• Embed Doc
  • Copy Link
  • Readcast
  • Collections
  • CommentGo Back
Download
 
Regeneration traits are structuring phylogenetic diversity in cork oak(
Quercus suber
) woodlands
Coca, Manuel
1
& Pausas, Juli G.
2
Ã
1
Empresa de Gestio´ n Medioambiental (EGMASA), Junta de Andalucia, C/Johan G. Gutenberg, no. 1, ES–41092Seville, Spain; E-mail mcocap@gmail.com;
2
Centro de Investigaciones sobre Desertificacio´ n (CIDE, CSIC), ApartadoOficial, ES–46470 Albal, Valencia, Spain;
Ã
Corresponding author; Fax
1
34 96 1270967; E-mail juli.g.pausas@uv.es
Abstract
Question:
What factors determine the deviations from therelationship between species richness (which considersspecies as independent entities) and phylogenetic diversity(PD) (which considers species relatedness)? What are theimplications for community composition and phyloge-netic structure?
Location:
Los Alcornocales Natural Park (36
1
03
0
 – 36
1
45
0
N and 5
1
20
0
 –5
1
45
0
W), in southern Iberian Peninsu-la (Spain).
Methods:
We recorded all woody species and geographicalfeatures on 94 (20m
Â
20m) plots of cork oak woodlands.Disturbance information was obtained from the Parkrecords; precipitation was estimated from local maps. PDwas computed as the minimum total length of all thephylogenetic branches spanning the set of species on eachsite. Then, PD was regressed against species richness totest to what extent the unexplained variance in thisrelationship could be accounted for by environmentalvariables and disturbances, and against the representationof species with different regeneration strategies.
Results:
Species richness and PD are strongly related;however, the remaining variability can be explained by:(1) precipitation and disturbance, and (2) the proportionof seeder species. Thus, the PD both of areas with lowprecipitation and high disturbance, and of areas with ahigh representation of seeder species, is lower than whatwould be expected from the species richness.
Conclusions:
Regeneration traits are important in struc-turing plant community composition; specifically, theycontribute to shaping biodiversity in Mediterranean eco-systems. Species richness tends to overestimate bio-diversity in highly disturbed systems.
Keywords:
Biodiversity; Disturbance; Phylodiversity; Phy-logenetic Community Structure; Resprouters; Seeders.
Nomenclature:
Valde ´s et al. (1987).
Introduction
One of the most important topics in ecology isunderstanding biodiversity. And the most tradi-tional and widely used index of biodiversity isspecies richness (Huston 1994; Rosenzweig 1995;Pausas & Austin 2001). This measure assumes thatall species are equal no matter how similar theyare to one another. With the recent advancesin molecular data and phylogenetic techniques,phylogenetic diversity (PD) is becoming more wide-ly used for biodiversity assessments (Forest et al.2007; Helmus et al. 2007). As PD includes the evo-lutionary divergence information for speciesassemblages, the factors determining the differencesbetween this information and the species diversityon a given site are related to the historical processesstructuring the PD. In high-disturbance-prone sys-tems, such as Mediterranean plant communities, thepresence of regeneration traits may determine thepersistence of the species, thus contributing to thebiodiversity structure. Consequently, we hypothe-size that the different plant regeneration strategiesare responsible for assembling Mediterranean com-munities (Verdu´ & Pausas 2007), and thus theyshould explain the difference between species rich-ness and PD.In Mediterranean flora the two main plant traitsfor regenerating after disturbance are: post-fire re-sprouting (R) and post-fire seeding (P) (Bond & vanWilgen 1996; Pausas et al. 2004). Species with onlythe P trait (R
À
P
1
, obligate seeder species) re-generate after disturbance only from seeds stored inthe seed bank; species with only the R trait (R
1
P
À
,obligate resprouter species) regenerate after dis-turbance from vegetative buds. Some species mayhave both traits (R
1
P
1
, facultative species), thusregenerating facultatively from seeds or vegetativebuds. It has been suggested that disturbance favoursthe P
1
strategy (Bond & Van Wilgen 1996; Pausas
Journal of Vegetation Science 20: 1009–1015, 2009
&
2009 International Association for Vegetation Science
 
et al. 2004; Lloret et al. 2005; Pausas & Bradstock2007) leading to an over-representation of seederspecies on high-disturbance sites (Pausas & Verdu´2005, 2008; Verdu´ & Pausas 2007). Because seedtraits, including dormancy, are heritable (Baskin etal. 2000) and taxonomically aggregated (i.e. they arecommon at a given taxonomic level), seeder speciestend to be phylogenetically aggregated (i.e. with acommon ancestry; Verdu´ & Pausas 2007). As ex-amples of this aggregation, many Cistaceas andFabaceae species are seeders (Thanos et al. 1992;Herranz et al. 1998, 1999). Indeed, the loss of theresprouting character has been related to diversifi-cation (Cowling & Pressey 2001; Verdaguer & Ojeda2005) and thus to a greater increase in richness thanin PD. Consequently, we predict that PD (i.e. con-sidering the relatedness of the species) should belower in relation to species richness (i.e. phyloge-netic clustering; Webb et al. 2002) where seeders aredominant. In addition, because seeder species aremore drought tolerant than resprouters (Paula &Pausas 2006; Pratt et al. 2007), we also predict thatPD should be lower in relation to species richness inthe driest parts of the landscape.Cork oak forests are an emblematic Mediterra-nean ecosystem of the western Mediterranean Basin(Pausas et al. 2009). They have been used for cen-turies to obtain acorns (for human and animalconsumption) and to produce cork. These wood-lands are still in use, especially for cork production,and shrub clearing to facilitate cork extraction andreduce fire risk is a common practice. Under-standing the structure of this ecosystem biodiversityis especially important in Los Alcornocales NaturalPark (southern Spain), the largest existing protectedCork oak forest (Aronson et al. 2009). Given itscomplex disturbance history, the Cork oak forest isa good system to evaluate whether disturbance andregeneration strategies are structuring its biodi-versity. Specifically, we aim to test whether havingseed traits that allow populations to persist throughdisturbances lowers the PD one would expect fromthe species richness.In this framework, we carried out field samplingto quantify woody species richness in different con-ditions (disturbance and climate) in the LosAlcornocales Natural Park. Using available phylo-genies, we computed PD for each site. We thentested whether the unexplained variance in the re-lationship between PD and species richness could beexplained by climate and disturbance factors, andby the representation of the different regenerationstrategies (seeding and resprouting).
Methods
Study area
We studied the Cork oak forests from Los Al-cornocales Natural Park (36
1
03
0
 –36
1
45
0
N and5
1
20
0
 –5
1
45
0
W; 167767ha), located just north of theStraits of Gibraltar, in the southern Iberian pe-ninsula (Spain; Fig. 1). This Park harbours thelargest cork oak forest in the Iberian Peninsula andis probably the best conserved in the whole dis-
Fig.1.
Location of Los Alcornocales Natural Park, on the Iberian side of the Strait of Gibraltar.
1010 Coca, M. & Pausas, J. G.
 
tribution area of the species (western Mediterraneanbasin; Aronson et al. 2009). The landscape is definedby a series of mostly north-south oriented mountainranges with a maximum altitude of 1092m. Soils areacidic and poor in nutrients, and originated fromOligocene-Miocene sandstones. The temperaturesare mild and highly predictable: annual average
5
15.7
1
C; average maximum
5
34.3
1
C; average mini-mum
5
2.78
1
C (CMA 2004). Total annualprecipitation varies between 700 and 2000mm, withmost occurring between September and May. In thestudy area, altitude and rainfall are positively cor-related (altitude and rainfall:
r
5
0.79,
P
5
0.012,based on nine meteorological stations with morethan 17 years of data).
Sampling
To sample a broad spectrum of cork oak forestsinthe studyarea, a stratified sampling was performedbased on a previous classification of the cork oakland properties in the Park (Coca 2007). Fourteenrepresentative properties were selected, ranging from150 to 650ma.s.l. Each land property was dividedinto homogeneous areas from the point of view of management and physiognomic features, yielding atotal of 32 land units. In each land unit, we randomlyselected three (rarely two or four) 20m
Â
20m plots (atotal of 94 plots) to account for the additional varia-bility within the land units (e.g. slope, aspect and soilcharacteristics). On each of these plots all the rootedwoody species as well as the geographical features(aspect, location, and altitude) were recorded.
Data analysis
A plot was considered disturbed if it had beenheavily cleared (usually with the use of machineryand affecting the soil) at least once since 1960 (theearliest year with reliable disturbance history in-formation). The precipitation regime was estimatedfrom local maps (UTE 1990) and classified in twocategories: low (
o
1300mm) and high (
4
1300mm).As previous analyses with more precipitation cate-gories showed very similar results, we used only twocategories for the sake of simplicity.We computed not only woody plant species rich-ness but also the richness of species presenting thedifferent post-disturbance regeneration traits (re-sprouting and seeding). Regeneration traits for eachspecies were assigned on the basis of field experienceandfromatraitdatabase(Paulaetal.2009;Fig.2).Tocompute the PD for each site we first assembled aphylogenetic tree with branch lengths (million years)for the species in all plots with the help of the Phylo-matic software implemented in
PHYLOCOM
3.41 (Webbetal.2007)andusingtheangiospermmegatreeversionR20050610.new (Fig. 2). Then, from this tree, and foreach site, the minimum total length of all the phylo-genetic branches to span the set of species on the site(Faith 1992) was computed using
APE
software (Para-dis et al. 2004).First, PD was regressed against species richness;then, we used this relationship as a null model andtested to what extent the remaining variance couldbe explained by: (1) environmental variables (pre-cipitation and aspect) and disturbance (yes/no), and(2) the representation of the species with differentregeneration strategies. To evaluate whether someof the unexplained variance can be attributed tospatial factors, the land units (nested in land prop-erty) were added to the model. To visualize theinfluence of the studied variables in explaining PD,the residuals from the PD-species richness regres-sion were plotted against the significant variables. Abinomial GLM model evaluated with a
w
2
test wasused test the different representations of seeder spe-cies in relation to precipitation and disturbance.
Results
The total number of woody species on the sam-pled plots was 69, and ranged from 7 to 30 speciesper plot (plot size
5
400m
2
). Obligate seeders(R
À
P
1
) ranged from being absent to representingup to 35% of the total species (mean
5
18%); ob-ligate resprouters (R
1
P
À
) ranged from 50% to96% (mean
5
71%); and facultative species(R
1
P
1
) ranged from absent to up to 17% of thespecies (mean
5
6%).Both PD and species richness were strongly re-lated (
r
5
0.91,
F 
1,92
5
467.7,
P
o
0.0001). Afterincluding species richness in a regression model, theproportion of species of any of the four regenerationgroups (tested independently) explained a significantproportion of the unexplained variance (Table 1). Therelation was negative both with the proportion of ob-ligate seeder species (R
À
P
1
), and with theproportion of all seeders (R
À
P
1
and R
1
P
1
). Con-versely, the relation was positive with the resprouters(all resprouters or obligate resprouters; Table 1).Thus, seeder species (P
1
) contribute to reduce PDfrom the values expected from the species richness(Fig. 3), while resprouting species increase PD.The initial model (PD against species richness)was also significantly improved by including pre-cipitation regime and disturbance (Table 2). Aspect- Regeneration traits and phylogenetic diversity - 1011
of 00

Leave a Comment

You must be to leave a comment.
Submit
Characters: ...
You must be to leave a comment.
Submit
Characters: ...