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Vegetation on limestone versus phyllite soils: a case study in the west Iberian Peninsula

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  • CITEUC -Department of Earth Sciences - University of Coimbra
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Vegetation on limestone versus phyllite soils: a case study in the west Iberian Peninsula

Abstract and Figures

The influence of soil on plant cover was investigated in two different soil types, on limestone and on phyllite, in the Coimbra peri-urban area. Two areas were selected in each soil type. Soil was analysed for colour, pH, electrical conductivity, moisture, organic matter content, mineralogy , texture and chemical composition. Floristic composition was assessed and abundance was calculated using DAFOR methodology. Data were statistically analysed in Canoco for Windows 4.5. The two soil types have distinct texture and distinct mineralogical, physical and chemical properties. Soil on limestone had pH, electrical conductivity and moisture content higher than soil on phyllite; the latter had higher organic matter content. Soil on limestone had silt clay loam texture and the most abundant minerals were calcite and quartz. The texture of soil from phyllite is sandy loam, loam and silt loam and the most abundant minerals were quartz and mica. Also, most common oxides and trace elements are different. A total of 288 taxa in 61 families (Fabaceae, Asteraceae, and Poaceae are predominant) were identified, showing the enormous diversity of the peri-urban vegetation of Coimbra. A clear distinction was found between the vegetation of the areas of limestone and phyllite; the main environmental factors influencing the ordering of species are pH, mineralogy and anthropic impact.
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F. Covelo, F. Sales, M. M. V. G. Silva & C. A. Garcia
Vegetation on limestone versus phyllite soils: a case study in the west
Iberian Peninsula
Abstract
Covelo, F., Sales, F., Silva, M. M. V. G. & Garcia, C. A.: Vegetation on limestone versus phyllite
soils: a case study in the west Iberian Peninsula. — Fl. Medit. 27: 159-173. 2017. — ISSN:
1120-4052 printed, 2240-4538 online.
The influence of soil on plant cover was investigated in two different soil types, on limestone
and on phyllite, in the Coimbra peri-urban area. Two areas were selected in each soil type. Soil
was analysed for colour, pH, electrical conductivity, moisture, organic matter content, mineral-
ogy, texture and chemical composition. Floristic composition was assessed and abundance was
calculated using DAFOR methodology. Data were statistically analysed in Canoco for
Windows 4.5. The two soil types have distinct texture and distinct mineralogical, physical and
chemical properties. Soil on limestone had pH, electrical conductivity and moisture content
higher than soil on phyllite; the latter had higher organic matter content. Soil on limestone had
silt clay loam texture and the most abundant minerals were calcite and quartz. The texture of
soil from phyllite is sandy loam, loam and silt loam and the most abundant minerals were quartz
and mica. Also, most common oxides and trace elements are different. A total of 288 taxa in 61
families (Fabaceae, Asteraceae, and Poaceae are predominant) were identified, showing the
enormous diversity of the peri-urban vegetation of Coimbra. A clear distinction was found
between the vegetation of the areas of limestone and phyllite; the main environmental factors
influencing the ordering of species are pH, mineralogy and anthropic impact.
Key words: multivariate analysis, Portuguese vegetation, soil on limestone, soil on phyllite, soil
properties vs vegetation.
Introduction
Soils are natural bodies and their properties derive from the combined effect of physical,
chemical and biological processes acting on the parent rock material over time. The result-
ing mineral and organic material is essential to living beings (Blum & al. 2006). Often, soil
provides a satisfactory environment for plant growth and long term establishment of plant
populations. Root growth, development and distribution across the soil profile are affected
by soil chemical composition (Gregory 2006). There is evidence that in the process of land
colonisation, plant lineages adapted differently to the variety of soils (Osaki & al. 2003;
Kenrick & Strullu-Derrien 2014).
Fl. Medit. 27: 159-173
doi: 10.7320/FlMedit27.159
Version of Record published online on 18 September 2017
Plants prefer simple elements, easy to absorb by osmosis. N, P, K, Ca, Mg are the main
nutrients and Fe, Mn, Zn, Cu, Mo and B are essential micro nutrients (Johnston 2005). Soil
with high clay content is more fertile because of its ability to adsorb nutrients; sandy soil
is well drained and tends to have higher temperature. Organic matter corrects nutrient defi-
ciencies, but in high concentration can lower the soil pH due to increase in organic acids
during decomposition (Rheinheimer & al. 2000). Soil pH indicates the activity of the ion
hydrogen in solution. Low pH (<4.5) promotes the dissolution of elements such as Al, Fe,
Mn that may attain toxic concentrations for plants; high pH (>8.0) turns other elements less
absorbable, e.g. Fe, Mn, Zn. Salts in high concentration are also toxic and their content is
assessed by determination of the electric conductivity (Brady & Weil 2012).
In the area of Coimbra (central Portugal, c. 40 km from the Atlantic coast) (Fig. 1), there
are two different types of soil exceptionally close to each other: (1) one developed in the
Jurassic carbonate sediments (limestone, marls, marly limestone, limy marls) belonging to
the Mesocenozoic Lusitanian Basin, and (2) the other developed in the phyllite of Coimbra
– Espinhal – Alvaiázere sector of the Ossa Morena zone, of Neoproterozoic age, belonging
to the Iberian Massif (Serviços Geológicos de Portugal 1992). These carbonate sediments
and phyllite run NNW-SSE almost parallel to each other and are separated by a narrow strip
(400 – 1300 m wide) of a Triassic sandstone (Grés de Silves) that widens south of Coimbra
(Serviços Geológicos de Portugal 1992). The soils in the area are poorly developed (C-hori-
zon is about 40-50 cm deep) and they are classified as calcaric cambisols in the limestone
areas and chromic cambisols in the phyllite areas (Soil Atlas of Europe 2005).
160 Covelo & al.: Vegetation on limestone versus phyllite soils: ...
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The climate of the area is within the Csb of the Köppen-Geiger climate classification
(warm temperate with dry and warm summer; Pell & al. 2007; AEMET & IPM 2011). The
vegetation falls within the Western Mediterranean sub-region, Mediterraneo-iberoatlantic
superprovince of Rivas-Martinez (1987).
The main objective of this investigation was to compare the floristic composition in two
soil types, on limestone and on phyllite, by selecting sites as near as possible to the spatial limit
between the soils. Due to their proximity, climate should not be a factor causing variation in
the floristic composition. Therefore, this area constitutes a model to study the influence of soil
type on plant cover. The area also has a rich native flora in spite of locally anthropic.
Materials and methods
Area selection
The criteria to select the areas for the study in limestone and phyllite were: proximity
between the two soil types and similar altitude in both soil types. In addition, the choice had
to fall on sites with the least possible anthropic disturbance. Four areas (Fig. 1) were selected
and they total 33,600 m2 for each of the two soils: on limestone, (1) Relvinha (13,600 m2;
altitude: 52-91 m; Lat.: 40°13'57.28"N, Long.: 8°26'2.88"W), well-developed herb, shrub and
arboreal strata; long-abandoned olive grove; and (2) Souselas (20,000 m2; alt. 62-106 m; Lat.:
40°16'50.59"N, Long.: 8°24'49.28"W), herb and small shrub strata; on phyllite, (3) Bostelim
(13,600 m2; alt. 200 m; Lat.: 40°16'34.21"N, Long.: 8°23'19.76"W), herb, shrub and arboreal
strata present with the latter very well-developed height-wise; crossed by a path and partially
occupied by a fallow-field; and (4) Torres do Mondego (20,000 m2; alt. 53-88 m; Lat.:
40°11'16.91"N, Long.: 8°23'40.22"W), very dense shrub and well-developed arboreal strata.
Soil sampling and analysis
In each of the four areas, three soil samples were collected (e.g. Relvinha 1, Relvinha 2
and Relvinha 3), each being a composite sample of 4 sub-samples. The proceeding was as
follows: the O-horizont in an area of about 1m2was removed and 4 small excavations were
performed, at about 20-30cm depth, to access the B-horizon, which was sampled. These 4
sub-samples were mixed to create a composite sample.
The weight of each composite sample was c. 3 kg. The samples were oven dried at 40 oC,
sieved through a 2 mm nylon mesh sieve and quartered to obtain the laboratory samples, with
c. 200 g each. The samples were analysed for methodologies readily available at the Earth
Sciences Department, University of Coimbra: (1) colour, (2) pH, (3) electrical conductivity,
(4) soil moisture, (5) organic matter, (6) mineralogy by X-ray diffraction, (7) chemical com-
position by X-ray fluorescence and (8) texture.
Soil colour was determined comparing with the Munsell Soil Color Chart (1994).
Assessment of pH was in a 2:5 suspension of soil:water. Electric conductivity was deter-
mined in the supernatant liquid of a 1:5 mixture of soil:water. The humidity content was
determined by weighing the sample before and after drying at 105 oC for 24 hours. The
organic matter content was determined by loss-on-ignition in a muffle furnace at 360 oC
(Salehi & al. 2011). Replicas of one sample per area were analysed for assessment the ana-
Flora Mediterranea 27 — 2017 161
lytical error using the methodology of Gill & Ramsey (1997). For all these methodologies,
errors were better than 4%.
The portable X-ray techniques are now used by researchers in soil sciences, archaeolo-
gy and biology, environment and geology (Gazley & al. 2014; Lemiere & al. 2014) and in
this study the chemical composition of the <125 µm fraction of soil samples was deter-
mined with a portable Niton XL3t, XRF spectrometer, Thermo Scientific. In each sample
three X-ray “shots” were made and the final result was the average of these, therefore the
sample chemical composition is better constrained. The analytical error was estimated with
replicas using the methodology of Gill & Ramsey (1997) and it was better than 14 %.
Soil texture was determined according to the data from granulometric analysis, which
was performed by laser granulometry using a Coulter LS 230 granulometer. The granulo-
metric analysis was performed after the dissolution of the organic matter and carbonates
by H2O2and HCl, respectively.
For the mineralogy two samples were analysed per area and was determined by X-ray dif-
fraction using a Philips PW 3710 diffractometer, with a Cu tube and 40 kV and 20 mA as oper-
ating conditions. The software APD 3.6J from Philips was used for peaks identification.
Floristic surveys
Surveys took place every month during 2014 with two objectives: (1) a complete check-
list of the vascular plants present in the total size of the areas studied, and (2) a statistical
analysis of the vegetation. The latter was performed along three N/S random transects 20
m long drawn in each of the four areas; along each transect four squares 10 × 10 m were
outlined. Taxa abundance was assessed by the DAFOR scale (Rich & al. 2005).
Plant identification/nomenclature followed Flora iberica (Castroviejo & al. 1986-2014). For
families not yet published, we followed Nova Flora de Portugal: Asteraceae (Franco 1971-
1984); Poaceae (Franco & Afonso 1998). Taxa nomenclature was up-dated according to WCSP
(2014), for Asteraceae to Flann (2009). Family circumscription follows APG IV (2016).
Anthropic impact
The anthropic impact is one of the environmental variables for the statistical analysis of
the vegetation and its assessment was very simple. During the surveys all impacts were
identified and absence/presence was determined for each area.
Statistical analysis of the vegetation
From the 45 environmental variables considered, 19 were selected: pH, electrical con-
ductivity, soil moisture, organic matter, sand, clay, Al2O3, CaO, MgO, Zn, Cu, K2O,
Fe2O3t, quartz, calcite, mica, rain fall, insolation, and anthropic impact. The criteria for this
selection were: (1) relevance to plant development (Cerqueira 2001; Johnston 2005); (2)
the few climate variables with variation between sites (insolation [Souselas and Bostelim:
2500 – 2600 hrs; Relvinha and Torres do Mondego: 2600 – 2700 hrs) and rainfall
(Souselas: 1000 – 1200 mm; Bostelim: 1200 – 1400 mm; Relvinha and Torres do
Mondego: 800 – 1000 mm) (Agência Portuguesa do Ambiente 1931-1960)].
Statistics analysis of data used CANOCO for Windows 4.5 (ter Braak & Ṧmilauer
2002). Analysis started with a Detrended Correspondence Analysis (DCA) (Hill & Gauch
1980). Gradient length of axis 1 was 3.788 indicating high ϐ-diversity (McCune & al.
162 Covelo & al.: Vegetation on limestone versus phyllite soils: ...
2000) and, therefore, it was performed a Canonical Correspondence Analysis (CCA). To
better understand the stronger correlations taxa/ environmental variables, a weight range
of 30% for the most abundant species was applied to the same CCA. The 19 environmental
variables were analysed in a Monte Carlo test for significance p < 0.05 and a meaningful
interpretation from the ecological point of view. The matrix of the environmental variables
was created using mean values since the soil samples per area (three) were fewer than the
quadrats (12). It was performed a Correspondence Analysis (CA) for species and quadrat
ordination according to floristic affinity.
Results
Soil analysis
SOIL COLOUR. In Relvinha the soil is light yellowish brown (10 YR 6/4); in Souselas
very pale brown (10 YR 7/3) and pale brown (2.5 Y 4/7); in Bostelim yellow (10 YR
7/6; 2.5 Y 7/6); Torres do Mondego light reddish brown (2.5 YR 7/4) and light red (2.5
YR 6/8).
PH (Table 1). The soil in the two limestone areas were strongly alkaline (Souselas 8.77 ±
0.09; Relvinha 8.59 ± 0.06) and they were moderately acid to slightly acid in the two
phyllite areas (Bostelim 5.68 ± 0.49; Torres do Mondego 6.34 ± 0.39), according to the
Soil pH classes of the Natural Resources Conservation Service (USDA 2016).
ELECTRICAL CONDUCTIVITY (Table 1). Both soils are non-saline soils, according to
USDA (2011) and the soils on limestone areas had a higher electrical conductivity
(Souselas 71.40 ± 5.63 μS/cm; Relvinha 78.65 ± 14.08 μS/cm) than the soils on the
phyllite areas (Bostelim 30.63 ± 10.66 μS/cm; Torres do Mondego 51.27 ± 13.05
μS/cm).
SOIL MOISTURE (Table 1). Soils on limestone areas retained more humidity (Souselas
5.47 ± 0.84%; Relvinha 4.77 ± 0.40%) than the soils in the phyllite areas (Bostelim 2.24
± 0.48%; Torres do Mondego 2.37 ± 0.08%).
ORGANIC MATTER (Table 1). Soil in Souselas, a limestone area, had the lowest content
in organic matter (2.49 ± 0.55%), while the soil in Relvinha, another limestone area, had
a higher organic matter content (4.01 ± 0.18%), approaching that of the soils in the phyl-
lite areas (Bostelim 4.39 ± 0.33% and Torres do Mondego 4.41 ± 0.88%).
TEXTURE (Fig. 2). Soil texture of the limestone areas is silty clay loam. The soil texture
in the phyllite areas is more variable, with high sand content and little clay, being sandy
loam, loam and silt loam.
OXIDES (Table 1). The most abundant oxide in the soil of Souselas is CaO (31.96 ±
6.47%), but the most abundant oxide in Relvinha is SiO2 (42.93 ± 2.61%). SiO2is also
the most abundant oxide in the soils of the phyllite areas, being higher than in the soil
of Relvinha (Bostelim 43.78 ± 0.48%; Torres do Mondego 47.63 ± 3.70%). Other
oxides, e.g. TiO2, Al2O3, Fe2O3t, K2O and P2O5, follow the pattern of SiO2.
TRACE ELEMENTS (Table 1). The soil in the two limestone areas are also distinct in
trace elements. The most abundant trace elements in Souselas soil are Sc (302.36 ±
121.98 mg/kg) and Sr (262.50 ± 49.98 mg/kg) but in Relvinha soil are Zr (237.97 ±
20.11 mg/kg) and Cr (175.99 ± 9.97 mg/kg). In the phyllite soil, the most abundant trace
Flora Mediterranea 27 — 2017 163
164 Covelo & al.: Vegetation on limestone versus phyllite soils: ...
Table 1. Results of soil physico‐chemical analysis; three samples were collected in each area. - data below the detection limit of the chemical elements.
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^ƌ ϭϬϯ͘ϵϬ ϲϰ͘ϰϬ ϴϮ͘ϲϯ Ϯϲϱ͘ϭϬ Ϯϭϭ͘Ϯϲ ϯϭϭ͘ϭϮ ϭϮϳ͘ϮϮ ϭϲϬ͘ϴϰ ϭϮϮ͘ϲϱ ϭϬϱ͘ϵϯ ϭϬϯ͘ϳϱ ϭϯϭ͘ϰϯ Ϭ͘ϲϵ
Zď ϲϳ͘ϭϲ ϳϬ͘ϱϬ ϳϰ͘ϯϳ ϯϮ͘Ϭϵ ϰϭ͘ϵϮ Ϯϴ͘ϯϭ ϳϳ͘ϱ ϳ ϴϵ͘ϴϮ ϴϴ͘Ϭϳ ϴϲ͘ϯϱ ϴϰ͘ϱϭ ϴϱ͘ϬϮ ϭ͘ϭϳ
dŚ ϭϭ͘ϯϮ ϭϮ͘ϳϱ ϭϭ͘ϱϮ Ͳ ϲ͘ϭϮ ϱ͘ϯϳ ϭϳ͘ϭϮ ϭϴ͘ϳϲ ϭϴ͘ϲϬ ϭϱ͘ϱ Ϭ ϭϰ͘ϯϯ ϭϯ͘ϵϱ ϴ͘ϲϲ
Wď Ϯϲ͘Ϯϴ Ϯϲ͘ϭϰ ϯϱ͘ϲϳ Ͳ ϭϭ͘ϰ ϲ Ͳ Ͳ ϯϮ͘ϬϬ ϯϭ͘ ϳϳ ϰϳ͘ϴϯ ϯϭ͘Ϯϭ ϯϱ͘ϯϳ ϲ͘ϲϲ
Ɛ ϵ͘ϳϳ ϭϭ͘ϱϮ ϭϭ͘ϰϯ Ͳ Ͳ Ͳ ϯϲ͘ϴϬ Ϯϰ͘Ϭϭ ϭϲ͘ϭϲ ϭϳ͘ϯϲ ϭϯ͘ϯϯ ϭϯ͘ϲϭ ϭϯ͘ϰϵ
Ŷ ϴϴ͘ϴϬ ϭϬϬ͘ϴϮ ϲϮ͘ϱϮ ϯϮ͘ϭϵ ϯϯ͘ϭϳ Ϯϯ͘ϱϰ Ͳ ϯϲ͘ϯϵ ϯϵ͘ϵϬ ϰϰ͘ϲϵ ϯϭ͘ϯϮ ϰϱ͘ϰϲ ϳ͘ϯϴ
Ƶ ϯϵ͘Ϯϭ ϯϮ͘ϱϴ ϯϯ͘ϳϮ Ͳ Ϯϲ͘ϰ ϯ Ͳ Ͳ ϯϳ͘ϰϴ Ͳ Ϯϴ͘ϱϵ Ͳ ϱϴ͘Ϭϳ ϭϮ͘ϵϰ
ƌ ϭϴϳ͘ϰϮ ϭϳϭ͘ϰϭ ϭϲϵ͘ϭϯ Ͳ ϭϱϲ͘ϵϯ Ͳ Ϯϭϭ͘ϭϬ ϵϭ͘ϯϯ Ϯϭϰ͘ϴϲ ϭϲϱ͘ϲϬ ϭϰϲ͘ϳϲ ϮϬϯ͘ϰϰ ϴ͘Ϯϲ
s ϭϮϯ͘ϵϯ ϭϮϰ͘ϴϰ ϭϯϵ͘ϵϰ Ͳ ϲϱ͘Ϭϰ ϲϳ͘ϵϰ ϮϬϬ͘Ϯϰ Ϯϱϳ͘Ϯϱ ϮϮϰ͘ϳϬ ϭϳϵ͘ϲϵ ϭϳϴ͘ϭϳ ϭϲϳ͘ϱϱ ϳ͘ϰϭ
^Đ ϭϮϯ͘Ϯϴ ϳϵ͘ϱϲ Ͳ ϯϲϯ͘ϭϱ ϯϮϯ͘ϴϳ ϯϴϭ͘ϵϵ Ͳ Ͳ Ͳ Ͳ Ͳ Ͳ ϯ͘ϱϭ
Eď ϭϲ͘ϴϱ ϭϳ͘ϴϮ ϭϲ͘ϵϯ ϭϬ͘ϭϴ ϭϭ͘Ϯϲ ϳ͘ϲϲ Ϯϭ͘ϵϱ ϮϮ͘ϲϲ ϮϮ͘ϭϬ ϮϬ͘ϮϮ ϭϵ͘Ϭϱ Ϯϭ͘ϳϰ ϰ͘ϰϲ
ŝ ϴ͘ϳϱ ϭϮ͘ϰϯ ϵ͘ϳϭ Ͳ Ͳ Ͳ ϭϳ͘ϭϲ ϭϵ͘Ϭϱ ϭϴ͘ϴϵ Ͳ ϭϯ͘ϳϵ ϭϯ͘Ϯϭ ϭϬ͘Ϭϯ
elements are Zr (Bostelim 344.02 ± 68.34 mg/kg; Torres do Mondego 334.54 ± 1.24
mg/kg) and V (Bostelim 227.40 ± 28.60 mg/kg; Torres do Mondego 175.14 ± 6.62
mg/kg). The content in Zr, Rb, Th, Pb, V, Nb, Bi follows the pattern of the oxides SiO2,
TiO2, Al2O3, Fe2O3t, K2O and P2O5, i.e., being higher in soils of the phyllite areas than
in the soils of the two limestone areas; here they are lower in Souselas.
MINERALOGY (Table 2). The main minerals in the four areas were quartz, mica and clay
minerals in different degrees; feldspar was detected in the phyllite and in Relvinha. Soil
in the limestone area of Souselas is rich is calcite and it has some quartz and mica. In
contrast, soil in Relvinha has a mineralogy similar to that of the soils in the phyllite
areas, in that it has high content of quartz and mica, but it has calcite and dolomite. The
soils in the phyllite areas don´t have calcite. The soil in Bostelim also has a small con-
tent of hematite, and the soil in Torres do Mondego has traces of titanium oxides and
iron sulphides.
Flora Mediterranea 27 — 2017 165
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)LJ'LDJUDPRI VRLOWH[WXUHFODVVLILFDWLRQRIWKHLQYHVWLJDWHGDUHDV 5HOYLQKD 6RXVHODV  %RVWHOLP 
7RUUHVGR0RQGHJR
166 Covelo & al.: Vegetation on limestone versus phyllite soils: ...
/ŶǀĞƐƚŝŐĂƚŝŽŶĂƌĞĂ ZĞůǀŝŶŚĂ ^ŽƵƐĞůĂƐ ŽƐƚĞůŝŵ d͘DŽŶĚĞŐŽ
^ĂŵƉůĞƐ ZĞůǀŝŶŚĂϭ ZĞůǀŝŶŚĂϮ ^ŽƵƐĞůĂƐϮ ^ŽƵƐĞůĂƐϯ ŽƐƚĞůŝŵϭ ŽƐƚĞůŝŵϮ d͘DŽŶĚĞŐŽϭ d͘DŽŶĚĞŐŽϮ
DŝŶĞƌĂůƐ
YƵĂƌƚnj yyyy yyyyydž yyy y yyyyydž yyyydž yyyyydž yyyyyyy
ĂůĐŝƚĞ ydž yydž yyyy
ŽůŽŵŝƚĞ ydž
&ĞůĚƐƉĂƌ dž dž dž dž dž y
,ĞŵĂƚŝƚĞ dž
DŝĐĂ;ŵƵƐĐŽǀŝƚĞ͕ďŝŽƚŝƚĞͿ yyydž yyyydž yyy dž yyyyydž yyyy yyyydž yyyyyy
ůĂLJŵŝŶĞƌĂůƐ;ĐŚůŽƌŝƚĞ͕ǀĞƌŵŝĐƵůŝƚĞ͕ŬĂŽůŝŶŝƚĞͿ dž dž dž dž dž dž dž dž
dŝŽdžŝĚĞƐ dž dž
&ĞƐƵůƉŚŝĚĞƐ       dž dž
Table 2. Results of soil physico-chemical analysis; three samples were collected in each area.
Checklist
Plant diversity was high with 288 specific and subspecific taxa identified in 190 genera
and 61 families: Relvinha, 110 taxa; Souselas, 135; Bostelim, 143; and Torres do
Mondego, 95 (Electronic Supplementary File 1). Bostelim has 49.65% of the total diver-
sity in contrast with the other much larger area in phyllite, Torres do Mondego, this having
only 32.99%. The largest plant families were Fabaceae, Poaceae and Asteraceae; the next
most abundant were Lamiaceae and Orchidaceae. Families with more taxa in limestone
were Asteraceae (Relvinha, 10%; Souselas, 12.59%), Lamiaceae (Relvinha, 5.45%;
Souselas, 8.89%) and Orchidaceae (Relvinha, 4.55%; Souselas, 8.15%) (Table 3).
Families with more taxa in phyllite were Fabaceae (Bostelim, 14.69%; Torres do
Mondego, 16.84%), Poaceae (Bostelim, 13.99%; Torres do Mondego, 12.63%) and
Ericaceae (Bostelim, 4.20%; Torres do Mondego, 3.16%) (Table 3).
Flora Mediterranea 27 — 2017 167
 ZĞůǀŝŶŚĂ^ŽƵƐĞůĂƐ ŽƐƚĞůŝŵ DŽŶĚĞŐŽ dŽƚĂů
ŽĨϮϴϴƚĂdžĂ
&ĂŵŝůLJ ;йƚĂdžĂͿ;йƚĂdžĂͿ ;йƚĂdžĂͿ ;йƚĂdžĂͿ ;йƚĂdžĂͿ
ƉŝĂĐĞĂĞ ϯ͘ϲϰϱ͘ϭϵ Ϯ͘ϭϬ ϯ͘ϭϲ ϯ͘ϰϳ
ƐƉĂƌĂŐĂĐĞĂĞ Ϯ͘ϳϯϮ͘ϵϲ Ϯ͘ϭϬ ϯ͘ϭϲ Ϯ͘Ϭϴ
ƐƚĞƌĂĐĞĂĞ ϭϬ͘Ϭ ϭϮ͘ϱϵ ϭϭ͘ϭϵϴ͘ϰϮϭϬ͘ϰϮ
ĂƉƌŝĨŽůŝĂĐĞĂĞ ϰ͘ϱϱϮ͘ϮϮ Ϯ͘ϭϬ ϭ͘Ϭϱ ϭ͘ϳϰ
ŝƐƚĂĐĞĂĞ Ϯ͘ϳϯϮ͘ϵϲ Ϯ͘ϴϬ ϯ͘ϭϲ Ϯ͘ϰϯ
ƌŝĐĂĐĞĂĞ ϭ͘ϴϮϬ͘Ϭ ϰ͘ϮϬ ϯ͘ϭϲ Ϯ͘ϰϯ
&ĂďĂĐĞĂĞ ϭϬ͘ϵϭϴ͘ϴϵ ϭϰ͘ϲϵ ϭϲ͘ϴϰ ϭϰ͘Ϯϰ
&ĂŐĂĐĞĂĞ ϭ͘ϴϮϬ͘Ϭ Ϯ͘ϭϬ ϯ͘ϭϲ ϭ͘Ϭϰ
'ĞŶƚŝĂŶĂĐĞĂĞ Ϯ͘ϳϯϮ͘ϮϮ Ϯ͘ϭϬ ϭ͘Ϭϱ ϭ͘ϳϰ
>ĂŵŝĂĐĞĂĞ ϱ͘ϰϱϴ͘ϴϵ ϰ͘ϮϬ ϯ͘ϭϲ ϱ͘ϵ
KƌĐŚŝĚĂĐĞĂĞ ϰ͘ϱϱϴ͘ϭϱ Ϭ͘ϳϬ ϭ͘Ϭϱ ϰ͘ϱϭ
WůĂŶƚĂŐŝŶĂĐĞĂĞ Ϭ͘ϵϭϮ͘ϮϮ Ϯ͘ϭϬ ϭ͘Ϭϱ Ϯ͘Ϭϴ
WŽĂĐĞĂĞ ϭϭ͘ϴϮϴ͘ϭϱϭϯ͘ϵϵ ϭϮ͘ϲϯ ϭϭ͘ϭϭ
ZŽƐĂĐĞĂĞ ϱ͘ϰϱϰ͘ϰϰ ϰ͘ϮϬ ϱ͘Ϯϲ ϯ͘ϭϯ
ZƵďŝĂĐĞĂĞ ϭ͘ϴϮϮ͘ϵϲ Ϯ͘ϴϬ Ϯ͘ϭϭ ϭ͘ϳϰ
dŽƚĂůϳϬ͘ϵϮϳϭ͘ϴϱ ϳϭ͘ϯϲ ϲϴ͘ϰϮ ϲϴ͘Ϭϲ
Table 3. Analysis of plant diversity in the four areas. Only families with more than two taxa in at least
two areas were selected; abundance (%) at family level in each area is indicated. Families in bold are
those with the highest diversity.
Statistical analysis of the vegetation
The matrix of taxa abundance (DAFOR data) refers to 196 taxa in 48 squares in a
total of 1.585 entries.
168 Covelo & al.: Vegetation on limestone versus phyllite soils: ...
Table 4. Anthropic impacts identified in the areas investigated. Totals were used in the matrix of envi-
ronmental variables. X: presence of anthropic impact.
/ŵƉĂĐƚZĞůǀŝŶŚĂ ^ŽƵƐĞůĂƐ ŽƐƚĞůŝŵ ĚŽDŽŶĚĞŐŽ
ŐƌŝĐƵůƚƵƌĞy
ĞƐĨůŽƌĞƐƚĂƚŝŽŶy
&ĂůůŽǁĨŝĞůĚy y
'ƌĂnjŝŶŐy y
DŽƚŽƌƚƌĂĨĨŝĐy y
dŽƚĂůƐϭ Ϯ ϱ Ϭ
ŶǀŝƌŽŶŵĞŶƚǀĂƌŝĂďůĞƐ /ŶĨůĂƚŝŽŶĨĂĐƚŽƌ & W
Ɖ,ϱ͘ϴϱϰϭ ϭϬ͘Ϭϱ Ϭ͘ϬϬϮϬ
DŝĐĂϰ͘ϯϬϬϳ ϲ͘ϰϯ Ϭ͘ϬϬϮϬ
ŶƚŚƌŽƉŝĐŝŵƉĂĐƚ Ϯ͘Ϯϵϱϭ ϴ͘Ϭϱ Ϭ͘ϬϬϮϬ
Table 5. Values of F e P (test de Monte Carlo, with 499 simulations)
and inflation factor of environmental variables selected for the pre-
paration of the CCA.
 
ĂŶŽŶŝĐĂůĂdžĞƐϭϮ ϯ ϰ ϭ Ϯ ϯϰ
ŝŐĞŶǀĂůƵĞƐ Ϭ͘ϲϬϰ Ϭ͘ϯϵϵ Ϭ͘ϮϴϬ Ϭ͘ϯϴϵ  Ϭ͘ϲϭϴ Ϭ͘ϰϱϱ Ϭ͘ϯϰϰ Ϭ͘Ϯϵϰ
ĞĐŝĞƐͲĞŶǀŝƌŽŶŵĞŶƚĐŽƌƌĞůĂƚŝŽŶƐϬ͘ϵϵϭϬ͘ϵϴϬϬ͘ϵϳϴ Ϭ͘ϬϬϬ  Ϭ͘ϵϴϰ Ϭ͘ϳϲϭ Ϭ͘ϲϲϳ Ϭ͘ϵϯϭ
ƵŵƵůĂƚŝǀĞƉĞƌĐĞŶƚĂŐĞǀĂƌŝĂŶĐĞ͗
ŽĨƐƉĞĐŝĞƐĚĂƚĂϭϴ͘ϱϯϬ͘ϳ ϯϵ͘ϯ ϱϭ͘Ϯ ϭϴ͘ϵ ϯϮ͘ϴ ϰϯ͘ϯ ϱϮ͘ϰ
ŽĨƐƉĞĐŝĞƐͲĞŶǀŝƌŽŶŵĞŶƚƌĞůĂƚŝŽŶϰϳ͘Ϭ ϳϴ͘ϭ ϭϬϬ͘ϬϬ͘Ϭ  ϰϲ͘ϲϲϳ͘Ϯ ϳϵ͘ϭ ϵϵ͘Ϭ
^ƵŵŽĨĂůůĞŝŐĞŶǀĂůƵĞƐ     ϯ͘Ϯϲϴ ϯ͘Ϯϲϴ
^ƵŵŽĨĂůůĐĂŶŽŶŝĐĂůĞŝŐĞŶǀĂůƵĞƐ     ϭ͘Ϯϴϯ ϭ͘Ϯϴϯ
Table 6. CCA and CA ordination summary.
For the CCA ordination the environmental variables selected from the Monte Carlo test
were pH, mica content of soil and anthropic impact (Table 4), all with p = 0.002; a selec-
tion of 10 from the remaining 16 variables was used as supplementary variables in the
analysis. The inflation factor of the environmental variables that explain species ordination
is < 10 which indicates a desirable low multicollinearity between them (Table 5). The CA
explains 52.4% of the total floristic variation found amongst the four areas (Table 6). As
the eigenvalues of CA and CCA are similar in the first four axes, the environmental vari-
ables included in the CCA explain the floristic variation found in the CA.
The CCA ordination shows different groups along the first and second axes. Along axis
1 are separated the areas of limestone and phyllite. The two limestone areas, Souselas and
Relvinha, are very close to each other and are positively correlated with pH, electrical con-
ductivity, soil moisture, calcite and clay; the two areas of phyllite, Bostelim and Torres do
Mondego, are positively correlated with mica, K2O, organic matter and quartz. Along axis
2 are separated the two areas of phyllite, Bostelim having a positive correlation with both
anthropic impact and rainfall (Electronic Supplementary File 2).
In the CA ordination, species and quadrats have a similar relation to that in the CCA,
although the quadrats overlap in the CCA but split in the CA. The quadrats in Bostelim are
clearly split in two groups in the CA (Electronic Supplementary File 3).
Discussion
Soil analysis
Soil pH and soil electrical conductivity of soil influence the amount of salts available to
plants. The differences in pH found in the two soil types studied are mainly due to the min-
eral composition of soils, which is controlled by lithology. Electrical conductivity is pro-
portional to the content in soluble salts. In the limestone soils, both pH and electrical con-
ductivity are high, due to easy calcium carbonate dissolution that causes higher pH and
higher content in dissolved salts.
In the two limestone areas, the soil has higher clay content (c. 32.62%). This soil type
influences plant development in different ways. The little permeability of such soil leads
to the accumulation of water at the surface and reduces infiltration to lower horizons.
Water accumulated in small depressions explains the local presence of a few Cyperaceae.
On slopes, overland flow takes place and the soil becomes drier, which explains the pres-
ence of Carlina gummifera, a species adapted to water stress (Barceloux 2012). This soil
with small macro-porosity is compact and, therefore, with little oxygen.
The soils on the two phyllite areas are loamy and silt-loamy with lower clay and higher
sand content (36.61%) than in the limestone areas (10.02%). Because of their higher perme-
ability and macro-porosity, water runs through them quicker and, therefore, they are drier than
the soil of Relvinha and Souselas. Soils with a silty to sandy loam texture have a cation
exchange capacity (CEC) of 10-15 meg/100g, while clay loams have a CEC of 15-30
meq/100g (Donahue & al. 1977), but the soils in the phyllite areas have higher organic matter
content (4.39 – 4.41 %), than the soils in the limestone areas (2.49 – 4.01 %). As organic mat-
ter has a much higher cation exchange capacity e.g. CEC of 150-500 meq/100g (Birkeland
1974), the soils in the phyllite areas will have higher contents of available nutrients. In fact,
Flora Mediterranea 27 — 2017 169
essential elements to plant development, such as Zn, K, P and Fe (Johnston 2005), are in high-
er concentration in the soils of the phyllite areas than in the soils of the limestone areas.
The lower content in organic matter of the soils in Souselas may result from its mildly
alkaline pH, enough to favour the soil decomposers (Andrews & al. 2003). The higher con-
tent in organic matter in the soils from Relvinha and from the two phyllite areas (Bostelim
and Torres do Mondego) may also be explained by the vegetation itself, here with greater
development of the shrub stratum.
Soils from the phyllite areas have higher concentrations of SiO2, TiO2, Al2O3, total Fe2O3,
K2O and P2O5and lower concentrations of CaO, than soils from the limestone areas (Table 2).
These differences, evaluated with t-Student test, are statically significant and they are
explained by differences in the lithology of the parent material. In these cambisols the parent
material seems to be the main factor controlling the chemical composition of the soils. This
is also true for trace elements as the soils from the phyllite areas have statistically higher con-
tents of Zr, Rb, Th, V and Nb than soils from limestone areas (Table 1) and these have an aver-
age of 254.37 mg/kg of Sc, which was not detected in the soils from the phyllite.
Altogether, the results of soil analysis showed that in the two phyllite areas (Bostelim
and Torres do Mondego) the soils have similar composition but that there are various dif-
ferences between the soils in the two limestone areas (Relvinha and Souselas). The soil
from Relvinha has a chemical composition more similar to the soils from the phyllite areas,
than the soil from Souselas (Table 1). This can be explained by two factors: 1) the parent
material in Relvinha has an higher marl content than in Souselas; 2) in Relvinha the soil
has a transported component, derived from a terrace deposit of red sands (Areias
Vermelhas do Ingote, Soares & al. 1985), located at higher altitude, above up-stream
Relvinha, as quartz pebbles were identified during field work in this soil.
Checklist
The most abundant families are Fabaceae (14.24%), Poaceae (11.11%) and Asteraceae
(10.42%), the typical pattern in the Mediterranean (Table 3), which compares very well
with a nearby area (Barrico & al. 2012). An indicator used for the degree of the
Mediterranean influence is the index of Cistaceae (Cueto & al. 1991). It is considerably
high (2.43%) (Table 3) in the areas here studied, even higher than in Southern Iberian
Peninsula (2.30% in Almeria; Cueto & al. 1991).
Proportionally to the size of the areas, species number in the soils from the limestone
areas is quite similar, whereas it is very disparate in the soils from phyllite areas. Bostelim
has the highest species number (143) of the four areas and Torres do Mondego has the low-
est (95) (Electronic Supplementary File 1). Torres do Mondego has very steep topography
were slope is 40% and, therefore, anthropic impact is the lowest of all areas (Table 4).
Native vegetation has developed well, mainly trees and shrubs, e.g. Arbutus unedo, Cistus
salviifolius, Erica arborea, Myrtus communis, Quercus suber (Electronic Supplementary
File 2) leaving little space for annuals. The high species number in Bostelim is discussed
in the sections below.
Statistical analysis of the vegetation
Species ordination is not substantially altered in presence of the environmental variables
(compare CCA and CA, Electronic Supplementary File 2 and Electronic Supplementary
170 Covelo & al.: Vegetation on limestone versus phyllite soils: ...
File 3). This means that the environmental variables in the CCA are the explanation for the
floristic composition of the areas. The differences between CCA and CA result from the
use of averages for each environmental variable; this leads to the observed grouping of
quadrats in each area in the CCA.
The differences in mineralogy and physico-chemical composition of the soils explain
the clear separation of both the limestone and phyllite areas along the axis 1 in the CCA
(Electronic Supplementary File 2).
Along the axis 2 of the CCA are observed major contrasts between the soils in lime-
stone and phyllite areas. Whereas the two limestone areas are very close to each other,
the two phyllite are further apart (Electronic Supplementary File 2). Although the soil
analysis showed differences between the two limestone areas greater than those
between the phyllite, the vegetation composition does not seem to be affected (at least
in the analysis) by such differences. The major distance along the axis 2 between the
two phyllite areas does not result from the soil features (mica, K2O, organic matter,
quartz). In fact, the anthropic impact shows its highest strength in the direction of
Bostelim placing it away from its counterpart, Torres do Mondego. In fact, many of the
species restricted to Bostelim are ruderal taxa, e.g., Digitalis purpurea subsp. purpurea,
Medicago polymorpha, Raphanus raphanistrum subsp. raphanistrum, Rumex
bucephalophorus subsp. hispanicus and Vicia sativa subsp. sativa. In the CA ordina-
tion, it is clear the heterogeneity of Bostelim itself with one transect placed a long dis-
tance away in axis 2 (Electronic Supplementary File 3). This transect is a small fallow
field and the human impact here is evident in the higher number of annuals.
The analysis showed a number of taxa with stronger affinities to particular environ-
mental variables. These taxa are densely placed around the respective squares
(Electronic Supplementary Files 2 & 3). To better understand the stronger correlations
taxa/kind of soil, the same CCA is also given with a weight range of 30% for the most
abundant species (Electronic Supplementary File 4).
Conclusions
Plant diversity was dissimilar in the two soil types studied, one developed from lime-
stone, the other from phyllite. Different parent materials have determined different soil
mineralogy, thus different soil proprieties that, in turn, influenced the vegetation structure
and floristic composition.
The variability of soil proprieties in the limestone areas did not substantially alter the
floristic composition.
The decisive environmental variable for the differences in floristic composition between
the two phyllite areas was the stronger anthropic impact observed only in one of them.
Acknowledgements
We thank Prof. José P. Sousa, Dpt Life Sciences, University of Coimbra, for his valuable advice
on the statistical treatment. We would also like to thank Ian C. Hedge, Honorary Associate of the
Royal Botanic Garden Edinburgh, for revising the English. We are grateful for the comments of the
referees to the manuscript.
Flora Mediterranea 27 — 2017 171
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Addresses of the authors:
Filipe Covelo1, Fátima Sales1, 2, Maria Manuela da Vinha Guerreiro da Silva3, César
Augusto Garcia4,
1Centre for Functional Ecology, Department of Life Sciences, University of
Coimbra, Calçada Martim de Freitas, 3000-456 Coimbra, Portugal. E-mail:
filipe.covelo@student.uc.pt
2Royal Botanic Garden Edinburgh EH3 5LR, Edinburgh, Scotland.
3Centre for Mechanical Engineering, Department of Earth Sciences, University of
Coimbra, Rua Sílvio Lima, Univ. Coimbra - Pólo II, 3030-790 Coimbra, Portugal.
4Museu Nacional de História Natural e da Ciência, Centre for Ecology, Evolution
and Environmental Changes, Universidade de Lisboa, Rua da Escola Politécnica, 58,
1250-102 Lisboa, Portugal.
Flora Mediterranea 27 — 2017 173
... Because of the great diversity of edaphic conditions and topography, limestone vegetation in this region is rich in endemic taxa (Zhu, Wang, Li, & Sirirugsa, 2003), which makes it remarkably different from the representative vegetation type on acid soils in the same latitudinal zone (Guo, Liu, & Dong, 2011). It has been well documented that vegetation types on limestone contrast, sometimes dramatically so, with those on other bedrock types in other regions of the world (Covelo, Sales, Silva, & Garcia, 2017;Goldin & Nimlos, 1977;Wentworth, 1981;Whittaker & Niering, 1968). However, the species composition and species diversity of plant communities in the limestone region of China are still unclear due to limited vegetation plot data collected in its rugged topography Tang, Lü, Yin, & Qi, 2011;Zhu, Wang, & Li, 1998). ...
... Basin. Other comparative studies have reported the distinct features of limestone vegetation (Covelo et al., 2017;Goldin & Nimlos, 1977;Wentworth, 1981;Whittaker & Niering, 1968). ...
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