Woodlands : Ecology, Management and Conservation [1 ed.] 9781612090443, 9781611225426

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Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved. Woodlands : Ecology, Management and Conservation, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved. Woodlands : Ecology, Management and Conservation, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

ENVIRONMENTAL SCIENCE, ENGINEERING AND TECHNOLOGY

WOODLANDS: ECOLOGY, MANAGEMENT AND CONSERVATION

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ENVIRONMENTAL SCIENCE, ENGINEERING AND TECHNOLOGY

WOODLANDS: ECOLOGY, MANAGEMENT AND CONSERVATION

ERWIN B. WALLACE

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EDITOR

Nova Science Publishers, Inc. New York Woodlands : Ecology, Management and Conservation, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

Copyright © 2011 by Nova Science Publishers, Inc. All rights reserved. No part of this book may be reproduced, stored in a retrieval system or transmitted in any form or by any means: electronic, electrostatic, magnetic, tape, mechanical photocopying, recording or otherwise without the written permission of the Publisher. For permission to use material from this book please contact us: Telephone 631-231-7269; Fax 631-231-8175 Web Site: http://www.novapublishers.com NOTICE TO THE READER The Publisher has taken reasonable care in the preparation of this book, but makes no expressed or implied warranty of any kind and assumes no responsibility for any errors or omissions. No liability is assumed for incidental or consequential damages in connection with or arising out of information contained in this book. The Publisher shall not be liable for any special, consequential, or exemplary damages resulting, in whole or in part, from the readers‘ use of, or reliance upon, this material. Any parts of this book based on government reports are so indicated and copyright is claimed for those parts to the extent applicable to compilations of such works.

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Independent verification should be sought for any data, advice or recommendations contained in this book. In addition, no responsibility is assumed by the publisher for any injury and/or damage to persons or property arising from any methods, products, instructions, ideas or otherwise contained in this publication. This publication is designed to provide accurate and authoritative information with regard to the subject matter covered herein. It is sold with the clear understanding that the Publisher is not engaged in rendering legal or any other professional services. If legal or any other expert assistance is required, the services of a competent person should be sought. FROM A DECLARATION OF PARTICIPANTS JOINTLY ADOPTED BY A COMMITTEE OF THE AMERICAN BAR ASSOCIATION AND A COMMITTEE OF PUBLISHERS. Additional color graphics may be available in the e-book version of this book. LIBRARY OF CONGRESS CATALOGING-IN-PUBLICATION DATA Woodlands : ecology, management, and conservation / editor, Erwin B. Wallace. p. cm. Includes index. ISBN 978-1-61209-044-3 (eBook) 1. Forest management. 2. Forest ecology. I. Wallace, Erwin B. SD431.W57 2010 634.9--dc22 2010041380

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CONTENTS Preface Chapter 1

Chapter 2

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Chapter 3

Chapter 4

Chapter 5

Chapter 6

Chapter 7

vii Colonization of Post-Agricultural Black Alder (Alnus Glutinosa (L.) Gaertn.) Woods by Woodland Flora Anna Orczewska Diversity Patterns, Adult Resource Use and Conservation of Butterfly Communities in and around a Primeval Woodland of Mount Fuji, Central Japan Masahiko Kitahara Guidelines For Sustainable Management of Degraded Lands: Experiences on Caatinga and Semi-arid Mediterranean Woodlands Miguel Ángel Herrera Machuca, Rinaldo Luiz Caraciolo Ferreira, Juan Ramón Molina Martínez and Mércia Virginia Ferreira dos Santos Partial Harvesting in Old-Growth Boreal Forests and the Preservation of Animal Diversity from Ants to Woodland Caribou Daniel Fortin, Christian Hébert,, Jean-Philippe Légaré,, Nicolas Courbin, Kyle Swiston, James Hodson, Mélanie-Louise LeBlanc, Christian Dussault,, David Pothier, Jean-Claude Ruel and Serge Couturier Ecology and Management of Natural and Reforested Canary Island Pine Stands José Ramón Arévalo, Agustín Naranjo Cigala, José María Fernández-Palacios, and Silvia Fernández-Lugo A Landscape History Approach to the Assessment of Ancient Woodlands Ian D. Rotherham Miombo Woodland Productivity: The Potential Contribution to Carbon Sequestration and Payment for Environmental Services in East and Southern Africa Stephen Syampungani and Paxie W. Chirwa

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13

49

91

115

137

163

185

vi Chapter 8

Chapter 9

Contents British Bluebells: The Potential of Using a Protected Species as a Provider of Fine Chemicals to Enhance its Conservation Vera Thoss African Management of Woodland Vimbai Chaumba Kwashirai

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Index

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203 215 233

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PREFACE Ecologically, a woodland is a low-density forest forming open habitats with plenty of sunlight and limited shade. Woodlands may support an understory of shrubs and herbaceous plants including grasses. Woodlands may form a transition to shrubland under drier conditions or during early stages of primary or secondary succession. This book presents research in the study of woodlands, including temporal and quantitative changes in the proportion of different land-use forms in the landscape of woodlands; butterfly communities in the woodland of Mount Fuji, Japan; experiences of the Caatinga and semiarid Mediterranean woodlands; ecology and management of natural and reforested Canary Island pine stands; and the ecological distribution of miombo woodlands productivity. Chapter 1- A survey on the colonization of the herb layer of post-agricultural black alder woods by woodland flora and on the edaphic, hydrological and light conditions responsible for the colonization mechanisms present in such woods was carried out in the Oleśnica Plain and Żmigród Valley (SW Poland) in the habitats of an oak-hornbeam community, alder-ash carrs and wet alder woods. In the 33 transects (80 m long by 4 m wide) consisting of 10 quadrats (16 m2 each) laid out perpendicularly across the ancient-recent border, data were collected on herb layer composition, chemical soil properties and illumination level. Furthermore, the groundwater level was recorded in piezometers. Migration rates were calculated for 51 woodland plant species. The mean migration rate for typical wet alder woods based on the maximum cover reached 1.20 m yr-1 and the one based on the farthest individual 1.60 m yr-1, for oak-hornbeam forests were 1.17 and 1.63 m yr-1 respectively and for alder-ash carrs 0.79 and 1.26 m yr-1. Among the woodland herbs recorded there was a group of species unable to migrate into recent alder woods and a group of slow-colonizing species (AAWS – ancient alder woodland species). On the other hand, many herbs were able to colonize recent woods very efficiently (OAWS – other ancient woodland species); some of them migrated at a pace exceeding 2 or 3 m yr-1. Thus, true woodland species differ in their dispersal potential. In wet and fertile recent forests adjacent to ancient source woodlands, herb layer recovery proceeds faster than in poorer and drier habitats. Forest age, pH, humus type and groundwater level were the variables having the greatest influence on the distribution pattern of woodland species in recent woods. A competitive species, vigorously growing in the herb layer of recent woods – Urtica dioica, avoided sites with a high level of groundwater combined with poor illumination. To create the best conditions that allow for effective forest recovery in alder woods, it is essential to maintain a high level of water and shade in the forest floor. This reduces the competitive exclusion of woodland flora by

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Erwin B. Wallace

aggressive herbs and facilitates the immigration of forest species. In conclusion, forest management should give priority to either the restoration or maintenance of the natural water regime. If necessary natural water conditions should be restored prior to afforestation. In addition, in order to ensure the successful recovery of a herb layer rich in woodland flora, recent forests should be located in direct proximity to ancient forests. Chapter 2- In central Japan, Aokigahara primeval woodland is considered to be one of the most natural areas around Mount Fuji and a core area in the conservation of the biodiversity of Mount Fuji. First, I chose butterflies as an indicator species of biodiversity and examined six communities in and around the woodland in 2000 using transect counts to examine and search for diversity and rarity hotspots and their associated landscapes. The results showed that butterfly species richness and species diversities H‟ and 1/λ were significantly higher in woodland-edge sites than in woodland-interior and/or open-land sites, and variation in the total number of species among these three landscape types was well accounted for by ecologically specialist species, such as landscape specifics, oligo-voltines, narrow diet feeders and low density species. Thus, the species regarded as vulnerable to extinction, including Red List species, were observed more often in woodland-edge sites than in woodland-interior and/or open-land sites. As a result, in the study area, diversity and rarity hotspots were found in woodland-edge landscapes. The reasons why butterfly diversity and rarity hotspots were established in forest-edge landscapes were analyzed and interpreted from several points of view, including disturbance level, landscape elements and plant species richness. From these results, and the fact that some species were confined to woodland-interior sites, I conclude that it is very important to conserve and manage woodland-edge habitats (considered to be semi-natural) as well as woodland-interior habitats (considered to be the most natural) to maintain the diversity of butterfly communities and preserve the various types of threatened species in and around the Aokigahara primeval woodland. Second, I examined the relationships between the diversities of vegetation, adult nectar plants, and butterflies in and around the Aokigahara woodland. The results showed that the nectar resource utilization by adult butterflies was significantly biased to herbaceous plants, especially to perennials, compared to woody species, although most of the study area was in and near the woodland. There were greater nectar plant species in sites with greater plant species richness. Among the butterfly community indices analyzed, the strongest correlation was detected between butterfly species richness and nectar plant species richness at each site. Another close correlation was detected between the species richness of nectar plants and herbaceous plants at each site. These results suggest that herbaceous plant species richness in a habitat plays a central role in its nectar plant species richness, and the nectar plant richness is a highly important factor supporting its adult butterfly species richness. Consequently, I propose that the maintenance and management of herbaceous plant species richness in a butterfly habitat, which lead to those of its nectar plant species richness, are very important for conservation of butterfly diversity even in and around woodland landscapes of temperate regions. Chapter 3- Land degradation and soil erosion are perceived as important and worldwide problems in dryland areas. In semi-arid climate conditions, interactions between vegetation, rainfall and soil properties have been related to the effectiveness in reducing the degredation processes. Landscape structure can be characterized by different woodlands based on the structure and physiognomy of the vegetation communities. From the perspective of water availability, the shortage of rainfall during summer is the main limitation for livestock development. Soil properties limitations are vital in order to the soil erosion and soil water

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Preface

ix

availability. On the other hand, climate change, socio-economic changes and cultural evolution have been associated with the gradual loss of ecosystem productivity due to biophysical stress limiting the environmental and human sustainability. In Brazil, the semi-arid climate zone is located on the Northeastern of the country, an area of 845,000 km2 showing about of the 11% of the Brazil area. In the case of Spanish semi-arid area, woodlands tend to be shrublands because of the limited environmental conditions and the recent changes in land use. Brazil and Spanish semi-arid landscapes have some common characteristics. The socio-economic development of their societies have required aggressive practices such as land use change, land occupation, burning, deforestation and overgrazing. In semi-arid ecosystems, species are adapted to environmental and subsistence activities on a local level. In this paper, field experiences associated to Brazil and Spain woodlands are analyzed in order to find technical alternatives on the urgent need to semi-arid landscape conservation. Chapter 4- Current forest management must maintain biodiversity, an objective that has led to the rapid development of new forestry practices in recent years. However, empirical evaluation of the impact that these practices have on biodiversity has not kept the same pace. For example, small merchantable stems are now frequently protected during the harvest of old-growth boreal forests in eastern Canada. This silvicultural practice (referred to as CPPTM in Québec) ends up protecting all stems with a diameter at breast height of 9-15 cm, and is therefore expected to maintain some of the irregular attributes of old-growth forest structure. Whether or not this approach is sufficient to maintain local biodiversity remains unclear. The authors evaluated the short-term impact of CPPTM harvesting, mostly 2-3 years after logging, on a broad range of animal species differing largely in size and resource requirements. More specifically, the authors estimated the abundance, occurrence or local intensity of habitat use by ants, beetles, forest birds, snowshoe hare (Lepus americanus), moose (Alces alces) and woodland caribou (Rangifer tarandus caribou) in a boreal ecosystem. The 19,000 km2 study region is dominated by >270 year old stands with irregular structure mostly comprised of black spruce (Picea mariana) and balsam fir (Abies balsamea). Harvesting by CPPTM caused a 75-85% reduction in tree basal area. This decrease was sufficient to alter animal assemblages of all taxonomic groups. Ants and beetles associated with open areas were more abundant in CPPTM sites than in uncut stands. Conversely, species associated with mature forests were much lower in CPPTM than in uncut stands. Most birds associated with latesuccessional habitats were also less likely to be observed in CPPTM sites than in uncut stands. Snowshoe hares significantly decreased their use of harvested stands following CPPTM. Woodland caribou displayed a strong avoidance for CPPTM sites, while moose did not display any such aversion. CPPTM thus alters species assemblages, potentially reshaping trophic interactions with consequences for wildlife conservation. For example, moose did not avoid CPPTM cuts as much as they avoided stands that were harvested more intensively. Wolves that generally focus their hunting on moose might become attracted to CPPTM cuts, which could result in a higher concentration or even in a numerical response of wolves. At a regional scale, the outcome might lead to an increase in local predation risk for woodland caribou, a threatened species. While CPPTM could still be useful in an ecosystem-based management context, the authors study shows that the protection of small merchantable stems is often insufficient for the short-term maintenance of species composition which characterizes old-growth boreal forests.

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Chapter 5- Pinus canariensis Chr. Sm. Ex DC in Buch is an endemic conifer tree of the Canary Islands archipelago and its stands occupied much larger areas in the past. Plantation programs have been very common in the Canary Islands since the 1940s. The main objective of the plantations analyzed in this chapter is to restore the canarian pine forest which was heavily disturbed and eliminated during the last 5 centuries after the European colonization of the Canary Islands and reforestation with exotic species. There is not a general agreement in the plantation technology or the management of these plantations, probably due to the low number of studies related with the ecology of this forest stand. In this chapter, the authors will evaluate the ecological studies carried out with P. canariensis and analyze the management practices followed by the authorities. They will analyze the valuable aspects of this management and suggest, if possible, new alternatives for forest restoration. The impact of fire and the introduction of exotic species in the potential area for P. canariensis is also analyzed and evaluated. The authors finish the chapter with some concluding remarks based in the information provided by different sources found during this review. Chapter 6 - There is considerable interest in the idea of either ‗ancient‘ woods or ‗old growth‘ forest and in relating this to contemporary management. However, many such approaches to wooded landscapes are fundamentally flawed by the absence of an understanding of the historic context of the sites and of their ecologies. Recent research in the UK and across Europe seeks to address these issues and to provide a robust interrogation of forest and woodland dynamics that can better inform contemporary management and conservation. This chapter presents the evidence base for these assertions and a model into which historical and ecological information can be placed in order to critically assess issues of woodland antiquity and ecological continuity. The work seeks to synthesize problems of cultural severance in wooded landscapes, with the potential for both dynamic change in ecology and the contradictions of long-term spatial stability too. Progress so far represents an attempt to reconcile to ideas of scholars such as Fran Vera, of an open savannah-like primeval landscape in Europe, with mediaeval and contemporary woodland. A key idea to emerge from the studies is that of the Act of Commons or Statute of Merton representing watershed in the spatial fixing of ‗woods‘ in their landscape context. These named woods are today marked out and identifiable by socalled botanical indicators. Examples to test the ideas have been drawn from wooded landscapes in England in order to relate known historic time-lines to demonstrable changes in ecology, in woodland structure, and in pedology. The methodology described helps to facilitate an improved understanding of woodland landscape history but it also informs future management strategies for forest and woodland areas. Chapter 7- The chapter describes the ecological distribution of miombo woodlands, and gives an account of miombo woodland productivity under different land use systems based on the available data and literature. Additionally, the chapter shows that even with high levels of deforestation in the miombo ecoregion, degraded sites can still be managed for carbon sequestration through coppice or natural regeneration management as the regrowth stands have very high rates of photosynthetic process and therefore high uptake of carbon dioxide. By unravelling the carbon sequestration potential of miombo woodlands, and drawing lessons from other ecosystems where carbon trading projects have contributed to rural livelihoods, the chapter tries to show the appropriateness of the Payment for Environmental Services (PES) schemes as a strategy to bridge the perceived disconnect between forest conservation

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Preface

xi

and rural livelihoods in the miombo ecoregion. The chapter therefore, intends to show how carbon sequestration potential of the miombo woodlands may be exploited to enhance and improve natural woodland management and rural livelihoods in the ecoregion. Chapter 8- Native British bluebells (Hyacinthoides non-scripta) are an iconic plant in woodlands within the British Isles holding an estimated half the world‘s population of the species. A combination of habitat loss, plant removal, hybridisation with Spanish bluebell (H. hispanica) and grazing or trampling have resulted in British bluebells being less abundant and now protected as a species under Schedule 8 of the Wildlife and Countryside act 1981 (as amended 1998). The sale of wild British bluebells, whole plants, bulbs, seeds or derivatives, is only permitted through license. Deciduous woodlands are carpeted with British bluebells with thousands of individuals per square meter being possible. These carpets are considered an indicator for ancient woodlands as the spread of populations is slow, equalling the length of the scape, about 0.3 metre per year. Propagation is by seed and vegetative reproduction through axillary daughter bulbs. Seeds are heavy and not windborne. They fall close to the mother plant with germination rates of 80% recorded for covered and chilled seeds. Predation of seeds is rare. Herbivory by small mammals, muntjac and roe deer has been reported, however, damaging the emerging leaves has the most deleterious effect on the plant and can occur as a result of trampling. In general British bluebells are considered ―poisonous‖. As a society there needs to be a shift from crude oil to renewable resources to provide for the material needs, in other words shift from the hydrocarbon to the carbohydrate economy. Bluebell seeds contain iminosugars, which are highly biologically active. The mixture of iminosugars found within bluebells is the chemical reason for their toxicity. The main metabolite 2R, 3R, 4R, 5R,-2,5-dihydroxymethyl-3,4-dihydroxypyrrolidine (DMDP) has been patented as a nematocide and has been associated with, for example, anti-HIV, antituberculosis and anti-cancer activity. Early results for chemically breaking apart bluebell seeds showed that the seeds also contain oil and a carbohydrate fraction. Bluebell seeds could be providers of fine chemical which may give an economic incentive for their collection and in turn would also provide an incentive for conservation of existing populations. In addition, the seed collection for chemical fractionation could also supply seeds for new populations to be established. Bluebell seeds ripen gradually on the scape, making it impossible to remove all seeds and thus assuring that seeds still remain within the ecosystem. In summary, this chapter suggests to combine conservation with renewable resource use that goes beyond utilising woodland biomass for burning. Chapter 9- African forest provide the focus for a growing body of historical research. This chapter draws on economic and environmental history approaches in exploring the exploitation and conservation of woodland, respectively. The main focus of the investigation is the consumption–conservation relationship between pre-colonial African people and the forest zone, an interaction viewed by colonial foresters in Zimbabwe as wasteful and based on religious superstition. In spite of the open criticism of rapacious timber cutting by mining companies and poor farming techniques by settlers, colonial perceptions over time stressed the notion of ‗improvident Africans‘ as the prime cause of environmental destruction, in particular, deforestation and erosion. Within the African context, historical forest literature is bound to reject colonial misconceptions regarding the scope of indigenous woodland management. Customary forest practice in the Zambezi teak or Baikiea woodland points towards a better understanding on the subject, informed by a wide range of sources; oral

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tradition, missionary records, travel accounts and colonial documents. In reconstructing precolonial resource use from interviews and archival data, this chapter adopts a multi-source approach, while guarding against an overly romanticised view of indigenous practice. Teak or Kalahari sand forests were the largest commercial and most important indigenous hardwood forests in Zimbabwe, and indeed, in the whole of Southern Africa. Called gusu by the Ndebele people, they constituted an important natural resource in nineteenth-century North Western Matabeleland. Gusu directly and indirectly provided food, medicine, water, firewood, timber and ornaments. It enhanced soil fertility, sheltered game and was the site for sanctuaries and holy shrines. Various gusu inhabitants conserved woody species most significantly for food provision in an agriculturally precarious region, especially given the phenomena of unreliable rainfall, cyclical droughts and famine. Many nineteenth-century African communities would not have survived in an environment stripped of trees, bush and grass.

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Chapter 1

COLONIZATION OF POST-AGRICULTURAL BLACK ALDER (ALNUS GLUTINOSA (L.) GAERTN.) WOODS BY WOODLAND FLORA Anna Orczewska Department of Ecology, Faculty of Biology and Environmental Protection, University of Silesia, Katowice, Poland

ABSTRACT

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A survey on the colonization of the herb layer of post-agricultural black alder woods by woodland flora and on the edaphic, hydrological and light conditions responsible for the colonization mechanisms present in such woods was carried out in the Oleśnica Plain and Żmigród Valley (SW Poland) in the habitats of an oak-hornbeam community, alderash carrs and wet alder woods. In the 33 transects (80 m long by 4 m wide) consisting of 10 quadrats (16 m2 each) laid out perpendicularly across the ancient-recent border, data were collected on herb layer composition, chemical soil properties and illumination level. Furthermore, the groundwater level was recorded in piezometers. Migration rates were calculated for 51 woodland plant species. The mean migration rate for typical wet alder woods based on the maximum cover reached 1.20 m yr-1 and the one based on the farthest individual 1.60 m yr-1, for oak-hornbeam forests were 1.17 and 1.63 m yr-1 respectively and for alder-ash carrs 0.79 and 1.26 m yr-1. Among the woodland herbs recorded there was a group of species unable to migrate into recent alder woods and a group of slow-colonizing species (AAWS – ancient alder woodland species). On the other hand, many herbs were able to colonize recent woods very efficiently (OAWS – other ancient woodland species); some of them migrated at a pace exceeding 2 or 3 m yr1 . Thus, herbaceous woodland species differ in their dispersal potential. In wet and fertile recent forests adjacent to ancient source woodlands, herb layer recovery proceeds faster than in poorer and drier habitats. Forest age, pH, humus type and groundwater level were the variables having the greatest influence on the distribution pattern of woodland species in recent woods. A competitive species, vigorously growing in the herb layer of recent 

ul. Bankowa 9, 40-007 Katowice, Poland, telephone: +48 32 359 15 48. e-mail: [email protected].

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Anna Orczewska woods – Urtica dioica, avoided sites with a high level of groundwater combined with poor illumination. To create the best conditions that allow for effective forest recovery in alder woods, it is essential to maintain a high level of water and shade in the forest floor. This reduces the competitive exclusion of woodland flora by aggressive herbs and facilitates the immigration of forest species. In conclusion, forest management should give priority to either the restoration or maintenance of the natural water regime. If necessary natural water conditions should be restored prior to afforestation. In addition, in order to ensure the successful recovery of a herb layer rich in woodland flora, recent forests should be located in direct proximity to ancient forests.

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INTRODUCTION In current studies on the ecology and dynamics of forest communities, their biological diversity and the mechanisms behind these processes and patterns, knowledge of the history of forested areas has become essential. Therefore, temporal and quantitative changes in the proportion of different land-use forms in the landscape have recently become an important area of interest among woodland ecologists. Knowledge about the temporal aspects in forest cover is pivotal in understanding many phenomena observed in nature. Primeval and natural woodlands, i.e., those which date back to the very beginning of European forest formations after the last glaciation period, are characterized by the highest biological diversity, especially when they grow on fertile habitats. The continuous existence of such forests in the countryside should be confirmed by thorough palinological, archaeological and dendrochronological studies. In practice, however, such detailed evidence is often unavailable due to the relatively scarce grid of archaeological and paleobotanical study sites. Thus, the concept of so-called ancient woodlands, proposed by Peterken (1974; 1977) and Rackham (1980), has become widely applied by woodland ecologists in their biodiversity studies. Forests are regarded as ancient when they have been growing continuously for a long time. The threshold understood as the minimum age of a forest to be regarded as an ancient one depends on the availability of the oldest cartographic sources documenting woodlands‘ existence in the landscape. Thus, in the case of many countries in continental Europe, these are usually maps dating back to 200–250 years ago; 1770–1778 in Belgium (Bossuyt et al. 1999), 1765–1780 in many regions of Germany (Wulf 2004), 1779–1783 in some areas of southern Poland (Dzwonko and Loster 2001), 1750 in Scotland, but 1600 in England (Goldberg et al. 2007) (see Goldberg et al. 2007 for more examples of such maps). Hence, ancient woodlands include either remnants of primeval forests with their soils least modified by human activities (Peterken 1983) or of secondary woods that have existed in the landscape at least prior to the date of the oldest land-use maps available for the region under study. Thus, the concept of ancient forests does not place emphasis on the age of a stand but on the continuous existence of a forest habitat in the landscape for at least the last two centuries. Although this concept is a compromise used in situations where there is a lack of evidence of a centuries‘ long existence of forest cover in the countryside, it is also a valuable tool in ecologists‘ hands, since it allows for better explanations of the reasons for the current distribution of species observed in the landscape. Most studies in this respect concentrate on the mechanisms responsible for the contemporary species composition recorded in forests that differ in their management history, i.e., ancient vs. recent ones. Apart from floristic comparative investigations, those aimed at the estimation of the pace of recolonization of the

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Colonization of Post-Agricultural Black Alder (Alnus Glutinosa (L.) Gaertn.)…

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herbaceous layer by true forest species have also been carried out. Since situations where there is a spatially direct proximity of ancient and post-agricultural woods in similar habitats are rather scarce, surveys on species‘ recovery rates have not been very common. Investigations on the migration of woodland flora into recent woods have concentrated on recent stands on habitats of either mesotrophic deciduous forests, on pine woods on habitats of broadleaved forests, or on very poor sandy soils (Dzwonko and Loster 1992; Dzwonko 1993; Dzwonko and Gawroński 1994; Matlack 1994; Brunet and von Oheimb 1998a, b; Bossuyt et al. 1999; Bossuyt and Hermy 2000; Brunet et al. 2000; Dzwonko 2001a, b; Orczewska and Fernes 2011, in press). The data presented in this chapter are the results of studies on the colonization of the herb layer of post-agricultural black alder woods, which represent very fertile and wet forest types, by woodland herbs. Furthermore, the research also focused on the edaphic, hydrological and light conditions responsible for the colonization mechanisms present in such woods.

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THE HISTORY OF FORESTS OF SOUTHWESTERN POLAND The growing awareness of the importance of old maps in biodiversity studies and in surveys on vegetation changes over time has resulted in many cartographical surveys aimed at a reconstruction of history of the countryside, especially forests (for example Cousins 2001; Moszkowicz 2005, Majchrowska and Woziwoda 2009, Wulf et al. 2009). The Oleśnica Plain (510 04‘ N; 170 43‘ E) (part of the Opole Silesia Region) and the Żmigród and Milicz Valleys (510 28‘ N; 160 54‘ E) (belonging to Lower Silesia), two regions of southwestern Poland, are among those areas for which the age and origin of forests existing in the countryside have been thoroughly recognized and described. In the case of this part of the country, the year 1780 should be treated as a threshold date for forests to be regarded as ancient and the Schmettausche map as the first, reliable cartographic document illustrating the former distribution of forests in the countryside. Thus, ancient forests in those regions are at least 230 years old. According to the results of those reconstructions (Orczewska 2009a), it is known that the forested cover of the Oleśnica Plain and the Żmigród and Milicz Valleys reaches 34% and 41%, respectively, exceeding the average for Poland, which was estimated at 28.8% in 2005. Over 77% and 70%, respectively, of today‘s woods in those regions are ancient ones (present on the Schmettausche map, dated 1780, as was already mentioned), which is a high share, especially when compared with some regions in Europe or eastern North America where forests of post-agricultural origin represent as much as 80% of current forest cover (Flinn and Vellend 2005). In the surveyed regions, ancient forests coexist with woods that have been growing for a much shorter period of time, i.e., with those existing since 1880 and with much younger ones that have been growing for only several decades. Recent woods are of a post-agricultural origin since they were planted either on former arable land or in abandoned meadows and pastures (Orczewska 2009a). The history of the environment of the Oleśnica Plain and Żmigród Valley was described in more detail in a separate paper (Orczewska 2009a), thus only the most important facts regarding how the proportion of different land-use forms have changed over time are presented here. Special emphasis was placed on the development of human-induced transformations on the forest cover in these regions. After the last glacial age and the full

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development of vegetation cover, deciduous woodlands dominated the landscape of this part of Europe (Hensel 1988; Ralska-Jasiewiczowa et al. 2003). Neither Mesolithic groups of hunter-gatherers, the first inhabitants of those regions, nor the successive Neolithic tribes, had a negative impact on woodlands (Kulczycka-Leciejewiczowa 1993). A predominant part of these areas was covered by extensive, untouched broadleaved and mixed forests, which were difficult to access mainly due to their location in the marshy, waterlogged valleys of the tributaries of the main river – the Oder (Żabko-Potopowicz 1959). In subsequent times, although a gradual deforestation associated with human settlement occurred, many historic records confirm that inaccessible forests in the Oder valley made the deforestation process more difficult than in other regions. The first historical mentions of forest clearance and burning associated with settlement and agriculture dated from the thirteenth and fourteenth centuries. The deforestation process continued over the next few centuries until the first part of the nineteenth century when it slowed down. At that time the knowledge of forest management techniques began to increase, especially those concentrating on continuous forest management and maintenance of the forest habitat. It was an important period when modern forestry started, since at that time forest management policy and planning regime were also implemented. This therefore resulted in reforestation in accordance with the conditions of the natural habitat (Nyrek 1992). Such historical records mentioning the changing attitude to forests, which at first led to a decrease and then a successive increase in woodland cover, were also confirmed in cartographic studies. According to these the forest cover in the Oleśnica Plain and Żmigród Valley in the eighteenth century reached 54% and 56%, respectively. In 1880 it dropped to 29% and 33% and then it increased to 34% and 41% in 1980 (Orczewska 2009a). Thus, the current forested landscape of those regions, like most of Europe or eastern North America (Flinn and Vellend 2005; Matlack and Leu 2007), is a mosaic of woodlands of a different age and origin. Although ancient woods predominate, there is also a high share of woodlands which were present in 1780 and in 1980, some present continuously since 1880 and eventually the youngest ones, i.e. present exclusively in 1980 or even younger. The presence of the youngest generation of recent woods of a post-agricultural origin is strictly related with the recent history of these regions after the 2nd World War. Political changes that took place after 1945, which involved alterations in the alignment of the Polish-German border, led to a mass migration of Germans followed by the abandonment of agricultural land as a consequence of a decrease in the human population. Thus, the period after the 2nd World War was a time of huge changes in land-use management, especially afforestation (Orczewska 2009a). At that time almost one million hectares of abandoned agricultural land in Poland were converted into forests (Tuszyński 1990). The second wave of afforestation on a bigger scale in western Poland was recorded in the last two to three decades. It was of economic origin since there was pressure to convert those agricultural sites whose management was unprofitable due to their poor, infertile, or over-damp soils into forests. Scots pine (Pinus sylvestris L.) is the main tree species used for afforestation in poor and dry sites (previously managed as arable fields or pastures), whereas on sites with a high level of groundwater (cultivated as damp meadows, Figure 1), too damp to be easily cultivated without permanent amelioration and drainage, black alder (Alnus glutinosa (L.) Gaertn.) is planted.

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Photo by Anna Orczewska. Figure 1. An abandoned meadow prepared for planting black alder (Namysłów forest district, Minkowskie, Oleśnica Plain).

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Black alder belongs to the group of trees that have a great influence on the habitats which they occupy. It has some very specific features, which make it a very good pioneer species. The most important ones are listed below. 1. Alder, as a species of an open management type of nutrition, does not withdraw its nutrients from leaves before leaf fall (McVean 1953). Therefore, much foliar nitrogen is retained, contributing to a substantial increase in its content in the litter (Karkanis 1975; Zimka and Stachurski 1976). 2. Leaf litter of alders decomposes very quickly (Dilly and Munch 1996). Decomposition rate k for alder leaves was estimated at -0.908 yr-1, which means a mass loss of 66% during a period of 16.5 months (Pereira et al. 1998). 3. Unlike most European trees, black alder tolerates heavy, waterlogged soils (McVean 1953). 4. Due to its symbiosis with actinobacteria – Frankia alnii, which assimilates atmospheric nitrogen, the alder contributes to the enrichment of the soil with this nutrient by approximately (10) 50-300 kg ha-1 yr-1 (Schaede 1967; Pancer-Kotejowa and Zarzycki 1980; Binkley 1986; Sprent and Sprent 1990). 5. Alders form a symbiosis with ectomycorrhizal fungi, which provide them with nutrients, especially phosphorus (Laganis 2007), necessary for nitrogen fixation and the growth of nodules (Roy et al. 2007). Although maintenance of symbiosis with bacteria and fungi requires a lot of energy from alders (Roy et al. 2007), it also gives them the potential to colonize very poor soils. This in turn makes alders a key element in the cycling of N, P, C and other nutrients (Compton and Cole 1998; Laganis 2007; Roy et al. 2007), thereby contributing to an increase in their content in soils (Binkley and Giardina 1998). For detailed reviews of black alder biology, ecology and the types of forest ecosystems which it is a component of, see for example McVean (1953), Solińska-Górnicka (1987), Prieditis (1997), Sienkiewicz et al. (2001), and Laganis (2007).

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Photo by Anna Orczewska

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Figure 2. A fish pond which has existed in the landscape at least since the Schmettausche map dated 1780 (Oborniki Śląskie forest district, Prusice, Żmigród Valley).

Photo by Anna Orczewska. Figure 3. Typical landscape of the Żmigród Valley dominated by agricultural and forested areas.

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Photo by Anna Orczewska

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Figure 4. Typical landscape of the Żmigród Valley with a mosaic of agricultural land and forests (Żmigród forest district, Niezgoda, Żmigród Valley).

Photo by Anna Orczewska. Figure 5. Ancient forest representing the wettest type of the Tilio cordatae-Carpinetum betuli community, rich in ancient woodland species (Namysłów forest district, Minkowskie, Oleśnica Plain).

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CHARACTERISTICS OF ALDER-DOMINATED WOODLANDS Alnus glutinosa-dominated forests are on the decline in many areas of Europe as a result of forest drainage and agricultural activities (Dembek et al. 2002; Kowalska 2009), mainly in the western part of the continent. However, in the east, especially in the countries around the Baltic region like Estonia, Latvia, Lithuania and also in Russia, Byelorussia and Ukraine, a relatively large part of forested landscape is still occupied by alder forests that have enjoyed a long continuity (Prieditis 1997). Although the proportion of such woodlands in Poland is not large, there are still some regions of the country, including those described in this chapter, where alder woods occupy a relatively large part of the forests. Today‘s landscape of the Oleśnica Plain and Żmigród Valley is free of any evidence of industry and is occupied by agricultural land, forested areas and fish ponds (Figures 2-4). The forests here are dominated by meso- and eutrophic deciduous woodlands composed mainly of Quercus robur, Tilia cordata, Carpinus betulus with an admixture of many other tree species, like Fraxinus excelsior, Fagus sylvatica, Acer pseudoplatanus, Ulmus ssp. and Alnus glutinosa. Less fertile, sandy soils are covered by coniferous and mixed stands composed of Pinus sylvestris and Quercus robur (Matuszkiewicz 1993). Among deciduous forests those with stands dominated by black alder have a special position and relatively high proportion compared to other regions of Poland, mainly due to the dense network of tributaries of the Oder river that provide proper habitats for these species. Forests whose main component is black alder are associated with sites with a high level of groundwater, i. e. places which are damp and wet. Therefore such communities are of an azonal character and include riverside carrs (with black alder, ash and elm) and typical wet alder woods (usually with pure black alder stands). Finally, black alder can be also found in the dampest and most fertile types of oak-hornbeam forests, which represent a zonal type of vegetation in this part of Europe (Figure 5). According to the phytosociological classification by Braun-Blanquet (Matuszkiewicz 2001), the communities which were studied belong to the following syntaxa: 



Class: Alnetea glutinosae Br.-Bl. et R. Tx. 1943 Order: Alnetalia glutinosae R. Tx. 1937 Alliance: Alnion glutinosae (Malc. 1929) Meijer Drees 1936 Association: Ribeso nigri-Alnetum Sol.-Górn. (1975) 1987 Class: Querco-Fagetea Br.-Bl. et Vlieg. 1937 Order: Fagetalia sylvaticae Pawł. in Pawł., Sokoł. et Wall. 1928 Alliance: Alno-Ulmion Br.-Bl. et R. Tx. 1943 Suballiance: Alnenion glutinoso-incanae Oberd. 1953 Association: Fraxino-Alnetum W. Mat. 1952 Alliance: Carpinion betuli Issl. 1931 em. Oberd. 1953 Association: Tilio cordatae-Carpinetum betuli Tracz. 1962 Subassociation: T-C stachyetosum Association: Galio sylvatici-Carpinetum betuli Oberd. 1957 Subassociation: G-C stachyetosum

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ones. Riverside carrs (including alder-ash carr community) and the wettest types of oakhornbeam community belong to this class. The Alnetea glutinosae class contains azonal wet alder woods.

Photo by Anna Orczewska.

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Figure 6. Ancient forest representing a Fraxino-Alnetum community (alder-ash carr) (Namysłów forest district, Minkowskie, Oleśnica Plain).

Photo by Anna Orczewska. Figure 7. Ribeso nigri-Alnetum in early spring with its typical physiognomy with hummocks and hollows (Żmigród forest district, Niezgoda, Żmigród Valley).

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Photo by Anna Orczewska

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Figure 8. Ribeso nigri-Alnetum community (typical alder wood) with its mosaic structure of the herb layer (Oborniki Śląskie forest district, Prusice, Żmigród Valley).

Although both the Fraxino-Alnetum (Figure 6) and Ribeso nigri-Alnetum forests are azonal and associated with habitats of high humidity, they differ distinctively in their water management type. Riverside carrs are temporarily flooded with moving subsoil streams or brooks without an evident bed. Thus, their water regime is based on lateral movement without a tendency to stagnate for longer periods. Typical wet alder woods are enriched by water originating from rainfall or by underground water outflow, thus with no tendency for lateral movement but rather with a tendency to stagnate. Until late spring or even early summer wet alder woods remain flooded with snowmelt water or water from rainfall. Thus, this leads to the development of a very peculiar physiognomy in this community with hummocks formed by elevated alder roots around their trunks and with hollows surrounding the hummocks filled with water in spring and after heavy rains (Figure 7). These two totally different microhabitats, which create a mosaic in a forest, contain a specific flora. Hummocks provide a habitat for woodland species which do not tolerate soils permanently saturated with water, whereas plants characteristic for marshes and swamps occupy the spaces in the hollows (Figure 8) (Solińska-Górnicka 1987; Prieditis 1997; Matuszkiewicz 2001). Such a water regime is characteristic for forest sites which are free from the heavy impact of humans, i.e. those which escaped from drainage, usually either undertaken intentionally in these types of forests or related to protecting the neighbouring meadows from inundations. Considerable changes in hydrological regimes over the centuries led to the disappearance of wet, alder-dominated forests from many parts of Europe including Poland (Sienkiewicz et al. 2001). Furthermore, those that remain in the landscape have also suffered the detrimental effects of past drainage. However, the unique value of waterlogged, swampy black alder forests was recognized in Poland recently and such ecosystems were included in the European network of areas of great ecological importance – the Nature 2000 network. Wet, waterlogged alder forests located in the Żmigród and Milicz Valleys also received the status of areas of great international ecological value and were included into the international Nature 2000 system as a part of the ―Dolina Baryczy‖ (―The River Barycz Valley) region.

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a.

b.

c. Figure 9. Continued Woodlands : Ecology, Management and Conservation, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

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24

d. Figure 9. Changes in the groundwater level within the hydrological year in a) oak-hornbeam community; b) alder-ash carr; c), d) typical wet alder wood.

Table 1. Ranges of mean spring and annual levels of groundwater in the communities studied

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Spring level (mean from three spring months) Average annual level

Water depth [cm] Oak-hornbeam From –16 to –70

Alder-ash carrs From –1 to –54

Wet alder woods From +18 to –29

From –43 to –79

From –24 to –108

From +3 to –83

Sign ‗+‘ refers to water stagnating on the surface; ‗–‘ indicates the level of water below the ground.

The above-described gradient in the water regime of the alder-dominated forests surveyed was confirmed in the course of the observations of the groundwater level. In over 30 study sites piezometers (ø = 5 cm; h = 50-200 cm) were installed that recorded the changes in the water level at monthly intervals over the whole hydrological year. The average spring and annual levels of water are presented in Table 1, and Figures 9a-d illustrate water oscillations over the entire hydrological year recorded for some sample sites located in the habitats of a oak-hornbeam community, an alder-ash carr and a typical wet alder wood. This data confirms that oak-hornbeam communities have a good water supply with a lack of extremes; neither inundations nor periods with conditions that are too dry occur there (Figure 9a). Alder-ash carrs have an intermediate position since in spring there is no stagnating water but the groundwater table is close to the surface, whereas in other periods of the year it may lie deeper (Figure 9b). Wet alder woods occupy the most waterlogged habitats with a high fluctuation in water level with stagnant water persisting above the ground surface at least in hollows in the winter and spring, but dropping off in the summer and autumn, in extreme cases to even –83 cm (Figures 9a,b). These fluctuations in the water level are in accordance with the ranges for alder-dominated wetland forests reported by other authors (from Germany and Latvia) and cited by Prieditis (1997). Great oscillations in the water table, which may

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range from situations where water reaches the surface to cases where it is 3 m below ground level, were also confirmed by Sienkiewicz et al. (2001) in floodplain forests. According to these authors in alder-ash carrs the groundwater level in summer is at a depth of 0.3-0.6 m. Such specific water regimes of the surveyed sites (with stagnating water leading to spatial and temporal changes of aerobic and anaerobic conditions) are associated with the soil types which alder forests occupy. Most woods, both ancient and recent ones, develop on humic gleysols and ferri-umbric gleysols (according to the WRB nomenclature, 1998). Thus, mesoand eutrophic soils on moist, wet or periodically waterlogged sites with wet or moist mull humus predominate among them (Orczewska 2009c) (Tables 2-3). Table 2. Soil types present in ancient and post-agricultural alder woodlands (nomenclature after WRB 1998) Ancient woodlands M o i s t u r e

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s c a l e

Fertility scale Mesotrophic

Eutrophic

Moist

Humic Gleysols

Humic Gleysols Stagni-Eutric Gleysols Stagni-Haplic/Stagni-Umbric Gleysols Mollic Gleysols Saprihistic Gleysols Haplic Gleysols Ferri-Umbric Gleysols

Wet

Humic Gleysols

Humic Gleysols Sapric Histosols

Very wet or submerged

Humic Gleysols

Post-agricultural woodlands M o i s t u r e s c a l e

Mesotrophic

Eutrophic

Moist

Luvic Gleysols

Humic Gleysols Stagni-Eutric Gleysols Stagni-Haplic/Stagni-Umbric Gleysols Umbric Gleysols Mollic Gleysols Ferri-Umbric Gleysols

Wet

Humic Gleysols

Humic Gleysols Ferri-Umbric Gleysols Sapric Histosols

Very wet or submerged

Humic Gleysols

Eutrophic – exchangeable cations >50%; mesotrophic – exchangeable cations = 20-50%.

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Table 3. Chemical characteristics of the soils from four woodlands that differ in their length of existence in the landscape – ancient (sites 2, 4) and recent (sites 1, 3) Site

Soil type

Horizon

Depth [cm]

pHKCl

Corg [%]

Ntot [%]

C/N

1 1 1 1 2 2 2 3 3 3 3 4 4 4 4

GLum-fr

Aep Box Gor Gr AOM Gor Gr AOMp A/Cg Gor IIGr Ae Gor Gor IIGr

23 38 95 150 25 85 150 25 48 70 170 24 46 95 170

3.86 5.62 6.36 4.31 4.91 3.85 4.09 6.13 6.55 6.65 7.11 5.83 7.62 7.01 6.80

6.23 0.05 0.02 n.m. 17.98 0.04 n.m. 2.90 0.42 0.04 n.m. 3.24 0.08 0.062 n.m.

0.58 0.01 0.01 n.m. 1.60 0.01 n.m. 0.31 0.04 0.00 n.m. 0.27 0.01 0.004 n.m.

10.7 8.1 1.8 n.m. 11.2 2.5 n.m. 9.4 10.2

GLhu

GLhu

GLhu

n.m. 11.8 9.3 15.5 n.m.

Mg+2 [cmol kg-1] 0.60 0.67 0.03 0.06 2.90 0.11 0.07 0.79 0.29 0.17 1.62 0.73 0.41 0.22 1.31

GLum-fr – Ferri-Umbric Gleysols; GLhu – Humic Gleysols; n. m. – not measurable.

Ca+2 [cmol kg-1] 17.52 3.16 1.02 1.69 50.90 1.42 1.17 18.25 4.93 2.76 26.26 22.92 5.76 2.41 18.66

Na+ [cmol kg-1] 0.07 0.03 0.04 0.02 0.39 0.03 0.03 0.18 0.07 0.05 0.19 0.15 0.08 0.03 0.12

K+ [cmol kg-1] 0.14 0.01 0.01 0.01 0.36 0.02 0.02 0.02 0.01 0.01 0.27 0.03 0.03 0.03 0.24

P2O5 [mg kg-1]

MgO [mg kg-1]

K2O [mg kg-1]

231.00 137.00 11.90 1.12 198.00 121.00 0.63 8.51 7.67 5.81 16.00 15.20 19.20 31.00 22.00

72.33 52.54 50.50 39.56 277.72 51.66 47.59 165.40 80.08 64.07 198.54 133.88 94.04 66.23 186.42

46.08 16.00 14.81 14.52 136.44 20.22 15.50 15.12 12.94 17.06 120.77 22.10 22.35 24.25 118.00

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Table 4. Total number of species and ancient woodland species in ancient and recent alder woods Oak-hornbeam community (Tilio-Carpinetum, Galio-Carpinetum) Total number of species Number of ancient woodland species Forest type Mean (±SD) Min Max Median XMean (±SD) Min Max Median Ancient (N=43) 31.8 (±10.7) 12 52 32 14.1(±5.4) **** 5 23 12 Recent (N=66) 28.7 (±9.3) 8 47 28 7.5 (±4.3) 1 20 7 Alder-ash carr (Fraxino-Alnetum) Ancient (N=48) 29.4 (±10.1) 11 47 32 13.4 (±5.1) **** 6 24 14 Recent (N=72) 29.3 (±10.8) 10 54 28.5 8.9 (±4.4) 1 20 8 Typical wet alder woods (Ribeso nigri-Alnetum) Ancient (N=43) Recent (N=72)

32.3 (±8.0) 31.7 (±7.7)

17 19

53 50

28.5 31

11.2 (±3.8) **** 8.3 (±0.0)

5 1

21 15

11 8

X

(data from Orczewska 2009c); N = number of sample plots. **** Significance level according to t-test, p < 0.00001; * Significance level according to the t-test, 0.01 < p < 0.05.

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RECOVERY OF HERBACEOUS LAYER IN RECENT ALDER WOODS Although the share of Alnus glutinosa-dominated woodlands (pure alder stands or mixed with other species, mainly ash) in the total forest cover in Poland reaches only ca. 2.6% (Lasy Państwowe w liczbach 2009), these forests are very rich in woodland herbs. The mean number (and median) of species recorded in the herb layer studied in plots of 16 m2 in the ancient woods varied between 29 to 32 with the maximum number recorded exceeding 50 species (Table 4). Similar species richness was observed in the recent woodland plots. However, plots from the ancient and recent woods differed in the number of ancient woodland species, which were significantly higher in ancient woodlands regardless of habitat type (Table 4). Thus, the herb layer composition in ancient and recent woodlands differed. These alterations had a purely qualitative character since they were expressed by a higher share of woodland species in ancient rather than in recent woods. Similar trends were recorded previously by other authors (e. g. Matlack 1994; Brunet and von Oheimb 1998b; see Flinn and Vellend 2005 for more reference). The above results on species richness confirm that alder wetland forests that occupy the most eutrophic habitats belong to the communities that maintain the greatest biodiversity among the European temperate woodlands (Prieditis 1997; Matuszkiewicz 2001). Furthermore, many species, especially of the alder-ash carr community, as emphasized by Prieditis (1997), are indicators of long-continuity forests. All investigations focusing on the process of natural recovery of true woodland species in recent forests in landscapes with highly fragmented forest cover indicate that spatial isolation from ancient woods limits colonization. In such situations this process may last for many centuries (Peterken 1977; Peterken and Game 1984; Faliński 1986; Matlack 1994) and the herb layer of recent woods located far from colonization sources may never reach a biodiversity and composition similar to ancient woods. Only the direct proximity to ancient woodlands enhances (allows for) the effective colonization of post-agricultural woods by

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woodland species (Peterken and Game 1984; Dzwonko and Loster 1992; Dzwonko 1993; Dzwonko and Gawroński 1994; Matlack 1994; Brunet and von Oheimb 1998a, b; Bossuyt et al. 1999; Bossuyt and Hermy 2000; Dzwonko 2001a, b). The existence of such situations in nature where recent black alder stands border ancient source woodlands rich in species provides a great opportunity to study the colonization process of woodland herbs in these eutrophic habitats.

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Field Methods and Data Analyses Much of the data presented in this review refers to already published results. Also the methods used in the studies on the migration rates of woodland flora from ancient into adjoining recent alder woods were described in detail in other papers (Orczewska 2009b; 2010). Thus, only the most important information is given here to allow the reader to become familiar with how the data were obtained. In order to study the colonization process of herbaceous species, 33 transects (11 in the habitat of Tilio-Carpinetum or Galio-Carpinetum, 12 in Fraxino-Alnetum and 10 in Ribeso nigri-Alnetum) laid out perpendicularly across the border between ancient and recent woods were set up. Recent alder stands differed in age, which ranged from 6 to 69 years (median = 27 yrs). Transects were approximately 80 meters long and consisted of four 4x4m plots located in ancient woods and six plots in the recent woods. Sample quadrats were laid out at intervals of 4 m. Thus, each transect was 28 meters in length in the ancient part and 44 in the recent one. Additionally, if the boundary between the woods was wider than 4 m, additional quadrats adjoining the edge of the ancient wood were sampled. In each sample plot (131 in ancient woods, 198 in recent stands and 34 in the boundary between them) the percentage cover of all herb layer species (1%, 5%, 10% and then at 10% intervals) was recorded twice per year; in April (to focus on the early spring herbs) and in June. The survey on herb layer composition was accompanied by investigations of the abiotic factors, including edaphic and light conditions in each plot of a transect. Therefore, soil samples were collected in order to determine the chemical properties (Lityński et al. 1976). Moreover, the canopy cover, using a densiometer and the illumination level (PAR – photosynthetically active radiation) in the ground floor were measured. For more details on the methods used in estimating habitat conditions see Orczewska (2009c). Matlack‘s (1994) method, successively used by other authors, i.e. Brunet and von Oheimb (1998b), Bossuyt et al. (1999), Dzwonko (2001b), and Wulf and Heinken (2008), was used to calculate the colonization rates of woodland species. Thus, both, the rate based on the occurrence of the farthest individual and on the most distant occurrence of the maximum cover of species were calculated. The migration rates were calculated for all of the species which were significantly more frequent in ancient woods (according to the Fisher exact probability test) and for all other species which are regarded as ancient woodland indicator species for Poland (according to Dzwonko and Loster 2001), even though some of them did not occur significantly more often in ancient alder woods. Since the recent alder stands differed in age, again following Matlack‘s (1994) method, the standardized distances counted by dividing the distance of each plot from the ancient forest by the age of the recent forest were taken into account. This was to allow for comparisons of migration rates regardless of the age of the recent woodland. The rates were calculated for each transect where a species was recorded and then the mean rates were calculated for a species separately in each of the

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three habitats investigated. For more details on the methods of calculating migration rates in alder woodlands see Orczewska (2009b). Alder forests were situated within big forest complexes where they occupied scattered localities associated with a high level of water. The patches of ancient alder woods located within these large forests ranged in size from 0.73 ha to 15.54 ha, whereas recent woods varied in size from 0.72 ha to 8.6 ha. As the next step in the floristic studies the entire forest patches where transects were situated were surveyed in order to make complete lists of the species present in their herb layer. Then, based on their frequency woodland species were divided into two groups. The first included the species which were more frequent in the patches of ancient alder woods and belonged to the list of ancient woodland indicators for Poland (AAWS = ancient alder woodland species). The second group included those ancient woodland indicators for Poland which did not occur significantly more frequently in ancient alder woods (OAWS = other ancient woodland species). For more details see Orczewska (2010).

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RESULTS In total, there were 51 species for which the migration rates were calculated; 44 species in Tilio-Carpinetum and Fraxino-Alnetum habitat and 37 herbs in Ribeso nigri-Alnetum forest types (Table 5). Woodland species differed in their behavior along the ancient/recent ecotones, thus they could be divided into a few groups. Among them 11 species did not migrate at all in any of the habitats investigated (not included in Table 5). This group included: Carex sylvatica, Gagea lutea, Hepatica nobilis, Luzula pilosa, Melica nutans, M. uniflora, Pulmonaria obscura, Convallaria majalis, Daphne mezereum, Sanicula europaea and Mycelis muralis. All of the species listed belong to ancient woodland indicators for Poland. Over 60% of them are myrmecochorous, therefore possessing a very poor dispersal capacity, whereas the remaining four herbs have effective (epizoochorous, endozoochorous and anemochorous) modes of seed dispersal. As was already mentioned, the group of species which are unable to migrate into recent alder woods includes C. majalis, an endozoochorous herb. According to Bossuyt et al. (1999), as an herbaceous plant of a small size, C. majalis has a reduced migration capacity and its colonization pattern is more like that of a barochorous species. Even its ability to spread vegetatively through its rhizomes did not help it to colonize the recent woods. A similar observation was also reported by Brunet and von Oheimb (1998b), who noted a sporadic lack of migration of C. majalis in broadleaved forests in southern Sweden. Similarly, it did not migrate into pine woods on poor sandy soils in southern Poland (Orczewska and Fernes 2011, in press), whereas it managed to colonize recent pine woods in southern Poland (Dzwonko 2001b).

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Table 5. Mean migration rates [m yr-1] of the species in the recent black alder woods adjacent to communities of oak-hornbeam, alderash carrs and wet alder woods based on the farthest individuals (FI) and the farthest occurrence of maximum cover (MC)

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Species

Dispersal mode

Adoxa moschatellina Aegopodium podagraria Ajuga reptans Anemone nemorosa Anemone ranunculoides Anthriscus nitida Asarum europaeum Athyrium filix-femina Brachypodium sylvaticum Bidens frondosa Calamagrostis canescens Carex acutiformis Carex elongata Chrysosplenium alternifolium Circaea lutetiana Dryopteris carthusiana Equisetum sylvaticum Eupatorium cannabinum Festuca gigantea Ficaria verna Galeobdolon luteum Galium palustre Geranium robertianum Geum urbanum Glechoma hederacea

Endo Baro Myrm Myrm Myrm Epi Myrm An1 Epi Epi Hy/An1 Hy Hy Hy Epi An1 An1 An2 Epi Myrm Myrm Hy Ballo Epi Blasto

Humulus lupulus Impatiens noli-tangere Impatiens parviflora Iris pseudacorus

An2 Ballo Ballo Hy

Oak-hornbeam forest type

Alder-ash carr

Typical wet alderwood

B and O 1998b

B et al. 1999

MC 0.55 0.07 1.57 0.95 3.45 1.19

MC 0.49 0.34 1.25 0.72 2.29 0.83 0.01 / 0.15 / 0.70 3.54 0.54 1.85 0.30 0.00 1.71 1.06 0.85 0.01

MC 0.01 / 1.55 0.60 / 0.17 / 0.85 1.29 1.74 0.98 0.01 2.94 0.88 / 1.97 1.94 0.69 / 1.22 1.14 1.47 2.04

FI 1.53 / 2.17 1.63 / 1.03 / 1.61 1.05 1.22 1.24 1.45 2.05 1.63 / 2.26 1.95 2.05 / 2.22 1.46 1.90 1.77

FI

1.69 0.85 0.85

FI 1.49 0.94 3.07 2.16 1.68 1.25 0.44 / 1.03 / 1.72 4.11 1.19 1.39 1.59 0.00 1.94 1.07 1.96 0.34 2.17 1.00 1.51 1.37

MC 0.42

0.67 2.00 0.80 0.66 0.31 0.90 1.01 2.30

FI 1.19 0.64 1.90 1.39 1.73 1.63 0.01 1.31 1.21 / / / / 1.92 1.86 1.48 0.14 3.92 1.31 2.99 0.20 1.82 0.40 2.21 2.22

1.34 1.46 2.05 /

0.93 1.91 2.52 /

0.55 0.80 0.24 0.84

0.76 1.74 1.21 0.69

2.89 0.81 0.94 0.83

2.17 1.86 1.58 1.29

0.01 / / / / 1.16 0.70

FI 0.59

0.20 0.11

0.85 0.72

0.25 0.77

0.42 0.93

0.43 0.38

0.73 0.44

0.51 0.28 0.22

0.75 0.63 0.50

0.55

1.15

D 2001a

D 2001b

FI

FI

0.95 2.09

0.00 0.00 0.21

1.57

>0.53

2.28

0.27 >0.53

Table 5 (Continued)

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Species

Lamium maculatum Lycopus europaeus Lysimachia vulgaris Maianthemum bifolium Mercurialis perennis Milium effusum Moehringia trinervia Myosoton aquaticum Oxalis acetosella Paris quadrifolia Ranunculus auricomus Rubus caesius Rubus hirtus Rubus idaeus Rumex sanguineus Scrophularia nodosa Solanum dulcamara Stachys palustris Stachys sylvatica Stellaria holostea Stellaria nemorum Viola reichenbachiana

Dispersal mode

Myrm Hy Hy/An2 Endo Myrm An2 Myrm Hy Ballo Endo Myrm Endo Endo Endo An2 An1 Endo Epi Epi Baro Hy Myrm

Oak-hornbeam forest type

Alder-ash carr

Typical wet alderwood

B and O 1998b

B et al. 1999

MC / 0.44 0.14

MC 0.49 0.91

MC / 1.26 0.74 / 0.01 0.34 1.51 1.91 0.65

MC

FI

FI

0.33 0.36 0.40

0.34 0.73 0.55

0.01 0.81 0.82 1.84 0.65 2.92 0.97 0.54 1.33 2.38 0.89 / 3.59 1.06 0.75

FI / 0.54 0.84 1.33 0.67 1.05 2.24 1.52 0.81 1.41 2.43 1.96 1.37 2.29 3.24 1.78 1.48 / 1.84 0.48 1.63 0.01

/ 0.38 0.76 0.65 0.75 0.31 1.40 0.10 / 1.18 0.92 / 1.13 / 0.08 0.20 0.61 0.00

FI 0.79 1.02 1.09 / 0.41 0.89 1.07 1.02 0.66 0.86 2.04 1.07 / 1.42 1.41 / 1.39 / 0.91 0.93 0.81 0.30

2.70 / / 1.34 1.37 2.64 0.94 0.47 0.45 /

FI / 1.62 2.00 / 0.22 0.80 1.95 1.24 1.09 0.28 2.85 / / 2.03 1.16 1.61 1.88 0.63 1.27 /

D 2001a

D 2001b

FI

FI

0.05 1.29

0.26 0.33

>0.53

0.43 0.37

>0.53

0.58

0.58

0.30 0.46 0.12 0.44

0.30 0.91 0.23 0.67

1.00

0.24

For comparison, the mean migration rates of species in recent deciduous woodlands in southern Sweden (Brunet and von Oheimb 1998b––B and O 1998b), central Belgium (Bossuyt et al. 1999 – B et al. 1999), southern Poland (Dzwonko 2001a – D 2001a) and in secondary pinewoods in southern Poland (Dzwonko 2001b – D 2001b) are given. Dispersal modes are given after Van der Pijl (1982): Endo – endozoochores; Epi – epizoochores; Myrm – myrmecochores; Hy – hydrochores; Baro – barochores; Ballo – ballochores; Blasto – blastochores; An1 – unwinged anemochores with small diaspores; An2 – anemochores with diaspores with pappus or wings. Bold fonts signify ancient woodland indicator species (after Dzwonko and Loster 2001), the ‗/‘ sign indicates that speci –‘ mark means that a species was absent in a particular habitat. Original source: Plant Ecology, 204, 2009, 83-96. Migration of herbaceous woodland flora into post-agricultural black alder woods planted on wet and fertile habitats in south western Poland. Orczewska A. Table 1. With the kind permission of Springer Science and Business Media

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32

Table 6. Mean (± standard deviation) cover of selected ancient woodland species in ancient and recent alder-dominated woods (calculations based on the data from transect plots)

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Species a) Convallaria majalis Gagea lutea Hepatica nobilis Melica nutans Asarum europaeum Viola reichenbachiana Galeobdolon luteum Equisetum sylvaticum b) Brachypodium sylvaticum Anemone nemorosa Chrysosplenium alternifolium Circaea lutetiana Rumex sanguineus c) Ficaria verna Impatiens noli-tangere Ranunculus auricomus Impatiens parviflora* Glechoma hederacea Moehringia trinervia Ajuga reptans

Mean cover (± SD) ancient recent

Frequency ancient recent

3.0 (±2.8) 1.9 (±1.8) 1.0 (±0.0) 1.0 (±0.0) 6.3 (±9.7) 1.1 (±0.7) 18.4 (±14.4) 10.4 (±20.0)

0.0 0.0 0.0 0.0 2.3 (±2.3) 1.0 (±0.0) 16.3 (±11.1) 4.0 (±5.2)

2 9 12 7 28 32 36 28

0 0 0 0 3 2 4 3

3.2 (±4.0) 25.6 (±19.9) 3.4 (±4.2) 3.1 (±3.4) 2.0 (±2.7)

1.3 (±1.2) 11.5 (±14.7) 4.3 (±7.1) 3.0 (±3.6) 1.5 (±1.6)

36 75 42 38 34

12 71 52 57 58

26.8 (±21.9) 24.8 (±25.3) 2.1 (±2.1) 10.1 (±13.1) 7.3 (±11.3) 2.2 (±2.7) 3.1 (±5.3)

22.5 (±23.2) 23.3 (±24.5) 2.7 (±3.2) 6.6 (±9.8) 10.0 (±14.1) 3.0 (±4.5) 2.3 (±2.3)

117 106 40 75 73 69 15

162 128 109 74 73 102 48

a) non-migrating species and slow colonizers; b) species with a moderate pace of migration, exceeding the rate values given by other authors; c) fast-colonizing species. * - species not regarded as ancient woodland indicator for Poland.

Out of the 51 species for which colonization rates were calculated many herbs from the country‘s list of ancient woodland indicators, mainly myrmecochorous, migrated at a slow pace, up to 0.7 m yr-1 (for example Asarum europaeum – Figure 10, Viola reichenbachiana and Galeobdolon luteum) and their mean cover in recent woods was usually lower compared to ancient forests (Table 6). On the other hand, slow colonizers also included species with a potentially very effective dispersal mode, like anemochorous Equisetum sylvaticum. All of the species from the groups mentioned above, i.e. either not migrating or with low mobility in alder woods were also listed as ancient alder woodland species – AAWS (Table 7). For explanations of the calculations of species‘ migration rates and frequency see Table 7.

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Table 7. Species present significantly more often in ancient alder woods (AAWS) and the data on their dispersal mode, migration rate and the status for ancient woodlands according to different sources. Data on species’ migration rates were obtained from 33 transects. Species’ frequency is based on the species’ lists (N = 33 in ancient forests; N = 33 in recent forests) obtained from the entire area of forest sections where transects were located

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Species

Dispersal mode

Number of occurrences

P level

Mean migration rate [m yr-1] in alder woods studied

H et al. 1999

D and L 2001

Asarum europaeum Athyrium filix-femina

Myrm An1

Ancient N=33 11 27

Recent N=33 3 15

0.0163 0.0022

x x

x x

7

0

0.0055

x

x

11 13 12 13 25 13 4 20 8 27 10 25

1 5 5 4 17 4 0 7 2 19 1 14

0.0014 0.0257 0.0448 0.013 0.0361 0.0113 0.0568 (*) 0.0012 0.0412 0.0297 0.0030 0.0058

x x x x x x x x -

x x x x x x -

Auto

5

0

0.0266

x

x

Hy/An2 Endo Myrm Myrm An2 Auto Endo

29 21 4 15 6 29 17

19 5 0 7 0 21 9

0.0058 0.0001 0.0568 (*) 0.0332 0.0122 0.0212 0.0385

0.34 1.17 Absent in transects 0.30 0.82 1.12 0.07 1.35 1.50 0.00 0.27 0.45 0.95 0.00 0.99 Absent in transects 1.31 1.33 0.00 0.43 0.00 0.85 0.85

Campanula trachelium

An2

Carex sylvatica Corylus avellana Crataegus monogyna Equisetum sylvaticum Euonymus europaea Fagus sylvatica Gagea lutea Galeobdolon luteum Galium odoratum Geranium robertianum Hepatica nobilis Iris pseudacorus

Myrm Baro Endo An1 Endo Baro Myrm Myrm Epi Auto Myrm Hy

Lathyrus vernus Lysimachia vulgaris Maianthemum bifolium Melica nutans Mercurialis perennis Mycelis muralis Oxalis acetosella Paris quadrifolia Polygonatum multiflorum Pulmonaria obscura

x x x x x x

x x x x x x x

Endo

13

2

0.0012

0.94

x

x

Myrm

7

0

0.0055

x

x

Rorippa amphibia

Hy

4

0

0.0568 (*)

-

-

Rubus caesius Sanicula europaea Scutellaria galericulata

Endo Epi Hy

17 6 23

9 0 12

0.0385 0.0122 0.0065

x -

x -

Tilia cordata

An2

8

2

0.0412

x

-

Viola reichenbachiana

Myrm

15

1

0.0000

0.00 Absent in transects 1.52 0.00 1.55 Absent in transects 0.15

x

x

H et al. 1999 – Hermy et al. 1999; D and L 2001 – Dzwonko and Loster 2001. Symbols: ‗x‘ – listed as associated with ancient woodland; (*) – significance level between 0.05 and 0.1 according to the Fisher exact probability test Myrm – myrmecochores; Auto – autochores; Baro – barochores; Epi – epizoochores; Endo – endozoochores; An1 – unwinged anemochores with small diaspores; An2 – anemochores with diaspores with pappus or wings; Hy – hydrochores. Original source: Polish Journal of Ecology, 58, 2010, 297-310. Colonization capacity of herb woodland species in fertile, recent alder woods adjacent to ancient forest sites, Orczewska A. Table 1. With the kind permission of the Centre for Ecological Research, Polish Academy of Sciences

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Table 8. Ancient woodland indicator species for Poland (after Dzwonko and Loster 2001) which did not reach such a status in the alder woods studied (OAWS) and the data on their dispersal modes and migration rates.

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Species occurring in both forest types with a similar frequency

Dispersal mode

Number of Number of Mean migration rate occurrences in occurrences in [m yr-1] in alder ancient woods recent woods woods studied N=33 N=33 Adoxa moschatellina Endo 23 18 1.40 Aegopodium podagraria Baro 23 20 0.79 Ajuga reptans Myrm 17 21 2.38 Anemone nemorosa Myrm 27 23 1.73 Anthriscus nitida Epi 14 15 1.44 Brachypodium sylvaticum Epi 19 18 1.12 Carex elongata Hy 18 14 2.68 Carex remota Hy 9 7 0.31 Chrysosplenium alternifolium Hy 19 22 1.52 Circaea lutetiana Epi 21 19 1.77 Convallaria majalis Endo 6 2 absent in transects Dryopteris carthusiana An1 31 30 1.57 Dryopteris filix-mas An1 9 5 0.69 Festuca gigantea Epi 31 28 3.70 Ficaria verna Myrm 32 32 1.44 Geum urbanum Epi 28 27 1.87 Impatiens noli-tangere Auto 32 30 1.83 Milium effusum An2 28 24 0.91 Moehringia trinervia Myrm 32 29 1.75 Ranunculus auricomus Myrm 23 26 2.44 Ranunculus lanuginosus Epi 9 7 1.50 Ribes nigrum Endo 9 4 0.67 Ribes spicatum Endo 3 7 0.99 Rumex sanguineus An2 22 24 1.94 Scrophularia nodosa An1 20 15 1.70 Stachys sylvatica Epi 17 12 1.38 Stellaria holostea Baro 13 9 0.71 Stellaria nemorum Hy 11 12 1.24 Species with low frequency in ancient woods or either with low frequency or absent in recent woods Anemone ranunculoides Myrm 5 3 Bromus benekenii Epi 0 1 Circaea intermedia Epi 1 2 Daphne mezereum Endo 2 0 Dentaria bulbifera Baro 1 1 Epilobium montanum An2 1 3 Galium sylvaticum An2 2 0 Listera ovata An1 2 3 Luzula pilosa Myrm 3 0 Melampyrum nemorosum Myrm 2 0 Melica uniflora Myrm 3 0 Poa nemoralis An2 5 1

Original source: Polish Journal of Ecology, 58, 2010, 297-310. Colonization capacity of true woodland species in fertile, recent alder woods adjacent to ancient forest sites, Orczewska A. Table 2. With the kind permission of the Centre for Ecological Research, Polish Academy of Sciences Myrm – myrmecochores; Auto – autochores; Baro – barochores; Epi – epizoochores; Endo – endozoochores; An1 – unwinged anemochores with small diaspores; An2 – anemochores with diaspores with pappus or wings; Hy – hydrochores.

A large number of species migrated into recent alder woods very efficiently, much faster than in other post-agricultural forests in Europe and their migration rates exceeded the values given by other authors (Brunet and von Oheimb 1998b; Bossuyt et al. 1999; Dzwonko 2001b; Woodlands : Ecology, Management and Conservation, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

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Orczewska and Fernes 2011, in press) (Table 5; Figure 11). Circaea lutetiana, Stachys sylvatica, Festuca gigantea, Adoxa moschatellina, Scrophularia nodosa, Dryopteris carthusiana and Ajuga reptans are among them. All of these species, except for the last one mentioned, have dispersal modes that allow them to be very mobile (epizoochorous, anemochorous, or endozoochorous). They were also mentioned as good colonizers of recent, post-agricultural woods in other parts of Europe (for example Verheyen and Hermy 2004; Brunet 2007) and were classified as OAWS – other ancient woodland species, i.e. as those ancient woodland species for Poland which did not reach such a status in the alder woods (Table 8).

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Photo by Anna Orczewska. Figure 10. Asarum europaeum – ancient woodland species in many European forests, exhibiting a lowcolonizing capacity in recent alder woods.

Photo by Anna Orczewska. Figure 11. Migration of Anemone nemorosa into a recent black alder wood adjoining a Tilio cordataeCarpinetum betuli community (Namysłów forest district, Minkowskie, Oleśnica Plain).

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Anna Orczewska

Finally, there was a small group of species that can be identified as very effective colonizers of recent alder stands since not only did their migration rates exceed 2.0 m yr-1, but their mean cover in the herb layer of recent woods was also comparable with that recorded in ancient woods, or sometimes even exceeded it (Table 6). Ficaria verna, Ranunculus auricomus (ant-dispersed species) and Impatiens noli-tangere (autochorous) showed such behavior in each of the three habitats investigated. The mean migration rate for F. verna based on the farthest individual ranged between 1.96 m y-1 in alder-ash carrs and 2.99 m y-1 in oak-hornbeam communities; for R. auricomus from 2.04 m y-1 in alder-ash carrs to 2.85 m y-1 in wet alder woods, whereas in the case of I. noli-tangere it varied between 1.74 m y-1 in alder-ash carrs and 1.91 m y-1 in oak-hornbeam communities (Table 5). The potential reasons of such a great success of F. verna, R. auricomus and I. nolitangere in recent alder woods should be discussed further. I. noli-tangere is an annual ballochorous species and its seeds can be thrown a maximum distance of 2-3 m (Hiratshuka and Inoue 1988; cited after Hatcher 2003). However, in shady woodland conditions almost 90-96% of flowers are cleistogamous and the relatively small number of 360 seeds produced per plant can be dramatically reduced to one or no seeds in the deep shade of the forest floor (Falińska 1979; cited after Hatcher 2003). Thus, one may assume that the more open character of the recent alder stands with higher illumination before the canopy closure promotes the reproduction and spread of I. noli-tangere. Another, potential factor facilitating the migration of this species is the very intensive wild boar activity in recent alder woods. According to Jankowska-Błaszczuk (1998) this contributes to soil disturbance and the disturbance of the forest floor, a reduction in root competition from trees and herbs and consequently to the successful establishment of the population of I. noli-tangere. Wild-boarinduced disturbances are much more intensive in recent alder stands than in the bordering ancient forests. In the latter ones, additionally, the competition from other woodland species seemed to be higher, which might also be due to the limited number of patches of open space created by wild boars compared to recent sites. A similar mechanism may be involved in the distribution of another successful colonizer of recent alder woods, that is F. verna. This spring geophyte spreads mostly via vegetative propagation as it produces a large number of root tubers and bulbils (axillary tubercles called potato rain) (Taylor and Markham 1978). Wild boars feed on F. verna bulbils, thereby contributing to dispersal of this species, especially in recent alder woods. The growth of F. verna is favoured in open sites. In such conditions the size of the root system is bigger and more flowers are produced. The activity of wild boars, for which this species is an important part of their diet, contributes to the disturbance of the ground and the appearance of temporary patches of bare soil, especially in summer. One may expect that in such situations colonization of the open spaces may be at least partly through the seeds for which ants are the dispersers. Jung et al. (2008) discovered that although it has been maintained that F. verna seldom produces seeds (many flowers remain sterile) (Taylor and Markham 1978), it may produce them in some situations (0-18 ripe seeds per plant). At intermediate levels of disturbance it invests most into reproduction by seeds and seeds in such cases are beneficial for rapid recolonization of open sites after a disturbance (in the study by Jung et al. 2008 inundation was regarded as a disturbance factor). However, such effective and rapid colonization of F. verna observed in the herb layer of recent alder woods suggests that vegetative dispersal plays a more important role than ants. F. verna, among others, was also classified as a good colonizer of recent woodland sites on the habitats of broadleaved forests

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in Belgium by Bossuyt et al. (1999) and Verheyen and Hermy (2004) and in Sweden by Brunet (2007). By contrast, according to Honnay et al. (2009) F. verna belonged to the functional group called ‗small geophytes‘ which contain reliable indicators of the restoration status of post-agricultural woods since their abundance increases with the age of a recent forest. In general the members of this group fail to quickly colonize recent forests. Honnay et al. (2009) maintain that in the Belgian forests the reasons for the failure of small geophytes lies in the high content of soil nutrients and the high illumination level at the forest floor. Nevertheless, at least in the case of F. verna in recent black alder woods such an explanation seems to be doubtful. The vernal phenology of F. verna allows it to complete both the vegetative and generative growth stages before the full development of the competitive species – Urtica dioica or Solidago gigantea – that dominate in the herb layer of alder woods in summer. Thus, a high illumination level should not inhibit its development in recent woods. Finally, the possible reasons for the success of R. auricomus in recent woods need to be discussed. This is an apomictic species and this feature is heritable and genetically controlled (Nogler 1984; cited after Hörandl 2008). Such a trait and self-compatibility appear to be very advantageous in temporally and spatially unstable environments (moderately disturbed habitats) and also for colonization events because of its uniparental reproduction. Such a strategy allows a new population to be started from even a single individual (in the situation when there is a limitation of pollen and/or pollinators) (Hörandl 2008). This is especially important after an occasional long-distance dispersal (Baker 1967; cited after Hörandl 2008). R. auricomus is ant-dispersed so its dispersal mode is not effective. Nevertheless, Hörandl (2008) considered whether it is also capable to migrate with flood waters. In alder woods this dispersal mode could be a method of a long-distance dispersal event. Hence, apart from the group of species which were either unable to colonize recent alder woods or were very slow in this respect, many herbs migrated into the herb layer at a pace exceeding 2 or 3 m yr-1. The mean migration rate of species into recent woods adjacent to typical wet alder woods based on maximum cover reached 1.20 m yr-1 and the one based on the farthest individual 1.60 m yr-1. In the case of species in oak-hornbeam forest habitats it was 1.17 m yr-1 and 1.63 m yr-1, respectively and for alder-ash carrs 0.79 m yr-1 and 1.26 m yr-1. These ranges of migration rates indicate that the restoration of the herbaceous layer in woods of such high fertility and moisture proceeds faster than in habitats of deciduous woodlands in southern Sweden, where it was calculated for 0.3-0.5 m yr-1 by Brunet and von Oheimb (1998b). The rates in alder woods also exceeded the ranges given by Bossuyt et al. (1999) for mesotrophic, deciduous woods in central Belgium (0.5-1.0 m yr-1) and by Dzwonko (2001b) for pine plantations in a habitat of deciduous woods in southern Poland where they reached 0.18-0.38 m yr-1. The differences between the study results of alder woods and those reported by other authors were also significant when the ranges of migration rates of individual species were compared. The biggest ones were between alder woods and the results reported by Dzwonko (2001b), who calculated them at 0-0.38 m yr-1. The individual migration rates for species in broadleaved woods in Sweden varied from 0 to 1.25 m yr-1 and in Belgium from 0 to 1.15 m yr-1. A relatively faster pace was observed by Matlack (1994) in northeastern America, where species migration rates varied from 0 to 2.5 m yr-1 (Orczewska 2009b). Successful migration of F. verna and R. auricomus, two species with potentially low colonizing capacity due to their myrmecochory on the one hand and a low migration rate of

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the species with potentially very effective dispersal mode, like Equisetum sylvaticum and a few others, on the other hand, has provoked a discussion on the role of the dispersal mode in the migration of woodland herbs. Most surveys on the process of the recovery of the herbaceous layer in post-agricultural woods indicate the importance of dispersal limitation in the colonization process. Numerous studies have shown that it is regarded as a pivotal factor hampering the migration of forest herbs, especially in landscapes with highly fragmented forest cover (Peterken and Game 1984; Whitney and Foster 1988; Dzwonko and Loster 1990; 1992; Dzwonko 1993; Matlack 1994; Brunet and von Oheimb 1998b; Bossuyt et al. 1999; Honnay et al. 1999; Ehrlén and Eriksson 2000; Dzwonko 2001a, b; Singleton et al. 2001; Verheyen and Hermy 2001a; Bellemare et al. 2002; Verheyen et al. 2003; Takahashi and Kamitani 2004; Matlack 2005; Brunet 2007; for an extensive review of literature in this respect see also Honnay et al. 2002). Despite the fact that recent alder woods were established adjacent to the ancient forests on the same type of habitat, there was a group of woodland species which did not take advantage of such a spatial configuration and persisted in ancient woods. Many of the above-mentioned authors who have observed the species which were limited by dispersal also emphasized that the number of woodland herbs grew with the age of the recent woodland stand and decreased with its distance from the ancient, source woodlands. According to them this confirms the role of dispersal limitation in the distribution of woodland species in recent woods. In the case of recent black alder forest stands, age also accounted for most variation in the species composition of the herb layer (Orczewska 2009b). In addition, a comparison of the values of the mean migration rates of AAWS (Table 7) and OAWS (Table 8) groups confirms that the ability of species to colonize new, post-agricultural alder forests is highly associated with the dispersal mode. The majority of forest species rapidly colonizing recent alder woods (OAWS) are dispersed either by animals or the wind and their mean migration rate (1.54 m yr-1) is much higher than for the herbs classified as ancient alder woodland indicators (AAWS), for which it reached 0.68 m yr-1 (Orczewska 2010). Thus, it supports the observations made previously by many authors (Dzwonko and Loster 1992; Matlack 1994; Brunet and von Oheimb 1998b) that dispersal ability limits a species presence in recent woods. Poor colonizing capacity, among other traits, is typical for ancient woodland indicator species since about 25% of the herbs from that group are myrmeco-, auto- or barochorous (for detailed characteristics of the ecological profile of ancient woodland species see Hermy et al. 1999; Dzwonko and Loster 2001; Hermy and Verheyen 2007). However, other life-history traits should also be taken into account because the results concerning the migration rates of species in alder woods do not confirm their strict association with dispersal mode in some cases. The lack of a relationship between species colonization ability (migration rate) and its dispersal mode was also pointed out by Verheyen and Hermy (2001b) and Singleton et al. (2001). Similar to alder woods, they observed situations when ant-dispersed herbs were very good at colonizing recent woods (in the alder forests described in this chapter this refers, for example, to Ficaria verna, Ajuga reptans and Ranunculus auricomus), whereas species with effective modes of dispersal appeared to be poor colonizers (like Equisetum sylvaticum or Sanicula europaea in recent alder forests under study). Such observations allow a conclusion to be drawn that other species‘ traits may be more helpful in explaining the mechanisms behind the colonization potential of woodland species. A detailed classification and overview of such features is proposed by Honnay et al. (2009), who divided them into dispersal-related, establishment-related and persistence-related traits. The features related to the establishment and persistence of a species in a recently

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colonized habitat of a post-agricultural wood play equally important roles in the final success of a woodland species in a recent wood, as does its dispersal mode. Once a new site has been successfully reached by a woodland species other limiting factors may start to influence its further existence in a new site. The success of a species, understood as establishment and persistence, depends on both abiotic and biotic factors. Hence, recruitment limitation, like dispersal limitation, appears to be a very important mechanism which accounts for a slow migration rate of woodland species into recent woods. Species distribution in post-agricultural woods is a combined effect of both dispersal limitation and the availability of a suitable habitat (Flinn and Vellend 2005; Baeten et al. 2010). Former cultivation (agricultural use of soils) of today‘s secondary woods leads to undesirable alterations of soil properties (Flinn and Vellend 2005, Hermy and Verheyen 2007, and others). A detailed review of this topic that also focuses on the recovery pace of some of those parameters was done by Honnay et al. 2002).

a.

b. Figure 12. Relationship between the abundance of Urtica dioica and the a) number, and b) cover of woodland species in alder woods. ‗-‗ symbol with the numbers on axis X indicates plots located in ancient woodland

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Photo by Anna Orczewska.

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Figure 13. Recent alder wood (second age class – 11-20 years) with herbaceous layer dominated by Urtica dioica and Galium aparine (two competitive, nitrophilous species).

The influence of former agricultural use of the sites of recent alder woods on their soils was presented in a separate paper (Orczewska 2009c), thus only the most important mechanisms shaping current distribution of woodland herbs in these forests are described here. Among the abiotic parameters measured (groundwater level, canopy cover, illumination level, humus type and soil chemical properties) pH, type of humus and the level of the water table made the greatest contribution in explaining the distribution patterns of forest species in secondary alder woods. However, recruitment limitation is not related to a single factor but it is the combined effect of abiotic and biotic factors involving, as has been so often reported, a high nutrient and illumination level, especially in younger stands with an open canopy, the water regime and finally the influence of competitive species with broad ecological tolerances taking advantage of low canopy closure and the high availability of nutrients (Honnay et al. 1999; De Keersmaeker et al. 2004; Honnay et al. 2009). In alder woods Urtica dioica exhibits such behavior since its vigorous growth in recent alder stands leads to the competitive exclusion of woodland species from such sites (Orczewska 2009b, Figure 12). In the case of alder woods a similar behavior was also observed for Solidago gigantea, Galium aparine (Figure 13) and to a lesser extent also Poa trivialis. Excessive light may also promote the vigorous growth of grasses, which can have a negative impact on the successful establishment of woodland herbs, a mechanism which was reported by Brunet and von Oheimb (1998a). Furthermore, species with low germination rates and a small stature are prone to establishment limitation. Such features are disadvantageous especially in nutrient-rich recently planted woods with poor canopy closure where competition is a strong selecting (inhibiting) factor (Honnay et al. 2009). All of the authors who are studying relationships with U. dioica associate its abundant growth with an increased level of phosphorus in soils of a post-agricultural origin. Nevertheless, in alder woods the growth of this species depended, to a much bigger extent, on the groundwater level and light conditions than on the amount of

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P and N, which were apparently higher in ancient alder woods where stinging nettle was less abundant. U. dioica avoided sites with a high level of groundwater combined with poor illumination. Thus to create the best conditions allowing for effective forest recovery in alder woodlands, it is essential to maintain a good water regime and shade in the forest floor. This reduces the competitive exclusion of woodland flora by aggressive herbs and facilitates the immigration of forest species (Orczewska 2009c). The maintenance of stands with a closed canopy providing shady conditions also significantly enhanced the colonization of secondary oak and sycamore woods by shade-tolerant woodland herbs in southern Sweden (Brunet 2007).

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CONCLUSION Rapid and in many cases dramatic human-induced transformations in forest ecosystems due to their intensive exploitation and management have resulted in a decrease in the richness of woodland flora, a homogenization of the herbaceous layer, and even its invasion by aggressive, alien species, which are widely reported in numerous papers from all over the world every year. Alterations of the habitat and species composition that are observed in the young woods of a post-agricultural origin belong to the most severe instances. From the point of view of a woodland ecologist, such sites may be simply named as tree plantations since their soil and species composition do not resemble a true, natural forest. The most detectable feature of such a forest, apart from its simplified tree composition and uniform age, is the physiognomy of its herb layer. Herbaceous woodland species are very sensitive indicators of changes that take place in forest habitats. A wide range of life-history traits possessed by forest species (described in detail by Bierzychudek 1982; Whigham 2004, and others) and expressing their perfect adaptation to woodland environment have evolved over a long period of time. Thus, any rapid changes in the woodland habitat lead to transformations of the herb layer. Recent, post-agricultural woods are among those where such alterations are especially drastic. Furthermore, in these specific forests such changes may last for decades (Brunet and von Oheimb 1998b; Bossuyt et al. 1999; Dzwonko 2001b; Brunet 2007) or, when recent woods are spatially isolated from the ancient ones, even for centuries—350 years according to Faliński (1986) to 800 years as estimated by Peterken (1977). As has been presented in this chapter, in some habitats the herb layer recovery in recent woods may proceed faster than in other sites. In wet and fertile alder-dominated forests adjacent to ancient source woodlands, there is a chance of a reasonably fast forest restoration compared to poorer and drier habitats. The colonization capacity of many woodland herbs seems to be enhanced by the high humidity and fertility of these sites. However, many undesirable processes which take place in such woods, being a combined effect of abiotic and biotic factors, also make the migration of woodland flora into such habitats difficult. Among those factors, alterations in the water regime of such habitats—amelioration and drainage— have the most deteriorating effect on the composition of the herb layer. The results of the surveys in alder woods presented here, similar to some previous studies of other authors (De Keersmaeker et al. 2004; Hipps et al. 2005) show that a high water table reduces the growth rate of the aggressive herb – Urtica dioica in such sites. In addition, as was also reported by Honnay et al. (2002), high fertility combined with a high illumination level, typical for young

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woods before closure of the canopy, promotes the abundant growth of competitive species that take advantage of the availability of such conditions and therefore inhibit the immigration and successful establishment of woodland herbs. In order to prevent the competitive exclusion of woodland herbs, silvicultural practices focusing on fast canopy closure that provide shady conditions should be implemented. Forest management should also give priority to either the restoration or maintenance of the natural water regime. This refers both to ancient and recent alder woods. Only in such sites with a high water level is the conservation and sustainable management of ancient Alnus glutinosa-dominated forests focusing on maintaining their high biodiversity possible. Post-agricultural sites that are to be planted with black alder should be carefully selected. If necessary the natural water conditions should be restored prior to afforestation in order to avoid the expansion of nitrogen- and phosphorus-demanding species in the herb layer of recent woods. Dembek et al. (2002) emphasize that this is especially important in places with rich, organic soils where drainage contributes to decession (mineralization) and loss of organic matter and consequently to an increase of CO2 in the atmosphere rather than to carbon sequestration. In addition, black alders belong to the group of trees that have a great influence on their habitats. As an actinorhizal plant that also maintains a symbiosis with mycorrhizal fungi, it greatly contributes to the enhancement of the cycling of nutrients and to an increase in the N, P and C content in soils (Compton and Cole 1998; Binkley and Giardina 1998; Roy et al. 2007). However, drainage of hydrogenic habitats accelerates the mineralization of organic matter. Thus, altering hydrological regimes also contributes to global climate change, as was already mentioned. Therefore, implementation of the plans for increased afforestation on marginal agricultural land currently underway in Poland should take many different aspects into consideration in order to avoid unnecessary problems. Efforts should be made to select sites with a functioning natural water regime or the water regime should be restored prior to afforestation. Bearing in mind the fact that restoration of the herb layer in recent forests is possible only when post-agricultural woods border ancient forests, a wise afforestation plan that promotes the successful recovery of herb layer rich in woodland flora should be introduced. Even in such situations where recent woods are in direct proximity to ancient woodlands, some woodland species cannot fully take advantage of such a spatial configuration of forests. The results presented here indicate that many representatives of woodland herbs have a very poor colonization potential (see AAWS list, Table 7) and some of them are unable to migrate into adjacent recent woods. On the other hand, there are many species with a high colonization potential. Matlack and Monde (2004) stated that it is more likely that fast-moving forest species survive in dynamic landscapes and persist at higher frequencies than slow colonizers and the ability to cross gaps increases the chances of survival and persistence. Recent investigations by De Frenne et al. (2010, in press) focusing on the factors responsible for interregional differences in the recovery rates of herb layer species into recent woods on a European scale revealed that woodland species do not show a uniform behavior. Recovery of species with the highest colonization capacity (short-lived forest herbs) increased with the habitat availability in the landscape. By contrast, there was a group of the slowest species (small perennial forest herbs with heavy seeds) confined to ancient forests throughout Europe, hence not successful in colonizing recent woods even in relatively dense forested landscapes. Nevertheless, many forest herbs managed to colonize and establish their populations in post-agricultural alder woods. Furthermore, in many cases

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their colonization rates exceeded the values reported from other types of recent woodlands. Thus, black alder-dominated recent woods are very important elements in the spectrum of post-agricultural forests currently existing in human-dominated landscapes, and providing a good opportunity to study the forest restoration process.

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REFERENCES Baeten, L., De Frenne, P., Verheyen, K., Graae, B. J., and Hermy, M. (2010). Forest herbs in the face of global change: a single-species-multiple-threats approach for Anemone nemorosa. Plant Ecology and Evolution, 143, 19-30. Bellemare, J., Motzkin, G., and Foster, D. R. (2002). Legacies of the agricultural past in the forested present: an assessment of historical land-use effects on rich mesic forests. Journal of Biogeography, 29, 1401-1420. Bierzychudek, P. (1982). Life histories and demography of shade-tolerant temperate forest herbs: a review. New Phytologist, 90, 757-776. Binkley, D. (1986). Forest nutrition management. New York: Wiley and Sons, Inc. Binkley, D. and Giardina, C. (1998). Why do tree species affect soils? The Warp and Woof of tree-soil interactions. Biogeochemistry, 42, 89-106. Bossuyt, B. and Hermy, M. (2000). Restoration of the understorey layer of recent forest bordering ancient forests. Applied Vegetation Science, 3, 43-50. Bossuyt, B., Hermy, M., and Deckers, J. (1999). Migration of herbaceous plant species across ancient-recent forest ecotones in central Belgium, Journal of Ecology, 87, 628-638. Brunet, J. (2007). Plant colonization in heterogeneous landscapes: an 80-year perspective on restoration of broadleaved forest vegetation. Journal of Applied Ecology, 44, 563-572. Brunet, J. and von Oheimb, G. (1998a). Colonization of secondary woodlands by Anemone nemorosa. Nordic Journal of Botany, 18, 369-377. Brunet, J. and von Oheimb, G. (1998b). Migration of vascular plants to secondary woodlands in southern Sweden. Journal of Ecology, 86, 429-438. Brunet J., von Oheimb, G., and Diekmann, M. (2000). Factors influencing vegetation gradients across ancient-recent woodland borderlines in southern Sweden. Journal of Vegetation Science, 11, 515-524. Compton, J. E. and Cole, D. W. (1998). Phosphorus cycling and soil P fractions in Douglasfir and red alder stands. Forest Ecology and Management, 110, 101-112. Cousins, S. O. (2001). Analysis of land-cover transitions based on 17th and 18th century cadastral maps and aerial photographs. Landscape Ecology, 16, 41-54. De Frenne, P., Baeten, L., Graae, B. J., Brunet, J., Wulf, M., Orczewska, A., Kolb, A., Jansen, I., Jamoneau, A., Jacquemyn, H., Hermy, M., Diekmann, M., De Schrijver, A., De Sanctis, M., Decocq, G., Cousins, S. A. O., and Verheyen, K. (2010). Interregional variation in the floristic recovery of post-agricultural forests. Journal of Ecology, doi: 10.1111/j.1365-2745.2010.01768.x. (in press).De Keersmaeker, L., Martens, L., Verheyen, K., Hermy, M., De Schrijver, A., and Lust, N. (2004). Impact of soil fertility and insolation on diversity of herbaceous woodland species colonizing afforestations in Muizen forest (Belgium). Forest Ecology and Management, 188, 291-304.

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Dembek, W., Grzyb, M., Kloss, M., and Mikułowski, M. (2002). Łąki i lasy w dolinach – nowe zagrożenia i szanse (Forests and grasslands in the river valleys – new threats and opportunities). Postępy Nauk Rolniczych, 3, 87-119. Dilly, O. and Munch, J-C. (1996). Microbial biomass content, basal respiration and enzyme activities during the course of decomposition of leaf litter in a black alder (Alnus glutinosa (L.) Gaertn.) forest. Soil Biology and Biochemistry, 28, 1073-1081. Dzwonko, Z. (1993). Relations between the floristic composition of isolated young woods and their proximity to ancient woodland. Journal of Vegetation Science, 4, 693-698. Dzwonko, Z. (2001a). Effect of proximity of ancient deciduous woodland on restoration of the field layer vegetation in a pine plantation. Ecography, 24, 198-204. Dzwonko, Z. (2001b). Migration of vascular plant species to a recent wood adjoining ancient woodland. Acta Societatis Botanicorum Poloniae, 70, 71-77. Dzwonko, Z. and Gawroński, S. (1994). The role of woodland fragments, soil types and dominant species in secondary succession on the western Carpathian foothills. Vegetatio, 111, 149-160. Dzwonko, Z. and Loster, S. (1990). Vegetation differentiation and secondary succession on a limestone hill in southern Poland. Journal of Vegetation Science, 1, 615-622. Dzwonko, Z. and Loster, S. (1992). Species richness and seed dispersal to secondary woods in southern Poland. Journal of Biogeography, 19, 195-204. Dzwonko, Z. and Loster, S. (2001). Wskaźnikowe gatunki roślin starych lasów i ich znaczenie dla ochrony przyrody i kartografii roślinności (Ancient woodland plant species indicators and their importance for nature conservation and vegetation map ping). Prace Geograficzne, 178, 119-132. Ehrlén, J. and Eriksson, O. (2000). Dispersal limitation and patch occupancy in forest herbs. Ecology, 81, 1667-1674. Faliński, J. B. (1986). Vegetation dynamics in temperate lowland primeval forests. Geobotany, 8, 1-537. Flinn, K. M. and Vellend, M. (2005). Recovery of forest plant communities in postagricultural landscapes. Frontiers in Ecology and the Environment, 3, 243-250. Goldberg, E., Kirby, K., Hall, J., and Latham, J. (2007). The ancient woodland concept as a practical conservation tool in Great Britain. Journal for Nature Conservation, 15, 109119. Hatcher, P. E. (2003). Impatiens noli-tangere L. Journal of Ecology, 91, 147-167. Hensel, W. (1988). Polska starożytna. Wrocław, Warszawa, Kraków, Gdańsk, Łódź, POLAND: Ossolineum. Hermy, M., Honnay, O., Firbank, L., Grashof-Bokdam, C. J., and Lawensson, J. E. (1999). An ecological comparison between ancient and other forest plant species of Europe and its implications for forest conservation. Biological Conservation, 91, 9-22. Hermy, M. and Verheyen, K. (2007). Legacies of the past in the present-day forest biodiversity: a review of past land-use effects on forest plant species composition and diversity. Ecological Research, 22, 361-371. Hipps, N. A., Davies, M. J., Dodds, P., and Buckley, G. P. (2005). The effects of phosphorus nutrition and soil pH on the growth of some ancient woodland indicator species and their interaction with competitor species. Plant and Soil, 271, 131-141.

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Colonization of Post-Agricultural Black Alder (Alnus Glutinosa (L.) Gaertn.)…

45

Honnay, O., Bossuyt, B., Verheyen, K., Butaye, J., Jacquemyn, H., and Hermy, M. (2002). Ecological perspectives for the restoration of plant communities in European temperate forests. Biodiversity and Conservation, 11, 213-242. Honnay, O., Hérault, B., and Bossuyt, B. (2009). Opportunities and constraints of using understorey plants to set forest restoration and conservation priorities. In M.-A. Villard, and B. G. Jonsson (Eds.), Setting targets for managed forest landscapes (pp. 227-243). Cambridge University Press. Honnay, O., Hermy, M., and Coppin, P. (1999). Impact of habitat quality on forest plant species colonization. Forest Ecology and Management, 115, 157-170. Hörandl, E. (2008). Evolutionary implications of self-compatibility and reproductive fitness in the apomictic Ranunculus auricomus polyploid complex (Ranunculaceae). International Journal of Plant Science, 169, 1219-1228. Jankowska-Błaszczuk, M. (1998). Variability of the soil seed banks in the natural deciduous forest in the Białowieża National Park. Acta Societatis Botanicorum Poloniae, 67, 313324. Jung, F., Böhning-Gaese, K., and Prinzing, A. (2008). Life history variation across a riverine landscape: intermediate levels of disturbance favor sexual reproduction in the antdispersed herb Ranunculus ficaria. Ecography, 31, 776-786. Karkanis, M. (1975). Rozkład ściółki pochodzącej z różnych gatunków drzew liściastych i jej wpływ na środowisko glebowe (Decomposition of litter of various species of deciduous trees and its effect on soil environment). Fragmenta Floristica et Geobotanica, 21, 71-97. Kowalska, A. (2009). Zmiany sposobu użytkowania terenów rolniczych a zanikanie przyrodniczo cennych zbiorowisk roślinnych na przykładzie doliny środkowej Wisły (Changes in agricultural land-use practices leading to the disappearance of valuable natural vegetation communities – the example of Middle Vistula River Valley). In Polskie krajobrazy wiejskie dawne i współczesne. Prace Komisji Krajobrazu Kulturowego 12, 166-177. Kulczycka-Leciejewiczowa, A. (1993). Osadnictwo neolityczne w Polsce południowozachodniej: Próba zarysu organizacji przestrzennej (Neolithic settlement in southwestern Poland: An outline of spatial organization). Wrocław, POLAND: PAN, Instytut Archeologii i Etnologii. Laganis, J. (2007). Emergy analysis of black alder (Alnus glutinosa (L.) Gaertn.) floodplain forest growth. Dissertation: University of Nova Gorica Graduate School: SLOVENIA. Lasy Państwowe w liczbach. (2009). Warszawa, POLAND: Centrum Informacyjne Lasów Państwowych. Lityński, T., Jurkowska, H., and Gorlach, E. (1976). Analiza chemiczno-rolnicza. Warszawa, POLAND: Wydawnictwo Naukowe PWN. Majchrowska, A. and Woziwoda, B. (2009). Effects of forest history on the biodiversity of vascular plant flora in the Łask Upland (Central Poland). In J. Holeksa, B. BabczyńskaSendek, and S. Wika (Eds.), The role of geobotany in biodiversity conservation (pp. 165174). Katowice, POLAND: University of Silesia. Matlack, G. R. (1994). Plant species migration in a mixed-history forest landscape in eastern North America. Ecology, 75, 1491-1502. Matlack, G. R. (2005). Slow plants in a fast forest: local dispersal as a predictor of species frequencies in a dynamic landscape. Journal of Ecology, 93, 50-59.

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Matlack, G. R. and Monde, J. (2004). Consequences of low mobility in spatially and temporally heterogeneous ecosystems. Journal of Ecology, 92, 1025-1035. Matlack, G. R., and Leu, N. A. (2007). Persistence of dispersal-limited species in structured dynamic landscapes. Ecosystems, 10, 1287-1298. Matuszkiewicz, J. M. (1993). Krajobrazy roślinne i regiony geobotaniczne Polski (Vegetation landscape and geobotanical regions of Poland). Prace Geograficzne, 158, 1-107. Matuszkiewicz, W. (2001). Przewodnik do oznaczania zbiorowisk roślinnych Polski (A guide for the identification of Polish plant communities). Vademecum Geobotanicum. Warszawa, POLAND: Wydawnictwo Naukowe PWN. McVean, D. N. (1953). Biological flora of the British Isles: Alnus glutinosa (L.). Gaertn. (A. rotundifolia Stokes). Journal of Ecology, 41, 447-466. Moszkowicz, Ł. (2005). Pochodzenie i wiek lasów Ojcowskiego Parku Narodowego (Origin and age of woodlands in the Ojców National Park). Prace Muz. Szafera, 15, 275-281. Nyrek, A. (1992). Kultura użytkowania gruntów uprawnych, lasów i wód na Śląsku od XV do XX wieku (Methods of utilizing cultivable land, forests and water reservoirs in Silesia from the 15th to the 20th century). Acta Universitatis Wratislaviensis, 1361, Historia XCVII, 1-177. Orczewska, A. (2009a). Age and origin of forests in south-western Poland and their importance for ecological studies in man-dominated landscapes. Landscape Research, 34, 599-617. Orczewska, A. (2009b). Migration of herbaceous woodland flora into post-agricultural black alder woods planted on wet and fertile habitats in south western Poland. Plant Ecology, 204, 83-96. Orczewska, A. (2009c). The impact of former agriculture on habitat conditions and distribution patterns of ancient woodland plant species in recent black alder (Alnus glutinosa (L.) Gaertn.) woods in south-western Poland. Forest Ecology and Management, 258, 794-803. Orczewska, A. (2010). Colonization capacity of herb woodland species in fertile, recent alder woods adjacent to ancient forest sites. Polish Journal of Ecology, 58, 297-310. Orczewska, A. and Fernes, M. (2011). Migration of herb layer species into the poorest postagricultural pine woods adjacent to ancient pine forests. Polish Journal of Ecology (in press) Pancer-Kotejowa, E. and Zarzycki, K. (1980). Zarys ekologii. In S. Białobok (Ed.), Olsze – Alnus Mill. Nasze Drzewa Leśne. Monografie Popularnonaukowe 8 (pp. 229-258). Warszawa-Poznań, POLAND: Wydawnictwo Naukowe PWN. Pereira, A. P., Graca, M. A. S., and Molles, M. (1998). Leaf litter decomposition in relation to litter physico-chemical properties, fungal biomass, arthropod colonization and geographical origin of plant species. Pedobiologia, 42, 316-327. Peterken, G. F. (1974). A method for assessing woodland flora conservation using indicator species. Biological Conservation, 6, 239-245. Peterken, G. F. (1977). Habitat conservation priorities in British and European woodlands. Biological Conservation, 11, 223-236. Peterken, G. F. (1983). Woodland conservation in Britain, In A. Warren and F. B. Goldsmith (Eds.), Conservation in Perspective (pp. 83-100). Chichester, GREAT BRITAIN: Wiley.

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Colonization of Post-Agricultural Black Alder (Alnus Glutinosa (L.) Gaertn.)…

47

Peterken, G. F. and Game, M. (1984). Historical factors affecting the number and distribution of vascular plant species in the woodlands of Central Lincolnshire. Journal of Ecology, 72, 155-182. Prieditis, N. (1997). Alnus glutinosa - dominated wetland forests of the Baltic Region: community structure, syntaxonomy and conservation. Plant Ecology, 129, 49-94. Rackham, O. (1980). Ancient Woodland, its History, Vegetation and Uses in England. London, GREAT BRITAIN: Edward Arnold. Ralska-Jasiewiczowa, M., Nalepka, D., and Goslar, T. (2003). Some problems of forest transformation at the transition to the oligocratic/Homo sapiens phase of the Holocene interglacial in northern lowlands of Central Europe. Vegetation History and Archaeobotany, 12, 233-247. Roy, S., Khasa, D. P., and Greer, C. W. (2007). Combining alders, frankiae and mycorrhizae for the revegetation and remediation of contaminated ecosystems. Canadian Journal of Botany, 85, 237-251. Schaede, R. (1967). Symbiozy roślinne. Warszawa, POLAND: Państwowe Wydawnictwo Rolnicze i Leśne. Sienkiewicz, J., Kloss, M., and Grzyb, M. (2001). The floodplain forest ecosystems in Poland. In E. Klimo and H. Hager (Eds.), The floodplain forests in Europe: current situation and perspectives. European Forest Institute Report 10, 249-271. Singleton, R., Gardescu, S., Marks, P. L., and Geber, M. (2001). Forest herb colonization of postagricultural forests in central New York State, USA. Journal of Ecology, 89, 325338. Solińska-Górnicka, B. (1987). Bagienne lasy olszowe (olsy) w Polsce. Regionalna synteza syntaksonomiczna (Schwarzerlenbruchwälder in Polen. Regionale syntaxonomische Revision). Dissertationes Universitatis Varsoviensis 275. Warszawa, POLAND: Wydawnictwa Uniwersytetu Warszawskiego. Sprent, J. I. and Sprent, P. (1990). Nitrogen Fixing Organisms. London, GREAT BRITAIN: Chapmann and Hall. Takahashi, K. and Kamitani, T. (2004). Effect of dispersal capacity on forest plant migration at a landscape scale. Journal of Ecology, 92, 778-785. Taylor, K. and Markham, B. (1978). Ranunculus ficaria L. (Ficaria verna Huds., F. ranunculoides Moench). Journal of Ecology, 66, 1011-1031. Tuszyński, M. (1990). Właściwości gleb porolnych a gospodarka leśna (Properties of postagricultural soils and forest economy). Sylwan, 3, 41-51. Van der Pijl, L. (1982). Principles of dispersal in higher plants. Berlin, GERMANY: Springer. Verheyen, K. and Hermy, M. (2001a). An integrated analysis of the spatio-temporal colonization patterns of forest plant species in a mixed deciduous forest. Journal of Vegetation Science, 12, 567-578. Verheyen, K. and Hermy, M. (2001b). The relative importance of dispersal limitation of vascular plants in secondary forest succession in Muizen Forest, Belgium. Journal of Ecology, 89, 829-840. Verheyen, K. and Hermy, M. (2004). Recruitment and growth of herb-layer species with different colonizing capacities in ancient and recent forests. Journal of Vegetation Science, 15, 125-134.

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48

Anna Orczewska

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Verheyen, K., Guntenspergen, G. R., Biesbrouck, B., and Hermy, M. (2003). An integrated analysis of the effects of past land use on forest herb colonization at the landscape scale. Journal of Ecology, 91, 731-742. Whigham, D. F. (2004). Ecology of woodland herbs in temperate deciduous forests. Annual Review of Ecology, Evolution and Systematics, 35, 583-621. Whitney, G. C. and Foster, D. R. (1988). Overstorey composition and age as determinants of the understorey flora of woods of central New England. Journal of Ecology, 76, 867-876. World Reference Base for Soil Resources. (1998). FAO, ISSS, ISRIC. Rome, ITALY. Wulf, M. (2004). Auswirkungen des Landschaftswandels auf die Verbreitungsmuster von Waldpflanzen: Konsequenzen für den Naturschutz. Dissertationes Botanicae, Band 392 (J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung. Berlin, Stuttgart). Wulf, M. and Heinken, T. (2008). Colonization of recent coniferous versus deciduous forest stands by vascular plants at the local scale. Applied Vegetation Science, 11, 307-316. Wulf, M., Sommer, M., and Schmidt, R. (2009). Forest cover changes in the Prignitz region (NE Germany) between 1790 and 1960 in relation to soils and other driving forces. Landscape Ecology, 25, 299-313. Żabko-Potopowicz, A. (1959). Dotychczasowy stan badań nad rozmieszczeniem lasów i zasięgiem poszczególnych gatunków drzew w Polsce do połowy XII wieku (Hitherto position of research upon forest distribution and range of individual tree species in Poland till the early 12th century). Sylwan, 80, 1-35. Zimka, J. R. and Stachurski, A. (1976). Vegetation as a modifier of carbon and nitrogen transfer to soil in various types of forest ecosystems. Ekologia Polska, 24, 493-514.

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In: Woodlands: Ecology, Management and Conservation ISBN 978-1-61122-542-6 Editor: Erwin B. Wallace ©2011 Nova Science Publishers, Inc.

Chapter 2

DIVERSITY PATTERNS, ADULT RESOURCE USE AND CONSERVATION OF BUTTERFLY COMMUNITIES IN AND AROUND A PRIMEVAL WOODLAND OF MOUNT FUJI, CENTRAL JAPAN Masahiko Kitahara Department of Animal Ecology, Yamanashi Institute of Environmental Sciences (YIES), Kenmarubi, Fujiyoshida, Yamanashi, Japan

ABSTRACT Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved.

In central Japan, Aokigahara primeval woodland is considered to be one of the most natural areas around Mount Fuji and a core area in the conservation of the biodiversity of Mount Fuji. First, I chose butterflies as an indicator species of biodiversity and examined six communities in and around the woodland in 2000 using transect counts to examine and search for diversity and rarity hotspots and their associated landscapes. The results showed that butterfly species richness and species diversities H‟ and 1/λ were significantly higher in woodland-edge sites than in woodland-interior and/or open-land sites, and variation in the total number of species among these three landscape types was well accounted for by ecologically specialist species, such as landscape specifics, oligovoltines, narrow diet feeders and low density species. Thus, the species regarded as vulnerable to extinction, including Red List species, were observed more often in woodland-edge sites than in woodland-interior and/or open-land sites. As a result, in the study area, diversity and rarity hotspots were found in woodland-edge landscapes. The reasons why butterfly diversity and rarity hotspots were established in forest-edge landscapes were analyzed and interpreted from several points of view, including disturbance level, landscape elements and plant species richness. From these results, and the fact that some species were confined to woodland-interior sites, I conclude that it is very important to conserve and manage woodland-edge habitats (considered to be seminatural) as well as woodland-interior habitats (considered to be the most natural) to 

Correspondence: E-mail: [email protected].

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50

Masahiko Kitahara maintain the diversity of butterfly communities and preserve the various types of threatened species in and around the Aokigahara primeval woodland. Second, I examined the relationships between the diversities of vegetation, adult nectar plants, and butterflies in and around the Aokigahara woodland. The results showed that the nectar resource utilization by adult butterflies was significantly biased to herbaceous plants, especially to perennials, compared to woody species, although most of the study area was in and near the woodland. There were greater nectar plant species in sites with greater plant species richness. Among the butterfly community indices analyzed, the strongest correlation was detected between butterfly species richness and nectar plant species richness at each site. Another close correlation was detected between the species richness of nectar plants and herbaceous plants at each site. These results suggest that herbaceous plant species richness in a habitat plays a central role in its nectar plant species richness, and the nectar plant richness is a highly important factor supporting its adult butterfly species richness. Consequently, I propose that the maintenance and management of herbaceous plant species richness in a butterfly habitat, which lead to those of its nectar plant species richness, are very important for conservation of butterfly diversity even in and around woodland landscapes of temperate regions.

Keywords: adult nectar plants, butterfly community, diversity patterns, forest edge habitats, plant–butterfly relations, primeval woodland, species hotspots

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1. GENERAL INTRODUCTION To clarify the diversity patterns of biological communities and to search for the factors governing the patterns are one of the central aims of community ecology (MacArthur 1972; Pianka 1988; Begon et al. 1996). This kind of studies and information is also vital for the conservation of biodiversity (Primack 1993, 1995). At present, a remarkable decline of biodiversity is progressing on a global scale, and the conservation of biodiversity is being recognized as an international key problem. Also in Japan, a decrease in biodiversity has remarkably progressed after high economic growth period, and as a result, it is known that there are a lot of threatened species in the country (Biodiversity Center of Japan, 2010). The protection and maintenance of these threatened species thought to be a natural heritage of the nation are becoming the matters of the pressing need (Ministry of the Environment of Japan, 2007). On the other hand, the analysis of the characteristics and biotope of the threatened species are advancing, and it has been understood so far that many of threatened species of Japan are not the species to live in a primeval environment such as natural forest, but ones to live in a secondary environment that human moderately uses (Ministry of the Environment of Japan, 2001). However, the distribution and habitat type of threatened species are usually different in each region. Therefore, to consider the conservation and protection of threatened species, we should clarify their distribution pattern and biotopes at a regional level. In addition, we should search for a biological hotspot to maintain biodiversity in the region where threatened species inhabit. Until recently it has been stated that insects respond more rapidly to disturbance than vertebrates and, therefore, have potential as early indicators of environmental change (Kremen 1992; Kremen et al. 1993; Hamer et al. 1997). Among insects, butterflies are

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 51 believed to be the most suitable for indirect measures of environmental variation because of their high sensitivity to local weather, climate, light levels and other parameters that are affected by habitat change (Ehrlich et al. 1972; Weiss et al. 1987; Hill et al. 1995; Blair and Launer 1997; Wood and Gillman 1998). They have been well examined on their ecological attributes and biotope occupancy (e.g. Dennis et al. 2000; Shreeve et al. 2001; Dennis et al. 2004, 2005), and therefore, repeatedly used in the studies of conservation biology as a highly bioindicator for environmental assessment (e.g. Sakuratani 1991; Kremen 1992; Schmitt 2003). In addition, they are also very suitable for studies examining the structure and dynamics of populations and communities (Ehrlich 1992). Indeed butterflies have been well examined on the variation in diversity patterns along various environmental gradients (e.g. Ishii et al. 1991; Spitzer et al. 1993, 1997; Kitahara and Fujii 1994, Blair and Launer 1997; Natuhara et al. 1999; Kitahara et al. 2000; Kocher and Williams 2000; Natuhara 2000; Inoue 2003; Hogsden and Hutchinson 2004). In addition, the taxonomy and life histories of most Japanese butterfly species are already well known, and the adults of many species can be reliably identified in the field. Thus, diurnal Lepidoptera make ideal study subjects for diversity hotspot analyses of local biological communities. At the foot of Mount Fuji, central Japan, various types of environments such as primeval, semi-natural, and highly disturbed ones exist. In addition, it is well known that there are many Red Listed butterfly species in the area that the Ministry of the Environment made public (Kitahara, 1999; Kitahara and Watanabe, 2001, Kitahara, 2003; Kitahara and Watanabe, 2003). Thus, we chose Aokigahara woodland at the northwestern foot of Mount Fuji considered to be a primeval area and the surrounding semi-natural and human land use areas as study plots, and monitored adult butterfly communities for the purposes mentioned as follows. The aims of the present study are (1) to search for diversity and threatened species‘ hotspots and to clarify the diversity patterns of butterfly communities in and around the primeval woodland of Mount Fuji, central Japan, (2) to search for the factors affecting the patterns from the viewpoint of adult diet resource utilization of butterflies, and (3) to consider a better conservation plan to protect and maintain the butterfly diversity and the threatened species in and around the primeval woodland habitats.

2. DIVERSITY PATTERNS AND RARE SPECIES HOTSPOTS OF BUTTERFLY COMMUNITIES IN AND AROUND A PRIMEVAL WOODLAND OF MOUNT FUJI 2.1. Intoduction What we discover, identify and define as a biological hotspot, that is, a geographic area with high concentrations of species, endemic species, rare or threatened species and/or high levels of threat to species survival (Myers 1988; Mittermeier et al. 1998; Reid 1998), at a geographic location of concern is one of the most important steps in conserving and maintaining local biodiversity, in prioritizing areas for potential protection and in the establishment of nature reserves in local ecosystems (Primack 1993, 1995; Hunter 1996; Pressey 1996; Mittermeier and Forsyth 1997; Myers 1997). In the analysis of hotspots at a local scale/ level, it is very important to specify the type of natural environment or landscape

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52

Masahiko Kitahara

that is associated with these biological hotspots. In general, this attempt is very useful in assigning and setting conservation priorities to local hotspots, which produce a profound effect on the conservation of local biodiversity (Washitani 1999). The Aokigahara primeval woodland on the northwestern foot of Mount Fuji, central Japan, chosen as our study site, is considered to be one of the most natural areas around Mount Fuji and a core area in the conservation of biodiversity at Mount Fuji. In this section, we attempted to discover and identify butterfly hotspots (diversity and/or rarity (endangered species) hotspots) from among the representative landscapes present in and around the Aokigahara woodland. Many studies of butterfly communities have suggested that butterfly diversity and species richness is high at forest edges or transitional and ecotonal areas compared to open-land and woodland areas (e.g. Erhardt 1985; Leps and Spitzer 1990; Ishii et al. 1993, 1995; Spitzer et al. 1993; Kitahara and Fujii 1994; Ishii 1996a; Yata 1996; Tashita and Ichimura 1997; Natuhara et al. 1999; Balmer and Erhardt 2000; Schneider and Fry 2001). Thus, we aimed to confirm and clarify changing patterns in the structure of butterfly communities among selected representative landscapes, and to examine whether the abovementioned butterfly diversity pattern is observed in the present study area, with a view to formulating conservation strategies for community diversity.

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2.2. Methods Study Sites The study was carried out at an altitude of approximately 1000 m in and around the Aokigahara primeval woodland on the northwestern foot of Mount Fuji (altitude 3775.6 m) in central Japan. Three representative types of landscapes were selected in and around the woodland: (i) forest interior (FI); (ii) forest edge (FE) of the woodland; and (iii) open land (OL) outside the woodland. Two census sites were established (1 and 2) in each landscape type. Thus, all six census sites (named FI-1, FI-2, FE-1, FE-2, OL-1 and OL-2) were within the study area. This design made it possible to, at least in part, differentiate the effects of landscape or environmental differences on butterfly communities from the effects of other physical factors. That is, all six study sites were similar in terms of altitude and topography (almost flat or gently sloping land), and were located inside an area measuring 2.63 km east to west and 1.38 km north to south in or around the eastern part of the Aokigahara woodland. The characteristics of the six study sites, mainly their vegetation, are outlined in Table 1. Sites FI-1 and FI-2 were in the forest interior of the eastern part of the Aokigahara woodland. At site FI-1 we established a fixed census route (300 m in length) along a path crossing the site‘s interior to examine butterflies present mainly in the forest understory. At site FI-2, an artificial tower that stood in the forest was used. A fixed census point was established on the table of the tower, which was above the canopy of the forest (approximately 18 m above the ground), to examine butterflies present mainly above or near the forest canopy.

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Table 1. Characteristics and vegetation of the six study sites Study

Altitude

site FI-1

1030 m

Landscape and landscape element (open land)

Main plant (Phanerogamae) species Small trees and shrubs

Herbs

Type

Woodland

Trees Quercus mongolica var. crispula

No. plant (Phanerogamae ) species

Ilex pedunculosa

Polygonum cuspidatum

Trees

21

(forest understory)

Clethra barbinervis

Maianthemum dilatatum Oplismenus undulatifolius

Shrubs

15

Acanthopanax sciadophylloides

Acer micranthum Sorbus americana ssp. japonica

Perennials

9

Acer sieboldianum

Rhus trichocarpa

Corydalis incisa

Annuals

1

Pinus densiflora

Enkianthus campanulatus

Artemisia princeps

Others

3

Chamaecyparis obtusa

Rhododendron dilatatum

Total 49

Skimmja japonica f. repens

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FI-2

FE-1

1070 m

1025 m

Woodland

Tsuga sieboldii

Ilex pedunculosa

(forest canopy)

Chamaecyparis obtusa

Euonymns macropterus

Clethra barbinervis

Acer tschonoskii

Acer distylum Quercus mongolica var. crispula

Pieris japonica

Betula grossa

Prunus jamasakura

Pinus parviflora

Ilex macropoda

Quercus mongolica var. crispula

Prunus incisa

Quercus serrata

Lonicera japonica

Castanea crenata

Malus toringo

Woodland

Open land

unresearc h

unresearch

Acer micranthum

Miscanthus sinensis Boehmeria tricuspis ssp. paraspicata Cirsium nipponicum var. incomptum

Trees

16

Shrubs

15

Perennials

48

106

Table 1. (Continued)

Study

Altitud e

site

Landscape and landscape element (open land) secondary grassland conifer plantations

Main plant (Phanerogamae) species Trees

Small trees and shrubs

Herbs

Type

No. plant (Phanerogama e) species

Pinus densiflora

Rosa multiflora

Lysimachia clethroides

Annuals

18

Larix kaempferi

Deutzia crenata

Agrimonia pilosa

Others

9

vegetable plots abandoned arable land

Alnus hirsuta

Hydrangea paniculata

Sanguisorba officinalis

Magnolia obovata

Vicia cracca

sparse forest

Abies firma

Vicia unijuga

Total

Picris hieracioides ssp. japonica Erigeron annuus

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FE-2

1010 m

Woodland

Quercus serrata

Acer crataegifolium

Miscanthus sinensis

Trees

31

Pinus densiflora

Lindera obtusiloba

Oplismenus undulatifolius

20

Larix kaempferi

Enkianthus campanulatus

Campanula punctata

Shrubs Perennial s

Alnus hirsuta

Rhododendron dilatatum

Cirsium nipponicum var. incomptum

Annuals

20

Acer capillipes

Euonymns macropterus

Kalieris pinnatifida

Others

10

sparse forest

Zelkova serrata

Spiraea japonica

Lysimachia clethroides

bare site

Prunus maximowiczii

Ligustrum obtusifolium

Polygonum cuspidatum

Open land conifer plantations secondary grassland

Clethra barbinervis

Trifolium repens Astilbe microphylla Picris hieracioides ssp. japonica

55

136

Table 1. (Continued)

Study

Altitude

site OL-1

990 m

Landscape and landscape element (open land) Open land

Main plant (Phanerogamae) species Herbs Oxalis corniculata

Type Trees

No. plant (Phanerogamae) species 0

athletic fields and

Taraxacum officinale

Shrubs

1

open areas with

Geranium nepalense

Perennials

23

grassland

Vicia cracca

Annuals

27

Trifolium repens

Others

1

Trees

Small trees and shrubs

Total 52

Cerastium fontanum ssp. japonica Poa annua Ambrosia artemisiifolia var. elatior Kummerovia striata

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OL-2

1025 m

Open land

Morus australis

Rorippa indica

Trees

6

farmland consisting of

Pinus densiflora

Miscanthus sinensis

Shrubs

9

cabbage, potato, and

Cornus controversa

Plantago asiatica

Perennials

24

strawberry plots

Salix bakko

Oxalis corniculata

Annuals

18

Pieris japonica

Taraxacum officinale

Others

3

Celastrus orbiculatus

Rumex crispus ssp. japonicus

Rosa multiflora

Agrimonia pilosa Calystegia japonica Trifolium pratense Vicia unijuga Hemerocallis fulva var. Kwanso

From Kitahara and Watanabe, 2003.

60

Table 2. The values of various community indices and the numbers of several types of species vulnerable to extinction at each census site in the three landscape types in and around the Aokigahara woodland Landscape type Census site

Open land OL-1 OL-2

Mean

Forest edge FE-1 FE-2

Mean

Forest interior FI-1 FI-2

Mean

ANOVA or t-test

Bonferroni test

(a) Community index Total population density Species richness (S) Species diversity (H')

44.04 13 2.44

137.81 24 2.825

90.93 18.5 2.633

215.39 44 3.357

180.98 42 3.37

198.19 43 3.363

22.59 10 2.149

39.44 11 2.149

31.02 10.5 2.149

F = 8.389, P = 0.0591 F = 27.310, P = 0.0119 F = 30.232, P = 0.0103

FE, FI* FE, OL* FE, FI*

Species diversity (1/λ)

12.46

18.227

15.343

22.926

23.973

23.45

10.409

8.137

9.273

F = 15.360, P = 0.0265

FE, FI*

Evenness (J')

0.951

0.889

0.92

0.887

0.902

0.894

0.933

0.896

0.915

F = 0.405, P = 0.6985

0

3

1.5

4

4

4.0

0

0

0

t = – 1.667, P = 0.2375

–

3

7

5.0

20

18

19.0

6

5

5.5

F = 36.048, P = 0.0080

FE, FI* FE, OL*

3

5

4.0

9

10

9.5

2

2

2.0

F = 36.200, P = 0.0079

FE, FI* FE, OL*

0

0

0.0

6

6

6.0

4

0

2.0

t = 2.000, P = 0.1835

0

6

3.0

13

14

13.5

5

2

3.5

F = 9.152, P = 0.0528

–

1

3

2.0

16

13

14.5

5

3

4.0

F = 31.824, P = 0.0096

FE, FI* FE, OL*

(b) Species vulnerable to extinction† Red List spp‡ Univoltine spp.

‡

Larval-diet specialist spp.

‡

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Site-specific (endemic) spp. Narrow-geographic-range spp.‡ Low-population-density spp.‡

‡

From Kitahara and Watanabe, 2003. †Reference to Primack (1993). ‡See text for exact criteria. *P < 0.01.

Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 57 Table 3. Number and frequency (%) of butterfly species belonging to each of several ecological characteristics in the three landscape types at the census sites

(a) No. landscape types observed 3 2 1 (b) Voltinism (no. generations per year) ≥ 2 1 (c) No. potential hostplant species ≥30 10–30 1–10 (d) Average population density (no. month–1 ha–1) ≥5 2–5 0–2

Open-land sites (26 spp.)

Forest-edge sites (53 spp.)

Forest-interior sites (16 spp.)

r2

8 (30.8%) 18 (69.2%) 0 (0.0%)

8 (15.1%) 21 (39.6%) 24 (45.3%)

8 (50.0%) 4 (25.0%) 4 (25.0%)

– 0.666 0.834

14 (53.8%) 4 (15.4%) 8 (30.8%)

16 (30.8%) 13 (25.0%) 23 (44.2%)

5 (33.3%) 2 (13.3%) 8 (53.3%)

0.671 0.992 0.931

5 (19.2%) 8 (30.8%) 13 (50.0%)

6 (11.5%) 22 (42.3%) 24 (46.2%)

2 (12.5%) 6 (37.5%) 8 (50.0%)

0.736 0.978 0.998

12 (46.2%) 10 (38.5%) 4 (15.4%)

14 (26.4%) 19 (35.8%) 20 (37.7%)

5 (31.3%) 5 (31.3%) 6 (37.5%)

0.709 0.990 0.863

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From Kitahara and Watanabe, 2003.

These sites were both situated inside the Aokigahara woodland, which consisted of a continuous and extensive natural and primeval forest (3000 ha in area) that grew on the Aokigahara lava flow from the crater of Mount Nagao, which is located halfway up Mount Fuji and formed in the ad 864 eruption (Tsuya 1971). It is believed that this forest has never been subjected to large-scale human disturbance. The average tree age is approximately 150 years and the highest age ever recorded of the representative dominant tree species is 356 years for Tsuga sieboldii and 240 years for Chamaecyparis obtusa (Seido 1991). Almost no human land use or disturbance were found in these two woodland interior sites. Sites FE-1 and FE-2 were at the eastern forest edge of Aokigahara woodland. In each site, a fixed census route was established (300 m in length) along the forest boundary, bordering other landscapes (treeless areas). A small part of the route at site FE-1 ran through a grassland near the forest boundary. In general, both sites comprised natural forest (usually along one side of the census route), secondary (seminatural) grassland, conifer plantations, vegetable plots, abandoned farmland and scattered forest (usually on the other side of the census route). As these sites were in the same woodland, the forest structure and composition of the sites were basically similar to the two forest interior sites; however, they were further characterized by a greater presence of broad-leaved deciduous trees that resulted from good

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sunlight along the forest edge. There were no major differences in the landscape and environmental structure between FE-1 and FE-2, except for some differences in the component vegetation species. At these two sites, various types of human land use (e.g. plantations and vegetable plots) and disturbance (e.g. mowing and cultivation) were found in most places except the forested parts. Thus, approximately half the area at each site (the wooded parts) was relatively undisturbed, whereas the other half (the treeless parts) had moderate human disturbance. In general, the human disturbance level at both sites can be considered to be intermediate. Open land was located outside the eastern edge of the Aokigahara woodland. Most areas of this open land, except near the woodland edge, were reclaimed as farmland, recreational and resort areas with athletic fields, and a summer cottage zone. Sites OL-1 and OL-2 were located in the open land and a fixed census route was established (300 m in length) along a path running through each site. Site OL-1 comprised mainly athletic fields and open areas with tracts of turf for track and field sports and events. There were almost no large trees, small trees or shrubs at this site. Site OL-2 comprised farmland consisting of cabbage, potato, strawberry and Sanguisorba officinalis (for use in flower arrangements) fields. OL-2 contained some species of trees, shrubs and herbs in the grasses around the agricultural plots and along the farm roads. High levels of human land use (e.g. sports and recreational areas, farmland) and disturbance (e.g. trampling, mowing, pesticide spraying, fertilizer application, cultivation) were observed in most areas of these two reclaimed open-land sites. The human disturbance level at these two sites was considered to be the highest of all three landscape types.

Census Method Intensive regular censuses were carried out twice per month during the adult flight season (from April to November 2000), from 10:00 to 15:00 h local time under fair weather conditions. Data were collected at all sites except FI-2 using the line transect method (Pollard 1977, 1984; Thomas 1983; Pollard and Yates 1993), which is extensively used to survey and monitor butterfly populations and communities (e.g. Shreeve and Mason 1980; Erhardt 1985; Warren et al. 1986; Yamamoto 1988; Ishii 1993; Pollard and Yates 1993). It is a method of considerable value when investigating differences in species abundance among sites (Gall 1985; New 1991). Walking at a steady pace along the transect line, we recorded the number of adult individuals of each butterfly species sighted within a belt approximately 10 m wide and 300 m long (~3000 m2 in area). Individuals that could not be identified immediately on sight were often netted and released after identification. Data were collected from site FI-2 using the visual observation method in a fixed area (Southwood 1978). We covered approximately the same survey area (~3000 m2) as at the other sites and spent a standard time (~15 min) estimated from the surveys at the other study sites so that data obtained using this method were directly comparable with data from the other sites. We recorded all adult butterflies within a circular area, with a radius of approximately 30 m, near and above the forest canopy around the fixed census point on the table of the tower. We often used binoculars to help identify distant individuals. We recorded the weather conditions and human disturbance-related events, such as mowing and cultivation, at each site during each census.

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We conducted a vegetation survey along each transect route using the belt-transect method (Lincoln et al. 1998) at all sites surveyed using the transect method. Vegetation surveys were conducted twice (12 June and 27 August 1999) for each transect route, and we recorded all species of plants (belonging to Phanerogamae) sighted within a belt approximately 10 m wide along each route. The vegetation at site FI-2 was inferred from (T. Ohtsuka, unpubl. data, 2001).

Data Analysis We analyzed the butterfly community structure at each site using the following ecological parameters: population density, average population density, total population density, species richness, species diversity and species evenness. We used the population density of each butterfly species at each study site, which we obtained as follows, to analyze butterfly communities. First, we determined the monthly abundance of each species at each site during the study period (i.e. the mean number of adult individuals obtained from the two transect surveys in each month). However, in September, October and November, transect surveys were only conducted once per month. Thus, in these months monthly abundance was the number of adults from the one survey. To obtain a monthly mean abundance, we averaged the monthly abundance in months when a butterfly species was recorded. Excluding months when no butterflies were observed minimized the effect of different voltinism among butterfly species on the yearly abundance estimate (i.e. monthly mean abundance). Finally, we obtained the population density (no. adults month–1 ha–1) by dividing the monthly mean abundance by 0.3 ha (the area of each census site). Average population density was obtained by averaging the population densities of sites only where a specific species was found. Exclusion of study sites where no individuals were observed minimized the effect of different distribution patterns (widespread or restricted) of butterfly species on the average population density. As the application of different monitoring methods may restrict the comparison of results among the study sites, despite the fact that the areas of census was almost the same (0.3 ha) at all sites, we note that there may be a possibility that the population values recorded at FI-2 are more or less underestimated when compared to the other sites because of sampling errors related to the specific monitoring method (the visual point observation method in a fixed area) used at FI-2. The list of butterfly species observed in the present study and their population density values at each site and average population density are shown in Appendix I. Total population density was calculated as the population density of all component species at each site. Species richness was expressed as the number of species found at each study site during the observation period. Species diversity at each site was expressed by both the Shannon–Wiener function, H‟ = -Σ pi ln pi, where S is the number of species at each site and pi is the proportion of the population density of the ith species at each site, and the Simpson‘s index of diversity (Simpson 1949), 1/λ, λ = Σni(ni – 1)/N(N – 1), where ni is the population density of the i-th species at each site, and N is the total population density of all component species at each site. Species evenness was expressed by the Shannon equitability index, J‟ = H‟ ln–1 S, where H‟ is the Shannon–Wiener function and S is the number of species at each site. The ecological characteristics of the component species used in the analyses were as follows: average population density, local distribution pattern, voltinism, potential larval diet breadth, geographic range size in Japan, Red List category in Japan and the type of larval host

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plant. The local distribution pattern of each species was expressed by the number of census sites at which the species was recorded (i.e. 1–6). Voltinism was defined as the number of generations per year based mainly on Unno and Aoyama (1981), and complemented the actual data of seasonal fluctuations in the observed number of individuals of each species in the present study. Potential larval diet breadth was expressed as the number of all larval hostplant species ever recorded in Japan according to Endo and Nihira (1990). The geographic range size of each species in Japan was represented by the number of 10 km × 10 km squares in which the species was recorded in a report on the distribution patterns of Japanese animals and plants (Butterfly volume, Nature Conservation Bureau 1993). This report includes data on butterflies that were collected and recorded by 421 volunteer specialists, and plotted on a grid of 10 km × 10 km cells that cover almost the entire country. Most of the distribution records (89.33%) of butterflies used in the analyses were collected from 1980 onwards, with the rest (10.67%) completed before 1979. Component species observed in this study were compared to those on the Red Data list of Japan (Ministry of the Environment of Japan (hereafter M.E.J.) 2000) and species that corresponded to any of the Red List categories of Japan were determined. The type of larval host plant was classified as either grassy (herbaceous) or woody based on Unno and Aoyama (1981). The ecological characteristics of each component species are shown in Appendix I. We chose species vulnerable to extinction from the component species to examine which sites or landscapes were associated with vulnerable species. Referring to Primack (1993), we treated Red List species of Japan, univoltine species, diet-specialist species, site-specific species, narrow-geographic-range species and low-population-density species as species vulnerable to extinction. Among these species, diet-specialist species were defined as those whose larvae had been reported to feed on five or less plant species based on Endo and Nihira (1990), site-specific species were those recorded at only one census site in the present study, narrow-geographic-range species were those recorded in 500 or less 10 km × 10 km blocks in Japan based on a report on the distribution patterns of Japanese butterflies (Nature Conservation Bureau 1993), and low-population-density species were those with an average population density less than 2.00 (no. individuals month–1 ha–1) in the present study. To examine differences in the community indices and the observed numbers of species vulnerable to extinction among the landscape types, we used ANOVAS and Bonferroni tests on the annual values of the respective indices at each site. However, in cases where the mean value in a particular landscape type was zero, we used a t-test. To examine the impact on changes in species composition among landscape types, we calculated the coefficient of determination (r2) for the number of all species and for species belonging to each category associated with several ecological characteristics in each landscape. To clarify the relationships between community attributes and the threatened/vulnerable species, we conducted correlation analyses between species richness or diversity and the number of several threatened species among the communities at all six census sites. To investigate similarities and differences in species composition among census sites, we analyzed the community data using principal components analysis (PCA) based on the variance–covariance matrix, using the program EXCEL-multivariate analysis ver. 3.0 (Esumi 1998). A 57 (species) by six (census sites) matrix based on the population density of each species at each site was subjected to this analysis. To clarify the characteristics of site-specific species, we conducted a Spearman rank correlation analysis between the number of sites at

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 61 which a species was observed and the value of several ecological characteristics for all component species.

2.3. Results

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Community Indices and Species Vulnerable to Extinction at Each Site Table 2 shows the values of various community indices and the numbers of several types of species vulnerable to extinction at each census site. The mean values of species richness, species diversity H‟ and 1/λwere significantly different among the three landscape types. That is, species richness, species diversity H‟ and 1/λwere higher in forest-edge sites than in forestinterior and/or open-land sites. Whereas, the mean values of total population density and evenness were not significantly different among the landscape types. Red List species authorized by M.E.J. (2000) were only recorded in two forest-edge sites and at OL-2. From the other species vulnerable to extinction, univoltine, larval-diet specialist and low-population-density species were significantly more abundant in forest-edge sites than in forest-interior and open-land sites. Whereas, site-specific (endemic) species were only present in two forest-edge sites and at FI-1. Differences in Species Composition among the Landscape Types Table 3 shows changes in species composition among the three landscape types for several ecological factors. The values of the coefficient of determination (r2) and the maximum difference in the number of species among the landscapes indicate that variations in the total number of species among the three landscape types were accounted for more by the numbers of species observed in only one landscape type, oligovoltines (uni- or bivoltines), narrow diet breadth (corresponding to 1–9 or 10–29 species of larval potential host plants) and low population densities (corresponding to < 2 or from 2 to less than 5) than by the number of species observed in two landscape types, multivoltines (more than bivoltines), broad diet breadth (corresponding to ≥ 30 species of host plants) and high population densities (corresponding to ≥ 5), respectively. Thus, the high species richness in forest-edge sites was closely related to the high numbers of species observed in only one landscape type, and the number of oligovoltine species, narrow-diet-breadth species and low-populationdensity species. Relationship between Butterfly Species Richness and voltinism, Larval Host-Plant Type and Plant Species Richness As expected from the level of human disturbance at each landscape type, open-land (more disturbed) and forest-interior (less disturbed) communities had higher proportions of multivoltine (more than bivoltine) and univoltine species, respectively (Figure 1a). In forestedge communities on both less-disturbed (woodland parts) and more-disturbed (treeless parts) areas, the highest numbers of all categories of species (univoltines, bivoltines and multivoltines) were detected. In particular, marked increases in the number of univoltine or bivoltine species contributed to the high species richness in forest-edge communities. Similarly, as expected from landscape types, open-land (more grassy) and forest-interior (woodland) communities had higher proportions of species of larval grass feeders and larval

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tree feeders, respectively (Figure 1b). In forest-edge communities in both woodland and treeless grassland areas, the highest number of species of both tree and grass feeders were detected. In particular, marked increases in the number of species of grass feeders contributed to the high species richness in forest-edge communities.

From Kitahara and Watanabe, 2003. Figure 1. Relationship between species richness (total number of species) and (a) the number of species with different voltinism (univoltines: r = 0.961, P = 0.0007; bivoltines: r = 0.988, P = 0.0001; multivoltines (more than bivoltines): r = 0.844, P = 0.032), (b) with different host-plant types (tree feeders: r = 0.813, P = 0.0493; grass feeders: r = 0.969, P = 0.0003) and (c) with plants (belonging to Phanerogamae) (r = 0.924, P = 0.0225) in each butterfly community at each census site studied. The FI2 site is excluded from the analysis of (c) because of the lack of the data on the number of plant species.

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 63 Table 4. Correlation coefficient of species richness (total no. species) or species diversity (H) of each butterfly community with the number of the various species vulnerable to extinction at each of the six census sites

Species richness (total no. species) Species diversity (H)

(endemic) spp.

Narrowgeographicrange spp.

Lowpopulationdensity spp.

0.990***

0.734

0.936**

0.908**

0.986***

0.654

0.888*

0.831*

Red List

Univoltine

Larval-diet-

Site-specific

spp.

spp.

specialist spp.

0.957***

0.961***

0.965***

0.903**

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From Kitahara and Watanabe, 2003. *P < 0.05, **P < 0.01, ***P < 0.001.

From Kitahara and Watanabe, 2003. Figure 2. Scattergram of the six study sites along the first and second principal component axes of the principal components analysis (PCA) ordination.

Butterfly species richness was significant and positively correlated with plant (Phanerogamae) species richness at each census site (Figure 1c).

Relationship between Species Richness and Diversity and Species Vulnerable to Extinction All correlations of species richness and diversity (H‟) with numbers on the Red List, univoltine, larval-diet specialist, narrow-geographic-range and low-population-density species except site-specific species were positive (Table 4). This indicates that butterfly communities with high species richness and diversity had many more species vulnerable to extinction than communities with low species richness and diversity.

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64

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Ordination of Butterfly Communities The distribution of the butterfly communities at the six sites on major- and minor-axes planes using PCA based on the variance–covariance matrix is shown in Figure 2. There were two distinct groups: (i) group A distributed on the left side of the first axis; and (ii) group B distributed sparsely on the right side of the first axis. This suggests that the butterfly community in and around the Aokigahara woodland was represented by forest-edge and forest-interior communities, according to species composition and occurrence patterns. The cumulative contribution by the first and second principal components was 83.6%. In the first axis, the values of eigenvectors almost had a bias toward positive, and the sites FE-1, FE-2 and OL-2, which showed higher total population density, had higher eigenvector values (>0.5), whereas sites FI-1 and FI-2, which showed lower total population density, had lower eigenvector values ( 0.05) and (d) number of 10-kmblocks where the species was recorded in Japan (rs = 0.472, P < 0.001). Each dot and bar represents mean ± SD, respectively.

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2.4. Discussion Diversity, Structure and Hotspots of Butterfly Communities This study showed that butterfly species richness and species diversities, H‟ and 1/λ, were significantly higher in forest-edge sites than in forest-interior and/or open-land sites (Table 2), indicating that in and around Aokigahara woodland there were butterfly diversity hotspots at the forest edges. Relatively high species diversity and richness at forest edges or transitional and ecotonal areas from open land to woodland have been reported in many butterfly studies (e.g. Erhardt 1985; Leps and Spitzer 1990; Ishii et al. 1993, 1995; Spitzer et al. 1993; Kitahara and Fujii 1994; Ishii 1996a; Yata 1996; Tashita and Ichimura 1997; Natuhara et al. 1999; Balmer and Erhardt 2000; Schneider and Fry 2001). Thus, this trend appears to be a general and consistent one in butterfly diversity patterns in these landscapes. In general, this trend can also be understood as an edge effect (i.e. high species diversities and densities are recognized in boundaries between different habitat or landscape types) (Odum 1971; Fagan et al. 1999). In addition, this study also showed that variation in the total number of species among the three landscape types was well accounted for by species that were landscape specific, oligovoltine, and with a narrow diet breadth and low population density (Table 3). In other words, these types of species contributed to the differences in the number and composition of species among the landscape types, whereas other species types did not. When we examine the characteristics of the former species based on the r/K concept (MacArthur and Wilson 1967; Pianka 1970, 1988; Gadgil and Solbrig 1972; MacArthur 1972; Southwood et al. 1974; Southwood 1977, 1981, 1988) or the generalist/specialist concept (sensu Odum 1989; Novotny 1991; Kitahara and Fujii 1994, 1997; Leps et al. 1998), these species correspond to K-type (with lower r) or ecological specialist species. The tendency for K-type or ecologically specialist species to contribute to differences in the number and composition of species among communities has been shown in several butterfly community studies (Spitzer et al. 1993; Kitahara and Fujii 1994; Ishii 1996a; Kitahara et al. 2000; Kitahara and Sei 2001). Thus, this tendency also appears to be a general and consistent trend in the spatial changing patterns of butterfly communities. Why is butterfly species richness high in forest-edge landscapes? We answered this question from the following three points of view (Figure 1). First, forest-edge areas included more-disturbed (mainly treeless open land) and less-disturbed (mainly woodland) parts. Thus, it is predicted that forest-edge communities show high species richness because of the inclusion of both r- and K-type species, compared with communities in only open-land or woodland areas. Indeed, our study confirmed the high species richness of both multivoltines (r-type species) and oligovoltines (K-type species) in forest-edge sites and, therefore, verified the above prediction. Similarly, we confirmed that species richness in forest-edge sites, resulting from the inclusion of both grassland and woodland species, was high compared to the richness observed in open-land or woodland areas only. Second, butterflies are herbivores and their diet resources are almost entirely dependent on specific plants for both larval and adult stages. Thus, it is predicted that the number of species within a butterfly community in a certain area is strongly affected by the number of plant species. As this prediction suggests, our study revealed a strong positive correlation between butterfly and plant species richness at each census site and, thus, the high butterfly species richness at forest-edge sites was closely linked to a high plant species richness. Finally, the high butterfly species richness at forestedge sites probably led to the high butterfly species diversity.

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In the present study, the various types of species vulnerable to extinction, including Red List species, were more often observed at forest-edge sites than in forest-interior or open-land sites (Table 2). Thus, similar to the diversity hotspots, there were butterfly rarity hotspots in areas at the forest edges in and around the Aokigahara woodland and, therefore, both hotspots occurred in the same landscape area. Moreover, we suggest that because strong positive correlations were detected between the numbers of almost all types of species vulnerable to extinction and butterfly species richness and diversity (H‟) (Table 4), the high richness of species vulnerable to extinction in forest-edge sites significantly contributed to the high total species richness and diversity. Why were there butterfly rarity hotspots in forest-edge areas? In Japanese and British butterfly communities (Asher et al. 2001), species that are vulnerable to extinction are characteristic of semi-natural grasslands or coppice woodlands (Ishii 1996b). In particular, in and around Mount Fuji, most of the recorded Red List species authorized by M.E.J. (2000) are semi-natural grassland or open-forest species (Sei 1988; Kitahara 1999). In addition, various types of species treated as vulnerable to extinction in the present study are characteristic of semi-natural grassland or woodland habitats according to Fukuda et al. (1982, 1983, 1984a,b). Thus, we suggest that in the area of the present study, butterfly rarity hotspots formed at forest-edge sites, which included areas of semi-natural grassland with a low frequency of human impact and woodland areas with almost no human impact.

Conservation Implications for Butterfly Diversity and Rarity The present study showed that both butterfly diversity and rarity hotspots were detected in forest-edge landscapes (Table 2). In addition, site-specific and landscape-specific species, which are considered to have a high conservation value because of their endemism to specific habitats, were observed only in the areas of forest edges and interiors (Tables 2, 3). From these results we conclude that, at least in this area, the conservation of forest-edge habitats in addition to woodland habitats is very important for the maintenance of butterfly species diversity and richness and for the preservation of rare or threatened species. Moreover, the results of the PCA suggest that all six butterfly communities were divided into two community groups represented by forest-edge and forest-interior communities (Figure 2). It is likely that the communities at OL-1 and OL-2 belonged to a different group because the differences in their total population densities and species composition resulted from the different types of human disturbance at these sites. Thus, the need to conserve both forestedge and woodland habitats can be supported by the species grouping and composition of the butterfly communities observed in the present study. Accordingly, in and around the Aokigahara woodland, forest-edge and woodland areas have great potential value and, therefore, should be considered to be priority areas for butterfly conservation. In the present study, species observed at fewer sites (i.e. restricted species) were associated with traits that reflect a vulnerability to extinction, such as lower population densities, fewer generations per year and smaller geographic range sizes (Figure 3). Thus, these species appear to be representative of species prone to extinction and, therefore, can be thought of as species with high values of conservation or priority species of concern for conservation. The positive relationship between distribution and abundance has been well shown for many organisms, and is considered to be a general community pattern (Hanski 1982; Brown 1984; Bock 1987; Gaston and Lawton 1988, 1990; Hanski et al. 1993; Lawton 1993; Gaston 1994, 1996). In general, it is suggested that species prone to extinction have

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similar characteristics. This should be emphasized and is very important in considering and planning butterfly conservation. We need to monitor and manage extinction-prone species carefully to maintain local butterfly diversity.

3. RELATIONSHIP OF BUTTERFLY DIVERSITY WITH NECTAR PLANT SPECIES RICHNESS IN AND AROUND A PRIMEVAL WOODLAND OF MOUNT FUJI

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3.1. Introduction In general, as almost all butterflies utilize their species-specific plant resources in both larval and adult stages, it is believed that the diversity of plants influences the diversity of butterflies. In fact, the positive correlation between plant and butterfly diversities has been reported or pointed out in many previous studies (e.g. Erhardt 1985; Sparks and Parish 1995; Ishii 1996; Kitahara and Watanabe 2001, 2003; Simonson et al. 2001; Croxton et al. 2005). However, there have been a few studies (Väisänen 1992; Holl 1996; Kitahara 2004) that the correlation is weak between butterfly diversity and vegetational community composition or species richness. In another study (Hawkins and Porter 2003), it was pointed out that, although plant and butterfly diversities are positively correlated, plant diversity does not directly influence butterfly diversity but that both are probably responding to similar environmental factors. Thus, the actual relationship between butterfly and plant diversities and its produced mechanisms are not yet clear to perfection. On the other hand, it is generally thought that a greater diversity of resources should support a greater diversity of consumers. Indeed, it has been known that a greater abundance and/or diversity of nectar resources are associated with a greater abundance and/or diversity of butterflies (Murphy and Wilcox 1986; Kremen 1992; Munguira and Thomas 1992; Holl 1995; Ishii et al. 1995; Loertscher et al 1995; Steffan-Dewenter and Tscharntke 1997; Hardy and Dennis 1999; Clausen et al. 2001; Ries et al. 2001; Schneider and Fry 2001; Simonson et al. 2001; Pryke and Samways 2003; Pywell et al. 2004), although no associations between them has been known in a few studies (e.g. Sharp et al. 1974). Schneider and Fry (2001) advocated that the availability of both nectar sources and larval food plants are important in determining butterfly diversity. However, at least in temperate Japan, few studies have conducted on butterfly species and their nectar plant relationships, and almost no information is available on the conservation value of flowering nectar plants for butterfly abundance and diversity. Recently, the importance of a resource-based definition of habitat and approach based on the accurate identification of the spatial and temporal existence of resources in a landscape has been emphasized for butterfly conservation (Dennis et al. 2003, 2006). In addition to this, the need for the development of a resource database on butterfly biology necessary to adopt the resource-based approach has been proposed (Dennis et al. 2003). Thus, in the present study, considering the accumulation of new data on adult nectar resources, we examined the relationship between butterfly species and their nectar plant species richness in and around the Aokigahara primeval woodland at the northwestern foot of Mount Fuji, central Japan. Our goals of this study are (1) to clarify the relationships between the diversities of vegetation,

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 69 adult nectar plants, and butterfly communities, (2) to examine the role nectar resources play in determining butterfly diversity, and (3) to evaluate the conservation value of flowering adult nectar plants for butterfly diversity.

3.2. Materials and Methods

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Study Sites The study in this section was carried out at almost the same study sites in the section 2, and the outline of the study area is described in the previous section. However, two forest interior (FI) sites were different from those in the section 2. Therefore, the two FI sites in this section were described below. Sites FI-1 and FI-2 were in the forest interior, consisting of natural forest established on the Aokigahara lava flow, which originated in A.D. 864 eruption of Mount Nagao (located halfway up Mount. Fuji). Each census route was established along a path crossing the site‘s interior. It is believed that this forest has never been subjected to large-scale human disturbance. Average tree age is ca. 150 years and the highest ever ages recorded for the dominant tree species are 356 years for Tsuga sieboldii and 240 years for Chamaecyparis obtusa (Seido 1991). Comparison of FI-1 and FI-2 showed that the latter was dominated by evergreen coniferous trees such as Tsuga sieboldii and Chamaecyparis obtusa, with herb species present in some parts of the understory, while about half of FI-1 was dominated by broad-leaved deciduous trees such as Q. mongolica var. crispula, with very few herb species. Almost no human land use or disturbance was evident. The characteristics of the six study sites, mainly their vegetation, are outlined in Table 6. Census Methods Butterfly communities were monitored using the line transect method (Pollard 1977, 1984; Thomas 1983; Gall 1985; Pollard and Yates 1993). This method is now extensively used to survey and monitor butterfly populations and communities (e.g. Shreeve and Mason 1980; Erhardt 1985; Pollard and Yates 1993) and is of considerable value when investigating differences in species abundance between sites (Gall 1985; New 1991,1997). Transect counts were conducted twice a month usually during the adult flight season (from April to November 1999) and between 10:00 and 15:00 h local time under fair weather conditions. Walking at a steady pace along the transect line, the number of adult individuals of each butterfly species sighted was recorded within a belt approximately 10 m wide. Individuals that could not be identified immediately were netted, identified, and released. In the field, it is not possible to distinguish between Pieris melete and Pieris napi. Therefore, these two congeneric species complexes were treated as Pieris spp. for the analysis.

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Table 6. Characteristics of six study sites Study site

FI-1

Altitude

1030 m

Landscape and landscape element (open land) Woodland (forest understory)

Main plant (Phanerogamae) species Trees Quercus mongolica var. crispula

Small trees and shrubs

Herbs

Ilex pedunculosa

Polygonum cuspidatum

Clethra barbinervis Acanthopanax sciadophylloides

Acer micranthum Sorbus americana ssp. japonica

Maianthemum dilatatum

Acer sieboldianum

Rhus trichocarpa

Pinus densiflora

Enkianthus campanulatus

Chamaecyparis obtusa

Rhododendron dilatatum

Type of human disturbance (mainly in open land)

Skimmja japonica f. repens

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FI-2

FE-1

1020 m

1025 m

Woodland (forest understory)

Woodland

Open land

Tsuga sieboldii

Ilex pedunculosa

Oplismenus undulatifolius

Chamaecyparis obtusa

Callicarpa japonica

Plantago asiatica

Clethra barbinervis

Prunus incisa

Artemisia princeps

Acer distylum

Lindera obtusiloba

Maianthemum dilatatum

Cornus controversa Quercus mongolica var. crispula

Euonymns macropterus

Erigeron annuus

Skimmja japonica f. repens

Corydalis incisa

Quercus serrata Quercus mongolica var. crispula

Prunus incisa

Quercus serrata

Lonicera japonica

Castanea crenata

Malus toringo

Miscanthus sinensis Boehmeria tricuspis ssp. paraspicata Cirsium nipponicum var. incomptum

Mowing Cultivation Fertilization

Table 6. (Continued) Study site

Altitude

Landscape and landscape element (open land)

Main plant (Phanerogamae) species Trees

Small trees and shrubs

Herbs

secondary grassland

Pinus densiflora

Rosa multiflora

Lysimachia clethroides

conifer plantations

Larix kaempferi

Deutzia crenata

Agrimonia pilosa

vegetable plots

Alnus hirsuta

Hydrangea paniculata

Sanguisorba officinalis

abandoned arable land

Magnolia obovata

Vicia cracca

sparse forest

Abies firma

Vicia unijuga

Type of human disturbance (mainly in open land)

Picris hieracioides ssp. japonica Erigeron annuus FE-2

1010 m

Woodland

Quercus serrata

Acer crataegifolium

Miscanthus sinensis

Mowing

Pinus densiflora

Lindera obtusiloba

Oplismenus undulatifolius

Cultivation

Larix kaempferi

Enkianthus campanulatus

Fertilization

conifer plantations

Alnus hirsuta

Rhododendron dilatatum

Campanula punctata Cirsium nipponicum var. incomptum

secondary grassland

Acer capillipes

Euonymns macropterus

Kalieris pinnatifida

sparse forest

Zelkova serrata

Spiraea japonica

Lysimachia clethroides

bare site

Prunus maximowiczii

Ligustrum obtusifolium

Polygonum cuspidatum

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Open land

Clethra barbinervis

Trifolium repens Astilbe microphylla Picris hieracioides ssp. japonica

OL-1

990 m

Open land athletic fields and open areas with

Oxalis corniculata

Heavy trampling

Taraxacum officinale Geramium thumbergii

Intensive mowing Land readjustment

Table 6. (Continued) Study site

Altitude

Landscape and landscape element (open land)

Main plant (Phanerogamae) species Trees

Small trees and shrubs

grassland

Herbs

Type of human disturbance (mainly in open land)

Vicia cracca Trifolium repens Cerastium fontanum ssp. japonica Poa annua Ambrosia artemisiifolia var. elatior Kummerovia striata

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OL-2

1025 m

Open land

Morus australis

Rorippa indica

Intensive cultivation

farmland consisting of

Pinus densiflora

Miscanthus sinensis

Tilling

cabbage, potato, and

Cornus controversa

Plantago asiatica

strawberry plots

Salix bakko

Oxalis corniculata

Intensive mowing Intensive fertilization

Pieris japonica

Taraxacum officinale

Insecticide spraying

Celastrus orbiculatus

Rumex crispus ssp. japonicus

Rosa multiflora

Agrimonia pilosa Calystegia japonica Trifolium pratense Vicia unijuga Hemerocallis fulva var. Kwanso

From Kitahara et al. 2008.

Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 73

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Recently, several problems on butterfly distribution maps have been discussed (Dennis et al. 1999). In particular, it is pointed out that these maps fail to distinguish the status of records, that is, whether they are observations of breeding populations or vagrant individuals (Cook et al. 2001; Dennis 2001). Thus, it is very important to know whether records relate to breeding populations in favorable habitats or not, especially for conservation purposes. The same is true for transect data on recording adult individuals. In addition, it is pointed out that simultaneous collection of biotope, resources, and behavioural data is needed for monitoring affinities of butterflies to vegetation structures and using butterflies as indicators of environmental changes (Dennis 2004). Thus, in the present study, we recorded all adult feeding behaviors and their diet resource items (e.g. species name of nectaring plants) observed simultaneously with adult butterfly monitoring during each transect survey in every site. Weather conditions, light conditions, and human disturbance-related events such as mowing and cultivation were recorded at the same time during each transect survey. At all butterfly study sites surveyed using the transect method, the vegetation was surveyed within 10-m wide corridors along each transect route using the belt-transect method (Lincoln et al. 1998); to record as many plant species as possible, separate surveys were conducted on 12 June and 27 August, 1999. We recorded all species of plants (belonging to Phanerogamae) sighted along each route in the respective survey days. Only Phanerogammic species were surveyed because most butterflies in the study area utilize such plants in both the larval and adult stages.

Data Analysis We analyzed butterfly community structure at each site using the following ecological parameters: population density, total population density, species richness, and species diversity. The population density of each butterfly species at each study site was calculated as follows. The monthly count was determined as the mean of twice-monthly counts conducted in May – September or as the value of single counts in April, October, and November. The mean monthly count over the season was then calculated using only those months when the species was observed to minimize the effect of variable voltinism between species. Finally, the population density (number of adults /month /km) was obtained by dividing the mean monthly count by 0.3 km (the length of each census route). The total population density at each site was the sum of population densities of all component butterfly species observed in each site. The species richness at each site was the total number of butterfly species observed in each site during the study period. The species diversity at each site was expressed by both 

s

p

i

ln p i

Shannon-Wiener function, H‘= i 1 , where s is the total number of species recorded, and pi is the proportion of the population density of the i-th species, and Simpson's index of S diversity (Simpson 1949), 1-λ, whereλ = Σi =1 ni (ni - 1) / N (N -1), ni is the population density of the i-th species, and N is the total population density of all component species in each site.

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74

Table 7. The values of various indices of butterfly communities, vegetation, and nectar plants recorded in the six study sites Study site

OL-1

OL-2

FE-1

FE-2

FI-1

FI-2

Total population density

74.76

121.87

269.53

162.92

10.83

9.44

Total number of species

18

23

43

39

3

3

Species diversity (H')

2.633

2.659

3.240

3.364

1.058

1.028

Species diversity (1-λ)

0.920

0.900

0.930

0.960

0.700

0.700

No. all plant species

52

60

106

136

37

64

No. woody plant species

1

16

35

55

34

42

No. tree species

0

6

16

31

20

22

No. shrub species

1

9

15

20

13

17

No. herbaceous plant species

50

42

69

77

2

21

No. perennial species

23

24

48

55

2

17

No. annual species

27

18

18

20

0

3

No. all nectar plant species

17

19

28

28

3

8

No. woody nectar plant species

1

2

3

4

3

4

No. tree nectar species

0

1

1

1

2

2

No. shrub nectar species

1

1

2

3

1

2

No. herbaceous nectar plant species

16

17

25

24

0

4

No. perennial nectar species

11

12

17

16

0

3

No. annual nectar species

5

5

8

8

0

1

Butterfly community

Vegetation (Phanerogamae)

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Adult nectar plants

From Kitahara et al. 2008.

Concerning to the vegetation at each study site, we used the following seven parameters in the analyses: the numbers of (1) all plant species (belonging to Phanerogamae), (2) herbaceous plant species, (3) woody plant species, (4) annual plant species, (5) perennial plant species, (6) shrub plant species, and (7) tree plant species observed in the two vegetation surveys. To estimate the abundance of diet resources for adult butterflies, we gained the number of adult nectar plant species in each study site as follows. First, we listed up all species of nectar plants used by adult butterflies observed through all study sites during the study period. Second, we determined the presence or absence of these nectar plant species in each study site based on the results of the vegetation survey stated above. The distribution record of each

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 75 nectar plant species in each study site is shown in Appendix Ⅱ. We used the following seven parameters in the analyses: the numbers of species of (1) all nectar plants, (2) herbaceous nectar plants, (3) woody nectar plants, (4) annual nectar plants, (5) perennial nectar plants, (6) shrub nectar plants, and (7) tree nectar plants recorded in each study site. Table 8. Number of species of plants belonging to Phanerogamae, and the number of nectar plant species among them recorded across all study sites

Herbaceous plants

Woody plants

Other plants

Total

Annuals

Perennials

Total

Shrubs

Trees

Total

Plants belonging to Phanerogamae

43

84

127

40

44

84

10

221

Adult nectar plants

11

22

33

3

2

5

0

38

From Kitahara et al. 2008.

Table 9. Correlation coefficients of the number of species between various plant types and adult nectar plants recorded in each study site

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Nectar plants

All

Herbaceous

Annual

Perennial

Woody

Tree

Shrub

plants

plants

plants

plants

plants

plants

plants

0.843*

0.983***

0.767

0.958***

0.140

0.061

0.161

*** P < 0.001. * P < 0.05. From Kitahara et al. 2008.

Table 10. Correlation coefficients between various butterfly community indices and the numbers of nectar plant species in the six study sites Butterfly community index

All nectar plants Herbaceous nectar plants Woody nectar plants

Total population

Total number of species

Species diversity (H')

Species diversity (1-λ)

density 0.915**

0.979***

0.970***

0.926***

0.907**

0.968***

0.986***

0.958***

0.016

0.045

-0.198

-0.328

From Kitahara et al. 2008.

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3.3. Results

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Vegetation and Adult Nectar Plants The values of various indices of butterfly communities, vegetation represented by plants belonging to Phanerogamae, and nectar plants utilized by adult butterflies in each of all the six study sites are shown in Table 7. Table 8 shows the species composition of all plants (belonging to Phanerogamae) and adult nectar plants recorded all over the study sites. Out of all 221 plant species (Phanerogamae) recorded in this study, the utilization as a nectar resource by adult butterflies was observed in 38 plant species (17.2 %). While, out of all 127 herbaceous and 84 woody species, the proportion of utilization as a nectar resource by adult butterflies was extremely higher in herbaceous (33 spp., 26.0 %) than woody (6 spp., 5.9 %) plants. This trend is much remarkable in Figure 4. The chi-square test for goodness of fit showed that the proportions of number of woody and herbaceous plant species recorded in the study area (i.e. expected proportions) were significantly different from those of number of woody and herbaceous adult nectar plant species (i.e. observed proportions) (χ2 = 26.333, df = 1, P < 0.0001). In more detailed analysis, the chi-square test for goodness of fit showed that the proportions of number of species of trees, shrubs, perennials, and annuals recorded in the study area (i.e. expected proportions) were significantly different from those of number of adult nectar species of trees, shrubs, perennials, and annuals (i.e. observed proportions) (χ2 = 27.246, df = 3, P < 0.0001). These results indicate that, in the study area, the nectar resource utilization by adult butterflies deviated extremely to herbaceous plants especially perennials, compared to woody plants.

From Kitahara et al. 2008. Figure 4. Percent proportions of the number of species of the respective plant types in all plants belonging to Phanerogamae, and in adult nectar plants among them recorded in all the study area.

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 77 Similarly, a significant positive correlation of the number of species was detected between adult nectar plants and all, herbaceous, and perennial plants belonging to Phanerogamae recorded in each study site (Table 9). In particular, the correlation was strong in both herbaceous and perennial plants.

Butterfly Communities and Adult Nectar Plants All butterfly community indices (total population density, total number of species, species diversities H‟ and 1-λ) were positively and highly significantly correlated with the number of species of all adult nectar plants recorded in each study site. In particular, among them, the strongest correlation was detected with butterfly species richness (total no. of species) (Table 10). On the other hand, similar to the above results, all butterfly community indices were positively and highly significantly correlated with the number of species of herbaceous adult nectar plants recorded in each study site. However, no correlations were detected between all butterfly community indices and the number of species of woody adult nectar plants recorded in each study site.

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3.4. Discussion Resource Utilization Patterns of Adult Butterflies In the present study, we showed that the nectar resource utilization by adult butterflies deviated extremely to herbaceous plants, especially to perennials, compared to woody species, although most of the study area was in and near a primary woodland. Such deviated to herb resource utilization by adult butterflies has been observed by several previous authors (Kitahara 2000; Kamimura 2004; Mano 2004; Tiple et al. 2006). Also in arable field margins of Britain, the importance of perennial nectar sources rather than annual ones was pointed out for butterfly conservation (Dover 1996). In this study, the highest number of adult butterfly species was recorded in woodland edge study sites, intermediate in open land sites, and the lowest in woodland interior sites (Kitahara and Watanabe 2003). It is also a general observation that most adult butterflies avoid shade and are often encountered in open sunny places (Douwes 1975, Dennis and Bramley 1985; Warren 1985; Pivnick and McNeil 1987). Thus, one possible reason for deviated to herb resource use by adults is that most adult butterflies indeed prefer flowers of herbaceous plants to those of woody ones, or herb abundance and density are simply much and high in open sunny spaces such as woodland edge and open land sites with abundant adult butterflies. Concerning to butterfly conservation, the deviated to herb resource use by adults suggests that the maintenance of herbaceous plant species richness and diversity in its habitats is important for ensuring the nectar resources of adult butterflies. Relationship between Vegetation and Adult Nectar Plants The number of species of adult nectar plants in each site was significantly and positively correlated with that of all plants belonging to Phanerogamae. Thus, there were more nectar plant species in sites with more plant species richness. In more detail, the number of species of adult nectar plants in each site was more strongly correlated with that of herbaceous or perennial plants than that of all plants belonging to Phanerogamae, suggesting that the

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Masahiko Kitahara

numbers of species of herbaceous and perennial plants both are the best predictors for those of adult nectar plants in the respective study sites. Probably, the strong correlations of number of species between adult nectar plants and both herbaceous and perennial plants in each site were caused by the deviated to herb resource utilization by adult butterflies mentioned above. Then again, the relationships between vegetation and adult nectar plants stated above also suggest that the maintenance and management of habitats with the species richness and diversity of herbaceous plants is important for the supply and availability of adult nectar resources.

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Relationship between Butterfly Diversity and Adult Nectar Plants, and Conservation Implications In the present study, we showed that butterfly community indices were all positively correlated with the number of nectar plant species in each site. Among them, the strongest correlation was detected between butterfly species richness (total no. of species) and nectar plant species richness (r = 0.979, p < 0.001). This correlation with the number of species of nectar plants is highly stronger than that with the number of species of all plants (belonging to Phanerogamae) recorded in each site (r = 0.842, p < 0.05) shown in the previous study (Kitahara and Watanabe 2001). These results suggest that nectar plant species richness is an important factor governing adult butterfly species richness in each site rather than total species richness of plants (Figure 5). The importance of nectar plant abundance for butterfly abundance and diversity has been pointed out in many previous studies (Murphy and Wilcox 1986; Kremen 1992; Munguira and Thomas 1992; Holl 1995; Ishii et al. 1995; Loertscher et al 1995;

From Kitahara et al. 2008. Figure 5. Number of species of all plants belonging to Phanerogamae, adult nectar plants, and butterflies recorded in each study site.

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 79

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Steffan-Dewenter and Tscharntke 1997; Hardy and Dennis 1999; Clausen et al. 2001; Ries et al. 2001; Schneider and Fry 2001; Simonson et al. 2001; Pryke et al. 2003; Pywell et al. 2004), although the reverse has been known in a few studies (Sharp et al. 1974). In addition, a number of population and community studies of butterflies suggest that adult distribution patterns are more affected by the availability of nectar resources than the presence of larval host plants (Ehrlich and Gilbert 1973; Gilbert and Singer 1973; Murphy 1983; Grossmueller 1987; Feber et al. 1996; Hardy and Dennis 1999). In addition, it has been known in some studies (Thomas and Mallorie 1985; Holl 1996) that butterfly species richness is positively correlated with herbaceous cover. Our study also showed that butterfly community indices were all strongly correlated with the species richness of herbaceous nectar plants in each site, but perfectly not with that of woody nectar plants. This indicates that the richness of herbaceous nectar plant species is one of the most important factors determining the community structure and attributes of adult butterflies. On the other hand, it is suggested that, most of the present study area was in and near a primary woodland; nevertheless the richness of woody nectar plant species has almost no effects on the determination of adult butterfly community structure. The importance of herbaceous plant species richness even within woodlands for herbivore diversity is also known for moths (Usher and Keiller 1998). As a whole, a highly close correlation was detected between the species richness of butterflies and nectar plants, and another close correlation between the species richness of nectar plants and herbaceous plants in each site (Figure 6).

From Kitahara et al. 2008. Figure 6. Relationships between the number of nectar plant species and the number of butterfly species (a), and the number of herbaceous plant species (b) recorded in each study site.

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These results suggest that herbaceous plant species richness in a habitat has a central role in its nectar plant species richness, and the nectar plant richness is a highly important factor governing and supporting its adult butterfly species richness. Consequently, we propose that the maintenance and management of herbaceous plant species richness in a butterfly habitat, which lead to those of its nectar plant species richness, are very important for the maintenance and conservation of butterfly species richness and diversity even in and around woodland landscape of temperate region, as Tudor et al. (2004) claimed that management of woodland sites for butterfly conservation should give as much consideration to nectar sources as to host plant sources.

CONCLUDING REMARKS

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4.1. Butterfly Diversity Patterns in and around Temperate Primeval Woodland The present study showed that butterfly species diversity was higher in primeval woodland-edge sites thought of as medium successional stages than in the woodland-interior as later successional stages and/or open land sites as earlier successional stages, indicating that in and around the primeval woodland, there were butterfly diversity hotspots at the woodland edge landscape under semi-natural conditions. In general, relatively high species diversity and richness at woodland edges or transitional and ecotonal areas from open land to woodland have been reported in many butterfly studies (e.g. Erhardt 1985; Leps and Spitzer 1990; Ishii et al. 1993, 1995; Spitzer et al. 1993; Kitahara and Fujii 1994; Ishii 1996a; Yata 1996; Tashita and Ichimura 1997; Natuhara et al. 1999; Balmer and Erhardt 2000; Schneider and Fry 2001). In other words, butterfly species diversity is highest in intermediate successional stages such as secondary or semi-natural habitats (e.g., Blair and Launer, 1997; Inoue, 2003). Thus, our findings are almost consistent with and support these commonly held view concerning butterfly diversity patterns. That is, I emphasize that the butterfly diversity pattern obtained in the present study appears to be a general and consistent one detected in and around temperate woodland habitats.

4.2. Adult Resource Abundance Explains the Butterfly Diversity Patterns in and around Temperate Primeval Woodland The present study showed that the nectar resource utilization by adult butterflies was significantly biased to herbaceous plants, especially to perennials, compared to woody species, although most of the study area was in and near the primeval woodland. There were greater nectar plant species in sites with greater plant species richness. In addition, among the butterfly community indices analyzed, the strongest correlation was detected between butterfly species richness and nectar plant species richness at each site. Another close correlation was detected between the species richness of nectar plants and herbaceous plants at each site. These results suggest that herbaceous plant species richness in a butterfly habitat plays a central role in its nectar plant species richness, and the nectar plant richness is a highly

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 81 important factor supporting the adult butterfly species richness in and around the temperate primeval woodland.

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4.3. Implications for Conservation and Management of Butterflies Living in and around Temperate Primeval Woodland As stated above, this study showed that both butterfly diversity and rarity hotspots were detected in primeval woodland-edge landscapes. In addition, site-specific and landscapespecific species, which are considered to have a high conservation value because of their endemism to specific habitats, were observed only in the areas of the woodland edges and interiors. From these results, it is concluded that the conservation of woodland-edge habitats in addition to woodland habitats is very important for the maintenance of butterfly species diversity and richness and for the preservation of rare or threatened species. Accordingly, in and around the Aokigahara primeval woodland, the woodland-edge and woodland interior areas have great potential value and, therefore, should be considered to be priority areas for butterfly conservation. This study also showed that the strongest correlation was detected between butterfly species richness (total no. of species) and nectar plant species richness, suggesting that nectar plant species richness is an important factor governing adult butterfly species richness in each site. In addition, a highly close correlation was detected between the species richness of butterflies and nectar plants, and another close correlation between the species richness of nectar plants and herbaceous plants in each site, suggesting that herbaceous plant species richness in a habitat has a central role in its nectar plant species richness. Consequently, we propose that the maintenance and management of herbaceous plant species richness in a butterfly habitat, which lead to those of its nectar plant species richness, are very important for the maintenance and conservation of butterfly species richness and diversity even in and around woodland landscape of temperate region.

ACKNOWLEDGMENTS I am grateful to Dr. T. Kobayashi, Mss. M. Watanabe and M. Yumoto who are the coinvestigators of the present study. My thanks are also due to Drs. H. Imaki, Z. Jiang, H. Ueda and Y. Yoshida of the Lab. of Animal Ecology for their discussion, help and understanding of this study, and to Drs. T. Ohtsuka and T. Nakano of the Lab. of Plant Ecology for providing me with their unpublished data on the vegetation of the Aokigahara woodland. I express my gratitude to Ms. A. Kobayashi of the Lab. who made the diagram of this manuscript. I am deeply indebted to Prof. emeritus K. Fujii of the Univ. of Tsukuba, and Prof. emeritus T. Nakamura of the Univ. of Yamanashi for their continuous advice and encouragement throughout the study. This work was supported in part by both Grants-in-Aid for Scientific Research (B) (no. 17310138) and for Scientific Research (C) (no. 20510221) from the Japan Society for the Promotion of Science (JSPS) to M. Kitahara.

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REFERENCES Asher, J., Warren, M., Fox, R., Harding, P., Jeffcoate, G., and Jeffcoate, S. (2001). The Millennium Atlas of Butterflies in Britain and Ireland. Oxford, Oxford University Press. Balmer, O. and Erhardt, A. (2000). Consequences of succession on extensively grazed grasslands for central European butterfly communities: rethinking conservation practices. Conservation Biology, 14, 746–757. Begon, M., Harper, J. L., and Townsend, C. R. (1996). Ecology: individuals, populations, and communities (3rd edition). Oxford, Blackwell Science. Biodiversity Center of Japan (2010). Information on threatened species in Japan. Available at http://www.biodic.go.jp/rdb/rdb_top.html (In Japanese). Blair, R. B. and Launer, A. E. (1997). Butterfly diversity and human land use: species assemblages along an urban gradient. Biological Conservation, 80, 113-125. Bock, C. E. (1987). Distribution–abundance relationships of some Arizona landbirds: a matter of scale? Ecology, 68, 124–129. Brown, J. H. (1984). On the relationship between abundance and distribution of species. American Naturalist, 124, 255–279. Clausen, H. D., Holbeck, H. B. and Reddersen, J. (2001). Factors influencing abundance of butterflies and burnet moths in the uncultivated habitats of an organic farm in Denmark. Biological Conservation, 98, 167-178. Cook, L. M., Dennis, R. L. H. and Hardy, P. B. (2001). Butterfly-hostplant fidelity, vagrancy and measuring mobility from distribution maps. Ecography, 24, 497-504. Croxton, P. J., Hann, J. P., Greatorex-Davies, J. N. and Sparks, T. H. (2005). Linear hotspots? The floral and butterfly diversity of green lanes. Biological Conservation, 121, 579-584. Dennis, R. L. H. (2001). Progressive bias in species status in symptomatic of fine-grained mapping units subject to repeated sampling. Biological Conservation, 10, 483-494. Dennis, R. L. H. (2004). Butterfly habitats, broad-scale biotope affiliations, and structural exploitation of vegetation at finer scales: the matrix revisited. Ecological Entomology, 29, 744-752. Dennis, R. L. H. and Bramley, M. J. (1985). The influence of man and climate on dispersion patterns within a population of adult Lasiommata megera (L.) (Satyridae) at Brereton Hearth Cheshire (U.K.). Nota lepidopterologica, 8, 309-324. Dennis, R. L. H., Donato, B., Sparks, T. H. and Pollard, E. (2000). Ecological correlates of island incidence and geographical range among British butterflies. Biodiversity and Conservation, 9, 343-359. Dennis, R. L. H., Hodgson, J. G., Grenyer, R., Shreeve, T. G. and Roy, D. B. (2004). Host plants and butterfly biology: Do host-plant strategies drive butterfly status? Ecological Entomology, 29, 12-26. Dennis, R. L. H., Shreeve, T. G., Arnold, H. R. and Roy, D. B. (2005). Does diet breadth control herbivorous insect distribution size? Life history and resource outlets for specialist butterflies. Journal of Insect Conservation, 9, 187-200. Dennis, R. L. H., Shreeve, T. G. and Van, Dyck, H. (2003). Towards a functional resourcebased concept for habitat: a butterfly biology viewpoint. Oikos, 102, 417-426.

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 83 Dennis, R. L. H., Shreeve, T. G. and Van, Dyck, H. (2006). Habitats and resources: the need for a resource-based definition to conserve butterflies. Biodiversity and Conservation, 15, 1943-1966. Dennis, R. L. H., Sparks, T. H. and Hardy, P. B. (1999). Bias in butterfly distribution maps: the effects of sampling effort. Journal of Insect Conservation, 3, 33-42. Douwes, P. (1975). Distribution of a population of the butterfly Heodes virgaureae. Oikos, 26, 332-340. Dover, J. W. (1996). Factors affecting the distribution of satyrid butterflies on arable farmland. Journal of Applied Ecology, 33, 723-734. Ehrlich, P. R. (1992). Population biology of checkerspot butterflies and the preservation of global biodiversity. Oikos, 63, 6–12. Ehrlich , P. R., Breedlove, D. E., Brussard, P. F. and Sharp, M. (1972). Weather and the regulation of subalpine populations. Ecology, 53, 243–247. Ehrlich, P. R. and Gilbert, L. E. (1973). Population structure and dynamics of the tropical butterfly Heliconius ethilla. Biotropica, 5, 69-82. Endo, S., and Nihira, I. (1990). Larval Food of Japanese Butterflies. Tokyo, Group ‗Tamamushi‘ (in Japanese). Erhardt, A. (1985). Diurnal Lepidoptera: sensitive indicators of cultivated and abandoned grassland. Journal of Applied Ecology, 22, 849-861. Esumi (1998). EXCEL-Multivariate Analysis, Version 3.0. Tokyo, Esumi Inc (translated into Japanese). Fagan, W. F., Cantrell, R. S. and Cosner, C. (1999). How habitat edges change species interactions. American Naturalist, 153, 165–182. Feber, R. E., Smith, H. and Macdonald, D. W. (1996). The effects on butterfly abundance of the management of uncropped edges of arable fields. Journal of Applied Ecology, 33, 1191-1205. Fukuda, H., Hama, E., Kuzuya, T., Takahashi, A., Takahashi, M., Tanaka, B., Tanaka, H., Wakabayashi, M., and Watanabe, Y. (1982). The life histories of butterflies in Japan (Vol. 1). Osaka, Hoikusha (in Japanese with English summary). Fukuda, H., Hama, E., Kuzuya, T., Takahashi, A., Takahashi, M., Tanaka, B., Tanaka, H., Wakabayashi, M., and Watanabe, Y. (1983). The life histories of butterflies in Japan (Vol. 2). Osaka, Hoikusha (in Japanese with English summary). Fukuda, H., Hama, E., Kuzuya, T., Takahashi, A., Takahashi, M., Tanaka, B., Tanaka, H., Wakabayashi, M., and Watanabe, Y. (1984a). The life histories of butterflies in Japan (Vol. 3). Osaka, Hoikusha (in Japanese with English summary). Fukuda, H., Hama, E., Kuzuya, T., Takahashi, A., Takahashi, M., Tanaka, B., Tanaka, H., Wakabayashi, M., and Watanabe, Y. (1984b). The life histories of butterflies in Japan (Vol. 4). Osaka, Hoikusha (in Japanese with English summary). Gadgil, M. and Solbrig, O. T. (1972). The concept of r- and K-selection: evidence from wild flowers and some theoretical considerations. American Naturalist, 106, 14–31. Gall, L. F. (1985). Measuring the size of Lepidopteran populations. Journal of Research on the Lepidoptera, 24, 97-116. Gaston, K. J. (1994). Rarity. London, Chapman and Hall. Gaston, K. J. (1996). The multiple forms of the interspecific abundance–distribution relationship. Oikos, 76, 211–220.

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Masahiko Kitahara

Gaston, K. J.and Lawton, J. H. (1988). Patterns in the distribution and abundance of insect populations. Nature, 331, 709–712. Gaston, K. J. and Lawton, J. H. (1990). Effects of scale and habitat on the relationship between regional distribution and local abundance. Oikos, 58, 329–335. Gilbert, L. E. and Singer, M. C. (1973). Dispersal and gene flow in a butterfly species. American Naturalist, 107, 58-72. Grossmueller, D. W. (1987). The role of nectar source distribution in habitat use and oviposition by the tiger swallowtail butterfly. Journal of the Lepidopterists' Society, 41, 159-165. Hamer, K. C., Hill, J. K., Lace, L. A. and Langan, A. M. (1997). Ecological and biogeographical effects of forest disturbance on tropical butterflies of Sumba, Indonesia. Journal of Biogeography, 24, 67–75. Hanski, I. (1982). Dynamics of regional distribution: the core and satellite species hypothesis. Oikos, 38, 210–221. Hanski, I., Kouki, J., and Halkka, A. (1993). Three explanations of the positive relationship between distribution and abundance of species. In D. Schluter (eds.), Species Diversity in Ecological Communities: Historical and Geographical Perspectives (pp. 108–116). Chicago, University of Chicago Press. Hardy, P. B. and Dennis, R. L. H. (1999). The impact of urban development on butterflies within a city region. Biodiversity and Conservation, 8, 1261-1279. Hawkins, B. A. and Porter, E. E. (2003). Does herbivore diversity depend on plant diversity? The case of California butterflies. American Naturalist, 161, 40-49. Hill, J. K., Hamer, K. C., Lace, L. A. and Banham, W. M. T. (1995). Effects of selective logging on tropical forest butterflies on Buru, Indonesia. Journal of Applied Ecology, 32, 754–760. Hogsden, K. L. and Hutchinson, T. C. (2004). Butterfly assemblages along a human disturbance gradient in Ontario, Canada. Canadian Journal of Zoology, 82, 739-748. Holl, K. D. (1995). Nectar resources and their influence on butterfly communities on reclaimed coal surface mines. Restoration Ecology, 3, 76-85. Holl, K. D. (1996). The effect of coal surface mine reclamation on diurnal lepidopteran conservation. Journal of Applied Ecology, 33, 225-236. Hunter, M. L. Jr. (1996). Fundamentals of Conservation Biology. Cambridge, Blackwell Science. Inoue, T. (2003). Chronosequential change in a butterfly community after clear-cutting of deciduous forests in a cool temperate region of central Japan. Entomological Science, 6, 151-163. Ishii, M. (1993). Transect counts of butterflies. In O. Yata, and K. Ueda (Eds.), Decline and Conservation of Butterflies in Japan II (pp. 91–101). Osaka, The Lepidopterological Society of Japan and the Nature Conservation Society of Japan (in Japanese with English summary). Ishii, M. (1996a). Species diversity of butterfly communities in different environment of forests in southern Osaka. In B. Tanaka, and Y. Arita (Eds.), Decline and Conservation of Butterflies in Japan IV (pp. 63–75). Osaka, The Lepidopterological Society of Japan (in Japanese with English summary).

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 85 Ishii, M. (1996b). Decline and conservation of butterflies in Japan. In S. Ae, T. Hirowatari, M. Ishii and L. P. Brower (Eds.), Decline and Conservation of Butterflies in Japan III (pp. 157–167). Osaka, The Lepidopterological Society of Japan. Ishii, M., Hirowatari, T. and Fujiwara, S. (1995). Species diversity of butterfly communities in ‗Mt Mikusa coppice for Zephyrus‘. Japanese Journal of Environmental Entomology and Zoology, 7, 134–146 (in Japanese with English abstract). Ishii, M., Ueda, K., and Shigematsu, T. (1993). Conservation of the Nature of ‗Satoyama‘. Tokyo, Tsukiji Shokan Publishers (in Japanese). Ishii, M., Yamada, M., Hirowatari, T. and Yasuda, T. (1991). Diversity of butterfly communities in urban parks in Osaka prefecture. Japanese Journal of Environmental Entomology and Zoology, 3, 183-195 (in Japanese with English summary). Kamimura, Y. (2004). Nectar-source plants and their usage patterns by an adult butterfly community in a riparian biotope. Natural Environmental Science Research, 17, 107-115 (in Japanese with English summary). Kitahara, M. (1999). Species diversity of butterfly communities in a variety of woodlands at the northern foot of Mt Fuji, central Japan. Japanese Journal of Environmental Entomology and Zoology, 10, 11–29 (in Japanese with English abstract). Kitahara, M. (2000). Food resource usage patterns of adult butterfly communities in woodland habitats at the northern foot of Mt. Fuji, central Japan. Japanese Journal of Environmental Entomology and Zoology, 11, 61-81 (in Japanese with English summary). Kitahara, M. (2004). Butterfly community composition and conservation in and around a primary woodland of Mount Fuji, central Japan. Biodiversity and Conservation, 13, 917942. Kitahara, M. and Fujii, K. (1994). Biodiversity and community structure of temperate butterfly species within a gradient of human disturbance: an analysis based on the concept of generalist vs. specialist strategies. Researches on Population Ecology, 36, 187-199. Kitahara, M. and Fujii, K. (1997). An island biogeographical approach to the analysis of butterfly community patterns in newly designed parks. Researches on Population Ecology, 39, 23–35. Kitahara, M. and Sei, K. (2001). A comparison of the diversity and structure of butterfly communities in semi-natural and human-modified grassland habitats at the foot of Mt Fuji, central Japan. Biodiversity and Conservation, 10, 331–351. Kitahara, M., Sei, K. and Fujii, K. (2000). Patterns in the structure of grassland butterfly communities along a gradient of human disturbance: further analysis based on the generalist/specialist concept. Population Ecology, 42, 135-144. Kitahara, M. and Watanabe, M. (2001). Relationships of butterfly community diversity to vegetational species richness in and around the Aokigahara woodland at the northern foot of Mt. Fuji, central Japan. Japanese Journal of Environmental Entomology and Zoology, 12, 131-145 (in Japanese with English abstract). Kitahara, M. and Watanabe, M. (2003). Diversity and rarity hotspots and conservation of butterfly communities in and around the Aokigahara woodland of Mount Fuji, central Japan. Ecological Research, 18, 503-522. Kitahara, M., Yumoto, M. and Kobayashi, T. (2008). Relationship of butterfly diversity with nectar plant species richness in and around the Aokigahara primary woodland of Mount Fuji, central Japan. Biodiversity and Conservation, 17, 2713–2734.

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86

Masahiko Kitahara

Kocher, S. D. and Williams, E. H. (2000). The diversity and abundance of North American butterflies vary with habitat disturbance and geography. Journal of Biogeography, 27, 785-794. Kremen, C. (1992). Assessing the indicator properties of species assemblages for natural areas monitoring. Ecological Applications, 2, 203-217. Kremen, C., Colwell, R. K., Erwin, T., Murphy, D. D., Noss, R. F. and Sanjayan, M. A. (1993). Terrestrial arthropod assemblages-their use in conservation planning. Conservation Biology, 7, 796–808. Lawton, J. H. (1993). Range, population abundance and conservation. Trends in Ecology and Evolution, 8, 409–413. Leps, J. and Spitzer, K. (1990). Ecological determinants of butterfly communities (Lepidoptera, Papilionoidea) in the Tam Dao Mountains, Vietnam. Acta Entomologica Bohemoslovaca, 87, 182–194. Leps, J., Spitzer, K. and Jaros, J. (1998). Food plants, species composition and variability of the moth community in undisturbed forest. Oikos, 81, 538–548. Lincoln, R., Boxshall, G., and Clark, P. (1998). A Dictionary of Ecology, Evolution and Systematic (2nd edition). Cambridge, Cambridge University Press. Loertscher, M., Erhardt, A. and Zettel, J. (1995). Microdistribution of butterflies in a mosaiclike habitat: the role of nectar sources. Ecography, 18, 15-26. MacArthur, R. H. (1972). Geographical Ecology: Patterns in the Distribution of Species. New York, Harper and Row. MacArthur, R. H., and Wilson, E. O. (1967). The Theory of Island Biogeography. Princeton, Princeton University Press. Mano, T. (2004). Butterfly communities in the center of Toyota City. Yahagi-River Research, 8, 115-121 (in Japanese). Ministry of the Environment of Japan (2000). Red Data List of Insects in Japan. [http://www.biodic.go.jp/], Biodiversity Center of Japan (In Japanese). Ministry of the Environment of Japan (2001). The analysis of biodiversity in SATOCHI and SATOYAMA areas of Japan. Available at http://www.env.go.jp/nature/satoyama/ chukan.html (In Japanese). Ministry of the Environment of Japan (2007). The Third National Biodiversity Strategy of Japan. Available at http://www.biodic.go.jp/convention. Mittermeler, R. A., and Forsyth, A. B. (1997). Setting priorities for biodiversity conservation. In G. K. Meffe, C. R. Carroll, and Contributers (Eds.), Principles of Conservation Biology (2nd edition, pp. 141–142). Sunderland, Sinauer Associates Inc. Mittermeler, R. A., Myers, J. B., Thomsen, G. A. De Fonceca B. and Olivieri, S. (1998). Biodiversity hotspots and major tropical wilderness areas: approaches to setting conservation priorities. Conservation Biology, 12, 516–520. Munguira, M. L. and Thomas, J. A. (1992). Use of road verges by butterfly and burnet populations, and the effect of roads on adult dispersal and mortality. Journal of Applied Ecology, 29, 316-329. Murphy, D. D. (1983). Nectar sources as constraints on the distribution of egg masses by the checkerspot butterfly, Euphydryas chalcedona (Lepidoptera: Nympharidae). Environmental Entomology, 12, 463-466. Murphy, D. D., and Wilcox, B. A. (1986). Butterfly diversity in natural habitat fragments: a test of the validity of vertebrate-based management. In J. Verner, M. Morrison, and C. J.

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 87 Ralph (Eds.), Wildlife 2000: modeling habitat relationships of terrestrial vertebrates (pp 287-292). Madison, University of Wisconsin Press. Myers, N. (1988). Threatened biota: ‗hotspots‘ in tropical forests. Environmentalist, 8, 1–10. Myers, N. (1997). Global biodiversity II: losses and threats. In G. K. Meffe, C. R. Carroll, and Contributers (Eds.), Principles of Conservation Biology (2nd edition, pp. 123–158). Sunderland, Sinauer Associates Inc. Natuhara, Y. (2000). Changes in butterfly assemblage along the urban-forest gradient. Journal of the Japanese Institute of Landscape Architecture, 63, 515-518 (in Japanese with English summary). Natuhara, Y., Imai, C. and Takahashi, M. (1999). Pattern of land mosaics affecting butterfly assemblage at Mt Ikoma, Osaka, Japan. Ecological Research, 14, 105–118. Nature Conservation Bureau (1993). Report on the 4th Basic Research in Natural Environmental Conservation, with Special Reference to the Distribution Patterns of Japanese Animals and Plants (Volume Butterflies). Tokyo, Environmental Agency Government of Japan (in Japanese). New, T. R. (1991, 1997). Butterfly Conservation (1st edition, 2nd edition). Melbourne, Oxford University Press. Novotny, V. (1991). Effect of habitat persistence on the relationship between geographic distribution and local abundance. Oikos, 61, 431–433. Odum, E. P. (1971). Fundamentals of Ecology (3rd edition). Philadelphia, W. B. Sanders Co. Odum, E. P. (1989). Ecology and Our Endangered Life-Support Systems. Sunderland, Sinauer Associates Inc. Pianka, E. R. (1970). On r- and K-selection. American Naturalist, 102, 592–597. Pianka, E. R. (1988). Evolutionary Ecology (4th edition). New York, Harper and Row. Pivnick, K. A. and McNeil, J. N. (1987). Diel patterns of activity of Thymelicus lineola adults (Lepidoptera:Hesperiidae) in relation to weather. Ecological Entomology, 12, 197-207. Pollard, E. (1977). A method for assessing changes in the abundance of butterflies. Biological Conservation, 12, 115-134. Pollard, E. (1984). Synoptic studies on butterfly abundance. In R. I. Vane-Wright, and P. K. Ackery (Eds.), The Biology of Butterflies (pp. 59-61). London, Academic Press. Pollard, E., and Yates, T. J. (1993). Monitoring Butterflies for Ecology and Conservation. London, Chapman and Hall. Pressey, R. L. (1996). Protected areas: where should they be and why should they be there. In I. F. Spellerberg (Eds.), Conservation Biology (pp. 171–185). Essex, Longman. Primack, R. B. (1993). Essentials of Conservation Biology. Sunderland, Sinauer Associates Inc. Primack, R. B. (1995). A Primer of Conservation Biology. Sunderland. Sinauer Associates Inc. Pryke, S. R. and Samways, M. J. (2003). Quality of remnant indigenous grassland linkages for adult butterflies (Lepidoptera) in an afforested African landscape. Biodiversity and Conservation, 12, 1985-2004. Pywell, R. F., Warman, E. A., Sparks, T. H., Greatorex-Davies, J. N., Walker, K. J., Meek, W. R., Carvell, C., Petit, S. and Firbank, L. G. (2004). Assessing habitat quality for butterflies on intensively managed arable farmland. Biological Conservation, 118, 313325. Reid, W. V. (1998). Biodiversity hotspots. Trends in Ecology and Evolution, 13, 275–280.

Woodlands : Ecology, Management and Conservation, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved.

88

Masahiko Kitahara

Ries, L., Debinski, D. M. and Wieland, M. L. (2001). Conservation value of roadside prairie restoration to butterfly communities. Conservation Biology, 15, 401-411. Sakuratani, Y. and Fujiyama, S. (1991). Influence of highway construction on butterfly communities. Japanese Journal of Environmental Entomology and Zoology, 3, 15-23 (in Japanese with English summary). Schmitt, T. (2003). Influence of forest and grassland management on the diversity and conservation of butterflies and burnet moths (Lepidoptera, Papilionoidea, Hesperiidae, Zygaenidae). Animal Biodiversity and Conservation, 26, 51-67. Schneider, C. and Fry, G. L. A. (2001). The influence of landscape grain size on butterfly diversity in grasslands. Journal of Insect Conservation, 5, 163-171. Sei, K. (1988). The butterflies unable to have inhabited in the Mt. Fuji area. Tokyo, Tsukiji Shokan. (in Japanese). Seido, K. (1991). Age of the hinoki (Chamaecyparis obtusa) natural stands and their distribution on the Aokigahara Woodland, Mt. Fuji. Transactions of the Meeting in Kanto Branch of the Japanese Forestry Society, 42, 33-36 (in Japanese). Sharp, M. A., Parks, D. R. and Ehrlich, P. R. (1974). Plant resources and butterfly habitat selection. Ecology, 55, 870-875. Shreeve, T. G., Dennis, R. L. H., Roy, D. B. and Moss, D. (2001). An ecological classificationof British butterflies: ecological attributes and biotope occupancy. Journal of Insect Conservation, 5, 145-161. Shreeve, T. G. and Mason, C. F. (1980). The number of butterfly species in woodlands. Oecologia, 45, 414-418. Simonson, S. E., Opler, P. A., Stohlgren, T. J. and Chong, G. W. (2001). Rapid assessment of butterfly diversity in a montane landscape. Biodiversity and Conservation, 10, 13691386. Simpson, E. H. (1949). Measurement of diversity. Nature, 163, 688. Southwood, T. R. E. (1977). Habitat, the templet for ecological strategies? Journal of Animal Ecology, 46, 337–365. Southwood, T. R. E. (1978). Ecological Methods, with Particular Reference to the Study of Insect Populations. London, Chapman and Hall. Southwood, T. R. E. (1981). Bionomic strategies and population parameters. In R. M. May (Eds.), Theoretical Ecology: Principles and Applications (2nd edition, pp. 30–52). Oxford, Blackwell Scientific Publications. Southwood, T. R. E. (1988). Tactics, strategies, and templets. Oikos, 52, 3–18. Southwood, T. R. E., May, R. M., Hassell, M. P. and Conway, G. R. (1974). Ecological strategies and population parameters. American Naturalist, 108, 791–804. Sparks, T. H. and Parish, T. (1995). Factors affecting the abundance of butterflies in field boundaries in Swavesey Fens, Cambridgeshire, UK. Biological Conservation, 73, 221227. Spitzer, K., Jaros, J., Havelka, J. and Leps, J. (1997). Effect of small-scale disturbance on butterfly communities of an Indochinese montane rainforest. Biological Conservation, 80, 9-15. Spitzer, K., Novotny, V., Tonner, M. and Leps, J. (1993). Habitat preferences, distribution and seasonality of the butterflies (Lepidoptera, Papilionoidea) in a montane tropical rain forest, Vietnam. Journal of Biogeography, 20, 109-121.

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Diversity Patterns, Adult Resource Use and Conservation of ButterflyCommunities… 89 Steffan-Dewenter, I. and Tscharntke, T. (1997). Early succession of butterfly and plant communities on set-aside fields. Oecologia, 109, 294-302. Tashita, M. and Ichimura, T. (1997). Environmental evaluation by butterfly communities according to altitude in Nagano Prefecture. Japanese Journal of Environmental Entomology and Zoology, 8, 73–88 (in Japanese with English abstract). Thomas, C. D. and Mallorie, H. C. (1985). Rarity, species richness and conservation: butterflies of the Atlas Mountains in Morocco. Biological Conservation, 33, 95-117. Thomas, J. A. (1983). A quick method for estimating butterfly numbers during surveys. Biological Conservation, 27, 195-211. Tiple, A. D., Deshmukh, V. P. and Dennis, R. L. H. (2006). Factors influencing nectar plant resource visits by butterflies on a university campus: implications for conservation. Nota lepidopterologica, 28, 213-224. Tsuya, H. (1971). Topography and geology of Volcano Mt. Fuji. Report on the synthetic academic surveys of Mt. Fuji (pp. 1–149). Fujiyoshida, Yamanashi, Fujikyuko (in Japanese with English summary). Tudor, O., Dennis, R. L. H., Greatorex-Davies, J. N. and Sparks, T. H. (2004). Flower preferences of woodland butterflies in the UK: nectaring specialists are species of conservation concern. Biological Conservation, 119, 397-403. Unno, K. and Aoyama, J. (1981). The Butterflies of Japan. Tokyo, Shougakukan (in Japanese). Usher, M. B. and Keiller, S. W. J. (1998). The macrolepidoptera of farm woodlands: determinants of diversity and community structure. Biodiversity and Conservation, 7, 725-748. Väisänen, R. (1992). Distribution and abundance of diurnal Lepidoptera on a raised bog in southern Finland. Annales Zoologici Fennici, 29, 75-92. Warren, M. S. (1985). The influence of shade on butterfly numbers in woodland rides, with special reference to the wood white Leptidea sinapis. Biological Conservation, 33, 147164. Warren, M. S., Pollard, E. and Bibby, T. J. (1986). Annual and long-term changes in a population of the wood white butterfly Leptidea sinapis. Journal of Animal Ecology, 55, 707–719. Washitani, I. (1999). Introduction to New Ecology: Ecology for Biological Conservation. Tokyo, Kyouritsu Shuppan Co (in Japanese). Weiss, S. B., White, R. R., Murphy, D. D. and Ehrlich, P. R. (1987). Growth and dispersal of larvae of the checkerspot butterfly Euphydryas editha. Oikos, 50, 161–166. Wood, B. and Gillman, M. P. (1998). The effects of disturbance on forest butterflies using two methods of sampling in Trinidad. Biodiversity and Conservation, 7, 597–616. Yamamoto, M. (1988). On the research method for butterfly assemblages. Special Bulletin of the Lepidopterological Society of Japan, 6, 191–210 (in Japanese with English abstract). Yata, O. (1996). Butterfly community in laurel-leaved forest of Yamada park, Kitakyushu. In B. Tanaka, and Y. Arita (Eds.), Decline and Conservation of Butterflies in Japan IV (pp. 49–56). Osaka, The Lepidopterological Society of Japan (in Japanese with English summary).

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90

APPENDIX I. LIST OF BUTTERFLY SPECIES OBSERVED IN THE PRESENT STUDY, THEIR POPULATION DENSITIES, AND THEIR CHARACTERISTICS (KITAHARA AND WATANABE, 2003)

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Species

OL-1

OL-2

Population density FE-1 FE-2

FI-1

Hesperiidae Erynnis montanus 3.33 5.83 4.17 1.67 Daimio tethys 3.33 Leptalina unicolor 1.11 11.11 3.33 Thymelicus sylvaticus 1.67 1.67 Ochlodes venatus 1.67 Ochlodes ochraceus 7.41 9.72 Potanthus flavus 1.67 Pelopidas jansonis 1.67 1.94 Parnara guttata 2.50 15.00 8.89 15.83 Papilionidae Parnassius glacialis 4.72 20.83 18.89 Graphium sarpedon 1.67 Papilio machaon 2.50 Papilio macilentus 2.22 Papilio protenor 1.11 Papilio bianor 1.67 1.39 2.5 Pieridae Leptidea amurensis 2.59 4.17 Eurema hecabe 3.33 5.00 7.04 9.03 Gonepteryx rhamni 3.33 Gonepteryx aspasia 1.67 Colias erate 9.37 11.76 4.44 4.63 Anthocharis scolymus 1.67 Pieris rapae 3.15 13.49 2.92 2.41 Pieris melete or napi 3.33 15.78 15.19 13.67 2.22 Lycaenidae Narathura japonica 2.22 1.67 Antigius attilia 1.67 Chrysozephyrus smaragdinus 5.00 Favonius jezoensis 0.83 Rapala arata 1.39 1.48 Callophrys ferrea 1.67 1.67 Lycaena phlaeas 1.94 6.56 5.37 2.01 Lampides boeticus 6.67 6.67 Pseudozizeeria maha 6.67 3.33 1.67 Celastrina argiolus 1.67 3.06 1.32 1.67 Celastrina sugitanii 1.67 Everes argiades 2.22 1.39 2.50 2.92 Plebejus argus 4.44 15.83 1.53 Lycaeides subsolanus 1.39 11.11 Curetis acuta 1.39 Libytheidae Libythea celtis 4.17 1.39 1.48 3.06 4.81 Danaidae Parantica sita 1.11 Nymphalidae Argyronome ruslana 1.67 2.50 Argynnis paphia 3.33 3.33 5.83 4.44 Nephargynnis anadyomene 2.78 1.11 Fabriciana adippe 2.78 2.04 Limenitis camilla 1.57 3.15 Limenitis glorifica 1.67 Neptis sappho 1.48 8.19 6.39 1.11 Neptis philyra 1.11 2.22 1.11 Neptis rivularis 1.67 Polygonia c-aureum 1.67 11.98 7.66 9.44 Polygonia c-alubum 1.67 Kaniska canace 1.39 3.33 1.94 Inachis io 1.67 2.50 1.25 Cynthia cardui 4.03 1.48 0.56 Vanessa indica 0.83 Satyridae Ypthima argus 3.61 3.75 2.78 Minois dryas 14.17 22.50 13.33 † Number of larval potential host-plant species cited from the literature (see text). ‡ W (wood), H (herbaceous). § Number of study sites observed. ¶ Number of 10–10 km2 in which the species was recorded (see text). †† VU: vulnerable species, NT: semivulnerable species.

FI-2

Average

Voltinism

Potential larval diet breadth†

1.67

3.33 3.33 5.19 1.67 1.67 8.56 1.67 1.81 10.56

1 2 1 1 1 1 1 2 3

6 6 7 6 14 12 21 7 28

W H H H H H H H H

5 1 3 2 1 2 1 2 4

636 781 256 177 352 500 494 285 898

14.81 1.67 2.50 2.22 1.11 1.85

1 2 3 2 2 2

5 14 42 5 15 10

H W H W W W

3 1 1 1 1 3

599 738 1098 639 871 1152

3.38 5.55 3.33 1.67 7.55 1.67 5.49 10.04

2 3 1 1 3 1 3 3

1 30 1 4 21 18 25 16.5

H W W W H H H H

2 5 1 1 4 1 4 5

144 1301 47 377 1179 704 1209 902

10.00

1.94 1.67 5.00 1.11 1.44 1.67 3.97 6.67 3.89 1.99 1.67 2.26 7.27 6.25 5.69

1 1 1 1 2 1 3 3 3 3 1 3 1 1 2

13 11 14 8 25 18 5 41 2 32 3 31 38 9 10

W W W W W W H H H H W H H H W

2 1 1 2 2 2 4 2 3 5 1 4 3 2 2

556 561 345 260 695 597 1179 461 905 1276 325 946 274 86 672

3.06

2.99

1

4

W

6

820

8.33

4.72

?

12

H

2

660

2.08 4.06 1.94 2.41 2.36 1.67 3.77 1.39 1.67 6.82 1.67 2.22 1.81 2.02 0.83

1 1 1 1 2 2 3 1 1 3 2 2 2 3≤ 3

1 6 2 – 8 4 22 10 7 2 9 13 7 9 13

H H H H W W W W W W W H H H H

2 5 2 2 2 1 5 4 1 5 1 3 3 3 1

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Larval Local Geographic Status in hostplant distribution range Red Dat type‡ pattern§ size¶ list of Japan††

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In: Woodlands: Ecology, Management and Conservation ISBN 978-1-61122-542-6 Editor: Erwin B. Wallace ©2011 Nova Science Publishers, Inc.

Chapter 3

GUIDELINES FOR SUSTAINABLE MANAGEMENT OF DEGRADED LANDS: EXPERIENCES ON CAATINGA AND SEMI-ARID MEDITERRANEAN WOODLANDS Miguel Ángel Herrera Machuca1, Rinaldo Luiz Caraciolo Ferreira2,† Juan Ramón Molina Martínez1,‡ and Mércia Virginia Ferreira dos Santos3#

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1.

Departamento de Ingeniería Forestal, Universidad de Córdoba, Campus de Rabanales. Edificio Leonardo da Vinci. Córdoba 14071, Spain 2. Department of Forest Science; Universidade Federal Rural de Pernambuco, Rua Manoel de Medeiros, s/n, Dois Irmãos, Recife, Pernambuco, Brasil 3. Department of Zootechnics; Universidade Federal Rural de Pernambuco, Rua Manoel de Medeiros, s/n, Dois Irmãos, Recife, Pernambuco, Brasil

ABSTRACT Land degradation and soil erosion are perceived as important and worldwide problems in dryland areas. In semi-arid climate conditions, interactions between vegetation, rainfall and soil properties have been related to the effectiveness in reducing the degredation processes. Landscape structure can be characterized by different woodlands based on the structure and physiognomy of the vegetation communities. From the perspective of water availability, the shortage of rainfall during summer is the main limitation for livestock development. Soil properties limitations are vital in order to the soil erosion and soil water availability. On the other hand, climate change, socio

E-mail: [email protected]. E-mail: [email protected]. ‡ E-mail: [email protected]. # E-mail: [email protected]. †

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M. Á.Herrera Machuca, R. L. C. Ferreira, J. R. M. Martínez et al. economic changes and cultural evolution have been associated with the gradual loss of ecosystem productivity due to biophysical stress limiting the environmental and human sustainability. In Brazil, the semi-arid climate zone is located on the Northeastern of the country, an area of 845,000 km2 showing about of the 11% of the Brazil area. In the case of Spanish semi-arid area, woodlands tend to be shrublands because of the limited environmental conditions and the recent changes in land use. Brazil and Spanish semi-arid landscapes have some common characteristics. The socio-economic development of their societies have required aggressive practices such as land use change, land occupation, burning, deforestation and overgrazing. In semi-arid ecosystems, species are adapted to environmental and subsistence activities on a local level. In this paper, field experiences associated to Brazil and Spain woodlands are analyzed in order to find technical alternatives on the urgent need to semi-arid landscape conservation.

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INTRODUCTION Thirty-five percent of earth‘s people live in arid and semi-arid lands. Drylands cover forty-one percent of the planet and closely follow the world‘s map of poverty. While already exposed to climate extremes, according to IPCC, drylands are likely to be severely hit by climate change (ICID, 2010). Currently, about one-third of terrestrial surface is likely to affect of arid and semi-arid climate conditions in both North and South hemispheres. Different degradation processes (physical, chemical and biological) have an effect on these zones increasing the possibilities to the apparence of new desertization lands (Verstraete and Swchartz, 1991). Landscape and ecological processes are important sources of international environmental problems, both developing and industrialized countries. Arid and semi-arid areas assemble complex natural formations, disperse in several points of the planet and very diverse among them, but presenting common identification features that make them singular when compared to other ecosystems. Vegetation is adapted to the amount precipitation (Rivas Martínez., 1987). Rainfall changes would have consequences for vegetation (Lázaro et al., 2001) as well as the management recommendations. Climatic conditions such as temperature, moisture and annual precipitation, and soil characteristics such as horizon thickness, organic matter content, depth and structure, are important for ecosystem health and hillslope hydrology. In this sense, the environmental limitations of semi-arid areas have configured the vegetation cover that tends to be low and sparse (Peter et al., 1996). Most of the semi-arid ecosystems are characterized by a spatially discontinuous vegetation cover that can be randomly or clumped distributed. The distribution of runoff and erosion is directly related to this spatial distribution of the vegetation. Changes in the spatial pattern can be used as an indicator of desertification and applied to developing rehabilitation strategies. Some authors have assumed that arid and semi-arid ecosystems are driven more by external such as soil moisture than by internal factors (Noy-Meir, 1973). There is growing evidence that increasing concentrations of carbon dioxide in the atmosphere will have a longterm effect on the world climate. Global circulation models allow simulating the future evolution of climate according to different forcing scenarios related to human activities (Gutiérrez and Pons, 2006). Average temperature will increase some degrees based on the

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geographical position over the next 50 years. Climate conditions induce significant changes of vegetation floristic composition. In this sense, recent climate change influences a variety of ecological and socio-economic processes (Steenseth et al., 2002). The magnitude of these climate changes could be sufficient to bring about long-term changes in agricultural processes. Under these circumstances together with temperature increase lead to less water availability and soil organic matter content, and as a consequence, soil permeability and infiltration rate decrease increasing runoff process. These climate and socio-economic changes have increased the frequency and severity of wildfires (Flannigan et al., 2006, Vélez, 2009). Brazilian drylands are concentrated in the semi-arid region of NE Brazil. It presents the most diverse landscape in Brazil, both in geomorphological features and vegetation types. This environmental diversity must reflect in increasing biodiversity, complicated taxonomy and complex biogeographical patterns occurring in a relatively small scale. This situation of landscapes, vegetations and biodiversity happening in mosaics are a huge challenge not only for taxonomic and ecological studies but, especially, for conservation purposes (Queiroz et al., 2010). Desertification problem affects 31.49 % of the Spanish covering about 160,000 km2 (Molina et al., 2010). In semi-arid Spain is difficult to know which landscape was present previous to agricultural practices (Bonet, 2004). Although the changes in land use and population decline during the last 50 years have led to extensive revegetation with an increase in shrub cover in the upper mountain areas (Alados et al., 2004; Lasanta et al., 2006), agricultural lands have break the exploitation because of the low profits. Slope and altitude appears as important factors in the abandonment process due to the low benefits. Agricultural lands characterized by these different conditions for farming are the first to be abandoned (Bonet, 1997). This abandonment of agricultural practices in semi-arid areas during the last decades has promoted slow changes in cover of the different species. Finally, pines plantations have been used for land restoration in the Mediterranean area since 19 th century due to its stress-tolerant and pioneer features (Pausas et al., 2004). Landscape of agricultural lands and plantations is incorporated towards enhancing the ecological, cultural and aesthetic character of semi-arid regions (Makhzoumi, 1997). In semi-arid regions the patch-size distribution of the vegetation follows a power law (Kéfi et al., 2007) concerning the need to understand relationships between vegetation and hydrological processes (Imeson and Prensen, 2004). Fragmentation and aridity are the main determinants of vegetation recovery and distribution (Pueyo and Alados, 2007). In semi-arid areas, conservation and management guidelines are very important due to soil degredation and water availability. This paper attempts to analyze woodlands management in two different semi-arid scenarios: Brazilian and Mediterranean landscapes, emphasizing the technical actions to reach better results on the conservation and/or restoration of these limited ecosystems.

ENVIRONMENTAL CONDITIONS This paper was conducted in two different countries: Brazil and Spain where semi-arid climate zones are relevant. In the first case because of its area; and in the second according to

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its special vulnerability by climate change (Figure 1). These two semi-arid regions constitute an assembly of complex natural formations, very differentiated among them. Human activities have made use of the natural land in many different ways, from land occupation to land use. In spite of the socio-economic differences, some common recomendations and conclusions are presented. Semi-arid technical experiences developed in each country could be used or adapted in another country.

Brazilian Semi-Arid Area

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The Northeast Brazil is characterized by the diversity of landscapes, and the Caatinga stands out as the only biome exclusively Brazilian, with an area of 844,453 km2 (IBGE, 2004) fully inserted into the semi-arid climate. The Caatingas zone lies between 3° and 15° S, 34° and 45° W. There are several types of Caatingas, which are generally described as a dwarfed vegetation formation, deciduous in the dry period, comprising many different types of cactus and thorn-bushes which are perfectly suited to drought conditions (Bellefontaine et al. 2000).

Figure 1. Semi-arid areas of the two study countries.

In Brazil, tropical semi-arid climate is characterized by the amount of precipitation, with a very hight interannual and intraannual variability, strong annual cycles and a summer drought. Although in some years the isohyetal 1,000 is showed, the mean annual rainfall is between 400 and 800 mm (Brasil, 2005). The rain volume is concentrated on an important seasonal time including between three and six interannual and intraannual months. Potential evapotranspiration is always high, between 1,500 and 2,000 mm per year, indicating an independence of the amount and fluctuaction in rainfall.

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On the other hand, Brazilian semi-arid soils have a complex spatial distribution modelling a mosaic landscape with different characteristics. The spatial soil horizon attributes vary based on the soil type (sandy, clay), structure, depth, organic matter content, rock cover and texture. Using this information and topographic characteristics it is possible to describe the spatial fertility and productivity (Velloso et al., 2002). The agroforestry system is widely practiced in the semiarid region, formed by livestock under extensive grazing on Caatinga vegetation, timber resources exploitation and shifting cultivation with the use of burning techniques for the production of subsistence crops, under or not a spatial or sequential arrangement (Santos et al., 2010). Generally, there is a tendency of high pressure of Caatinga utilization because of the land tenure estructure composed by private small properties that are smaller than 100 ha on 90% of the its total area (IBGE, 1997). The agroforestry system would be the factor that exerts most pressure upon Caatinga and this effect varies on intensity as a function of location, estructure and size of the forest remainings according to Andrade et al. (2005). In this sense, Kumazaki (1992), studing the pressure of human actions upon forest remainings, emphasized that as smaller the area, the greater the impacts of human actions and usually conservation becomes unfeasable. Caatinga has a large number of plant species that varies according to stratum, year, methodology used, and human actions, but most surveys refer to shrubs and trees. This vegetation is characterized as a mosaic of thorny shrubs and dry seasonal forests, with more than 2,000 species of vascular plants, fish, reptiles, amphibians, birds and mammals. The endemism in these groups ranges from 7 to 57% in the Caatinga (Leal et al., 2005). Plant species from the Brazilian Semi-arid are an important resource to human populations, mainly to the ones who live in the Caatinga. The ‗sertanejos‘, native inhabitants of Caatinga, are able to survive the hard conditions imposed by the Caatinga because of their ability to interact with the environment, taking from it a substantial part of their bare necessities. Most of the animal farming in the Caatinga is extensive and the animals, particularly goats and cattle, feed on several native species of legumes, grasses and ‗velames‘ (Croton species). Besides that, the main source of energy is the firewood from several species of plants (Queiroz, 2010).

Mediterranean Semi-Arid Area The Mediterranean climate has a limited capacity in southeastern Spain. In this sense, sub-tropical sub desert mesotype covers about 14,000 km2 of the Spanish area (Allue, 1990). This climatic area lies between 36º and 40º N, 0º and 4º W. Annual mean temperature is between 16ºC and 21ºC and highest monthly temperature is between 24ºC and 33ºC. Mean annual precipitation varies between 200 and 500 mm based on the geographical position on this climatic mesotype. The study was conducted in the area of Almeria and Granada provinces that is characterized by the above mesotype. The sector selected is between 36º40´ and 37º30 latitude north and 1º30´ and 3º30´longitude west. The area (approx. 425,000 ha) has an important drought period and a very high interannual variability. Calcareous soils are dominant along semi-arid hillslopes. Rock cover varies spatially showing higher on southfacing slopes than on north facing slopes (Poesen et al., 1998). Soil water content was found to be hightly spatially heterogenous within small areas, controlled mainly by soil surface

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cover and soil properties (Cantón et al., 2004). Semi-arid species have acess to low reserves of water depp in the soil which enabled them to survive. The current vegetation is dominated by Stipa tenacissima L. that is presented on 36.07% of the Mediterranean study area. Pure Stipa tenacissima steppes cover about 19.87% resulting of shrubland degradation (Rivas Martínez, 1987) and abandoned lands (González Bernaldez, 1991). Other important formations are shrublands of Anthyllis cytisoides L., Genista umbellata Poiret., Cytisus fontanesii Spach., Thymus spp. and/or Arthrocnemum spp.; and dry grasslands dominated by Brachypodium retusum Pers. Beauv. and Hypauhenia hirta (L.) Stapf. Dwarf and sparse presence of Olea europarea var. sylvestris L., Quercus coccifera L. and Chamaerops humilis L. can be found. These dominant species are mixed with afforested Pinus halepensis Miller mainly on north-facing areas slopes.

CULTURAL EVOLUTION Modelling the impact of human activities on semi-arid ecosystems requires a multitemporal perspective. In order to understand the landscape in semi-arid areas, land use change and current socio-economic conditions must be analyzed over time: short and long term consequences. In this sense, we proposed these two temporal analyses for each study area.

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Brazilian Semi-Arid Area a) Long Term Unsustainable forest practices are intensifying day to day across the Brazilian history. Similar to other areas such as tropical countries and Spain, land use can be found in relation to hillslope gradient. The occupation of northeastern semi arid, specially Caatinga, began in a period called cattle cicle, named like this because cattle raising was the main economic complementary activity to the most wealthy activity of that time, the sugar cane, and developed along São Francisco river (―Rio dos Currais‖) during XVI, XVII and XVIII centuries (Prado Júnior, 1969). This activity was performed into an extensive system without adequate care of the soil; therefore cattle breeding became not only the responsible for the semi arid population, but also one of the main causes of Caatinga vegetation degradation. Land use was in the context of tangible assets such as timber, non-timber forest products and agroforestry systems. Caatinga woodlands changes were caused by production costs leading to grasslands and croplands (Viana et al., 1998). These land use changes affected all Brazilian biomes, but in semi-arid climate zone Caatinga was reduced to 50% by farming, soil salinity, soil erosion and urban activities. Caatinga is one of the most degraded biomes in Brazil by human activities and pressures (Sampaio et al., 2003). For centuries, overgrazing has exploited semi-arid ecosystems wih serious effects on soil and vegetation degradation, and as a consequence, increasing the degredation land (Velloso et al., 2002). Agribusiness is the main economic activity in clearly relation with the annual rainfall cycle. Agroforesty systems are used in two systems of policulture using agricultural crop, farming and timber harvest.

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Traditional agricultural revenue crop (between two and five years) based on burning practices is opposite to native vegetation and soil restoration that vary between ten and twenty years according to human pressure and the intensification of agricultural land use.

b) Short Term The monitoring of the Caatinga shows that 0.33% of its biomass is annually transformed into charcoal to be used as a source of energy, in the region where the biome is present as well as in others. This situation, which shows a growing demand of charcoal, leads to deforestation levels comparable only to those of the Amazon when the prevention programs began to be implemented in the forest (MMA, 2010). Combating deforestation in the Caatinga is justified by the need to control, as soon as possible, the deforestation in the biome, the only exclusively Brazilian, rich in biodiversity, but that has already lost 45% of its forest cover. The northeast Brazil degredation is a specific social problem that is directly in relation to the balance between the region population (quantum) and the required needs to food and to preserve environment conditions under the cultural standards and the limitations of its current system production (Ab'Sáber, 1995). In other words, there is a necessity to develop and improve the production system reaching the sustainability goals. One important point of view, it is the population density of the Northeast region (33 people per km2) that is one of the world´s highest density on semi-arid zones. In relation with this important population, economic activities are subsistence agriculture in small scale where agricultural and farming practices are developed by a low technological level leading to the aggressive agricultural practices, and as a consequence, the degradation of drylands.

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Mediterranean Semi-Arid Area a) Long Term Some studies show that during the Holocene period no significant forest cover existed in the Spanish study area (Pantaleon et al., 2003). This period (7,000 and 4,500 yr BP) modified the landscape that reflects the establishment of the steppe conditions that persist today. The dominance of steppe ecosystems leads to impact of high erosion processes and the marginalization of the canopy landscapes. Economic growth in this area was based mainly on the explotation of natural resources. Traditional land use activity such as sheep and goat farming has shaped the landscape for centuries (Barberá et al., 1997). This, by itself, need not be a problem, except that grazing resource had used to maximize revenues without consideration for long term sustainability of the semi-arid ecosystems. In the last century, aggressive practices were used causing increasing of productivity loss. This grassland condition is worsened by climatic change that increases the landscape vulnerability. b) Short Term Land use history determines the current mosaic landscape and the effects in semi-arid areas influence the dynamic succession. One research assessed land cover and landscape patterns using 1957, 1985 and 1994 years as reference. The results show that steppes and arid

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garrigues increased by 6% and 4%, respectively, while crop fields decreased by 15% (Alados et al., 2004). The transition from natural shrubland to steppe appeared to be found by gradual slopes. In Spain, forest management policies have traditionally encouraged land cover changes, with the establishment of tree cover in strategic or degraded ecosystems (Chirino et al., 2006). These actuations lead to reduce soil erosion and to increase the vertical structure and landscape diversity. In our study area, the forest policies have shaped a mosaic landscape of a mixed formation of rangelands and shrublands with Pinus halepensis cover (1.55% of the total study area). Mediterranean landscapes had influenced by a variety of disturbances of natural and human origin. In semi-arid area the interannual variation in rainfall is the factor that most affects net income. However, the effect of climate change on annual precipitation is a phenomenon whose impacts need be investigared in depthly. The application of general models of atmosphere circulation suggests a high probability of a temperature increase. Under these climatic change scenarios, evapotranspiration increases reducing water availability (Domingo et al., 1999). The economic development of society requires modifications of the natural environments. In the study area, it is possible to see the difference between the coastal buffer area and rest of the semi-arid zone. Steppes increase in calcareous substrate and at higher elevations within relation with anthropogenic influence. Attention should be given to vegetation conservation because of the relationship between natural remnant fragmentation and human activities. As a consequence, shrublands must be favoured by integral management planning in relation with bare soil and steppes areas where could develop human activities.

GLOBAL CHANGE AND CURRENT TRENDS Brazilian Semi-Arid Area Amongst the Brazilian biomes, the Caatinga is one of the least known from scientific standpoint and has been treated with low priority for purposes of biodiversity conservation. In spite of it, the Biome of Caatingas is highly endangered because of inadequate use of natural resources and the little area (less than 1%) under the protection of conservation units (FrancaRocha et al., 2010). Dantas et al. (2010) estimated that around 70% of Caatinga is already altered by human actions and that only 0.28% of its area is protected, as conservation units. Deforestation and use of fire in Caatinga areas are associated to climate changes with an estimated elevation of temperature around 2°C and 5°C, and as a consequence, they can be the cause of the Caatinga replacement by a more arid vegetation. Estimates indicate that one third of the economy of the Northeast, where 80% of the biome is, may disappear (MMA, 2010). In this sense Nobre et al. (2007) concluded that, as a function of the scenery from IPCC, Caatinga area will retreat and undergo desertification. For CEDEPLAR/UFMG and FIOCRUZ (2008), the climate projections for 2000-2050 point out that semi arid is one of the most vulnerable areas to climate changes in Brazil, because within forecasts there are the

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decrease of rain occurence, low fertility of soil, vegetation with less biological diversity and some places may even turn to be uninhabitable. The use of fire is common practice in preparing land for agriculture that destroy vegetal covering and compromise water quality and the balance between climate and soil (Fernandes et al., 2009). The use of fire is usual in semi arid region for pasture formation, mainly because of its low cost. However, the decrease of soil humidity due to this practice, is a negative aspect into a system that already suffer of water shortage. Besides, its use along time decrease floristic diversity and then grasses usually are predominant, as can be observed at areas that have been historically managed like this. The deforestation within 2002-2008 period was 2,763 km2/yr, considering the average amount of carbon in Caatinga of 25 t/ha, the estimated emission of CO2 was 25 million of t/ha (MMA, 2010). On the other side, standing biomass of native Caatinga shows nearly the full global range with 2-50 Mg C/ha. Litter fall around 1-2 Mg C/ha/yr is partly decomposed and partly consumed by animals, resulting in low average soil C levels near 8 g/kg, or 20 Mg C/ha. Under cultivation, C sequestration decreases, and soils losses approximately half their C stocks before being abandoned (Tiessen et al., 1998). Even thus, according to FAO (2007), the impact of degradation upon global carbon cycle and the potential impact of degradation control upon carbon capture in the semi-arid ecosystems have not been broadly investigated, there are few case studies and information and, consequently, there is few scientific evidence concerning the impact of degradation upon carbon emissions to the atmosfere. Considered one of the most vulnerable to climate change, the biome Caatinga is one of the planet's regions that will be most affected by global warming. In Caatinga conservation, the biggest threats come from fuelwood exploitation, grassland opening for cattle and goat farming and irrigation techniques which increase desertification processes such as soil erosion, deforestation and landscape degradation (Leal et al., 2005). The use and the lack of technical information about these procedures can become irreversible in fragile ecosystems (Santana and Souto, 2006).

Mediterranean Semi-Arid Area Two global and regional climatic models (CGCM and ECHAM) have been used to assess ecological and socio-economic effects at the landscape level. The application of these models on the Almeria province would predict changes in temperature between two and four degress (Molina, 2008). On the other hand, precipitation prediction is not understood even qualitatively, much less with any confidence for quantitative prediction. Andalusia territory has changed over 40% between 1956 and 2003. In the study area, recent changes in land use have occurred because of the climatic warning and growing demand on recreation activities among the natural resources of the forest. The development of tourist on the coast has led to the appearance of new land uses tending in some cases to the degradation of woodlands. The desertification processes and development of tourist infrastructure has expanded, with no-woodland areas increase from 32.95% to 34.21% of the study area during only five years (1999-2003 period). Desertification of semi-arid land can be the most threatened landscapes undergoing a degradation processes. In fragile ecosystems, desertification can become irreversible decreasing permanently the capacity to traditional sustain human activities. In last decades,

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the abandonment of traditional grazing areas plays an important role in the vegetation dynamics and fire risk. Land abandonment under condition of drought seems to promote soil erosion. However, about 10 yr after abandonment the site conditions produce almost no runoff and erosion (Cerda, 1997). In the semiarid Andalusian area, drought periods must be understood as part of its environment. In order to reduce the drought impacts, policies would be focused to improve the water management and to establish special plans. Wildfires have been implicated as an agent of desertification in semi-arid regions. Fire managers recognize the importance of the climate conditions in explaining the fire ignition and spread. Low precipitation (drought periods) and relative humidity coupled with high and prolonged temperatures increase the wildfire risk. Climate change causes long term droughts contributing to the increased in fire risk (Piñol et al., 1998; Pausas, 2004). Fire frequency and intensity have increased dramatically in the Mediterranean study area, resulting in major ecological and socio-economic losses on forests and on rural and urban people and economies. As an example, we show the official statistic of the last decade in Almeria province emphasizing the 8,571 ha of the last year and the 103 forest fires occurred in 2008 on semi-arid area. The variety of fire management strategies employed in wilderness areas has elicited much public concern, especially in the last decade with critical budget limitation. Prescribed fires may be a strategy to mitigate wildfire impacts due to benefits in terms of hazard reduction. Prescribed fire activities must be avoided under wide circumstances. Andalusian Government with fire management responsibilities is designing a specific law about the use of fire to involve reduction of fuel hazards and surface fuels accumulating. Increasing urban residents, in the so-called urban wildland interface, became a socioeconomic problem because of the potential fire impacts on homes and commercial enterprises. Urban interface areas present unique challenges to fire management. These areas required a completely different suppression and mitigation techniques whe fires burns surrounded forest fuels. In this sense, Environmental Andalusia Government increasingly concerned with impacts associated to these specific areas (Andalusian Law 5/1999).

WOODLAND MANAGEMENT ALTERNATIVES The research addresses the knowledge on conservation management and ecological restoration based on the ecological and socio-economic vulnerability of the semi-arid ecosystems on the above study areas. Field experiences, both Brazil and Spain, provide a technical framework to evidence guidelines for forest managers. Under similar semi-arid conditions field experiences of Brazil could be used to Spain, and viceversa, reaching better ways on the urgent need of these ecosystem conservation. Management alternatives are tested on a landscape design based on the economic limitation to have higher probability of successfully semi-arid actuations. The sustainable forest management of the semi-arid region must be developed according to the potential environmental conditions under cultural standards and the limitations of its current system production. Any other way to develop the rural economies could increase intangible assets deterioration such as environmental services and landscape goods similar to other well known systems.

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We have assessed the different strategies for the conservation of the different semi-arid landscapes taking into account the following: afforestation or reforestation practices, soil erosion, grazing resource and mycorrhizae associations.

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Afforestation or Reforestation Practices Over 2% of consumed firewood in the Brazilian semiarid is obtained by reforestation practices. Among several species, Eucalyptus spp, Prosopis juliflora, Leucaena leucocephala, Anadenanthera macrocarpa, Tabebuia sp., Mimosa caesalpiniifolia and M. tenuiflora presents great importance. As Poggiani (1982) affirmed, the most direct benefit is the shorttime production for wood energy and other activies (buildings, sawmills, etc.) from small forest areas that could be planted near urban centers. Mendes (1990) argued that reforestation in this region should be targeted primarily to the rural development and desertification control, giving more priority to social and ecological aspects than economic. The discussion about the introduction of fast growing species or the reintroduction of tree species in Caatinga areas implies a good knowledge of the region that must be seen as a whole and, at the same time, analyzed about its great regional diversity and ecosystem (Ab‘Saber, 1990). There is still much to be done for a secure reccomendation of which species, where, how and for what purpose must be planted in the different areas of Brazilian semi arid. In this sense, semi-arid experiences developed in Spain could be adapted to define Brazilian policies in afforestation or reforestation practices. Spanish forest policies in semiarid areas have encouraged land cover changes. Aleppo pine (Pinus halepensis Miller) plantation was the most widespread policy to reduce the effects of soil erosion. Although Aleppo pine is the dominand semi-arid species, other species such as Juniperus thurifera, Juniperus oxycedrus, Rhamnus lycioides and Quercus coccifera have been used in field experiences with satisfactory results (Jiménez et al., 2004). These experiences contribute to the debate on the suitability of mono-specific extensive P. halepensis plantations (Maestre and Cortina, 2004). In the semi-arid areas, environmental limitations, mainly drough period, influence on the afforestation outcome. The soil preparation is very important in semiarid regions for plant establishment due to the low growth in extreme water stressed conditions. Besides traditional soil preparation techniques, new techniques such as micro-watershed can be used to increase the water infiltration rate. In Spain, micro-watershed technique has been used since 1984 (De Simon et al., 2004) for rainwater harvesting. This technique suggests an increase of water availability, and as a consequence, the survival of the plants. In other words, the infiltration rate goes up and the surface runoff decreases.

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Figure 2. Box-Plot analysis between traditional soil preparation and micro-watershed.

Interaction between plant-soil characteristic and water availability due to microwatershed design modifies the plant microhabitat. A study in Almeria province showed the water availability for plant in relation with soil preparation technique: traditional soil preparation (backacter) and micro-watershed. The annual precipitation ranged between 200 and 600 mm due to the year and geographical position. Micro-watershed technique increased the amount of water available for plant at 84.74% (± 11.02) in relation to traditional technique (Figure 2). According to these results, the use of micro-watershed can be fundamental to conservation and restoration of degraded lands in semiarid regions. The increase of abandoned land due to its low profit has changed the semi-arid Mediterranean landscape. The development of a specific policy about the restorarion of the above abandoned patches has pointed to aditional tangible assets, environmental services and landscape goods. This forest policy, known as ―Land Forestation Planning‖, provides incentives to afforestation practices on degraded areas. However, the difficulty of the policy pursuit has led to bad practices such as incentives to areas where today don´t exist afforestations and if exist they don´t have both intermediate treatments and repositioning fail.

Soil Erosion The impact of over-grazing is even greater in semi-arid zones, considering that it can be associated to water stress, compromising the production of new sprouts, leading to low vegetation cover, decrease of water infiltration and increase of evaporation. According to Le Houérou (1996), over-grazing is among the main causes of soil degradation in arid and semiarid regions around the world, following only deforestation (Table 1). El Aich and Waterhouse (1999) have commented that over grazing in semi-arid environments is responsible for 29% of soil erosion by water and 60% of soil erosion by wind.

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Table 1. Main causes of soil degradation in dry areas of the world (103 km2) Causes Region Deforestation

Over-grazing

Agriculture

Africa

186

1,846

622

Asia

1,115

1,188

967

Australasia

42

785

48

Europe

398

413

183

North America

43

277

414

South America

322

262

116

Table 2. Environmental degradation scale and affected areas in northeast region Levels of environmental degradation

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Severe

High

Moderated

Low

Soil types and associations

Sensibility of erosion

Period of occupation

Semi-arid (%)

High

Long (cotton)

12.80

Strongly ondulated and Hill

Very high

Recent (Subsistenc e activity)

3.40

Podzolic; Eutrophic; ―Terra Roxa Estruturada‖ Cambissols

Ondulated and Strongly ondulated

Moderate

Long (Commerci al activity)

3.40

Planossols

Plain and Slightly ondulated

Moderate

Medium (Pasture and Subsistence activity)

2.35

Non-calcic Brown

Litholic

Relief

Slightly ondulated and Ondulated

Sá et al., 2004.

Human action in search of more fertile soils for agriculture and farming practice has been considered one of the main causes that lead Caatinga areas to devastation, exposing soil and turning it susceptible to erosive process, desertification and the disapearing of unnumbered vegetable and/or animal species. Degradation by human actions ends up compromising even more the Caatinga natural resources and sustainability.

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M. Á.Herrera Machuca, R. L. C. Ferreira, J. R. M. Martínez et al.

The replacement of Caatinga by agriculture has led to a generalized impoverishment of the soil, particularly relating to carbon and nitrogen (Tiessen et al., 1992; Fraga and Salcedo, 2004; Santos et al., 2009). In semi-arid conditions, water erosion is the most important event depending fundamentally of rain intensity, water infiltration, topography, type of soil and the existing vegetal cover. (Wischmeier and Smith, 1978). For the Brazilian semi-arid zone, Sampaio and Salcedo reported values of soil loss by pluvial erosion, reaching 100 t/ha/year. This soil loss is associated, mainly, to the traditional cultive cycle of five years followed by a period of regeneration under fallow of 10 to 20 years. It is worth to emphasize that this period is dependent of land availability, thus, as smaller the area for cultivation fewer this period will be. Sá et al. (2004) affirmed that semi-arid area affected by environmental degradation at high levels is greater than 20 million of hectares, and what worries the most is that this critical area reaches most part of the driest region. They also observed that the most devasted areas have soils with high fertility, that were and/or are being intensively explored and that degradation is not only manifested by sensibility of erosion but, above all, by its imposed use (Table 2). Mediterranean semi-arid area is generally characterized by discontinuous vegetation covers reflecting the soil erosion problem. Vegetation covering the soil reduces the kinetic energy of the raindrops before hitting the soil, thereby protecting the soil structure. The soil erosion could lead to the degradation of drylands, the major environmental worldwide problem. EU Communal Agricultural Policy (CAP) encouraged the agricultural abandonment process insighting for the soil erosion control and desertification impacts in Mediterranean regions. Recently, more attention has been given to semi-arid regions that has resulted in different forest management strategies. These strategies minimize the environmental, economic and social impacts during the drough period. Desertification affects 28 % of the Andalusian territory with 17.6% inside of the Almeria province. Other 68% of the Andalusia area is under high risk of desertification, mainly Granada and Jaén provinces. Desertification as the degradation of the natural and productive ecosystems can break the own exploitation of the agricultural lands. Agricultural lands have been altered in depthly due to the explotation during many continuous years. The dynamic succession on these abandoned areas is slow and not effective to reduce soil erosion. Afforestation practice is an effective tool for erosion control and the rehabilitation of degradaded lands. The Universal Soil Loss Equation (USLE) is a model that permits the prediction of longterm average soil erosion as a result of overland flow. USLE indicator has been proved useful with the help of GIS because of the easy application to make decisions. The afforestation policies and natural vegetation communities have been assessed to suggest the effectively of the different ecosystem to the control runoff. In this sense, one semi-arid public area covers over 1,720 ha have been used to perform the effects of vegetation cover increase in potential soil erosion. The vegetation is dominated by Aleppo pine and discontinuous understory. Most of the study area was classified by moderate loss (10-25 t/ha) according to current vegetation (Figure 3). However, this soil erosion increased considerably under potential conditions (discontinuous shrubland) without afforestation policies over 25 t/ha.

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Figure 3. Annual soil erosion rate according to afforestation practices

It is concluded that the potential soil erosion must be used to understand the interactions between landscape patterns and the design of soil erosion policies. The erosion induced loss of soil productivity may represent an economic cost to the society (Molina et al., 2009). Several methods for calculating the economic damages to society of a given loss of soil have been proposed in the literature (Afsen et al., 1996). We think that the cost of soil erosion must be calculated as the future income loss due to soil degradation. Different forest policies of the European Commission offer compensation payments between 40-120 euros/ha for soil conservation in semi-arid Mediterranean areas. In Spain, National Policy gave incentives about 7 euros per hectare and year to the proyects pointed to the soil erosion control in the most necessary regions. Later, Environmental Andalusia Government increased the cast payments to 17.26 euros per ha (Andalusia Plan to combat the Desertification). The environmental degradation scale, used in Brazilian semi-arid zone, could improve the Spanish incentives allocation to soil control based on the integral ecosystem consideration (soil type, relief, sensitibility of erosion and period of occupation). Brazilian and Spanish environmental policies must provide alternatives to sustainable management of the natural resources, including the soil control as another resource from their territories.

Grazing Resource Brazil presented populations of cattle, goats and sheep of 169,900,049, 7,109,052 and 13,856,747 heads according to Census of Agriculture (IBGE, 2006). In the Northeast region there are 15.32 % of cattle, 90.8 % of goats and 55.9 % of sheep. Thus, sheep/goat raising is a prominent activity in the northeast region, noticeable in the municipalities located in the semiarid and these animals graze in Caatinga area, usually without adequate stocking rate. Caatinga is the most important biome for the livestock in the Brazilian semi-arid region. Irregular rainfall distribution reduces possibility of annual crops and livestock production is one of the most important options for Northeast Brazil, and the Caatinga is an important forage resource for this region. Management methods are useful to increase herbage mass for ruminant feeding in the Caatinga (Araújo Filho et al., 2002). Higher phytomass values occurred in years with higher rainfall at the end of the rainy season (Table 3).

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M. Á.Herrera Machuca, R. L. C. Ferreira, J. R. M. Martínez et al. Table 3. Herbage mass (kg/ha) of the herbaceous stratum of managed Caatinga, in the end of rainy season Year

Cutting

Thinning

Mean

1981 (494 mm/year) 1982(328 mm/year) 1983(244 mm/year) Mean

3,835Aa 2,780Bb 1,156Bc 2,590B

3,826Aa 3,621Aa 1,816Ab 3,088A

3,830a 3,200b 1,486c -

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Means followed by the same capital letter within a row and small letter within a column do not differ (P>0.05) by Tukey test.

Moreover, Caatinga has low stocking rate, because of seasonal characteristics of plants, adverse conditions of climate and soil, and above all, the vegetation that is mostly composed by non-forage plants (Santos et al., 2010). Guimarães Filho et al. (1995) reported values of 12-15 ha/AU/year (1AU = 450 kg animal live weight) for the stocking rate of Caatinga and 68 kg of weight gain ha/year. When only the rainy season was considered, the stocking rate reached 4-5 ha/AU/year. Botanic composition is one of the pasture traits mostly modified by grazing. Caatinga species preferred by animals, according to the grazing pressure caused by the animal and the plant resistance to grazing, are those that will present more variability of occurence along time. In this sense, Leal et al. (2003) affirmed that herbivory by goats is an important factor of natural selection able to affect abundance and geographical distribution of woody species in Caatinga. Considering the importance of caatinga biodiversity management, the animal is an element of the system that can cause nonreversible effects to the botanic compostition of the vegetation, according to the grazing pressure used. Silva et al. (1999) observed that sheep grazing in caatinga under different managing systems, unbalaced the composition of herbaceous stratum and the grass disapeared from pasture in two years. In Spain, socio-economic changes in the last decades have led to an abandonment of marginal agricultural lands. In consequence, both landscape and socio-economic processes have suffered great changes in Mediterranean regions. The development of tourism on the coast has led to an increase in dry grasslands and dwarf shrublands in marginal lands. However, the quantity and quality of the pasture establishes limitations to the carrying capacity for grazing. The semi-arid rangelands in southeastern Spain show low carrying capacity and short term revenues based on forage yield and energy. Forage yield varies between 100 and 2,995 kg DM/ha/year and carrying capacity between 0.4 and 1.1 goat /ha/year) (Robles et al., 2001). The rangelands must be controlled by forest managers based on the lower level of forage quantity and quality, intake and digestibility. The overgrazing usually is found in terms of defoliation and seed consumption. The carrying capacity should include livestock and wildlife. In this sense, new hunting policies incorporated both resources to estimate the potential carrying capcity. Grasslands could be subjected to a disturbance as a consequence of grazing by livestock and wild herbivores. Management methods could be useful to increase herbage mass for livestock feeding in Mediterranean semi-arid zone similar to Caatinga experiences. Some leguminous plants associated to semiarid regions like as Anthyllis

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cytisoides could improve soil conditions by the fixation of atmospheric N and the erosion control in degraded areas.

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Mycorrhizae Association Mycorrhizae have the potential to contribute significantly to the success or failure of agro-ecosystems. Reeves et al. (1979) reported that in a semi-arid environment more than 99% of the plant over in a natural community was VA mycorrhizal, while in a disturbed soil less than 1% of the plant cover was mycorrhizal. Ecologic interactions have been estudied using species of tree legume and tree mycorrhizal fungi to establish plants into arid environments (Duponnois et al., 2005). The species of mycorrhizal fungus present in the soil can influence the competitive abilities of plant species. In the absence of mycorrhizal fungi; only the species which do not need mycorrhizae will be able to grow. Therefore, nonmycorrhizal species are most likely to dominate plant communities on poor soils, containing no or few mycorrhizal fungi propagules (Bagyaraj, 1989). The presence of these fungi is essencial for ecosystem regeneration from disturbances because they contribute for survival, growth and reproduction of tree species, mainly, when fertility is low (Caproni et al., 2005). The varied dependence of different plant species on different types of mycorrhizae can in fact influence the composition of both seral and mature plant communities (Bagyaraj, 1989). Among the several types of mycorrhizal fungi, the arbuscular assembled by Glomales (Zygomycetes) fungi, are of particular importance in the tropics, where they are better distributed and occur in more frequency (Smith and Read 1997). The fungi of VAM are present in almost all undisturbed soils but may be lost following mechanical disturbance and removal of the vegetation (St. John, 1990). Drought and soil erosion are other factors which may reduce or eliminate the VAM fungi propagules (Powell, 1980). Observations of the succession pattern of plants in semi-arid regions indicate that the arbuscular mycorrhizal fungi perform an important ecologic role in composing and establishing plants comunity (Marx 1980). Studies about associations between mycorrhizal fungi and Caatinga plants are scarce. However, some studies, like the ones of Souza et al. (2003) pointed out that more than 95% of the plants, among the 71 examined, formed arbuscular mycorrhiza (5-80% colonization). Despite the non-specificity of AM fungi with respect to host plant, certain fungus-plant combinations are more efficient than others. Siqueira et al. (1998, 2001) describe the importance of arbuscular mycorrhiza in forest ecology and land rehabilitation in the tropics associated to some Brazilian native woody species. Andrada et al. (2000) gave some results showing the predominancy of AM in South American forests. In this sense AM fungi colonization has shown importance in the ecophysiology and successional status in Brazilian woody species (Zangaro et al. 2000, 2003, 2005, 2007). Within botanic composition of Caatinga a high participation of Leguminosea species which have the N2-air fixation ability can be observed, therefore, the importance of these plants to maintain soil fertility in this region, as well as the large amount of crude protein concentration in the forage, has to be considered (Santos et al., 2010). Those authors affirmed that quantitative aspects of biological N2 fixation by Caatinga species have not been extensively studied.

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CONCLUSIONS Legislation about land use and woodland management in semi-arid areas needs reformulation when considering the effective potentialities of rural physical area, within cultural patterns of its population and the limits imposed by dominant relations of production. Any other way to perform it may lead to high risks of leaving the environmental liabilities that are already known in some systems. Into another line of action, changes are needed to adjust what are called ―Caatinga Sustainable Management Plan‖ and ―Andalusia Plan to Combat Desertification‖, by incorporating the indicators to be performed specifically for the semi-arid region. Nevertheless, it is fundamental, in terms of woodland management in the semi-arid, that a new paradigm, based in conservation and sustainability, must evolve, in order to minimize costs and environmental impacts associated to new projects. Conservation must be promoted with adequate management programs concerning demand, environmental education and sustainability directed to supply management, searching for alternative sources of supply, including agroforestry systems, cultivate fast growth species, as well as, the effective participation of the comunity involved in decision making.

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REFERENCES Ab' Sáber, A.N. (1995). Os sertões: a originalidade da terra. Ciência Hoje 3, 42-52. Ab‘Sáber, A.N. (1990). Projeto FLORAM: Nordeste seco. Estudos avançados 4(9), 49-174. Afsen, K., De Franco, M., Glomsrod, S. and Johnsen, T. (1996). The cost of soil erosion in Nicaragua. Ecological Economics 16, 129-145. Alados, C.L., Pueyo, Y., Barrantes, O., Escós, J., Giner, L. and Robles, A.B. (2004). Variations in landscape patterns and vegetation cover between 1957 and 1994 in a semiarid Mediterranean ecosystem. Landscape Ecology 19, 543-559. Allue, J.L. (1990). Atlas Fitoclimático de España. Taxonomías. Ministerio de Agricultura, Pesca y Alimentación. Instituto Nacional de Investigaciones Agrarias. Madrid, 221 pp. Andrade, AC.., Queiroz, M.H., Hermes, R.A.L and Oliveira, V.L. (2000). Mycorrhizal status of some plants of the Araucaria forest and the Atlantic rainforest in Santa Catarina, Brazil. Mycorrhiza. 10 :131–136. Andrade, L. A., Pereira, I. M., Leite, U. T. and Barbosa, M. R. V. (2005). Análise da cobertura de duas fitofisionomias de caatinga, som diferentes histórico de uso, no município de São João do cariri, estado da Paraíba, Revista Cerne 11(3), 253-262. Araújo Filho, J. A., Gadelha, A. G., Crispim, S. M. A. and Silva, N.L. da. (2002). Pastoreio misto em caatinga manipulada no Sertão Cearense. Revista Cientifica de Produção Animal 4(1-2): 9-21. Bacha, C. J. C. (2008). Análise da evolução do reflorestamento no Brasil. Revista de Economia Agrícola 55(2), 5-24. Bagyaraj, D.J. (1989). Mycorrhizas. In: H. Liet and M.J.A. Werger (Eds.) Tropical Rain Forest Ecosystems. Elsevier Press. Amsterdam, pp 537-546. Barberá, G., López, F. and Romero, A. (1997) Cambios del uso del suelo y desertificación en el Mediterráneo: el caso del Sureste Ibérico. In: Garcia, J.M and López (Eds.) Acción

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109

humana y desertificación en ambientes mediterráneos. Instituto Pirenaico de Ecología, Zaragoza, 9-39. Bellefontaine, R., Gaston, A. and Petrucci, Y. (2000). Management of natural forests of dry tropical zones. Roma: FAO Conservation Guide 32. FAO, 311pp. Bonet, A. (1997). Efectos del abandono de los cultivos sobre la vegetación en la cuenca del Alt Llobregat (Barcelona). Relación con factores ambientales y de usos del suelo. Ecología 11, 91-104. Bonet, A. (2004). Secondary sucesión of semi-arid Mediterranean old-fields in south-eastern Spain: insights for conservation and restoration of degraded lands. Journal of Arid Environments 56, 213-233. Brasil. Ministério da Integração Nacional/SDR. (2005). Plano estratégico de desenvolvimento sustentável do semiárido - PDSA. Ministério da Integração Nacional/IICA, Brasília. Cantón, Y., Solé-Benet, A. and Domingo, F. (2004). Temporal and spatial patterns of soil moisture in semiarid badlands of SE Spain. Journal of Hydrology 285, 199-214. Caproni, A.L., Franco, A.A., Berbara, R.L.L., Granha, J.R.D.O. and Marinho, N.F. (2005). Fungos micorrízicos arbusculares em estéril revegetado com Acacia mangium, após mineração de bauxita. Revista Árvore 29, 373-381. CEDEPLAR/UFMG; FIOCRUZ (2008). Mudanças climáticas, migrações e saúde: cenários para o nordeste brasileiro, 2000-2050. CEDEPLAR/UFMG FIOCRUZ, 47pp. Cerda, A. (1997). Soil erosion after land abandonment in a semiarid environment of southeasthern Spain. Arid Land Research and Management 11(2), 163-176. Chirino, E., Bonet, A., Bellot, J. and Sánchez, J.R. (2006). Effects of 30-years-old Aleppo pine plantations on runoff, soil erosion and plan diversity in a semi-arid landscape in south eastern Spain. Catena 65, 19-29. Dantas, J. G., Holanda, A. C., Souto, L. S. and Japiassu, A. (2010). Estrutura do componente arbustivo/arbóreo de uma área de Caatinga situada no município de Pombal-PB. Revista Verde 5, 134-142. De Simón, E., Bocio, I., Ripoll, M.A., Navarro, F.B., Jiménez, M. and Gallego, E. (2004). Microcuencas: una técnica de preparación del suelo en zonas semiáridas. In: De Simón, E., Ripoll. M.A., González, J.L. (Ed.). Forestación en paisajes agrarios. Junta de Andalucia. 95-119. Domingo, F., Villagarcía, L., Brenner, A.J. and Puigdefábregas, J. (1999). Evapotranspiration model for semi-arid shrub-lands tested against data from SE Spain. Agricultural and Forest Meteorology 95, 67-84. Duponnois, R., Paugy, M., Thioulouse, J., Masse, D. and Lepage, M. (2005). Functional diversity of soil microbial community, rock phosphate dissolution and growth of Acacia seyal as influenced by grass, litter and soil-feeding termite nest structure amendements. Geoderma 124, 349-351. El Aich, A., Waterhouse, A. (1999). Small ruminants in environmental conservation. Small Ruminant Research 34, 271-287. FAO (2007). Secuestro de carbono en tierras áridas. Roma, FAO. Informes sobre Recursos Mundiales de Suelos, 102, 120 pp. Fernandes, L. R., Almeida, A. M. and Duarte, C. R. (2009). Evolução da “cobertura vegetal” no município de Carnaúba dos Dantas – região do Seridó/RN. In: Simpósio Brasileiro de Sensoriamento Remoto, 16, 2009, Natal. Anais... INPE, Natal, 2721-2728.

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Flannigan, M.D., Amiro B.D., Logan, K.A., Stocks, B.J. and Wotton, B.M. (2006). Forest Fires and Climate Change in the 21st Century. Mitigation and Adaptation Strategies for Global Change 11(4), 847-859. Franca-Rocha, W., Silva, A. de B., Chaves, J. M., Nolasco, M. C., Accioly, L., J. de O.; Sá, I., B., and Pareyn, F.G.C. (2010). Plant Coverage and Soil Usage in the Caatinga Biome. In: Queiroz, L.P. de; Rapini, A. and Giulietti, A.M. (Eds.). Towards greater knowledge of the brazilian semi-arid biodiversity. http://www.uefs.br/ppbio/cd/english/chapter27.htm. (27/04/2010). González Bernáldez, F. (1991). Ecological consequences of the abandonment of traditional land use systems in Central Spain. Options Mediterranéss 15, 23-29. Guimarães Filho, C., Soares, J. G. G. and Riché, G. R. (1995). Sistema caatinga-buffelleucena para produção de bovinos no semiárido. Petrolina: Embrapa-CPATSA. 39 p. (Embrapa-CPATSA. Circular Técnica, 34). Gutiérrez, J.M., Pons, M.R. (2006). Modelización numérica del cambio climático: Bases científicas, incertidumbres y proyecciones para la Península Ibérica. Rev. C and G 20 (34), 15-28. Haase, P., Pugnaire, F., Clark, S.C. and Incoll, L.D. (1996). Spatial patterns in a two tiered semi-arid shrubland in southeastern Spain. Journal of Vegetation Science 7, 527-534. IBGE – Instituto Brasileiro de Geografia e Estatística. (2004). Mapa de Biomas do Brasil, primeira aproximação. IBGE, Rio de Janeiro. IBGE – Instituto Brasileiro de Geografia e Estatística. (1997). Anuário estatístico do Brasil. IBGE, Rio de Janeiro. IBGE – Instituto Brasileiro de Geografia e Estatística. (2006). Censo Agropecuário: 2006. Rio de Janeiro, RJ: IBGE. Disponível em: . Available in: 15 de maio de 2010. ICID. Second International Conference on Climate, Sustainability and Development in Semiarid Regions. Presentation. Available in: http://www.icid18.org/index.php? locale=enandm=conteudoanda=presentation. (26/04/2010). Imeson, A.C. and Prinsen, H.A. (2004). Vegetation patterns as biological indicators for identifying runoff and sediment source and sink areas for semi-arid landscapes in Spain. Agriculture, Ecosystems and Environment 104, 333-342. Jiménez, M., Ripoll, M.A., Navarro, F.B., Bocio, I. and De Simón, E., (2004).Forestación con especies arbustivas y de matorral en ambientes semiáridos. In: De Simón, E., Ripoll. M.A., González, J.L. (Ed.). Forestación en paisajes agrarios. Junta de Andalucia. 95-119. Kéfi, S., Rietkerk, M., Alados, C., Pueyo, Y., Papanastasis, V., ElAich, A. and De Ruiter, P. (2007). Spatial vegetation patterns and inminent desertification in Mediterranean arid ecosystems. Nature 449, 213-217. Kumazaki, M. (1992). A devastação florestal no sudoeste asiático e suas lições. Revista do Instituto Florestal 4, 46-52. Lasanta, T., González J.C., Serrano, S.M. and Sferi E. (2006). Using landscape ecology to evalúate an alternative management scenario in abandoned Mediterranean mountain areas. Landscape Urban Planning 78(1), 101-114. Lázaro, R., Rodrigo, F.S., Gutiérrez, L., Domingo, F. and Puigdefábregas, J. (2001). Analysis of a 30-year rainfall record (1967-1997) in semi-arid SE Spain for implications on vegetation. Journal of Arid Environments 48, 373-395.

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Leal, I. R., Silva, J.M.C. da, Tabarelli, M. and Lacher Júnior, T.E. (2005). Mudando o curso da conservação da biodiversidade na Caatinga do Nordeste do Brasil. Megadiversidade 1, 139-146. Leal, I.R., Vicente, A. and Tabarelli, M. (2003). Herbivoria por caprinos na Caatinga da Região de Xingó: Uma análise preliminar. In: Leal, I.R.; Tabarelli, M. and Silva, J.M.C. da (Eds). Ecologia e conservação da Caatinga. Editora da UFPE, Recife-PE, 695-716. Maestre, F. and Cortina, J. (2004). Are Pinus halepensis plantations useful as a restoration tool in semiarid Mediterranean areas? Forest Ecology and Management 198, 303-317. Makhzoumi, J.M. (1997). The changing role of rural landscapes: olive and carob multi-use tree plantations in the semiarid Mediterranean. Landscape and Urban Planning 37, 115122. Mamede, M. de A. and Araújo, F.S. de. (2008). Effects of slash and burn practices on a soil seed bank of Caatinga vegetation in Northeastern Brazil. Journal of Arid Environments 72, 458–470. Marx D.H. (1980). Ectomycorrhizal fungus inoculations: a tool for improving forestation practices. In: Mikola P. (Ed.). Tropical mycorrhiza research. Oxford University Press, New York, 13-71. Mendes, B.V. (1990). Observações sobre o Projeto FLORAM. Estudos avançados 4(9): 259262. MMA. (2010) Combating deforestation in the Caatinga is a priority to the MMA. http://www.mma.gov.br/sitio/en/index.php?ido=ascom.noticiaMMAandidEstrutura=8and codigo=5711. (26/04/2010). Molina, J.R. (2008). Integración de Herramientas para la Modelización Preventiva y Socioeconómica del Paisaje Forestal frente a los Incendios Forestales en relación con el Cambio Climático. Tesis Doctoral. Universidad de Córdoba, Spain. 411 pp. Molina, J.R., Rodríguez y Silva, F., Herrera M.A. and Zamora R. (2009). A Simulation Tool for Socio-Economic Planning on Forest Fire Suppression Management. In: Columbus, F. (Ed.). Forest Fires: Detection, Suppression, and Prevention. Nova Science Publishers. New York, 33-88. Molina, J.R., Herrera M.A. and Rodríguez y Silva, F. (2010). Consideraciones Técnicas para la Gestión del Paisaje Mediterráneo. Aplicación a la Región Mediterránea. In: Herrera, M.A. (Ed). Gestión Sostenible del Paisaje y de los Recursos Naturales (In Press). Naveh, Z. (1982). Mediterranean Landscape Evolution and Degradation as Multivariable Biofunctions: Theoretical and Practical Implications. Landscape Planning 9(2), 125-146. Nicolau, J.M., Solé-Benet, A., Puigdefábregas, J. and Gutiérrez, L. (1996). Effects of soil and vegetation on runoff along a catena in semi-arid Spain. Geomorphology 14, 297-309. Nobre, C.A., Salazar, L.F., Oyama, M., Cardoso, M., Sampaio, G. and Lapola, D. (2007). Mudanças Climáticas e Possíveis Alterações para os Biomas da América do Sul. MMASBioF-DCBio, Brasília. 25p. (ProBio Relatório 6) Noy-Meir, I. 1973. Desert ecosystems : environment and producers. Annual Review of Ecology and Systematics 4, 25-52. Pantaleón, J., Imanol, E., Pérez, R. and Roure, J.M. (2003). Palynological evidence for vegetational history in semi-arid areas of the western Mediterranean (Almería, Spain). The Holocene 13(1), 109-119.

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Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved.

112

M. Á.Herrera Machuca, R. L. C. Ferreira, J. R. M. Martínez et al.

Pausas, J. (2004) Changes in Fire and Climate in the Eastern Iberian Peninsula (Mediterranean Basin). Climate Change 63(3), 337-350. Pausas, J., Bláde, C., Valdecantos, A., Seva, J., Fuentes, D., Alloza, J.A., Villagrosa, A., Bautista, S., Cortina, J. and Vallejo, R. (2004). Pines and oaks in the restoration of Mediterrean landscapes of Spain: New perspectives for an old practice-a review. Plant Ecology 171, 209-220. Peter, H., Pugnaire, F., Clark, S.C. and Incoll, L.D. (1996). Spatial patterns in a twotiered semi-arid shrubland in southeastern Spain. Journal of Vegetation Science 7, 527-534. Piñol, J., Terradas, J. and Lloret, F. (1998). Climate Warming, Wildfire Hazard, and Wildfire Occurrence in Coastal Eastern Spain. Climatic Change 38(3), 1480-1573. Poesen, J., Wesemael, B., Bunte, K. and Solé-Benet, A. (1998). Variation of rock fragment cover and size along semiarid hillslopes: a case study from southeast Spain. Geomorphology 23, 323-335. Poggiani, F. (1982). O reflorestamento no nordeste brasileiro: consequências ecológicas. Série Técnica IPEF 3(10): 85-98. Prado Júnior, C. (1969). História Econômica do Brasil. 9ª ed. Editora Brasiliense, São Paulo. Available in: http://desvendandoaeconomia.blogspot.com/2009_01_01_archive.html (25/06/2010) Pueyo, Y. and Alados, C.L. (2007). Effects of fragmentation, abiotic factors and land use on vegetation recovery in a semi-arid Mediterranean area. Basic and Applied Ecology 8, 158-170. Queiroz, L.P. de (2010). Flowering plants of the brazilian semi-arid. In: Queiroz, L.P.de, Rapini, A. and Giulietti, A.M. (Eds.). Towards greater knowledge of the Brazilian semiarid biodiversity. http://www.uefs.br/ppbio/cd/english/chapter6.htm. (22/04/2010). Queiroz, L.P. de., Rapini, A. and Giulietti, A.M. (2010). Towards greater knowledge of the Brazilian semi-arid biodiversity. http://www.uefs.br/ppbio/cd/english/editorial.htm. (22/04/2010). Reeves, F.B., Wagner, D., Moorman, T. and Bkiel, J. (1979). The role of endomycorrhyzae in revegetation practices in the semiarid west. I A comparison of incidence of mycorrhizae in severely disturbed vs. natural environments. American Journal of Botany 66, 6-13. Rivas Martínez, S. (1987). Memoria del Mapa de Series de Vegetación de España. Serie Técnica. Ministerio de Agricultura, Pesca y Alimentación. Madrid, 268 pp. Robles, A.B., González-Rebollar, J.L., Passera, C.B.and Boza, T. (2001). Pastos de Zonas Áridas y Semiáridas del Sureste Ibérico. Arch.Zootec. 50, 501-515. Sá, I. B.; Riché, G. R. and Fotius, G. A. (2004). As paisagens e o processo de degradação do Semi Árido nordestino In: Biodiversidade da Caatinga: áreas e ações prioritárias para a conservação. MMA-UFPE, Brasília, DF, 17-36. Sampaio, E.V.S.B. and Salcedo, I.H. (1997). Diretrizes para o manejo sustentável dos solos brasileiros: região semi-árida. In: Congresso Brasileiro de Ciência de Solo, 26., Rio de Janeiro. Anais, Rio de Janeiro. Sociedade Brasileira de Ciência do Solo, 33pp. Sampaio, Y., Batista, J.E.M., Silva, J.S.H.e, Pereira Neto, J.J., Aroucha, M.L., Pereira, P.G. do P., Alcântara, R., Cavalcanti. R.C., Jucá, R. and Sant‘Ana, S.R. (2003). Pressões antrópicas atuais e futuras no bioma Caatinga. In: Silva, J.M.C. da, Tabarelli, M.,

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Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved.

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Fonseca, M.T. da and Lins, L.V. (Org.). Biodiversidade da Caatinga: áreas e ações prioritárias para conservação. Ministério do Meio Ambiente, Brasilia, 325-328. Santana, J. A. S. and Souto, J. S. (2006). Diversidade e estrutura fitossociológica da Caatinga na Estação Ecológica do Seridó-RN. Revista de Biologia e Ciências da Terra 6, 232-242. Santos, A. C. Dos, Salcedo, I. H., Candeias, A. L. B. and Galvão, S. R. da S. (2009). Influência do uso e da posição do perfil no relevo na fertilidade de solos em áreas de encosta. Pesquisa Agropecuária Tropical 39(1), 31-37. Santos, M.V.F. dos, M. de A. Lira, J. C. B. Dubeux Jr, A. Guim, A. C. L. de Mello and M. V. da Cunha (2010). Potential of Caatinga forage plants in ruminant feeding. Revista Brasileira de Zootecnia 39, 204-215. Silva, N. L., Araújo Filho, J. A., Sousa, F. B. and Araújo, M. R. (1999). Pastoreio de curta duração com ovinos em Caatinga raleada no sertão cearense. Pesquisa Agropecuária Brasileira 34(1), 135-140. Siqueira J.O., Carbone-Carneiro M.A., Curi N., da Silva Rosado S.C. and Davide A.C. (1998). Mycorrhizal colonization and mycotrophic growth of native woody species as related to successional groups in Southeastern Brazil. Forest Ecology and Management 107, 241–252. Siqueira, J.O. and Saggin-Júnior, O.J. (2001). Dependency on arbuscular mycorrhizal fungi and responsiveness of some Brazilian native woody species. Mycorrhiza. 11, 245-255. Smith, S.E. and Read, D.J. (1997). The symbiontes forming VA mycorrhizas. In Smith, S.E.; Read, D.J. (eds.). Mycorrhizal symbiosis. Academic Press, San Diego, 33-80. Souza, R.G., Maia, L.C., Sales, M.F. and Trufem, S.F.B. (2003). Diversidade e potencial de infectividade de fungos micorrízicos arbusculares em área de Caatinga, na Região de Xingó, Estado de Alagoas, Brasil. Revista Brasileira de Botânica 26(1), 49-60. Stenseth, N., Mysterud, A., Ottersen, G., Hurrel, J., Chan, K. and Lima, M. (2002). Ecological Effects of Climate Fluctuations. Science 23, 1292-1296. St. John, T.V. and Coleman, D.C. (1990). Mycorrhizal inoculation of container stock for restoration of self-sufficient vegetation. In: J.J. Berger (Ed.) Environmental Restoration. Island Press, Washington D.C., 103-112. Tiessen, H., Feller, C., Sampaio, E.V.S.B. and Garin, P. (1998). Carbon sequestration and turnover in semiarid savannas and dry forest. Climatic Change 40, 105-117. Tiessen, H., Sampaio, E.V.S.B. and Salcedo, I.H. (2001). Organic matter turnover and management in low input agriculture of NE Brazil. Nutrient Cycling in Agroecosystems, 61, 99-103. Vélez, R. (Coord.). (2009). La Defensa contra Incendios Forestales. Fundamentos y Experiencias. McGraw-Hill, Madrid. Velloso, A.L., Sampaio, E.V.S.B. and Pareyn, F.G.C. (Eds). (2002). Ecorregiões propostas para o Bioma Caatinga. Associação Plantas do Nordeste/Instituto de Conservação Ambiental The Nature Conservancy do Brasil, Recife. 81pp. Verstraete, M.M, and Schwartz, S.A. (1991). Desertification and global change. Vegetatio 91: 3-13. Viana, V. M., Veríssimo, A. and Pinheiro, L. A. F. V. (1998). Estratégia nacional de diversidade biológica: utilização sustentável de componentes da diversidade biológica e incentivos. Ministério do Meio Ambiente, Brasília. Available in: http://www.bdt.org.br/ publicacoes/politica/gtt/gtt4. (22/11/2008).

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Wischmeier, W.H. and Smith, D.D. (1978) Predicting rainfall erosion losses: a guide to conservation planning. Washington: USDA. (Agricultural Handbook 537). 58 pp. Zangaro, W., Bononi, V.L.R. and Trufen, S.B. (2000). Mycorrhizal dependency, inoculum potential and habitat preference of native woody species in South Brazil. Journal of Tropical Ecology 16 (4), 603-622. Zangaro, W., Nisizaki, S.M.A., Domingos, J. C. B. and Nakano, E. M. (2003). Mycorrhizal response and successional status in 80 woody species from south Brazil. Journal of Tropical Ecology 19 (3), 315-324. Zangaro, W., Nishidate, F.R., Spago-Camargo, F.R., Gorete-Romagnoli, G. and Vandressen, J. (2005). Relationships among arbuscular mycorrhizas, root morphology and seedling growth of tropical native woody species in southern Brazil. Journal of Tropical Ecology 21 (5), 529-540. Zangaro, W., Nishidate, F.R, Vandressen, J. Andrade, G. and Nogueira, M.A. (2007). Root mycorrhizal colonization and plant responsiveness are related to root plasticity, soil fertility and successional status of native woody species in southern Brazil. Journal of Tropical Ecology 23 (1), 53- 62.

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Chapter 4

PARTIAL HARVESTING IN OLD-GROWTH BOREAL FORESTS AND THE PRESERVATION OF ANIMAL DIVERSITY FROM ANTS TO WOODLAND CARIBOU Daniel Fortin 1,a, Christian Hébert 2,b, Jean-Philippe Légaré 3,c, Nicolas Courbin 1, Kyle Swiston 1, James Hodson 1, Mélanie-Louise LeBlanc 1, Christian Dussault 4,d, David Pothier 3, Jean-Claude Ruel 3, and Serge Couturier 4 Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved.

1.

Département de biologie, NSERC–Université Laval industrial research chair in silviculture and wildlife, Laval University, Québec, Canada 2. Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, Québec, Canada 3. Département des sciences du bois et de la forêt, NSERC–Université Laval industrial research chair in silviculture and wildlife, Université Laval, Québec, Canada 4. Direction de l'expertise sur la faune et ses habitats, Ministère des Ressources naturelles et de la Faune, Québec, Canada

ABSTRACT Current forest management must maintain biodiversity, an objective that has led to the rapid development of new forestry practices in recent years. However, empirical evaluation of the impact that these practices have on biodiversity has not kept the same pace. For example, small merchantable stems are now frequently protected during the harvest of old-growth boreal forests in eastern Canada. This silvicultural practice a

1045 Av. de la Médecine, pavillon Alexandre-Vachon, Québec, QC G1V 0A6, Canada. 1055 du P.E.P.S., P. O. Box 10380, Stn. Sainte-Foy, Québec, G1V 4C7, Canada. c Sainte-Foy, Québec G1V 0A6. d 880 chemin Sainte-Foy, Québec, QC G1S 4X4, Canada. b

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(referred to as CPPTM in Québec) ends up protecting all stems with a diameter at breast height of 9-15 cm, and is therefore expected to maintain some of the irregular attributes of old-growth forest structure. Whether or not this approach is sufficient to maintain local biodiversity remains unclear. We evaluated the short-term impact of CPPTM harvesting, mostly 2-3 years after logging, on a broad range of animal species differing largely in size and resource requirements. More specifically, we estimated the abundance, occurrence or local intensity of habitat use by ants, beetles, forest birds, snowshoe hare (Lepus americanus), moose (Alces alces) and woodland caribou (Rangifer tarandus caribou) in a boreal ecosystem. The 19,000 km2 study region is dominated by >270 year old stands with irregular structure mostly comprised of black spruce (Picea mariana) and balsam fir (Abies balsamea). Harvesting by CPPTM caused a 75-85% reduction in tree basal area. This decrease was sufficient to alter animal assemblages of all taxonomic groups. Ants and beetles associated with open areas were more abundant in CPPTM sites than in uncut stands. Conversely, species associated with mature forests were much lower in CPPTM than in uncut stands. Most birds associated with late-successional habitats were also less likely to be observed in CPPTM sites than in uncut stands. Snowshoe hares significantly decreased their use of harvested stands following CPPTM. Woodland caribou displayed a strong avoidance for CPPTM sites, while moose did not display any such aversion. CPPTM thus alters species assemblages, potentially reshaping trophic interactions with consequences for wildlife conservation. For example, moose did not avoid CPPTM cuts as much as they avoided stands that were harvested more intensively. Wolves that generally focus their hunting on moose might become attracted to CPPTM cuts, which could result in a higher concentration or even in a numerical response of wolves. At a regional scale, the outcome might lead to an increase in local predation risk for woodland caribou, a threatened species. While CPPTM could still be useful in an ecosystem-based management context, our study shows that the protection of small merchantable stems is often insufficient for the short-term maintenance of species composition which characterizes old-growth boreal forests.

INTRODUCTION Current forest harvesting practices have to sustain our long-term wood supply needs, while also maintaining primary ecosystem attributes, including regional biodiversity (Côté and Bouthillier 2002). Forest management strategies are increasingly developed following the principles of ecosystem-based management by emulation of natural disturbances (Bergeron et al. 2007, Rosenvald and Lõhmus 2007). The premise is that biodiversity and essential ecological functions are most likely to be maintained by emulating the spatio-temporal patterns of natural disturbances (Hunter 1999, Fenton et al. 2009) because local populations have been shaped by the processes leading to these patterns. The natural disturbance regime responsible for the dynamics of local ecosystems varies widely across the boreal forest. For example, the average time between wildfire recurrences can be less than a century in some areas, while in others, it can exceed 500 years and even thousands of years (Bergeron et al. 2002, Bergeron et al. 2007, Bouchard et al. 2008, Fenton et al. 2009, Johnstone et al. 2009). Forest composition and structure strongly depend on the length of the regional fire cycle. Balsam fir (Abies balsamea) generally increases in abundance as black spruce (Picea mariana) dominated forests get older (De Grandpré et al. 2000). Approximately 80-90 years after the last fire, the local basal area and density of trees (>9 cm in diameter at breast height, dbh) cease to increase, and even start declining

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(Bouchard et al. 2008). Under a long fire cycle regime, tree mortality due to windthrow, insects, pathogens, and senescence lead to fine scale canopy gap dynamics creating landscapes dominated by irregularly structured old-growth stands (Boucher et al. 2003, Pham et al. 2004). In response to strong regional variations in natural disturbance regimes, an ecosystembased approach to forest management should prescribe different silvicultural practices across the boreal forest (Rosenvald and Lõhmus 2007). Currently, clearcutting (or closely related harvest techniques, such as cutting with protection of advanced regeneration and soils [CPRS], Groot et al. 2005, Courtois et al. 2008) is the most common logging practice in the boreal forest (Bergeron et al. 2007, Rosenvald and Lõhmus 2008, Ruel et al. 2007). Clearcutting brings stands back to an early seral-stage from which they develop with a regular structure (Groot et al. 2005). This practice thus appears particularly relevant in regions with relatively short fire cycles (Bergeron et al. 2007). In contrast, the emulation of natural disturbances in regions with long fire cycles, dominated by old-growth forests, should require the use of partial harvesting to maintain the irregular structure of forest stands (Bergeron et al. 2007, Ruel et al. 2007, Rosenvald and Lõhmus 2007, Drapeau et al. 2009). A large number of partial harvesting techniques have emerged in recent years (Groot et al. 2005, Ruel et al. 2007, Courtois et al. 2008, Rosenvald and Lõhmus 2008, Cimon-Morin et al. 2010, Poulin et al. 2010). Since the 1990s, the protection of small merchantable stems has become increasingly frequent when harvesting old-growth boreal forests in eastern Canada. This silvicultural technique (taking the acronym ―CPPTM‖ in Québec and HARP in Ontario, Groot et al. 2005) protects all stems with a dbh of 9-15 cm, thereby maintaining some of the irregular structure of old-growth forests (Ruel et al. 2007). Not spending time and money harvesting these relatively small stems can lead to financial gains because small merchantable stems are worth less to the forest industry than larger stems (Liu et al. 2007, Ruel et al. 2007). However, the impact of CPPTM on biodiversity has remained largely unexplored, and the industry still does not know what role CPPTM serves in maintaining local biodiversity and ecosystem functioning of old-growth boreal forests. Clearcutting of mature forests generally alters local animal communities by favouring species associated with early-successional habitats at the expense of late-successional species (Annand and Thompson 1997, Simon et al. 2000, Vanderwel et al. 2007). In contrast, partial harvesting can maintain species assemblages, depending on logging intensity (Vanderwel et al. 2009). Whether the protection of small merchantable stems during CPPTM suffices to maintain local biodiversity remains an open question. Because it is impractical to evaluate the impacts of forest harvesting on all species, investigators often rely on indicator species (Potvin et al. 1999, Thompson et al. 2008), which are generally selected from a single taxonomic group. This approach may not fully inform us of the potential contribution of a given silvicultural practice to sustainable forest management, because the impacts of logging vary among animal groups (Venier and Pierce 2004). To acquire a comprehensive understanding of the impact of forest harvesting, it is therefore critical to consider a diversity of taxa for adequate wildlife conservation and management. We investigated the short-term impact of CPPTM (mostly 2-3 years after harvesting) on target species chosen from multiple taxa. These species were selected because they varied broadly in movement capacity, home-range size, and habitat requirements, as well as in their associations with different stages of forest succession. First, we considered two saproxylic ant

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species: Camponotus herculeanus, which is generally more abundant in closed forests, and Formica neorufibarbis, which is typical of open areas (Jennings et al. 1986, Francoeur 2001, Higgins and Lindgren 2006). We also considered 10 beetle species, five of which have been found to be associated with closed coniferous forest stands (Silvanus bidentatus (Majka 2008), Enicmus tenuicornis (Jacobs et al. 2007b, Majka et al. 2009), Oxypoda grandipennis (Csy.) (Klimaszewski et al. 2006), Epuraea planulata (Saint-Germain et al. 2004, Janssen et al. 2009), Rhizophagus dimidiatus (Saint-Germain et al. 2004, Jacobs et al 2007a, Janssen et al. 2009), whereas the other five are typical of open areas (Leiodes spp. from the assimilisgroup (Baranowski 1993, Unpublished data, C. Hébert), Clypastraea fusca (Boulanger and Sirois 2007), Xylita laevigata (Boulanger and Sirois 2007), Hylobius congener (Welty and Houseweart 1985, Boulanger and Sirois 2007), Pseudanostirus triundulatus (Randall) (SaintGermain et al. 2004). For instance, the last four species are particularly abundant shortly after fire in the boreal forest (Saint-Germain et al. 2004, Boulanger and Sirois 2007). Boreal Chickadee (Poecile hudsonsicus), Brown Creeper (Certhia familiaris), and Ruby-crowned Kinglet (Regulus calendula) were selected to represent closed-canopy bird species because they are abundant on our study area (Lemaître 2009), and they should respond negatively to low tree retention levels (Vanderwel et al. 2007, 2009, St-Laurent et al. 2008). White-throated Sparrow (Zonotrichia albicollis) and Dark-eyed Junco (Junco hyemalis) are also locally abundant species (Lemaître 2009), and they are expected to respond positively to the openings of the tree canopy resulting from logging (Vanderwel et al. 2007, 2009). Finally, we evaluated the response of snowshoe hare (Lepus americanus) and moose (Alces alces), two species favouring rather early (10-30 years) successional forests (Pastor et al. 1999, Dussault et al. 2005, Fisher and Wilkinson 2005, Newbury and Simon 2005), and forest-dwelling woodland caribou (Rangifer tarandus caribou), a late-successional species (Fortin et al. 2008, Courbin et al. 2009). The field methods used varied among taxa, from an evaluation of changes in abundance (ant and beetle species) or probability of occurrence (bird species) of individual species, to an assessment of habitat use and selection by mammalian herbivores (snowshoe hare, moose and caribou).

STUDY AREA The study took place in the eastern spruce-moss subdomain of the boreal forest, in the Côte-Nord region of Québec, Canada (Figure 1). The 19,000 km2 study area is typical of the Canadian Precambrian Shield, with a rolling and hilly landscape, and an altitude varying between 300 and 1,000 m. Mean annual temperature ranges from -2.5 to 0.0 °C, and annual precipitation ranges from 1,000 to 1,400 mm (Boucher et al. 2003). The abundant precipitation results in a long fire cycle (average stand age: >270 years old, Bouchard et al. 2008). The study area is therefore characterized by old-growth forest stands that are irregular in structure and composition (Boucher et al. 2003). The dominant tree species are black spruce, balsam fir, white birch (Betula papyrifera Marsh.) and trembling aspen (Populus tremuloides Michx).

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Figure 1. Study area located in the Côte-Nord region of Québec, Canada. Harvested areas are displayed in white.

METHODS Estimation of Ant Abundance In 2004 or 2005, ants were sampled in 14 stands (>4 ha) that had been harvested by CPPTM 1 to 8 years earlier (average: 4.2 ± 2.1 years). Ants were captured concurrently in 70 uncut conifer stands (median age: 120 years; range: 70 to >120 years). All sites were >2 km

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from one another. At each site, ants were captured in four pitfall traps buried at ground level, 10 m apart (following Janssen et al. 2009). The traps had a diameter of 10 cm, and were screened with a wire mesh (1 × 1 cm) to avoid capture of vertebrates. Trapped insects were collected every two weeks, from 2 June until 17 August, and preserved in a 40% ethanol solution with a trace of acetic acid (5%), before being identified in the laboratory. The abundance of each of the two focal ant species was contrasted between CPPTM and uncut stands with one-sided, nonparametric Wilcoxon Two-Sample Tests, evaluating the specific prediction that CPPTM opens the canopy sufficiently so that the closed-habitat species, C. herculeanus, would be less abundant in CPPTM sites than in uncut conifer stands, whereas the open-habitat species, F. neorufibarbis, would be more abundant in CPPTM sites than in uncut stands. Because the time between CPPTM harvesting and ant sampling varied by as much as 8 years, we used a Spearman rank correlation to test whether the abundance of each of these ant species varied with the time since partial harvesting.

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Estimation of Beetle Abundance Beetles were sampled from 5 June to 22 August 2007 in 16 old-growth forest stands (>120 years), and in 16 CPPTM conifer stands (average size: 19 ha; range: 12-24 ha). Eight of these sites had been harvested in 2004 and eight in 2005. All sites were >100 m apart and >100 m from the stand edge. For logistical reasons, these sites were grouped in four sectors (e.g., each near a forestry camp or associated with different forestry companies). The potential non-independence of data within a sector was considered through the use of mixedeffects modelling (see below). Four pitfall traps were located at each site to capture ground-dwelling beetles, together with one flight-interception trap to capture flying beetles. The multi-directional flightinterception trap was located at the centre of each sampling site, 0.5-1 m above ground. This trap was built using four 15 × 40 cm panels (two made of Plexiglas and two of mosquito net) mounted into a cross pattern, along a 10-cm diameter black ABS cylinder, with two funnels located above and below the cylinder, and leading to collecting vials (Saint-Germain et al. 2004). As with the ants, beetles were collected every two weeks, and preserved in a solution of ethanol, acetic acid and water, before being identified in the laboratory at the species level. The log-transformed abundance (log [x+0.1]) of each of the 10 focal beetle species was compared between cut and uncut stands using mixed-effects models with a Gaussian distribution. The four sectors were considered as a random-effect term.

Estimation of the Occurrence Probability and Abundance of Bird Species Birds were surveyed in 2006 and 2007 between 3 June and 30 June at 12 point count stations (Verner, 1985) located in old-growth forests, and at 12 stations located in conifer stands harvested by CPPTM (average size: 19 ha; range: 12-24 ha) in 2004 or in 2005. Stations were aggregated into three sectors, a design controlled for by the use of mixedeffects models. Point count stations were >150 m from one another and >100 m from the stand edges.

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Point count stations were visited by four observers three times each year, between 05:00 and 10:00, in the absence of wind or heavy rain. To reduce the potential risk of observer bias, the four observers surveyed different sites during the three yearly visits. All birds seen or heard within a 50-m radius of the point count stations were recorded during 10 minutes. Observers trained together to become consistent in their evaluation of bird distances. A species was coded as 1 for a given point count station if it was recorded during any of the visits. Otherwise, the species was coded as 0 for that station. We used this approach because of low bird count at individual stations: only 0-2 individuals were observed per station, except for White-throated Sparrow which was represented by up to 4 individuals per station. We used mixed-effects logistic regression for each of the five focal bird species to evaluate whether it was more likely to be observed at CPPTM sites or in the uncut stands. The three sectors were considered as a random-effect term to allow for potential inter-sector variations in the probability of occurrence of the different species. We also carried out mixedeffects Poisson regression for the White-throated Sparrow, the most abundant bird species, to test whether its abundance was higher in CPPTM sites than in uncut stands.

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Evaluation of Habitat Use by Snowshoe Hare Habitat use by snowshoe hare was assessed in four uncut conifer stands paired with four CPPTM sites harvested in 2004 or 2005. In 2007, we evaluated browse history at all sites by identifying years in which white birch stems (an abundant and preferred browse species, Newbury and Simon 2005) were browsed by snowshoe hare. This approach allowed us to quantify the intensity of stand use by snowshoe hare prior to logging, even though field sampling was conducted only 2 to 3 years after harvesting. During winter, this herbivore generally clips the terminal leaders of the previous summer‘s growth, which kills the terminal bud (Pease et al. 1979). The following spring, vertical growth will resume from a dormant lateral bud further down the stem (Keigley and Frisina 1998). In contrast, vertical growth of non-browsed stems resumes from the terminal bud, leaving a bud scar for each year of uninterrupted growth. The number of years of growth following the mortality of a terminal leader due to browsing can then be determined by counting the number of terminal bud scars along the stem originating from a lateral bud which resumed vertical growth (Keigley and Frisina 1998). We can thus determine the year in which each twig was clipped by counting the bud scars backwards from the current year‘s growth to the browsed segment. Bud scars are generally discernable for up to five years of previous growth. We were interested in the previous 4 years of growth, which included the winter preceding the earliest cuts (2004) in the experimental blocks. We evaluated the browse history of 18 white birch stems distributed across six quadrats (6 × 75 m) located in each CPPTM and each uncut site (i.e., 36 stems per pair of cut-uncut sites). In each quadrat, we sampled the first three white birch stems that met the following criteria: 1) stems had to be at least 5 years old (determined by counting terminal bud scars) so that they could provide information on pre-treatment use by hare, 2) stems also had to be at least 50 cm in height 5 years prior to the survey so that they would be above the snow during a large part of each winter, and 3) stems had to have at least one browse mark so that they could provide a historical record of use. For each stem, we identified the year in which twigs were clipped by snowshoe hare starting from the winter before the harvest treatment took

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Daniel Fortin, Christian Hébert, Jean-Philippe Légaré, Nicolas Courbin et al.

place (i.e. the winter of 2003/2004 or 2004/2005) until the winter of 2006/2007 (2-3 years after harvesting took place). We then used the presence or absence of scars in each year to classify each stem as used or unused for each of the previous 4 years. To assess whether snowshoe hare used sites with similar intensity before harvesting took place, we first tested a model predicting the probability of white birch browse use by snowshoe hare in the winter before harvesting occurred. We used mixed-effects logistic regression with the dependent variable being whether a stem was browsed (coded as 1) or not (coded as 0) in a given year. To account for the hierarchical structure of our sampling design and the non-independence among the 18 stems selected within each site, we included random effects which consider that stems were nested within habitats, and that habitats (CPPTM versus uncut stands) were paired. To model temporal changes in the probability of birch stem browsing by snowshoe hare, we used a second mixed-effects logistic regression for repeated measures. We considered browsing from the winter before harvest treatments (year = 0) took place until 2 to 3 years after logging. Habitat (CPPTM = 1, uncut = 0), year and ―habitat × year‖ interaction were included as fixed-effects terms. Birch stems were considered the experimental units upon which repeated measures were taken (i.e. each stem was classified as used or unused during each year) and we used an autoregressive (order 1) correlation structure to account for the fact that the probability of stem use was more likely to be similar in successive years than 2 or 3 years apart in time. We used the same random effects structure as specified for the pre-harvest model.

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Evaluation of Habitat Selection by Moose and Caribou From mid-March 2005 to mid-March 2007, we followed the movement of 18 adult female caribou and 10 adult female moose equipped with global positioning system collars (Lotek Engineering Inc., Newmarket, Ontario or Telonics Inc., Mesa, Arizona) taking locations at 4-h intervals for caribou and every hour for moose. Due to deaths or battery exhaustion, each caribou was followed during an average of 13 months (range: 12.50 and P < 0.002 for each species). For example, Leiodes sp. was only found in CPPTM, whereas X. laevigata was 34 times more abundant in CPPTM than in uncut stands (Figure 3).

Forest Birds

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The Boreal Chickadee and the Ruby-crowned Kinglet, two closed-canopy species, were more likely to be observed in uncut forests than in CPPTM sites (Table 1). However, we did not detect a difference in the probability of occurrence of the Brown Creeper, also a closedcanopy species, between uncut and harvested stands. The White-throated Sparrow and Darkeyed Junco had a higher probability of occurrence in CPPTM than in uncut forests (Table 1). For example, we only observed 2 White-throated Sparrows at two point count stations in uncut conifer stands (i.e., 2/12 stations in uncut stands), while we recorded the species at all the CPPTM stations (i.e., 12/12 stations), totalling 26 individuals observed in the harvested stands. In fact, White-throated Sparrows were significantly more abundant in CPPTM sites than in uncut stands (Mixed-effects Poisson regression: F1,20 = 485.7, P < 0.001). Table 1. Coefficients () from a mixed-effects logistic regression testing for a difference between the probability of occurrence of closed- and open-habitat bird species in CPPTM sites versus in uncutconifer stands in the Côte-Nord region of Québec, Canada.Negative s imply a lower probability of occurrence in CPPTM cuts than in uncut conifer stands Species Closed-canopy species Boreal Chickadee Brown Creeper Ruby-crowned Kinglet Open-canopy species White-throated Sparrow Dark-eyed Junco

 ± SE

F1,18

P

-2.40 ± 1.01 -1.30 ± 1.09 -3.09 ± 1.08

5.66 1.42 8.18

0.03 0.25 0.01

21.23 ± 0.76 2.73 ± 1.05

785.97 6.75