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How alpine landscapes enhance contrasting
vegetation mosaics and flora in the Pyrenees
Josep M. Ninot, Estela Illa & Empar Carrillo
1Institut de Recerca de la Biodiversitat (IRBio) & Dept. de Biologia Evolutiva, Ecologia i Ciències
Ambientals, Universitat de Barcelona. jninot@ub.edu
Abstract
Alpine landscapes include strong abiotic gradients, which promote the coexistence along
short distances of varying plant communities. Here, we analyse the vegetation of the
alpine belt of the Catalan and Andorran Pyrenees through functional plant traits, to better
understand plant diversity as coupled with the varying alpine landscapes. For this purpose,
we used 800 phytosociological relevés to characterize the associations in terms of plant
traits, and to relate them to species diversity and functional properties.
Most of the community-types assessed reflect the distinct stressing conditions found
in alpine environments, and are species-poor at the plot level. This is more evident in
particularly limiting environments, such as scree, rocky areas or snowbeds which, however,
host many singular species in the biogeographic and functional aspects. Most grassland
vegetation reflects better ecological conditions; community-types are species-richer, and
include great regional and ecological variation. Alpine heaths respond to the local fitness
of a few woody species able to exert dominance through persistence. There is still a lack
of knowledge on the actual effect of grazing on the relative role played in contemporary
landscapes by alpine heaths and grasslands.
Resumen
Los paisajes alpinos incluyen notables gradientes ambientales, que conllevan la coexistencia
de variadas comunidades vegetales en áreas relativamente reducidas. En este trabajo
analizamos la vegetación del piso alpino de los Pirineos catalanes y andorranos a través
de rasgos funcionales de plantas, con el fin de comprender mejor la interrelación entre
diversidad vegetal y la variabilidad del paisaje alpino. Para ello, partimos de 800 inventarios
fitosociológicos para caracterizar las asociaciones en base a rasgos funcionales, y para
relacionarlas con diversidad específica y aspectos funcionales.
Actes del XII Col·loqui Internacional de Botànica Pirenaica - Cantàbrica (Bou, J. & Vilar, L., eds.): 87-99.
2020, Universitat de Girona, Girona.
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La mayoría de las comunidades reflejan algún tipo de condiciones de estrés, comunes
en ambientes alpinos, y resultan pobres en especies a escala de parcela. Esto se hace más
evidente en los ambientes más limitantes, como son roquedos, pedrizas o neveros, donde
en cambio habitan muchas especies singulares tanto en el aspecto biogeográfico como en
el funcional. Los pastos densos traducen en general condiciones más favorables; incluyen
asociaciones más ricas en especies, que muestran una gran variabilidad regional y ecológica.
Los matorrales alpinos reflejan adaptaciones locales de unas pocas especies leñosas, capaces
de ejercer dominancia por el hecho de ser persistentes. Un aspecto todavía poco conocido es
hasta qué punto los efectos del pastoreo han condicionado la extensión y el rol que pastos
y landas juegan actualmente en los paisajes alpinos.
Introduction
One of the most apparent properties of the alpine landscapes is the contrasting array of
plant communities that they include. Although most of these plant communities show
moderate species richness, the contrast from one to another in terms of physiognomy,
plant composition and ecologic function gives to these landscapes a high overall diversity
(Körner, 2003; Onipchenko 2004; Fig. 1). Moreover, they include a number of plant species
of biogeographical interest (Braun-Blanquet, 1948; Gómez et al., 2017).
The flora found in the Pyrenean alpine belt reflects the main constraints imposed by high
elevation environments through its life-form spectrum, where strong seasonality involving
a pleasant growth period favours dominance of hemicryptophytes (Illa et al., 2006). They
respond to seasonality renewing all or almost all aboveground structures yearly, a strategy
coupled with varying herbivory by farm or wild fauna (Grime, 2001). Other life-forms are
mainly found in contrasting habitats, from which those implying particular plant limitations
(such as dryness, soil scarcity or shortened growing season) are particularly place for a wide
array of chamaephytes (Illa et al., 2006).
Most of the Pyrenean alpine flora is shared with other great mountains of central and
southern Europe as a result of repeated interchange throughout Pleistocene cold periods.
Other orophytes are restricted to the Pyrenees, mainly secluded in rocky habitats and secondly
in grasslands; overall, Pyrenean endemics account for roughly 12% in the alpine belt. Other
important chorotypes in the alpine belt are Boreo-Alpine disjuncts, and medio-European
(= Euro-Siberian) taxa (Gómez et al., 2017).
The study of the Pyrenean alpine vegetation started in the mid twentieth century, and mainly
by Braun-Blanquet (1948). Following the phytosociological method, he established the
most common plant associations in the eastern Pyrenees and the regional syntaxonomical
scheme. Some decades later, a good deal of particular papers enlarged and deepened this
knowledge, mostly referred to the central and eastern sectors (Gruber, 1978; Rivas-Martínez
et al., 1991; Carrillo & Ninot, 1992; Carreras et al., 1993; Vigo, 1996; etc.). This has led to
a good knowledge of the alpine plant communities in this area, and has facilitated regional
syntheses of the Pyrenean vegetation (Ninot et al., 2007 and 2017b).
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At the same time, the cumulative knowledge on plant traits grew into a useful tool to address
comparative plant ecology (Grime et al., 1988) and led to acknowledge the role played
by plant species in structuring vegetation through contrasting plant strategies fitting to
distinct environments, and then co-occurring in recurrent plant assemblages (Grime, 2001).
Moreover, behind focusing on plant competence as the mechanism structuring vegetation,
positive interactions between plants took more relevance as alternative explanatory processes
(Pugnaire, 2010). Particularly in stressing environments (e.g., alpine tundra, fellfields)
facilitation effects would shape plant communities. Interestingly, this enriched view of
vegetation structure incorporates the temporal dimension, since plant-plant interactions
change along the ontogeny of individuals. Therefore, plant species play a relevant role in
structuring vegetation by means of their functional traits, which are not only response
to environment gradients, but also drive vegetation and ecosystem processes (Lavorel &
Garnier, 2002; Grime & Pierce, 2012).
Figure 1. Example of an alpine landscape at the Espot valley, central Pyrenees, where contrasting habitats
and plant communities may be found within short distances. Rocky areas (a) and scree (b) cover most of
the steeper relieves, together with staircase grasslands (c) and heaths (e); even grassland (d) settle on gentle
areas; and water-related ecosystems (f) and snowbeds (g) occur restricted to particular environments.
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In this paper we use the example of the alpine vegetation of the Catalan and Andorran
Pyrenees to illustrate how phytosociology, behind a classification tool, is a strong
framework for analysing the relationship between plant cover and environment, and thus
for understanding plant diversity through the varying alpine landscapes.
Materials and method
For this purpose, we used 800 phytosociological relevés gathered in a vegetation databank
(Banc de Dades de Biodiversitat de Catalunya; Font, 2009) which have been classified into
46 community-types (associations). Then we have characterized the associations assessing
the most frequent plant species by means of a wide scope of traits, summarized at the relevé
level, and then at the association level.
Then we selected nine plant traits covering different aspects of plant life, thus making a
good approach to a complete plant strategy, and feasible to be assessed for a high number
of species. Each trait was split into a number of discrete states or categories:
• Life-form (i.e., nanophanerophytes, suffruticose chamaephytes, diffuse chamaephytes,
prostrate chamaephytes, reptant chamaephytes, graminoid hemicryptophytes,
non-graminoid hemicryptophytes, geophytes, therophytes)
• Succulence (not at all, semisucculent, succulent)
• Evergreeness (not at all, semievergreen, evergreen)
• Woodiness (not at all, semiwoody, woody)
• Lateral expansion (not at all, few nearby resprouts, few far resprouts, caespitose)
• Dissemination mode (four categories, from closer to farther from the mother plant)
• Seed weight (<0.01 mg, 0.01-0.05 mg, 0.05-0.5 mg, 0.5-5 mg, >5 mg)
These traits were mainly assessed or inferred from standard floras and herbarium material,
but seed weight was mainly obtained from own material. Then we built a main table where
the 46 community-types were characterized by means of the relative cover achieved by the
group of plant species involving any trait state (thus, a table with 46 × 31 cells).
We evaluated the relationships between plant associations on the basis of their adaptive
traits through Principal Component Analyses, one for the whole vegetation, and other for
the main core group of associations (for more details, see Ninot et al., 2018).
Complementarily, we characterized the plant associations in terms of a few descriptors
related to plant structure and function (plant cover, diversity of plant forms), and to plant
diversity (species richness, number of endemics), as mean values of the corresponding
relevés, thus at plot level.
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Results
Vegetation groups based on functional traits
The multivariate ordinations evidenced a few vegetation groups defined by the dominant
plant strategies, which could be related with distinct alpine environments. Thus, the first
ordination performed showed three vegetation groups; from which one formed a dense cloud
contrasting with respect to two more disperse clouds (Fig. 2). One of these corresponded
to seven associations of dwarf to taller heaths and other layering communities, whereas
the other is formed by ten associations settling on rocky areas, surface waters or snowbeds.
The first axis is strongly correlated with woodiness towards its negative part, whereas
the second axis express a wide array of functionally unrelated traits (such as graminoids,
semisucculents or diffuse chamaephytes; data not shown).
Figure 2. Ordination of the 46 associations (black dots with abbreviated association manes) on the two first axes of
the first PCA (accounting for 0.209 and 0.162 of total variance, respectively), according to the weight performed
by the distinct plant traits. Two vegetation groups appeared detached from a dense cloud of other associations.
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The second analysis done within the denser cloud allowed distinguishing four ecological
groups (Fig. 3), from which three more peripheral. These peripheral groups correspond to
ruderal communities, sparse plant communities settling on gliding scree, and discontinuous
grassland adopting a staircase appearance. Finally, the larger group includes a scope of even
grassland, from medium-sized dense swards to short open pasture. The first axis is related
to plant structure, with graminoid habit in the negative part and creeping chameaphytes
and long-distance expansion in the positive part. The second axis expands the difference
between dense-turfed graminoids versus other hemicryptophytes (data not shown).
Emergent characteristics of vegetation groups
The vegetation groups above presented showed suggesting trends concerning functional
structure and plant diversity at the association level, as shown in Figs. 4 and 5. To better
Figure 3. Ordination of the 29 associations from the denser cloud in Fig. 2, on the two first axes of the
second PCA (accounting for 0.221 and 0.157 of total variance, respectively), according to the weight
performed by the distinct plant traits. Most of them may be distributed into four vegetation groups.
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illustrate these trends, we have split some of the vegetation groups evidenced after plant
traits into smaller groups, following phytosociological criteria, namely according to the
floristic contents of the plant associations.
In the case of structural descriptors (Fig. 4), some vegetation groups reflected the stress
conditions where they occur through low plant cover (rupicolous, scree, riverine). The other
groups achieved clearly higher values, with minor differences between groups exhibiting
distinct physiognomy (e.g., heaths, staircase grasslands, even grasslands). The co-occurrence
of different plant forms is a general trend in the alpine associations, given the high values
of the Shanon index at plot level. Only the ruderal vegetation and the dense grasslands
showed clearly lower values in this index.
Figure 4. Comparison of vegetation groups in terms of two structural descriptors, cumulative plant cover
(top) and diversity of growth forms (bottom) at plot level. These groups correspond partly to those evidenced
in figures 2 and 3, and partly to subgroups delimitated after phytosociological criteria. The boxplots reflect
the distribution of the values for the associations within each group: hth, heaths (5 associations); rup,
rupicolous (5); riv, riverine (2); snw, snowbeds (4); scr, scree (8); stg, staircase grasslands (4); rud, ruderal
(3); fen, fens (3); meg, dense medium-sized even grasslands (5); seg, open short even grasslands (6).
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Species richness and singularity (evaluated as percentage of endemic species) partly
followed parallel trends through the vegetation groups, namely richer plant associations
include more endemics (Fig. 5). This trend is not so in scree vegetation, where species-poor
communities include fair number of endemics; nor in ruderal vegetation, where moderately
rich communities are very poor in endemics.
Discussion
The vegetation groups built through general plant traits give support to a number of
phytosociological groups, mainly at class level. This works where particular environments
drive singular plant traits, thus plant assemblages (e.g., scree, ruderal or heath groups),
Figure 5. Comparison of vegetation groups in terms of two descriptors of plant diversity, species richness
(top) and relevance of endemic species (bottom) at plot level. These groups correspond partly to those
evidenced in figures 2 and 3, and partly to subgroups delimitated after phytosociological criteria. The boxplots
reflect the distribution of the values for the associations within each group: hth, heaths (5 associations); rup,
rupicolous (5); riv, riverine (2); snw, snowbeds (4); scr, scree (8); stg, staircase grasslands (4); rud, ruderal
(3); fen, fens (3); meg, dense medium-sized even grasslands (5); seg, open short even grasslands (6).
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involving physical particularities such as slope dynamics, rock dominance, stronger grazing
pressure, or enhanced development for woody habit. However, contrasting stress types
such as those found in rocky areas and in semi-submerged vegetation give similar plant
assemblages in terms of basic plant forms and traits.
Other main habitat characteristics, particularly those referring to soil richness in carbonate,
do not drive particularities in the plant traits assemblages. Thus, contrasting with strong
differentiation between calcicole and calciphuge associations, the underlying environmental
gradient does not translate into changes in basic plant traits. This is also the case of fens,
where particular flora adapted to hydromorphic soils does not bear apparent functional
distinction.
According to these findings, the alpine vegetation may be ordered following main stress
gradients in the way that Keddy (2005) named ‘Centrifugal model of dominant plants
according diverging stress factors’. In our alpine case (Fig. 6), the core habitats (i.e., the
milder habitats) give advantage to a few potentially dominant species (e.g. Festuca airoides,
F. eskia or Nardus stricta) able to structure rather stable plant communities under moderate
grazing pressure. These occur on flat or gently sloping areas, coupled with general alpine
bioclimate and, typically, neither subjected to singular stress conditions nor to particular
disturbance regimes. The dominant graminoids show higher competitive ability and
persistence through dense caespitose habit, which is coupled with generalized turnover
of above-ground structures. The community strategy here is the response to moderate
stress conditions and moderate disturbance by grazing, which results into fair species-rich
assemblages from plot to region levels (Gómez et al., 2003; Illa et al., 2006). Some of these
associations are taken as climax vegetation (Braun-Blanquet, 1948; Vigo, 1996).
Farther towards different stressing conditions, distinct plant communities hold in the whole
high plant diversity and, particularly, plant singularity, as the result of hosting stress-tolerant
species, which generally show low competitive ability. These are singular species (endemics,
disjuncts) and functional types (pulvinules, semelparous) coupled to one or another stress
category, forming species-poor communities at the plot level (Gómez et al., 2003). This is
clear for distinct stress categories (such as drought, hydromorphy or shortness of growing
season), but also for disturbance (typically, slope dynamics in scree, but also recurrent grazing
and manuring in selected pasture areas). The occurrence of woody species (from dwarf to
low shrubs) able to form dense stable carpets may be understood as a form of biotic stress
for the rest of the plant community, and thus these are similar to plant communities placed
in other stressing positions.
The case of woodiness is certainly particular. Although this would not be a functionally
relevant plant trait in the alpine belt, it enhances dominance at community level, mainly
for plants able to clonal growth. This gives an ecological opportunity for a few dwarf
shrubs, either evergreen or deciduous. Taller shrubs (such as Rhododendron ferrugineum or
Juniperus communis subsp. nana) seem doomed to evergreeness, a plant trait that involves
more investment in persistent leaves (Illa et al., 2017) but that is, together with taller
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canopies, the basis for higher competitive ability. Other evergreens (Arctostaphylos uva‑ursi,
Loiseleuria procumbens) clearly thrive under stressing conditions found in ridges and rocky
snowless slopes, grace to their xeromorphous leaves. Deciduous heaths, contrarily, have
their opportunity in snowy spots, where shorter growing season is compensated by better
soils (Braun-Blanquet, 1948).
In fact, heath vegetation is the only vegetation category within the alpine belt where a
few species (taller woody plants) clearly exert a competitive advantage over a wide species
pool, and thus may drive plant succession to more steady vegetation. The contemporary
prevalence of alpine grassland over heath could be related to very ancient equilibriums
driven by grazing pressure (Gassiot et al., 2017). Pre-humanized herbivores and anthropic
herds might have extended grassland over alpine heath. The contemporary occurrence of
heaths, mainly secluded in more or less rocky areas, could reflect the lower ability of woody
species to recover from grazing disturbance. Present encroachment seems to proceed very
slowly (or irregularly) in the true alpine belt (Montané et al., 2007; Ninot et al., 2017a)
—contrasting with the Arctic tundra (Myers-Smith et al., 2011, Björkman et al., 2018). In
spite of the slowness of clonal shrubs to expand, however, cumulative encroachment could
mean a deterministic recovery of an ancient ‘shrubby lower alpine belt’.
Figure 6. An explanatory model for alpine vegetation according to relevant stress gradients, or
disturbance regimes, represented as centrifugal shadowing going from a balanced community-type
(i.e., climacic alpine grassland) to diverging environments limiting plant life. The shorter sectors of
woodiness and ruderalization reflect limited development of these trends in the alpine belt.
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In some of the stress categories (i.e., slope dynamics, hydromorphy) there is a trend to occur
higher number of plant community-types towards growing stress intensity, namely there
is a number of associations settling on scree (according to distinct chemical and physical
properties of the scree), and fewer grassland types covering sliding slopes or in the core
habitat. This may be understood as more determinism in stressing conditions (where abiotic
properties drive species-poor communities), and higher prevalence of vegetation structure
on the ecosystem under milder conditions.
The strategic scheme of the alpine vegetation shown in Fig. 6 could help at previewing
vegetation trends related to the on-going global change, although clear responses are difficult
to forecast. On the one hand, rising temperatures and increase in rainfall irregularities
(López-Moreno et al., 2009) could affect plant communities presently located in drier
environments (rock crevices, shelves) and reduce the occupancy of mesic grassland and heath.
However, there is no evidence pointing to a real effect of drought on plant distribution in
the alpine. It could really affect the hygrophilous vegetation, by reducing water amounts
and dynamics, and more precisely those depending on direct rainfall (i.e., ombrotrophic
bogs). More clearly, reduced snowfall and warmer summers endanger snowbed vegetation,
as is already recorded in permanent plots of these sensitive plant communities (Illa, 2016).
On the other hand, changes in land use through decreasing extensive herding and allowing
the expansion of spontaneous or introduced herbivores may drive the alpine landscape to
not previewed situations.
Conclusions
The vegetation knowledge acquired through the phytosociology method and gathered
in data banks is a very appropriate foundation to functional attempts, since it consists in
a species-specific evaluation of plant assemblages, related to distinct environments. The
system used relies on the evaluation of relevés (i.e., plot communities) through their species
composition, not by their phytosociological adscription. This illustrates how two distinct
scientific approximations to plant life add particular, valuable knowledge each, and thus
may be used synergistically to improve actual plant knowledge.
High mountain environments involve distinct particular life conditions and processes, which
drive particular biological plants and plant assemblages. Processes such as slope dynamics,
snowpack accumulation in appropriate spots (i.e., snowbeds) or the maintenance of initial
stages of primary succession (i.e., bare rock) generate place for plant species particular in
the functional aspects and in their biogeographical status.
Steady alpine habitats, namely gentle slopes neither affected by noticeable slope dynamics nor
by other environmental particularities, are mostly place for grassland apparently well-coupled
with alpine macroenvironment (i.e., moderately short growing season, moderate grazing
pressure), taken as climax vegetation. However, there is still a lack of knowledge on the
actual effect of grazing on the spatial distribution of main vegetation types; heath could
be actually recovering part of these habitats, mostly in the lower alpine belt.
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