Floristic and structural variability of the central apennines beech forests in relation to natural and anthropogenic determinants

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(1)DOCTORALSCHOOLI NBI OLOGY. Sec t i on: Bi odi ver s i t yandEc os ys t emsAnal ys i s. XXVI ICyc l e A. A. 2013/2014. Fl or i s t i cands t r uc t ur alv ar i abi l i t yoft he c e nt r alApe nni ne sbe e c hf or e s t si n r e l at i ont onat ur alandant hr opoge ni c de t e r mi nant s Var i abi l i t àl or i s t i c aes t r ut t ur al ede l l ef agge t ec e nt r oappe nni ni c hei nr e l az i oneaide t e r mi nant inat ur al i eant r opoge ni c i. Andr e aS c ol a s t r i. Tut or : Dr . Ma ur i z i oCut i ni.

(2) DOCTORAL SCHOOL IN BIOLOGY Section Biodiversity and ecosystems analysis. XXVII cycle. Floristic and structural variability of the central Apennines beech forests in relation to natural and anthropogenic determinants Variabilità floristica e strutturale delle faggete centroappenniniche in relazione ai determinanti naturali e antropogenici. Andrea Scolastri Tutor: Dr. Maurizio Cutini. A.A. 2013/2014.

(3) Thesis defense on the 24th February 2015 in front of the following jury:. Prof.ssa Angela Stanisci Prof. Francesco Spada Dott. Giandiego Campetella.

(4) Table of contents. Abstract. I. Riassunto. IV. Papers published or prepared in the course of the PhD. VII. General introduction. 1. Chapter 1 Patterns of floristic variation on a montane beech forest in the central Apennines (central Italy). 5. Chapter 2 Old Coppice vs High Forest: the impact of beech forest management on plant species diversity in central Apennines (Italy). 21. Chapter 3 Old Coppice vs High Forest: effects of two management types on understory functional composition. 45. General conclusions. 72. References. 76. Ringraziamenti. 81.

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(6) Abstract. Forests are complex ecosystems, in which the floristic composition is the result of the combined effect of several factors, acting at different levels. Climate, lithology, geomorphology act at a broader scale, driving the forest's communities differentiation while, at a finer scale, the understory species composition is shaped by the forest stand structure that directly influences the environmental conditions (light, temperature, moisture) at the ground level. Fagus sylvatica forests are the most abundant broadleaved communities in central and southern Europe; in Italy they represent the typical montane vegetation in the Apennine chain, where these coenoses reach the highest elevations of their whole distributional range. Beech forests have been managed for centuries, prevalently as coppice, or coppicewith-standards (CWS), and high forest (HF). However, in recent decades deep socio-economical changes have led to the progressive abandonment of coppice cut and its conversion to high forest management. These two systems differ in cut intensity and severity (both higher in CWS): given that forest management impacts upon overstory structure and therefore determines the understory composition, management changes are likely to affect understory species composition and diversity. In this thesis I have analyzed the effect of the main environmental drivers on beech forests communities: I then focused on the management effects on understory species and 9210* Habitat diagnostic species richness and composition (sensu Habitat Directive 92/43 EEC), in order to understand the possible implications of the old CWS conversion on the floristic composition. The study was conducted in central Apennines, on the Montagne della Duchessa massif, where beech forests occupy a surface of more than 1200 ha; these forests have ceased to be managed since '60s, so the CWS have become older, and are destined to be converted to the HF cut. I studied the role of the environmental factors in differentiating the beech forests communities by using a dataset of 40 relevés randomly selected. The dataset obtained has been analyzed through a cluster analysis and a Indicator Species Analysis (ISA), in order to obtain groups of relevés and to characterize them in floristic terms; the groups obtained were then compared in terms of environmental and topographic variables, Ellenberg indicator values, life forms, Social Behaviour Types (SBT) and structural parameters. Results showed that the floristic and coenological variability is. I.

(7) shaped by a climatic and edaphic gradient, that both contribute to define two main communities: a microthermal one, placed at higher altitudes and cooler aspects (Cardamino kitaibelii - Fagetum sylvaticae), and a termophilous one, lying at lower altitudes and warmer aspects (Lathyro veneti - Fagetum sylvaticae). Social Behaviour Types and structural parameters were useful for detecting the effects of the progressive reforestation process occurring inside the microthermal community at higher elevation. In order to investigate the differences between old CWS and HF stands in terms of understory richness and composition, I used 66 relevés, selected through a random-stratified method, so as to have a comparable number of relevés for each management category; old CWS and HF stands were compared in terms of structural attributes and floristic richness through a U Mann-Whitney test: the results showed how management is the main factor responsible for differences in the tree layer's spatial aggregation pattern and vertical layering, and that therefore determines the amount of surface available for understory species. HF stands showed a higher mean richness of both understory and diagnostic species, these latter being more evenly distributed (higher species equitability) inside the community. This leads to the consideration that in HF stands the cut regime provided a constant canopy cover over time, and then maintained more stable microclimatic conditions favoring a higher abundance and evenness of the shade-tolerant and vernal species. Conversely, in old CWS the dense canopy negatively affected the understory richness by reducing the light-demanding species pool, while the shade-tolerant species have not yet had time to spread. In order to understand the implications of the old CWS conversion on beech forests ecosystem's ecological functions, the same dataset was analyzed through plant functional traits, by creating a relevés x traits matrix. A Redundancy Analysis (RDA) was performed to assess the relationship between traits states and management, while a U Mann-Whitney test was used to assess differences in traits states richness between the management types. The results were consistent with the previous findings based on habitat diagnostic species, as HF stands showed a higher affinity to traits typical of mature forests, while old CWS were more related to traits related to managed stands. Moreover, HF stands showed a higher abundance of those traits related to the natural forest's seasonal change, this indicating a good species distribution among the functional niches. Even in this case, old CWS was shown to be in a transitional stage, still represented by some traits related to management, where the mature forest traits are also present, but with lower abundance.. II.

(8) Finally, given that in a forest ecosystem understory represent more than 90% of species richness, and is the most sensitive to disturbance, the modern silvi-cultural strategies should take into account the management effects on forest biodiversity, and adopt sustainable interventions able to favor the typical biodiversity of the ecosystems we want to manage. This thesis provides useful information for management purposes, as our findings show that, in the forests studied, the old CWS conversion to HF could be a good management strategy if our aim is the conservation over time of the species typically related to mature forest conditions.. III.

(9) Riassunto. Le foreste sono ecosistemi complessi, la cui composizione floristica è il risultato dell'effetto combinato di diversi fattori, che agiscono a vari livelli. Il clima, la litologia e la geomorfologia agiscono a scala più ampia, determinando la differenziazione delle comunità forestali, mentre, a scala più fine, la composizione floristica del sottobosco è selezionata dalla struttura del popolamento forestale, che ha una influenza diretta sulle condizioni ambientali (luce, temperatura , umidità) a livello del suolo. Le foreste a Fagus sylvatica sono le più abbondanti comunità di latifoglie in Europa centrale e meridionale ed in Italia rappresentano la tipica vegetazione montana della catena appenninica, dove queste cenosi raggiungono le quote più elevate dell'intero areale. Le foreste di faggio sono state gestite per secoli, prevalentemente come cedui, o cedui matricinati, e come fustaie ma negli ultimi decenni, profondi cambiamenti socioeconomici hanno portato al progressivo abbandono del taglio ceduo e al successivo avviamento all' altofusto. Questi due sistemi si differenziano sia per intensità che severità del taglio, entrambi di maggiore entità nel ceduo; dato che la gestione forestale agisce direttamente sulla struttura del popolamento forestale, cambiamenti nella gestione del taglio possono avere effetti sulla composizione e sulla diversità floristica del sottobosco. In questa tesi ho analizzato gli effetti dei principali fattori ambientali sulle comunità di faggeta, concentrandomi poi sugli effetti della gestione forestale sulla composizione e ricchezza specifica sia delle specie del sottobosco che delle specie diagnostiche dell' Habitat 9210* (sensu Direttiva Habitat 92/43 CEE), al fine di comprendere le possibili implicazioni della conversione dei cedui invecchiati sulla composizione floristica. Lo studio è stato condotto in Appennino centrale, sul massiccio delle Montagne della Duchessa, dove le faggete occupano una superficie di più di 1200 ettari; queste foreste non sono state più gestite a partire dagli anni '60, quindi i cedui, ormai invecchiati, sono ora destinati all'avviamento all'altofusto. Ho studiato il ruolo dei fattori ambientali nella differenziazione delle comunità di faggeta utilizzando 40 rilievi, selezionati in maniera casuale. Il set di dati ottenuto è stato analizzato attraverso una Cluster Analysis ed una Indicator Species Analysis (ISA) al fine di ottenere gruppi di rilievi, e di caratterizzarli in termini floristici; i gruppi ottenuti sono stati confrontati sulla base delle variabili ambientali e topografiche, dei valori di indicazione. IV.

(10) di Ellenberg, forme biologiche, Social Behaviour Types (SBT) e parametri strutturali. I risultati hanno mostrato che la variabilità floristica e cenologica delle faggete studiate è modellata da due gradienti, uno climatico ed uno edafico, che contribuiscono entrambi nel definire due comunità principali: una microtermica, posta a quote maggiori ed esposizioni più fredde (Cardamino kitaibelii - Fagetum sylvaticae), ed una termofila, posta a quote più basse ed esposizioni più calde (Lathyro Veneti - Fagetum sylvaticae). L'utilizzo dei Social Behaviour Types, ed il loro incrocio con i dati sui parametri strutturali, ha permesso inoltre di rilevare gli effetti del processo di riforestazione che sta avvenendo all'interno della comunità microtermica, alle quote maggiori. Al fine di indagare le differenze tra cedui invecchiati e fustaie in termini di ricchezza e composizione floristica del sottobosco, ho utilizzato 66 rilievi, selezionati attraverso un metodo random-stratificato in modo da avere un numero paragonabile di rilievi per ciascuna categoria di gestione;. I cedui invecchiati e le fustaie sono state confrontate sia in termini di attributi strutturali che di ricchezza floristica attraverso un test U di MannWhitney. I risultati hanno mostrato come la gestione sia il principale responsabile delle differenze nella distribuzione spaziale e nella stratificazione verticale dello strato arboreo, che determinano quindi la quantità di superficie disponibile per le specie del sottobosco. Le fustaie hanno mostrato una maggiore ricchezza media per rilievo sia delle specie del sottobosco che di quelle diagnostiche, le quali hanno inoltre mostrato una maggiore eterogeneità ed equiripartizione all'interno della comunità. Questo porta a considerare che nelle fustaie il regime di taglio ha assicurato una copertura costante nel tempo, favorendo quindi il mantenimento di condizioni microclimatiche più stabili, che hanno determinato una maggiore abbondanza sia delle specie sciafile che primaverili. Al contrario, nei cedui invecchiati la densa copertura della volta arborea sembra avere influenzato negativamente la ricchezza del sottobosco riducendo le specie più eliofile, mentre le specie più sciafile, tipiche di foreste mature, sono presenti, ma con abbondanze minori. Al fine di comprendere le implicazioni della conversione dei cedui invecchiati sulle funzioni ecologiche dell'ecosistema di faggeta, lo stesso gruppo di rilievi è stato analizzato sulla base dei plant functional traits, creando una matrice di rilievi x traits. Attraverso una Redundancy analysis (RDA) è stata valutata la relazione tra traits e gestione selvicolturale, mentre è stato utilizzato un test U di Mann-Whitney per analizzare le differenze in ricchezza dei traits tra i due sistemi selvicolturali. I risultati emersi sono coerenti con quelli precedentemente ottenuti, relativi alle specie. V.

(11) diagnostiche dell' Habitat; le fustaie hanno mostrato infatti una maggiore affinità per i traits tipici di foresta matura, mentre i cedui invecchiati si sono rivelati maggiormente legati a caratteri relativi a foreste gestite. Inoltre, nelle fustaie è stata rilevata una maggiore abbondanza di traits legati al naturale cambiamento stagionale delle foreste, indice di una migliore ripartizione delle specie tra le nicchie funzionali. Anche in questo caso, i cedui invecchiati hanno dimostrato di essere in una fase transitoria, ancora rappresentata da caratteri legati alla gestione, in cui i traits legati ad aspetti maturi sono comunque presenti, ma con minore abbondanza. In conclusione, considerando che in un ecosistema forestale il sottobosco rappresenta più del 90% della ricchezza specifica, le moderne strategie selvicolturali devono tenere in considerazione gli effetti della gestione sulla biodiversità forestale, in modo da adottare interventi sostenibili, in grado di favorire la biodiversità tipica degli ecosistemi che si vogliono gestire. Questa tesi fornisce indicazioni utili per la gestione in quanto i risultati ottenuti mostrano che, nelle foreste studiate, la conversione del ceduo invecchiato può essere una buona strategia di gestione, se il nostro obiettivo è la conservazione nel tempo delle specie tipicamente legate a condizioni di foresta matura.. VI.

(12) Papers published or prepared in the course of the PhD. Paper 1 Scolastri A., Cancellieri L., Iocchi M., Cutini M., 2014. Patterns of floristic variation on a montane beech forest in the central Apennines (central Italy). Plant Sociology 51 (2): 69-82.. Paper 2 Scolastri A., Cancellieri L., Iocchi M., Cutini M. Old Coppice vs High Forest: the impact of beech forest management on plant species diversity in central Apennines (Italy). Biodiversity & Conservation, submitted.. Paper 3 Scolastri A., Bricca A., Cancellieri L., Cutini M. Old Coppice vs High Forest: effects of two management types on understory functional composition. Plant Ecology, submitted.. VII.

(13) VIII.

(14) General introduction. Forests as complex biological systems In broadleaved forest communities, the floristic composition is the result of the combined effect of several factors, acting at different levels: at a broader scale, environmental variables such as climate, lithology, geomorphology, and also successional dynamics, are great drivers of the forests' coenological differentiation (Van der Maarel, 2005). Moreover, at a finer scale, the major factor shaping the biotic and abiotic conditions is the forest stand, as it creates the specific microclimate and governs carbon and nutrient cycling, dictating habitat conditions for countless organisms (Durak, 2012). In fact, overstory structural features play an important role in modulating floristic composition and ecosystem functions (Neumann & Starlinger, 2001): tree canopy and size, age, stem diameters and density, have a direct influence on the environmental conditions (light, temperature, moisture) at the ground level, and can impact on understory species composition and competition processes (e.g. Thimonier et al.,1992, 1994; Du Bus de Warnaffe and Lebrun, 2004). Growing under the trees canopy for most of the year, understory species are well adapted to the short light supply and show adaptations that are reflected into vegetative and reproductive strategies, that allow them to persist, compete, grow, and reproduce under shady conditions. Understory species represent more than 90% of the floristic diversity in forest ecosystems (Gilliam, 2007) and, by the reasons mentioned above, they are also the most sensible to changes in overstory structure and composition.. Beech forests: an overview Among broadleaved forests, those dominated by Fagus sylvatica are the most abundant in central and southern Europe, thanks to the beech's physiological characteristics, allowing it to spread over a wide range of habitats, soil types and climatic conditions (Peters, 1997; Ellenberg, 1988). As well as in the Eastern Europe (Willner et al., 2009), beech forests in Italy also represent the typical montane vegetation in the Apennine chain and, more generally, in the Mediterranean area (Di Pietro, 2009), as beech is found continuously throughout the whole Peninsula, from the southern Alps. 1.

(15) to Sicily, with the exception of Sardinia and the smaller islands (Jalas & Suominen, 1972-1999); within this range, Fagus sylvatica dominates in deciduous forests, forming both pure and mixed communities from (400) 800 up to 2000 m a.s.l.. Along the Apennine chain, it is the uppermost forest type, showing a pronounced aptitude for expansion in different environmental conditions, including those moderately altered by human activities. Given their key position, Apennine beech forests in many cases comprise a mixture of species belonging to different biogeographical districts; these biogeographical interrelations occur throughout the entire Apennine range, but are particularly represented in its central sector (Lazio and Abruzzo) where the central position, the higher altitude and the complex arrangement of the massifs, all contributed in creating a high degree of environmental heterogeneity (Di Pietro, 2009).. Forest management and its impact on understory Italian beech forests, as well as other European broadleaved forests, have been managed for centuries, since they represent the most common woody resource. The most common management types already existed in the 19th century, when many forests were managed as single coppice or coppicewith-standards (CWS), and some as high forests (HF) harvested tree by tree (Oldeman, 1990; Piussi, 1994), prevalently on the basis of the product needed: firewood, charcoal or poles. Moreover, the consequent disturbance regime has probably modified over time the understory floristic composition, creating communities well adapted to the recurrent disturbance. In a CWS system, young shoots are cut down during short rotations, and new shoots re-sprout from dormant buds on the cut stumps; single-stemmed trees (standards) are retained in a sparse canopy for one or more rotation, in order to ensure genetic diversity and a certain amount of canopy cover. By contrast, HF systems are characterized by single-stemmed trees stands which originate from seed and have prolonged rotation times. The tree stand is strongly affected by the forest management method, where the spatial structure of the forest is shaped by the management method implemented (Burton et al., 2009; Scheller and Mladenoff, 2002); therefore it has a potential role in determining species diversity and ecological stability (Humphrey et al., 2000; Decocq et al., 2004). The effects of forest management depend mainly on its intensity and extent, both of which could induce a disturbance regime in the understory (Van Oijen et al., 2005). Thus, the forest understory diversity and the ecosystem functioning are. 2.

(16) likely to be affected not only by forest management/its lack, but also by the type of the forest management implemented (Decocq et al., 2004, 2005; Schmidt, 2005).. Forest management: changes currently ongoing One significant, though largely overlooked, environmental change that has occurred in recent decades is the large-scale abandonment of CWS management (Baeten, et al., 2009): from '60s, the changing economical and societal demands, caused the progressive decline of CWS management in favor of modern HF management regimes in many parts of Europe (Van Calster et al., 2008; Baeten et al., 2009). These processes are also widespread in Italy, where progressive depopulation along the mountainous areas of the Apennine chain has led to a pronounced drop in local demand for small size timber, firewood and charcoal. As a consequence, many CWS have been almost completely abandoned and most of them are destined to HF conversion (Ciancio et al., 2006; Coppini and Hermanin, 2007). CWS and HF management differ in cut frequency and severity (both higher in CWS): as vegetation composition is partially the result of the environmental conditions created by this management form, the conversion to HF is likely to have caused significant changes in the herb layer (e.g. Barkham 1992; Decocq et al., 2004; Van Calster et al., 2008). In the last decades, the conservation of forest biodiversity has become a key topic in the discussion on the conservation and sustainability of natural resources, given the forests' role as carbon sinks, and the effects of the intense exploitation by man. Even though Fagus sylvatica forests have been intensely studied from several viewpoints, such as ecological (Bartha et al., 2008), functional (Campetella et al., 2011; Canullo et al., 2011), populational (Magri et al., 2006), biogeographical (Willner et al., 2009), we still know little about the effects of forest abandonment and coppice conversion on the understory plant communities. Modern management practices should also consider the forest ecosystem’s diversity, favoring the conditions for the persistence and abundance of the habitat's species.. 3.

(17) Aims Given these premises, in this thesis I aim to analyze differences in floristic richness and composition between old CWS and HF stands, trying to understand which management type most favors the abundance of the understory species typical of beech forest habitat. To do this, I have structured my project into three objectives:. 1. To study the floristic and coenological variability of the beech forests in a district of central Apennines, in relation to the main environmental determinants (chapter 1). 2. To analyze the differences in structural features and floristic richness between old CWS and HF stands, focusing on understory species and 9210* Habitat indicator species (sensu Habitat Directive 92/43 EEC)(chapter 2).. 3. To analyze the differences between old CWS and HF stands in terms of functional composition using plant functional traits, as they reflect the species' adaptations to the environment (chapter 3).. 4.

(18) CHAPTER 1. Patterns of floristic variation on a montane beech forest in the central Apennines (central Italy). 5.

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