Applied Vegetation Science ]] (2011) 18

Fire severity affects vegetation and seed bank in a wetland

Hideo Kimura and Shiro Tsuyuzaki

Keywords Litter; Prescribed re; Reed swamp; Seed bank; Standing vegetation Abbreviations AIC = Akaikes Information Criteria; GLM = Generalized Linear Model; LMM = Linear Mixed-Effects Model Received 8 April 2010 Accepted 19 January 2011 Co-ordinating Editor: Juli Pausas

Abstract Questions: How does the severity of prescribed res affect vegetation and seed bank in a wetland? Location: A re-prone reed swamp in northern Japan (250 ha, 40149 0 N,
141122 0 E, o 10 m a.s.l.).

Methods: Vegetation, biomass and seed bank were monitored for the 2 yr after
annual prescribed res were discontinued. Plant communities were placed into three categories based on re severity: high (H) re consumed litter completely; moderate (M) re removed standing litter but left wet fallen litter; and low (L) re incompletely removed standing litter and did not remove fallen litter. Soil samples were collected in autumn 2007 and early summer 2008, and germinable seed bank was investigated by greenhouse trials.

Tsuyuzaki, S. (corresponding author, tsuyu@ees.hokudai.ac.jp): Graduate School of Environmental Earth Science, Hokkaido University, Sapporo 060-0810, Japan Kimura, H. (kimuhide0613@hotmail.co.jp): Graduate School of Environmental Science, Hokkaido University, Sapporo 060-0810, Japan .

Results: High re severity increased diversity in the next growing season by the establishment of short herbs in the standing vegetation. The biomass of forbs and grasses was greater in H where Phragmites australis biomass was reduced. The density of seed bank was 4 30 000 seeds m2 throughout all the treatments. Perennial plants were dominant in the vegetation, while annuals, biennials and rushes were dominant in the seed bank. Small seeds were more abundant in the soil than in the litter. Qualitative and quantitative similarities between seed bank and the vegetation were low, and tended to be higher in H. Conclusions: Fire contributed to the development of diverse standing vegetation via the positive effects on seed bank dynamics, and can be considered a tool to maintain species-rich marshes.

Introduction
Although prescribed re has been used for increasing productivity and biodiversity in various ecosystems (Iwakuma 1996, Hiers et al. 2003), it threatens the sustainability of the vegetation if the prescription is over-used (Pausas & Keeley 2009). The effectiveness of re differs with severity, scale, frequency and patchiness (Hochkirch & Adorf 2007; Cox & Allen 2008). For example, biomass removal by re decreases interspecic and intraspecic competition. Moderate re severity promotes seedling recruitment by increases in light and nutrients (Parker & Kelly 1989), while severe re may decrease seed bank and species diversity (Mamede & de Araujo 2008). Therefore, re severity affects post-re vegetation development and recovery. Fire removes not only standing vegetation but also litter that affects vegetation dynamics and the seed bank

(Owens et al. 2007; Allen et al. 2008; Egawa et al. 2009). In burned forests, the severity re affects the seed bank more than the vegetation (Johnson 1992). In heath, some re regimes that promote overstory development are in conict with persistence of understory species because of interactions between functional groups that are related to growth, reproduction and competition (Keith & Bradstock 1994). Therefore, the removal of litter or biomass, which is related to light environments, may be key for predicting the effects of re on ecosystem maintenance, and clarifying relationships between seed bank and the vegetation provides insight to the resilience of a community against re (Hopfensperger 2007). In Hotoke Swamp, northern Japan, where Phragmites australis is dominant, prescribed re had been conducted every early spring for about 30 yr. Prescribed re was not carried out in 2008, and this abeyance gave an opportunity to monitor changes in vegetation and seed bank for

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Effects of re in early spring on wetland

Kimura, H. and Tsuyuzaki, S.

2 yr after the last prescribed re. We expected that this temporary release from prescribed re would lead to signicant changes in seed bank and/or vegetation. The hypothesis was that re severity inuences the seed bank and the standing vegetation differently. To test this hypothesis, we measured the effects of prescribed re on vegetation structure and species composition of the standing vegetation and seed bank.

Materials and Methods

Study site Land reclamation was promoted for wetlands in Japan soon after World War II, because of increases in rice demand. Hotoke Swamp, located in northern Honshu Island, Japan, was prepared for a change in land-use after 1964 by the construction of drains (250 ha, 40149 0 N, 141122 0 E, o 10 m a.s.l.). Before the construction of drains the swamp was a shallow pond with low plant cover. After construction, groundwater was usually below the ground surface except during times of snow-melt, and a P. australis grassland subsequently developed. Such reclamation was widely applied in Japan and led to the dominance of P. australis (Iwakuma 1996). After the 1970s, the demand for rice declined and the government switched the policy from reclamation to restoration (Sugiura et al. 2003). Subsequently, the reclamation plan of Hotoke Swamp was abandoned in early 1970s. However, local farmers perform prescribed re and remove water by pumps every spring to keep reclaimed land from the abandonment. The annual prescribed res usually burn the entire area. An unplanned wildre occurred on 1 May 2007 before the prescribed re was performed, and thus prescribed re was not conducted in 2007 and 2008. Soon after the wildre, we explored the swamp and established monitoring plots. The study region is in a warmcool temperature zone. In the city of Misawa, 15 km from the study site, annual precipitation was 1165 mm in 2007 and 1000 mm in 2008 (ADMO 2009). Mean annual temperature in 2007 was 10.4 1C with a monthly minimum of 0.8 1C in Jan and maximum of 23.7 1C in Augu, and was 10.0 1C in 2008 (minimum = 2.1 1C, maximum = 20.6 1C in August). Snow-free period is usually from Apr to Dec. Vegetation, biomass and micro-environmental characteristics Three re severities were recognized: high (hereafter, H) completely consumed standing and fallen litter; moderate (M) completely removed vegetation and standing litter but did not burn fallen litter because the fallen litter was wet as a result of high water level; and low (L) incompletely removed standing litter and did not remove

fallen litter. There were no unburned sites for control plots, because the re burned the whole area, with the same patterns of ordinal prescribed re. Three locations for each severity were selected for monitoring the development of vegetation (i.e. three replicates in the same burn area). Vegetation was measured three times, midAugt 2007, mid-Jul 2008 and mid-Aug 2008, in 20 1 m 1 m permanent plots at each location. More than 5 m separated each plot. The percentage cover of each species was visually estimated. As Pilea mongolica withered earlier in 2008 than in 2007 and a few other species did not emerge in mid-Jul 2008 but they emerged by Augu, plant cover measured in 2008 was averaged across the two surveys for the analysis in order to compare more accurately with the data collected in 2007. Biomass was harvested from 3 to 10 Sept 2007 and from 5 to 12 September 2008 when P. australis ceased growing. The harvested biomass was separated into eight categories: P. australis leaf, P. australis stem, forb leaf, forb stem, grass leaf, grass stem, sedge and fern. The biomass was classied into photosynthetic parts (i.e. leaves, including the whole of sedges and ferns) and non-photosynthetic parts (non-photosynthetic organs, i.e. stems). Standing litter was also collected and weighed. The weight was measured after drying up at 80 1C in an oven for more than 3 d. Light intensity and temperature were recorded at 1-h intervals from Mar to Sep in 2008 using light/temperature loggers (HOBO; Onset, Bourne, MA, USA) placed in each of the three severities. Two loggers were set up on the surface and subsurface of litter in M and L, respectively, and a logger was only on the soil surface in H because of the lack of litter. The sensors placed on subsurface were located directly above the soil surface and were more than 8 cm below the surface of the litter.

Seed bank Seed bank sampling was conducted in early Nov 2007 (autumn) and mid-Jun 2008 (summer). A 100-cm3 steel tin (20 cm2 surface area and 5 cm depth) was used for sampling soil. Seed bank samples were collected separately from the litter and the soil, adjacent to the L and H plots except that litter was not collected in H in autumn 2007 because of the very small amount. The total number of samples was 20 in H and 40 in L in autumn 2007, and 40 in H and L in summer 2008. The soil collections were made directly underneath the litter where the litter covered the soil. To consider the vertical movements of seeds, litter on L and soil on H were analysed as surface, and soil on L was analysed as subsurface. Seed germination experiments were conducted in a greenhouse at Hokkaido University, Japan. The samples

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Kimura, H. and Tsuyuzaki, S.

Effects of re in early spring on wetland

were cold-stratied for 1 month at 3 1C in a dark refrigerator within plastic bags which kept the moisture in the sample. The samples were then spread to less than 5 mm thick on plastic trays lled with vermiculite. The trays were covered with white sheer nets to prevent contamination. Temperature was between 10 1C and 35 1C under natural light. Water was automatically sprinkled seven to 12 times in a day, depending on air moisture. Seedlings were marked by numbered ags everyday until no more seedlings emerged for more than 1 wk. When seedlings were not identied, the seedlings were transplanted to large trays lled with culture soil and grown. Seed germination was observed for 3 months in each greenhouse experiment.

Data analysis The alpha diversities of the vegetation and seed bank were evaluated by three parameters; species richness, ShannonWiener diversity (H 0 ) and evenness (J 0 ). H 0 and J 0 are P expressed as: H 0 = (pi log2pi), and J 0 = H 0 /log2S, where S is total number of species in the vegetation plot or seed bank sample, and pi is relative dominance of species i from 1 to S. Vegetation analysis was performed using a linear mixedeffects model (LMM) because of repeated measures, and analysis of the seed bank and standing biomass used a generalized linear model (GLM). The dependent variables were species richness, diversity, evenness, plot cover, P. australis cover, total seed number, and seed number of each dominant species. Species richness and seed number were assumed to follow a Poisson probability distribution, diversity and cover a Gaussian, and evenness a negative binomial. In GLM, the explanatory variables were re severity, year and the interaction term between them. The best models of GLM were selected by stepwise procedures of Akaikes information criteria (AIC). Plot and year were assigned as random effects in LMM. Litter biomass was compared between severity, year and their interaction by GLM with a Gaussian error distribution. For seed bank, two independent variables, substrate (litter and soil) and layer (surface and subsurface), were also examined. Mean monthly temperature, temperature difference and light illumination were examined between severities and between months by LMM with months used as random effects. The signicance level was adjusted to 0.01 to reduce the chance of Type I errors associated with multiple tests. Species in seed bank and the standing vegetation were summarized by growth form, longevity, and seed-dispersal type. The growth form was divided into grass, forb, rush and sedge and longevity was into annual, biennial and perennial. Seed size was obtained for each species from the literature (mainly Ohwi & Kitagawa 1983; Ishikawa 1995).

In H and L, respectively, similarities were calculated on a matrix made by (seed bank vegetation season). Two similarity indices, qualitative Srensens index and quantitative Goodalls percentage similarity index, were applied to plant cover and number of seeds with both transformed into relative percentages (Egawa et al. 2009). Ferns, for which dominance in seed bank were not estimated, and unidentied species were not used for the evaluations. The presence and absence of each dominant species in each plot and in each soil sample were investigated based on AIC after GLM assuming a binomial distribution. In the analysis, explanatory variables were growth form, longevity, seed dispersal, seed volume and seed shape. Seed volume was calculated by length and width, assuming an oval. As seed shape was expressed as length divided by width, the lower values indicated more circular shapes. All statistical analyses were made with the package R 2.9.0 (R Development Core Team, Vienna, Austria.

Results
Vegetation structures and environments A total of 30 species was recorded from the standing vegetation (see the Supporting Information, Appendix S1). Phragmites australis was predominant throughout, and was the tallest (220 to 300 cm). There were no shrubs or trees. Except for lower species diversity in L than in H, species richness, diversity and evenness did not differ between re severities, and did not change across the 2 yr (Table 1). These results imply that diversities evaluated by plant cover were modied less by severe re. Fire severity did not affect the cover of all common species, including P. australis (LMM, P 4 0.01). Standing litter biomass in 2007 was 224 g m2 67 (mean standard deviation) in H, 294 g m2 52 in M, and 695 g m2 103 in L, and was signicantly higher in L (GLM, P o 0.01) because the litter was burnt out in H and M. In 2008, the litter biomass did not differ between re severities (543 g m2 62 in H, 636 g m2 55 in M, and 833 g m2 124 in L; P 4 0.01), showing that litter recovered promptly if re was absent. Mean monthly temperature was nearly zero in the three re severities in Feb because of snow cover, and then gradually increased to Jul. From Jul to Sep, the temperature was higher than 15 1C. Depth below litter showed small temperature differences within a day, although mean temperatures did not differ between surface and subsurface for any re severity. Therefore, the major effect of litter was to reduce temperature uctuations. After snowmelt, light illumination was not different between months, and was lower in the subsurface throughout the year. As plot cover did not differ between

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Effects of re in early spring on wetland

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Table 1. Mean species richness, diversity and evenness in vegetation (n = 20 in each severity) received different re damages in 2007 and 2008. Each numeral shows mean with standard deviation. Each numeral shows mean with standard deviation. The signicance is investigated by linear mixedeffects model. Explanatory variables that are not adopted by Akaikes Information Criteria are not shown. signicant at P o 0.01. NS = not signicant. The difference was compared from H to L and M, and from 2007 to 2008. On the species diversity of vegetation, the interaction was observed between year and severity. Abbreviations: Y = year, H = high re severity, M = moderate, and L = low. = not measured. 2007 H M L 2008 H M L Signicance Intercept Severity M Species richness Species diversity Evenness Total plant cover 6.1 1.6 4.7 1.3 5.0 1.5 8.2 2.5 6.0 1.8 6.0 1.8 1.9 0.3 1.7 0.3 1.5 0.3 2.0 0.4 1.7 0.3 1.7 0.3 11.80 11.88 0.25
NS

Year L 0.20
NS

Interaction YM YL
NS

10.30

NS

0.10

0.05NS 0.05NS

0.22NS

0.39

10.11NS

10.08NS

0.7 0.1 0.8 0.1 0.7 0.1 0.7 0.1 0.7 0.1 0.7 0.1 11.05NS 10.20NS 0.28NS 0.32NS 10.30NS 0.13NS 72 8 80 13 60 14 85 12 80 11 74 12 193.2NS 143.9NS 51.6NS 180.3NS 78.7NS 15.9NS

Table 2. Coefcients of general linear model (GLM) that compares species richness, diversity, evenness, total number of seeds, and numbers seeds on re frequently germinated species in seed bank. Stepwise Akaikes Information Criteria (AIC) was used to select the best models, and explanatory variables not selected by AIC are not shown. Year was a comparison between 2007 and 2008, layer was between the surface and subsurface, and substrate was between litter and soil. = signicant at P o 0.01. NS = not signicant. Severity High (intercept) Species richness Species diversity Evenness Species Pilea mongolica Cardamine exuosa Stellaria alsine var. undulata Juncus effusus var. decipiens Juncus leschenaultia Total number of seeds germinated 11.911 11.531 10.919NS 14.285 13.465 11.245 13.121 12.209 14.927 0.205 0.262 10.329 0.566 0.310 Less 0.180 10.011NS 10.164 10.566 10.323NS 1.299 0.686 0.126NS 0.338NS 0.976 0.776
NS

Year

Layer

Substrate

Interaction Year Depth Severity Depth 0.352NS

0.062NS

10.379NS

0.265NS

1.081 1.991 10.406 11.801 10.252

0.927 0.447 0.380NS 0.599 0.870

10.956 11.964 10.524 10.603

10.632 11.186 10.997NS

severities (Table 1), shading of the subsurface soil was caused by litter. The above-ground biomass of P. australis was highest in M, and decreased from 2007 to 2008 (GLM, P o 0.01). NonPhragmites photosynthetic biomass, including sedges and ferns, was lower in L in 2007 than in 2008 (P o 0.01), but did not differ between H and M. The non-Phragmites nonphotosynthetic biomass did not differ between severities. Litter thickness was less in H and M than in L (P o 0.01). Photosynthetic non-Phragmites biomass was inuenced more by litter amount than by P. australis biomass, suggesting that other species increased promptly after litter removal by re but then declined in the second year.

Seed bank composition In total, 4903 and 4967 seeds germinated in autumn 2007 and summer 2008, respectively, equivalent to 39 469 m2 and 31 044 m2 (see the Supporting Information, Appen-

dix S2). A total of 39 species were recorded from the seed bank (13 annuals, three biennials, 22 perennials, and one unidentied). No shrubs and trees were detected in the seed bank. Seed bank species richness was higher in H than in L (Table 2), but did not differ between autumn 2007 and summer 2008: i.e. the richness in H was 6.7 1.7 (mean standard deviation) in 2007 and 8.0 1.8 in 2008, and in L was 5.7 1.5 and 6.6 1.6. Species diversity and evenness did not differ between severities (diversity; H = 1.9 0.5 in 2007, 2.1 0.4 in 2008, L = 1.5 0.5 in 2007, 2.0 0.4 in 2008; evenness; H = 0.7 0.2 in 2007, 0.7 0.1 in 2008, L = 0.6 0.2 in 2007, 0.7 0.1 in 2008). Seed bank species diversity was higher in summer 2008 than in autumn 2007, while evenness did not differ between the years. As diversity was determined by the combination of species richness and evenness, both of which increased but were statistically non-signicant, diversity increased signicantly over the 2 yr.

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High severity 2007 Vegetation 7.5 41.7 14.2 43.2 Seedbank 2007 86.3 63.8 2008 2007 1.2 43.9 1.8 53.8 1.1 30.3 95.9 85.7 2008 2007

Less severity 95.9 85.7 1.5 46.2 0.2 34.3 0.5 40.9 2008

80.7 73.9

2008

Fig. 1. Similarities (%) between seed bank and the standing vegetation in each season and between seed bank in autumn and summer. Upper and lower numerals show quantitative and qualitative similarities, respectively. Quantitative similarity is by Goodalls index, and quantitative one is by Srensens index. Thickness of lines connected with two circles indicates the intensity of similarities. Thick line 4 80% in quantitative similarity; medium line: 2% to 15%, thin line: o 2%.

Seed density decreased from autumn 2007 to summer 2008, litter to soil, H to L, and subsurface to surface (Table 2). Five seed bank species were recorded from all severities for 2 yr: Pilea mongolica, Cardamine exuosa, Juncus effusus var. decipiens, Stellaria alsine var. undulata, and Juncus leschenaultii. Most species that accumulated seeds in litter decreased seed densities in the second year. Juncus leschenaultii and J. effusus var. decipiens had higher seed densities in subsurface than in surface soil. The responses of seed density to re severity varied between the common species; the seeds of P. mongolica, C. exuosa and J. leschenaultia were fewer in L, S. alsine var. undulata more abundant and J. effusus var. decipiens unchanged. Positive interactions were detected between depth and year for three species C. exuosa, S. alsine var. undulata and J. effusus var. decipiens , and between severity and depth for C. exuosa and J. effusus var. decipiens. These interactions indicated that the seeds moved to deeper soil layers with time. However, P. mongolica, which produced the largest seeds of the ve common seed bank species (see the Supporting Information, Appendix S3), developed more of a seed bank in the litter in H (Table 2), but decreased in seed density with time. The two Juncus species produced small seeds, and accumulated seeds more at greater depth.

Relationships between seed bank and standing vegetation The total number of identied seed plant species in the standing vegetation and seed bank was 47 (Appendix S3). In addition, three ferns and three unidentied taxa were recorded. Perennial forbs were most common (i.e. 30 species, followed by 14 annuals and three biennials). Of these annual and biennial plants, only ve species ap-

peared in the vegetation. Most annuals and biennials including P. mongolica produced owers in 2007. Of the seed plants identied, 33 species (70% of the total) had gravity dispersal (Appendix S3). However, six and three of the 33 gravity-dispersed species also use another dispersal agent, wind or animal, respectively. Therefore, seed dispersal distances of these species would be short if secondary seed dispersal by water functioned weakly. Species occurring in both standing vegetation and seed bank produced larger seeds (GLM, P o 0.01). There were no other signicant factors identied by AIC in the relationship between standing vegetation and seed bank. Qualitative similarities in seed banks between the 2 yr were more than 60% in both L and H, and quantitative similarities ranged from 81 to 87% (Fig. 1), indicating that species composition and relative dominance did not differ greatly between years, or between severities. Qualitative similarities between seed bank and the standing vegetation ranged from 30 to 55%, while quantitative similarities between them were extremely low (i.e. less than 2%), except for 14.2% between standing vegetation and seed bank in 2007 and 7.5% between standing vegetation in 2007 and seed bank in 2008. These showed that H developed more similar species composition between seed bank and standing vegetation than L. In addition, seed bank in H in 2008 was inuenced more by the standing vegetation in the previous year (2007), indicating that seed bank development is affected more by severe re that removed litter.

Discussion
Litter reduced light intensity and temperature uctuation without changing mean temperature, while the standing

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Effects of re in early spring on wetland

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biomass did not. Therefore, the major environmental changes between re severities were light intensity and temperature uctuation when litter was removed. The severe re did not eliminate perennials in the standing vegetation. Most perennials that do not develop a seed bank should survive and recover by vegetative reproduction after a re. For example, P. australis can resprout and dominate from rhizomes after re (Thompson & Shay 1985), but it does not develop a persistent seed bank (Lenssen et al. 1999; Egawa et al. 2009). After disturbances, perennial grasses often invest more in vegetative reproduction than seed reproduction (Gonzalez & Ghermandi 2008). In general, the plant cover of a species that can recover by vegetative reproduction is unlikely to be inuenced by the severity of re. Furthermore, species diversity in the standing vegetation was higher in H and M than in L. The higher diversity is related to the greater photosynthetic biomass of non-Phragmites species that increased because of litter removal. These results indicate that species under a P. australis canopy contribute to high species diversity when litter on the ground surface was removed by re. Pilea mongolica, which produced the largest seed of the dominant seed bank species, developed its seed bank more in litter layer and/or on the surface than at greater depth. However, its seed density declined in the second year, probably because of seedling emergence in the rst year and difculties in vertical movements owing to large seeds. Seed sizes often determine vertical distribution of seed bank; for example small seeds are distributed more at greater depth (Tsuyuzaki & Goto 2001; Cerabolini et al. 2003). In 2007 P. mongolica showed the highest cover on M and established well in H. The signicant difference between the rst and second years after re was caused by litter accumulation, suggesting that litter removal by re has an important role on the seedling emergence and seed bank dynamics of this species. Both annuals and biennials producing small seeds made persistent seed banks, and recovered quickly after re (Thompson et al. 1997; Gonzalez & Ghermandi 2008). However, about two-thirds of the species in the seed bank did not establish well in the vegetation of Hotoke Swamp. In particular, two biennials, C. exuosa and S. alsine var. undulata, both of which require light for seed germination (Baskin & Baskin 1998), were frequent in the seed bank of both litter and soil for the 2 years but had low cover in the standing vegetation. These results suggest that the biennial species maintained persistent seed banks even where the severity of re was greatest. Rushes (Juncus) and annuals, most of which require light for seed germination, as well as the two biennials, are also dominant in the seed bank of various grasslands (Allen et al. 2008). In addition, Juncus did not establish well in the vegetation,

even though the seed bank was well developed (Milberg 1995). The seeds of Juncus are particularly small, and are likely to move downwards easily. Numerous small seeds should preserve the viability in the soil by the downward movements. Juncus leschenaultii and J. papillosus develop long-term persistent seed banks with the vertical movements of seeds (Jensen 2004). In these ways, high seed density of these species should be maintained. Similarities between seed bank and standing vegetation are low in Hotoke Swamp. In wetlands where seeddispersal distance is short the similarities are high (Jutila 2003). Short-distance gravity seed-dispersal was common in Hotoke Swamp, however, only four of 13 annual species in the seed bank were recorded from the standing vegetation. Furthermore, species producing small seeds such as Juncus did not appear in the vegetation but did in seed bank. Low similarities between seed bank and the vegetation are often derived from the predominance of Juncus in seed banks that do not emerge in the vegetation (Parker & Leck 1985). These inconsistencies should lead to low similarities on Hotoke Swamp. Disturbances increase the similarities between seed bank and the standing vegetation, because the resultant sparse vegetation and litter reduce competition for light and promote seed germination (Osem et al. 2006), and/or seedling emergence is promoted by disturbances (Grandin 2001). High severity showed the greatest similarity between seed bank and the standing vegetation in the rst year, while similarities between them became quite low in low severity for the 2 yr. Fire that removes litter completely in early spring promotes the link between seed bank and the standing vegetation. The most severe re treatment, H, showed higher species diversity and seedling emergence, indicating that the re was not catastrophic even when litter was completely removed. For successful conservation and restoration, suitable germination and establishment conditions should be pro+ vided for seed banks (Leck & Schutz 2005). In conclusion, high-severity prescribed re is adequate for maintaining diverse vegetation structure and high diversity via developing seed bank that contributes vegetation dynamics.

Acknowledgements
We thank all members in Plant Ecology Laboratory and S. Suzuki for support, staff members of Misawa City Ofce for permission and support, and F. Kobari in CAST for technical help. Cordial thanks also J. Pausas, D. Keith, J.H. Titus and two anonymous reviewers for their critical reading of the manuscript. This work is partly supported by grants from JSPS.

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Kimura, H. and Tsuyuzaki, S.

Effects of re in early spring on wetland

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Applied Vegetation Science Doi: 10.1111/j.1654-109X.2011.01126.x r 2011 International Association for Vegetation Science

Effects of re in early spring on wetland

Kimura, H. and Tsuyuzaki, S.

Supporting Information
Additional Supporting Information may be found in the online version of this article: Appendix S1. Percentage cover on each taxa in the summers of 2007 and 2008. Appendix S2. Seed density (m2) in soil and litter substrates on heavily and less disturbed sites.

Appendix S3. Growth form, longevity, seed dispersal type and seed size on identied seed plants in vegetation and seed bank. Please note: Wiley-Blackwell is not responsible for the content or functionality of any supporting materials supplied by the authors. Any queries (other than missing material) should be directed to the corresponding author for the article.

Applied Vegetation Science Doi: 10.1111/j.1654-109X.2011.01126.x r 2011 International Association for Vegetation Science