitching, R. oribatid (Acari: Oribatida) communities in a subtropical L., 2002. Are tree trunks habitats or highways? Acomparison Japanese forest. Journal ofTropical Ecology, 22: 213-222. oforibatid mite assemblages from hoop-pine bark and litter. Karasawa, S. and Hijii, N., 2006b. Determinants of litter Australian Journal ofEntomology, 41: 294-299. accumulation and the abundance of litter-associated Rodgers, D., 1999. Structural dynamics ofrainforest collembolan microarthropods in bird's nest ferns (Asplenium nidus assemblages in a leaflitter suspended in epiphytes and on complex) in the forest of Yambaru on Okinawa Island, the forest floor. Ph.D. Thesis, Griffith University, Nathan, southern Japan. Journal ofForest Research, 11:313-318. Australia. Karasawa, S. and Hijii, N., 2006c. Does the existence of bird's Rodgers, D. J. and Kitching, R. L, 1998. Vertical stratification of nest fems enhance the diversity oforibatid (Acari: Oribatida) rainforest collembolan (Collembola: Insecta) assemblages: communities in a subtropical forest? Biodiversity and description of ecological patterns and hypotheses concerning Conservation, 15: 4533-4553. their generation. Ecography, 21:392-400. Kimoto, S. and Takeda, H., 1989. An Introduction to Community Romero, G. Q. and Vasconcellos-Neto, J., 2005. The effects of Ecology. Kyoritsu Shuppan Co. Ltd, Tokyo, (in Japanese) plant structure on the spatial and microspatial distribution of Lindo, Z. and Winchester, N.N., 2006. Acomparison of a bromeliad-living jumping spider (Salticidae). Journal of microarthropod assemblages with emphasis on oribatid mites AnimalEcology, 74: 12-21. in canopy suspended soils and forest floors associated with Stuntz, S., Ziegler, C, Simon, U. and Zotz, G., 2002. Diversity ancient western redcedar trees. Pedobiologia, 50: 31-41. and structure ofthe arthropod fauna within three canopy Longino, J. T. and Nadkarni, N. M., 1990. Acomparison of ground epiphyte species in central Panama. Journal ofTropical and canopy leaf litter ants (Hymenoptera: Formicidae) in a Ecology, 18: 161-176. Neotropical montane forest. Psyche, 97: 81-94. Walter, D. E., Seeman, O., Rodgers, D. and Kitching, R.L. Murakami, N., Watanabe, M., Yokoyama, J., Yatabe, Y, Iwasaki, 1998. Mites in the mist: how unique is a rainforest canopy- H. and Serizawa, S., 1999. Molecular taxonomy study and knockdown fauna? Australian Journal ofEcology, 23: revision ofthe three Japanese species ofAsplenium sect. 501-508. Thamnopteris. Journal ofPlant Research, 112: 15-25. Yanoviak, S. P., Nadkarni, N. M., Rodrigo, S. J., 2006. Arthropod Nadkarni, N. M., Schaefer, D., Matelson, T. J. and Solano, R. assemblages in epiphytes mats of Costa Rican cloud forests. 2004. Biomass and nutrient pools ofcanopy and terrestrial Biotropica 36: 202-210. components in a primary and a secondary montane cloud Yoshida, T. and Hijii, N., 2005. Vertical distribution and seasonal forest, Costa Rica. Forest Ecology and Management, 198: dynamics of arboreal collembolan communities in a 223-236. Japanese cedar (Cryptomeria japonica D. Don) plantation. Paoletti, M. G., Taylor, R.A.J., Stinner, B. R., Stinner, D. H. and Pedobiologia, 49: 425-434. Benzing, D. H., 1991. Diversity of soil fauna in the canopy Soil arthropods in bird's nest fems 23
Appendix 1. Soil-dwelling invertebrates collected from the bird's nest ferns and the forest floor. Bird's nest ferns 20 June 2001 13 Nov. 2001 22 Nov. 2001 27 Feb. 2003 Family Species Litter Rootball Litter Rootball Litter Rooball Litter Soil Litter Soil Litter Soil ARANEAE Ctenizidae Latouchia sp. 0 Amaurobiidae Coelotes yambaruensis 2 Coelotes spp. Dictynidae Dictyna sp. 2 Oonopidae Gamasomorpha sp. 1 2 Gamasomorpha sp. 2 5 Orchestina spp. 10 Oonopidae spp. 2 Pholcidae Spermophora sp. 0 Theridiidae Episinus sp. 0 Nipponidion ok'mawense 10 Theridiidae sp. 0
Linyphiidae Cresmatoneta sp. 0 . - - 2
Hylyphantes tanikawai 1 . - - 0 3 2 1 9 Linyphiidae spp. - Mysmenidae Mysmenella sp. 0 2
Araneidae Neoscona scylla - - - 0
Tetragnathidae Leucange sp. - - - 0 1 0 Tetragnatha sp. "
Zodariidae Mallinella spp. 0 1 - 2 Pisauridae Dolomedes zatsun 0 Lycosidae Pirata spp. 0 2 . . 5 0 Clubionidae Clubiona kurosawai - " - Clubiona spp. 1 2 2 1 - Liocranidae Phrurolithus spp. - Ctenidae Anahita fauna 0 10
Sparassidae Sinopoda okinawana 2 - - - 0 Gnaphosidae Gnaphosidae spp. 0 4 5 18 0 Thomisidae Lysiteles miniatus - Oxytale hoshizuna 1 0 S 0 Takachihoa truciformis - Philodromidae Philodromus sp. 0 Salticidae Myrmarachne sp. 4 0
Onomastus kanoi 2 - - - 0 0 1 3 Salticidae sp. 1 - - Salticidae sp. 2 0 Salticidae sp. 3 - - - Salticidae sp. 4 0
Telamonia vlijmi - - - 0 2 7 Unidentified sp. - -
PSEUDOSCORPIONES Allochthonius sp. 2 2 Tyrannochlhoniusjaponicus 4 3 5 31 Tyrannochthonius sp. 1 2 3 Noebisiidae sp. 1 1 1 Noebisiidae sp. 2
ACARI (MESOSTIGMATA)
1 - - - - - 0 Sejidae Sejus sp. -
Triplogyniidae Funkoiriplogynium sp. 1 1 ------0
- - - - 3 4 Zerconidae Mesozercon plumatus - 0 1 Parasitidae cf. Phityogamasus sp. 1 1 5 3 4 1 14 Parasitidae sp. 1 0 13 8 7 2 18 6 54 -
Parasitidae sp. 2 0 - 6 - - - 1 7
Parasitidae sp. 3 0 - 6 - 7 - 5 18
Veigaiidae Gamasolaelaps sp. 0 - 1 - - 12 6 19 -
Veigaia cf. ashizuriensis 0 - 4 - 1 - - 5
Veigaia uenoi 30 74 1 2 5 - - - 8
Rhodacaridae Rhodacarus sp. - 0 - - - - - 3 3
- - - - Digamasellidae Dendrolaelaspis sp. - 0 2 2 5 39 0 Ologamasidae Gamasiphis sp. 1 - - - Gamasiphis sp. 2 11 3 1 4
Gamasiphis sp. 3 0 8 9 1 - 1 6 25 0 1 2 3 Gamasiphis sp. 4 - - Laelaptiella sp. 48 48 2 3 5
cf. Rhodacaroides sp. - 0 17 15 4 6 29 41 112
Parholaspididae Gamasholaspis sp. 1 0 - - - - - 1 1 •
Gamasholaspis sp. 2 0 - 1 - 3 - - 4 Holaspina sp. 1 2 3 5 6 4 I 7 3 . 26
Holaspina sp. 2 1 24 ------0
- - - - Holaspina sp. 3 - 0 2 7 9
- - - Holaspina sp. 4 - 0 4 4 18 26
Holaspina sp. 5 0 15 24 - 2 9 15 65
- - - Holaspina sp. 6 - 0 11 2 13
- - - cf. Holaspina sp. - 0 1 3 4 8 24 Shigenori Karasawa et al.
Holaspulusishigakiensis - - - - 0 - 2 7 9 7 12 37 Holaspulus sp. 67 15 26 1 2 3 47 -
Neparholaspis sp. 1 - - - . 0 1 1 Neparholaspis sp. 2 - 0 - - - 1 6 1 3 1 4 16 Neparholaspis sp. 3 - 1 - - 1 2 - - - - - 2 Parholaspididae sp. 1 2 6 " 4 - 10 Parholaspididae sp. 2 - - - - - 0 1 1 Eviphididae Evimirus cf. uropodinus 1 1 - - 2 1 2 - 1 13 2 19 Macrochelidae Macrocheles sp. 1 30 3 15 19 - - 67 1 - - . . . 1 Macrocheles sp. 2 2 2 6 4 ' 1 1 7 3 22
Macrochelidae sp. - - - 0 2 2 Pachylaelapidae Neoparasitus sp. 0 1 7 20 28 Pachylaelaps sp. - - - - 0 . 1 1 2 Ameroseiidae Ameroseius variolarius - - - - 0 - - 3 - 103 . 106 Melicharidae Proctolaelaps sp. 2 - - - 4 1 - - - . . 1 Ascidae Asca aphidioides - - - - 0 7 1 3 1 108 7 127 Asca nubes 38 17 55 " - 0 - • " -
Asca sculptrata - - . 12 0 Asca sp. 1 5 2 7 " ------. 0 Asca sp.2 1 1 " ------. 0 Gamasellodes sp. " 0 ------1 . 1 Xenoseius sp. 1 - - - 1 - - - - . . Phytoseiidae 0 Amblyseius sp. 1 2 1 5 0 ' Amblyseiussp. 2 0 - 1 - 3 - 4 - Chanteius sp. 1 ;" 0 Podocinidae Podocinum sp. 3 1 2 4 • 1 11 10 1 4 4 4 2 25 Blattisociidae Blattisocius sp. 1 • 1 ------. 0 Cheiroseius sp. 1 6 3 4 13 15 2 4 4 2 1 Cheiroseius sp. 2 28 1 50 13 Cheiroseius sp. 3 13 0 2 2 Cheiroseius sp. 4 1 0 Lasioseiusqueenslandicus 19 2 3 6 4 34 2 1 3 Lasioseius sp. 1 10 1 11 0 Lasioseius sp. 2 3 3 Platyseiustriangralis 0 8 2 2 Blattisociidae sp. 1
- - 0 - - - 3 - 5 - 6 Blattisociidae sp. 2 14 - 0 Laelapidae ------4 4 Cosmolaelaps sp. 1 6 38 4 14 82 0 Cosmolaelaps sp. 2 0 7 7 Gaeolaelaps queenslandicus 0 1 Laelaspis sp. - 1 0 1 1 Pseudoparasitus sp. 2 12 - 0 Laelapidae sp. 4 4 0 Uropodina Uropodinasp. 1 0 15 6 21 Uropodinasp. 2 0 - - - 7 11 2 2 1 16 39 Uropodinasp. 3 - - - - 0 - . . . 1 j Uropodinasp. 4 9 68 5 112 2 1 3 Uropodinasp. 5 0 2 1 3 Uropodinasp. 6 0 1 2 6 3 12 Uropodinasp. 7 4 4 0 Uropodina sp. 8 1 1 0 Uropodina sp. 9 1 1 2 0 Uropodinasp. 10 2 2 1 Uropodina sp. 11 S 250 38 901 - - . 1215 1 1 • 4 6 Uropodinasp. 12 - - - 0 10 23 9 20 . 3 65 Uropodinasp. 13 - - - 0 - 3 - . . . 3 Uropodinasp. 14 2 2 0 Uropodina sp. 15 0 1 I - Uropodinasp. 16 0 - 6 1 7 Uropodina sp. 17 0 1 9 10 - Uropodinasp. 18 27 27 0 Uropodinasp. 19 0 1 Uropodinasp. 20 0 - - - 8 - 9 13 12 20 62 Uropodinasp. 21 0 7 4 1 12 Uropodinasp. 22 0 7 7 Uropodina sp. 23 0 - - 39 " 5 44 Uropodina sp. 24 0 - 5 5
ACARI (ORIBATIDA) Palaeacaridae Palaeacarus sp. 0 1 1 Hypochthoniidae Eohypochihonius crassisetiger 0 20 - - 31 8 81 2 455 597 Eohypochihonius parvus ? 0 1 8 " 18 ' 27
Eniochthoniidae Hypochthoniella fukushimaensis - - . 0 2 1 3 Arborichthoniidae Arborichthonius styosetosus 0 " 9 9 - -
Brachychthoniidae Brachychthonius sp. - . 1 : ; 0 Liochthonius spp. 81 144 225 * 8 • 3 - 11 Sellnickochthonius spp. - : 27 27 10 1 11 Cosmochthoniidae Cosmochthonius lanatus 2 1 2 11 1 Haplochthoniidae Haplochthonius simplex 4 4 0 - Epilohmanniidae Epilohmannia pallida 0 - 1 Epilohmannoides sp. - 0 54 1 23 78 Soil arthropods in bird's nest ferns 25
Lohmanniidae Lohmannia corallium 1207 4871 . - 6078 ------0
Mixacarus exilis - - - - 0 - - - - - 6 6
Papillacarus hirsutus - - . - 0 - 12 - 10 - 61 83
Mesoplophoridae Apoplophora pantolrema - - - - 0 - - 1 - - 10 11
Archoplophora rostralis - - - - 0 - - - - - 1 I
Oribotritiidae Austrotritia saraburiensis 14 4 - - 18 ------0
Indotritia javensis - - - - 0 - 4 - - - - 4
Mesotritia maerkeli - - - - 0 - 1 - - - - 1
Oribotritia chichijimensis 7 2 - 1 10 - 1 - - - - 1
Oribotritia sp. - 1 - - 1 - 9 - - - 2 11
Oribotritiidae sp. 1 - - - - 0 - - - - - 1 1
Oribotritiidae sp. 2 - - - - 0 1 2 - - - - 3
Oribotritiidae sp. 3 - - - 2 2 ------0 Euphthiracaridae Acrotritia ardua 5 5 9 2 27 76 124 6 68 10 4 22 10 120
Phthiracaridae Hoplophorella cucullata - - - - 0 - - - - - 2 2
Hoplophthiracarus sp. 1 15 9 - 2 1 27 1 4 22 7 49 15 98
Hoplophthiracarus sp. 2 8 7 11 6 19 49 100 6 10 - - - - 16
Hoplophthiracarus sp. 3 - - - - 0 - 1 - - 6 11 18
Hoplophthiracarus sp. 4 - - - - 0 - - - - - 1 1
Phthiracarus parmatus - - 14 6 1 1 22 - - 1 5 3 3 12
Phthiracarus sp. - - - 1 1 ------0
Phthiracarus spp. - - 1 2 3 - - 5 5 3 7 20 0 - 2 - Trhypochthoniidae Trhypochthoniidae sp. - - 2 - - - - -
Malaconothridae Malaconothrus sp. - - - 4 4 ------0
Malaconothrus spp. - - 1 13 14 3 6 - 5 50 67 131
Nothridae Nothrus biciliatus - - - - 0 - - - - - 1 1
Nanhermanniidae Masthermannia hirsuta - - - - 0 - - - 12 - 2 14 1 52 110 Nanhermannia parallela - - - 1 1 21 1 6 29
Hermanniidae Hermannia kanoi 29 67 - 3 83 182 4 8 2 2 1 - 17 0 Hermanniellidae 1 " Hermanniella sp. 1 * " - - Hermanniella sp. 2 ;; 0 . . 1 1
Plasmobatidae Plasmobates asiaticus - - - 0 - - 1 - 2 10 13
Neoliodidae Neoliodidae sp. - - - - 0 - 1 - 1 1 2 5 0 Licnodamaeidae Pedrocortesallajaponica - - - i i 2 ------0 1 1 Damaeidae " " " Belba sp. 1 •
Belba sp. 2 . _ . 0 . 1 3 Cepheidae Sadocepheus yakuensis - - - - 0 1 2 - - - - 24 4 36 - - 1 Microtegeidae Microtegeus borhidii - - - 0 6 1
Microzetidae Berlesezetes ornatissimus - - 19 - 19 - - - - - 0
Microzetes lunaris - - - - 0 - - 2 2 2 40 46
Zetorchestidae Zetorchestes aokii - - - . 0 4 2 - 1 3 11 21 Astegistidae Cultroribula lata 0 1 4 26 31
Cultroribula sp. ;;;; _ 0 . 1 . 1 ------1 Ceratoppiidae Ceratoppia sp. 0 - -
Gustaviidae Gustavia microcephala - - - - 0 - - 2 - - 12 14 Xenillidae Xenillus heterosetiger 2 1 4 1 1 ' Xenillus tegeocranus ; . 0 . . . . 1 1 2
Tenuialidae Tenuiala nuda - - .- - 0 - 2 - - 1 - 3
Ctenobelbidae Ctenobelba sp. - - - - 0 - - - - - 1 1
Amerobelbidae Yambaramerus itoi - - - - 0 - - - - - 7 7 Eremulidae 0 2 2 Eremulus avenifer - - - - •
Eremulus flagellifer . 0 . . 6 6
Fenestrella japonica - - - - 0 - 7 - - - 10 17
Damaeolidae Fosseremus sp. - 3 - 3 3 - 1 1 2 31. 38 Eremobelbidae Eremobelba japonica 0 1 7 3 1 5 25 42 - - - Eremobelba minuta 7 7 0
Eremobelba Okinawa - - - - 0 - 16 - 1 1 16 34
Heterobelbidae Heterobelba stellifera - - - 0 - - - - - 6 6
Ameridae Caenosamerus spatiosus - - - 0 - - - - 2 I 3
Autognetidae Autogneta sp. - - - - 0 - 5 - - - 5 Oppiidae Acroppia viperea 5 67 1 73 3 5 1 22 31 Arcoppia sp. 1 0 1 23 6 30 -' - - Arcoppia sp. 2 0 2 2
Lasiobelba insulata - - - - 0 2 17 1 4 - - 24
Medioxyoppia actirostrata - - - 0 - - - 1 5 19 25
Medioxyoppia yuwana - - - - 0 5 49 - 12 1 31 98
Medioxyoppia sp. - - - - 0 - - - - - 6 6
Microppia minus 1 - - 3 4 - - - - - 158 158
Multioppia sp. - - - - 0 - - - - - 39 39
Oppiella nova - 2 - 2 4 - 5 2 3 8 47 65
Subiasella spp. - - 3 1 110 114 11 4 - - 23 1 39
Oppiidae sp. 1 35 5 16 - 57 ------0
Granuloppiidae Senectoppia pectinata - - - - 0 2 - - - 2 53 57
Machuellidae Machuella spp. - - 198 204 402 - - - - - 4 4
Quadroppiidae Quadroppia circumita 2 8 111 121 - - - - - 2 2
Quadroppia hammerae - 12 488 239 739 - - - - 1 7 8 Suctobelbidae Allosuctobelba grandis 0 1 6 11 18 • • Allosuctobelba tricuspidata ; ;; 0 ; ;; 1 1
Suctobelba sp. - - - - 0 1 2 - - 1 - 4
Suctobelbella sp. - - - . 0 - - - - - 3 3
Suctobelbila sp. - - - - 0 - - - - - 4 4 26 Shigenori Karasawa et al.
Suctobelbila spp. ------0 2 - - 2 - 107 Ill Suctobelbidae sp. 1 0 3 " " 4 7 • • - - Suctobelbidae sp. 2 . : . . 0 2 2 Suctobelbidae spp. 2 2 11 7 13 7 47 178 263 Oxyameridae Oxyamerus spathulatus . . 1 . . 0 Dampfiellidae Dolicheremaeusbaloghi i 16 16 5 7 22 66 Dolicheremaeus elongatus 1 1 1 3 1 12 22 39 - Fissicepheus sp. 1 0 - - 2 2 Fissicepheus sp. 2 5 1 2 5 13 0 Otocepheidae Acrotocepheus gracilis ------0 7 9 22 5 9 4 56 Megalotocepheusjaponicus 3 - - - - - 3 10 7 8 11 21 17 74 Trichotocepheus amamiensis ------0 - 3 1 - 4 8 16
Otocepheidae sp. - - - -. - - 0 - - - - - 1 1 Tokunocepheidae Tokunocepheus mizusawai 10 6 45 15 2 13 91 0 Carabodidae Austrocarabodes sp. 0 1 1 - Carabodes ikeharai 1 1 0 Carabodes palmifer 0 1 - - 1 Carabodes sp. 0 1 2 2 5 Diplobodes kanekoi 3 - - - - - 3 7 8 10 5 26 29 85 23 Gibbicepheusfrondosus 6 7 4 14 8 62 1 3 2 - 9 1 16 Yoshiobodes nakatamarii 17 4 1 3 - - 25 21 8 32 7 94 12 174 Carabodidaesp. 1 2 11 13 1 1 1 80 83 Carabodidae sp. 2 0 - - - - 5 5 Nippobodidae Nippobodes yuwanensis 0 4 4 1 1 7 13 30 Tectocepheidae Tectocepheus cuspidentatus 1 - - - - - 1 2 7 - 7 2 11 29 Tectocepheus velatus ------0 - - - - 2 . 2 Cymbaeremaeidae Scapheremaeus sp. 5 1 6 " 0 - - - • - - Licneremaeidae Licneremaeus spp. . 1 131 132 0 Achipteriidae Campachipteriadistincta - - - - - 2 2 ------0 Oribatellidae Oribatella sp. 1 - - - - - 1 ------0 Ceratokalummidae Cultrobates nipponicus - - - - 5 5 10 1 - - 11 2 24 Ceratozetidae Ceratozetella imperatoria ------0 - - - - . 1 1
Ceratozetes sp. 1 ------0 - - - 1 . . 1 Ceratozetes sp. 2 0 12 15 27 Ceratozetidae sp. 1 1 - - - 1 1 Mochlozetidae Unguizetes sp. 0 2 1 - - 3 Hemileiidae Hemileius singularis 11 11 1 1 Scheloribatidae Perscheloribates clavatus 115 238 353 6 " 11 5 113 - 135 - Perscheloribates lanceolatus . . 0 24 24 Scheloribates sp. 1 2 6 2 3 13 0 Scheloribates sp. 2 1 0 Scheloribates sp. 3 0 1 3 4 Scheloribates sp. 4 0 - 1 1 Oripodidae Brachyoripoda sp. 1 1 2 1 1 Cosmopirnoduspulcherrimus 2 2 1 1 Truncopes moderatus 11 1 12 4 1 5
Truncopes sp. 1 - 16 2 1 - 19 - - . . . 0
Truncopes sp. 2 3 - 1 2 1 - 7 - - - . . . 0 Protoribatidae Protoribates sp. 1 2 24 - - - 26 4 82 - - - 38 124 Protoribates sp. 2 28 - - - 658 686 5 238 - 18 26 274 561 Protoribates sp. 3 0 1 1 Protoribates sp. 4 0 - 7 - - 33 - - - 40 - Protoribates sp. 5 37 37 1 6 7 Protoribates sp. 6 0 14 15 17 46 Protoribates sp. 7 0 - - 36 - - - - 1 37 " - Protoribates sp. 8 0 4 1 29 34 Haplozetidae Lauritzenia sp. 7 7 1 1 Peloribates barbatus 0 1 " ------Peloribates levipunctatus 0 1 12 13 2 Peloribates longisetosus 1 1 2 - 6 - - - - - . 0 Peloribates ominei 100 43 122 91 - - 356 32 15 1 4 23 7 82 Trachyoribates ovulum 648 238 20 912 6 376 2200 109 298 29 51 64 557 1108 Haplozetidae sp. 3 298 45 9 8 363 " 0 - - Parakalummidae Neoribates rotundus 0 ; 4 2 2 8 Neoribates roubali 1 3 - - - 1 5 l 2 1 1 13 2 20 Neoribates sp. - - - - 1 1 - 4 - - 2 6 Allogalumnarotundiceps 0 1 1 2 Dimidiogalumna azumai 0 - 1 - Galumna hiroyoshii 1 1 2 - - 1 4 7
Galumna sp. 1 1 - - 0 Galumna sp. 2 0 1
Pergalumna sp. 1 121 180 9 56 - 14 380 . 1 . . . . 1 Pergalumna sp. 2 1 3 4 0 Pergalumna sp. 3 0 - - 2 2 4 - - - Trichogalumna sp. 0 l 1 3 .2 7 Trichogalumna spp. 5 2 1 8 17 9 7 5 28 4 70 Galumnidae sp. 1 1 2 1 ; 1 2 7 0 Galumnidae sp. 2 3 i 4 2 - - - 2 - - - - j j Galumnidae sp. 3 1
Galumnidae sp. 4 - - . . - 0 . . . 13 17 30
Galumnidae sp. 5 ------0 2 2 Soil arthropods in bird's nest fems 27
- 1 Galumnidae sp. 6 - 46 Galumnellidae Galumnella nipponica - 26 Unidentified Unidentified sp. 1 -
ISOPODA 1 1 10 Ligiidae Ligidium ryukyuense - 7 13 Philosciidae Burmoniscus okinawaensis - 2 1 2 Agnariidae Agnera ryukyuensis - 2 Armadillidae Spherillo brevipalma 1 Spherillo kunigamiensis 4 1 5 -
DIPLOPODA (Glomerida) 1 Glomeridae Hyleoglomeris yamashinai - (Julida) 3 Julidae Anaulaciulus yamashinai - 4 Nemasomatidae Sinostemmiulus sp. - I 3 Unidentified Julida sp. - (Siphonophorida)
Unidentified Siphonophorida sp. - (Polydesmida)
Pyrgodesmidae Ampelodesmus sp. - 7 23 986 0 214 331 • Thelodesmus sp. • 0 2 3 1 1 7 Cryptodesmidae Cryptodesmidae sp. -
- - - - - 1 14 17 Paradoxosomatidae Chamberlinius hualienensis - 0 2
Nedyopus sp. 1 159 233 4 3 399 - 1 - 1 - - 2 3 6 22 7 29 1 2 - Nedyopus sp. 2 - " Polydesmidae Epanerchodus sp. 0 10 10
Polydesmidae sp. . 0 2 4 6 0 1 2 27 1 1 32 Unidentified - - Polydesmida spp. -
PAUROPODA 0 6 2 8 Eurypauropodidae Samarangopus sp. -
Sphaeropauropus sp. 27 - - 40 - 4 - - 3 9 16
- - 0 - - - - 1 - 1 Trachypauropus sp. - 0 1 1 2 - " Pauropodidae Pauropus sp. - - - - -
GEOPHILOMORPHA Mecistocephalidae Arrup cf. holstii 0 1 . ,
Mecistocephalus karasawai - 19 66 1 1 1 - - 3
Takashimaia ramungula - - 0 - 4 - 1 - 8 13
Ballophilidae Ityphilus cf. tenuicollis - 1 10 - 2 - - - 2 0 2 1 1 5 - Linotaeniidae Strigamia sp. - - 1
PROTURA Eosentomidae Eosentomon sakura 0 17 1 23 41
Eosentomon cf. udagawai - - 0 - 1 - - - - 1
Eosentomon sp. 1 - - 0 - 2 - - - - 2
Eosentomon sp. 2 - - 0 - - - - - 2 2 Eosentomon sp. 3 0 2 2
COLLEMBOLA Hypogastruridae Hypogastrura communis 0 167 167
Hypogastrura tergilobata - - 0 5 12 - 2 - 5 24
Hypogastrura sp. 1 . - 0 - - - - - 1 1
Hypogastrura sp. 2 - - 0 4 7 44 - 1 56
Schaefferia cf. proserpinae . . 0 - - - 103 - - 103 Xenylla brevispina 2 3 56 2 1 3 6 5 1 1 - Xenylla sp. 1 - Xenylla sp. 2 2 0 Onychiuridae Lophognathella choreutes 0 1
Tullbergia yosii - 2 13 - 4 - - 9 13
Onychiurus folsomi . - 0 - - 3 - - - 3
Onychiurus izuruensis . - 0 - - - - 3 3 0 2 1 4 7 - - - Onychiurus yodai - - Onychiurus sp. 1 1 1
Onychiurus sp. 2 . 0 l" 2 . 2 8 13
Pseudachorutidae Superodontella distincta - - 0 - - - - - 1 1
Superodontella similis - 0 1 - - 2 1 - 4
Frieseajaponica 1 - 2 - 1 - - - 1
Pseudachorutes longisetis - - 0 1 2 - - - - 3
Pseudachorutidae sp. - - 0 2 3 - 3 - 14 22
Crossodonthina nipponica - - 7 47 10 3 1 4 3 68
Yuukianura pacifica - . 0 - - - 4 - 13 17 Morulina australis 2 1 1 6 1 9 "• - - - Neanura okinawana 0 2 5 1 8
Vitronura rosea >0 42 70 - - - - 15 16 31
Neanuridae sp. - 3 3 ------0
Ballistura stricta - - 0 - - 152 42 22 66 282
Folsomina wuyanensis 616 1 4264 5576 - - - 2 - 16 18
Folsomia minipunctata - 0 - 5 - - 3 379 387 28 Shigenori Karasawa et al.
Folsomia octoculata 311 - 218 479 12 37 95 959 1800 Folsomidesparvulus 0 - - 11 1 47 - 51 110 Jsotomiella minor - 1 11 70 5 16 . 198 300 Proisotoma sp. 21 336 2 4 - - 6 - Desoria sensibilis - 3 0 Desoria sp. - 0 - - - 1 . . 1 Isotomidae sp. - 0 - - - 23 . . 23 Tomoceridae Tomocerus ishibashii 6 ------0 Tomocerusliliputanus 0 - - - - - 3 7 10 Tomocerus ocreatus - 1 - - - - - . 0 Tomocerus violaceus 17 20 ------0 Tomocerus viridis 1 ------0 Tomocerussp. - 11 - 2 - - . . 2 Oncopoduridae Oncopodura sp. - 0 - - - . . 1 1 Cyphoderidae Cyphoderus javanus - 0 - 1 . . . . 1 Paronellidae Salina affinis 34 - 118 - - - - . 2 2 Salina bicincta 7 - 100 - - - - . . Entomobryidae 0 Sinella dubiosa 0 - 1 22 - 1 1 11 36 Sinellasubquadrioculata - 0 2 9 1 - . 1 13 Sinella sp. 102 2 104 - - - - . . 0 Entomobrya sp. 1 4 75 - 98 7 11 6 3 14 4 45 Entomobrya sp. 2 108 1 23 146 - - - - . . 0 Homidia sp. 0 - - 2 - - 3 . 5 Janetschekbrya sp. 1 3 - - - - - . 0 Seira sp. 6 6 ------. 0 Lepidocyrtus sp. 1 5 9 5 - 1 5 1 18 5 35 Pseudosinellaoctopunctata 21 173 149 458 131 42 32 2 159 20 386 Pseudosinella sp. 0 - 1 3 - - 3 . 7 Willowsia sp. 1 2 8 39 - - - . . . 0 Willowsia sp. 2 13 - 13 - - - - . . 0 Sminthuridae Afrosminthurus imadatei - 0 2 - - . . . 2 Allacma sp. 0 - - - - - 2 1 3 Arrhopalites alticolus - 0 - 1 - - . . 1 Arrhopalites sp. 0 - 1 8 8 5 3 . 25 Dicyrtominaformosana 0 10 - 1 2 - - - 13 Dicyrtominayaeyamensis 1 ------0 Neosminthurus amabilis 0 15 - 4 - 4 1 24 Sminthurinus modestus 0 - - - - - 7 . 7 Sminthurinus spp. 2 8 - - - - . . 0 Sphaeridiapumilis 0 - 3 - 5 - 1 . 9 Sphyrotheca sp. 38 52 - - 9 3 5 3 20 Temeritas summelongicornis 0 - - - 1 - . . Ptenothrix spp. 1 2 3 10 - - - - . . 0 Yosiides chinensis 0 - - - 18 3 4 1 26 Sminthuridaesp. 7 7- " - - - 0 - -
DIPLURA Campodeidae Lepidocampa weberi . 0 . 2 Parajapygidae 3 5 Parajapyx emeryanus 0 - - 1 - - - 6 7 Parajapyx isabellae - 0 - - - 1 . . Parajapyx sp. 1 0 3 " - - 3 - - -
HEMIPTERA Ortheziidae Nipponorthezia ardisiae 17 603 422 1046 3 14 3 2 31 53 Cixiidae Cixiidae sp. 3 5 - - - - - . 0 Issidae Gergithus okinawanus 2 0 Ceratocombidae Ceratocombus sp. 0 - - 1 - 1 Schizopteridae Kokeshia esakii - 0 17 1 1 19 Reduviidae Reduviidae sp. 0 " • - 1 - 1 Aradidae - Aradidae sp. 1 . 0 1 Aradidae sp. 2 0 1 Lygaeidae 1 Lygaeidae sp. 10 - - - - 0 -
COLEOPTERA Carabidae Colpodes buchanani 0 Colpodes rubriolus 2 2 ------0 Eustra crucifera - 1 - - 0 Synuchus sp. 0 " 1 1 Hydrophilidae Cercyon sp. 9 - 2 - 11 1 Hydrophilidae sp. - 0 1 3 6 10 Histeridae Bacanius sp. 1 1 6 . 7 2 5 7 Bacanius sp. 2 2 38 16 - 56 0 - Bacanius sp. 3 1 " - . . ; 0 Ptiliidae Acrotrichis spp. 90 25 62 30 2 209 - - n 11 4 10 36 Cissidium spp. 0 - - - 2 3 - • 8 13 Dipentiumjaponicum 1 . 191 61 - 253 10 7 - - 1 . 18 Sindosium sp. 0 - 1 - - 1 1 1 4 Nanoptilium sp. 0 - - - 3 26 - - 29 Skidmorellaquadrisulucia - 4 I 4 - ll" 4 41 61 Bambara sp. 263 478 26 777 " • - - 0 Leiodidae Agathidium sp. 1 - - 0 1 1 Soil arthropods in bird's nest fems 29
Agathidium sp. 2 - - - - - 0 - - 2 2
Colon sp. - - - - - 0 - 1 - 1
Pseudoliodes sp. - - - - - 0 - - 2 4 6 Staphylinidae 6 (Pselaphinae) Apharinodes papageno - - - - - 0 5 1 - 4 4 Aphilia sp. - 5 1 58 1 65 - -
Batriscenaulax kunigamensis - - - - - 0 - 2 - 2
- . Batrisodellus sp. - - - 3 - 3 - - 0
Batrisodes sp. - - 5 35 - 40 - - - 0
Bibloporus sp. - - 1 - 2 3 - - - 0
Bryaxis sp. 1 - - - - - 0 19 1 2 1 9 32 2 2 Bryaxis sp. 2 - - - - - 0 - -
Euplectomorphus ? sp. - - - - - 0 - - 2 2 0 Euplectus sp. 1 4 12 1 8 - 25 - - - 0 Euplectus sp. 2 - 1 - - - 1 - - -
Lasinus sp. - - - - - 0 1 - - 1 42 Morana deigo - - - - - 0 9 28 5
Morana sp. - - - - - 0 - - 2 2 2 Parapyxidicerus sp. 1 - 9 - - - 9 2 - -
Parapyxidicerus sp. 2 - - - - - 0 - - 1 1 2 Philiopsis sp. - - - - 0 1 1 - - 0 Plagiophorus fujiyamai - 4 - - - 4 - - -
Raphitreus sp. - - - 1 - 1 - - - 0
Saltisedes sp. - 3 - - - 3 - - - 0
Takaorites ? sp. - - - - 0 1 - - 1
Tribasodites sp. - 1 2 96 11 110 - - - 0
- 0 (Tachyporinae) Erchomus sp. 10 6 - 36 5 57 - - 0 (Aleocharinae) Atheta sp. - 1 - - - 1 - - - 1 Cypha nomurai - - - - - 0 - - 1
Myllaena sp. - - - 0 1 - - 1 0 Oxypoda sp. 1 61 11 18 - 91 - - -
Homalotini sp. 1 2 30 5 70 14 121 - - 0
Homalotini sp. 2 - 1 - - - 1 - - - 0 0 Oxypodini sp. - 1 - - - 1 - - -
Aleocharinae sp. 1 - 2 - - 1 1 4 - - - 0
Aleocharinae sp. 2 - - - - 0 1 - - 1
Aleocharinae sp. 3 - - 3 - - 3 - - - 0
Aleocharinae sp. 4 - - - 1 - 1 - - - 0 19 20 (Osoriinae) Lispinusjambar - - - - 0 - 1
Osoriinae sp. 1 - - - - 1 - - - 0
- - 4 (Oxytelinae) Anotylus lewisius - - - - 0 - 2 2 1 - - - 1 (Steninae) Stenus punctifer - - 0 - - 3 - 1 (Euaesthethinae) Edaphus sp. 1 - - - - 0 1 1 - 3 - 1 Edaphus sp. 2 - - - - 0 2
Euaesthetus sp. - - - - - 0 14 3 8 25
Octavius sp. - - - - - 0 3 - - 3 1 2 3 (Staphylinidae) Gabrius sp. - - - - - 0 - -
Gabronthus sp. 1 - - - - 1 - - - 0 1 2 4 (Scaphidiinae) Scaphisoma sp. 1 - - - - - 0 1 - 2 1 1 2 Scaphisoma sp. 2 - - 1 1 - - 0 Scaphobaeocera sp. - - 3 - - 3 - - - Scydmaenidae Cephennodes sp. 1 0 2 1 1 4 Cephennodes sp. 2 0 1 1 Cephennodes sp. 3 0 1 1 Cephennodes sp. 4 0 1 1 2 Euconnus sp. 1 1 1 0 Euconnus sp. 2 6 6 1 2 3 - Euconnus sp. 3 0 4 1 5 Euconnus sp. 4 0 1 - Euconnus sp. 5 0 1 I 2 Euconnus sp. 6 0 1
Microscydmus sp. 13 105 . 118 0 Neseuthia ? sp. 1 1 0 Neseuthia ? sp. 2 1 0 Stenichnus sp. I 0 0 1 1 Scarabaeidae Apogonia bicavata " - -
Sophrops kawadai 1 . . 1 .' 0 0 Elateridae Adelocera deficilis - - 1 6 7 - - - Nitidulidae Epuraea sp. 1 0 * *
Stelidota multiguttata . 0 . . 2 2 0 Aspidiphoridae Aspidophorus sp. - - . 1 - - 1 - 1 2 2 3 13 Cryptophagidae Cryptophagidae sp. - - - 1 3 3 1 0 Cerylonidae Cerylonidae sp. 1 " ' "
Cerylonidae sp. 2 1 . . 1 . . 0 Corylophidae Arthrolips sp. 1 1 7 2 11 0 ♦ 4 4 0 Corylophodes sp. - Lewisium sp. 4 1 1 6 0
Coccinellidae Coccinellidae sp. - - 1 - - 1 - - - 0
Lathridiidae Lathrididae sp. - - - 1 1 - - - 0
Colydiidae Penthelispa sp. 1 - - - - 1 - - 0
Tenebrionidae y4 Derispajaponica 1 0 Laena sp. 0 4 Anthribidae Coraginae sp. 0 1 Curculionidae Myosidesokinawanus 0 8 Otibazo sp. 1 0 4 Otibazo sp. 2 0 1 Acicnemidinae sp. 1 0 Chryptorhynchinae sp. 1 1 0 Chryptorhynchinae sp. 2 0 2 Chryptorhynchinae sp. 3 1 0 Cossoninae sp. 1 2 0 Cossoninae sp. 2 3 0 Cossoninae sp. 3 14 0 Hylobiinae sp. 1 38 0 Hylobiinae sp. 2 10 17 Scolytidae Scolytidae sp. 1 4 7 Scolytidae sp. 2 11 2 Scolytidae sp. 3 0 Scolytidae sp. 4 0 2 Scolytidae sp. 5 0 1 Total no. individuals 3371 8119 2281 5078 491 8527 27867 1058 2285 852 1024 1593 5440 12252 Total no. species 96 95 124 87 66 105 267 120 189 100 150 159 246 403