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Acknowledgements

A great number of people contributed to this final work, and to let their contributions go

without mentioning would be unthinkable.

Firstly, I cannot express how much I owe to my Supervisor, Dr. João Tereso. Not only

was he extremely supportive of my pursuit of this theme, he was a constant presence

throughout my whole thesis, providing valuable input whenever requested, guidance

where necessary and much silliness whenever the mood needed lightening. I could

truly not ask for a better supervisor, and would do it all over again if given the chance.

To my Co-supervisor, Dr. Rubim Almeida I also owe a great deal. His classes on plant

biology and taxonomy were quite decisive in determining my choice of scientifical area,

and even today I look back on them fondly. It was also he who introduced me to my

supervisor and in this sense was the driving force that ultimately led to this thesis.

To Cláudia Oliveira, Cristiana Maia and Paula Portela I also have to give my thanks. It

was thanks to their earnest enthusiasm and mostly to a quiet informal chat one lazy

summer afternoon that I chose this course at all. Without their input I might have ended

up in a completely different course altogether. Remember, if it weren’t for you I might

well be stuck in a molecular sciences laboratory somewhere!

To my colleagues in the MEAT, who interspersed these last two years with moments of

fun, camaraderie and madness. We could have been strangers, instead we were

comrades, brothers-in-arms, Tasqueiros. Thanks to you all.

I extend my thanks to those that had the willingness and patience to accompany and

assist me on my field trips. Ana Luísa Ramos, Cláudia Oliveira, Filipe Vaz, João

Tereso, Juliana Monteiro, Paula Portela. There was hard and sometimes difficult work

but also very good moments and I am only sorry that we couldn’t have gone out more

often.

A number of people also provided helpful input, in particular Dr. José Pissarra who

gratiously provided practical advice as well as a borer for use in field work and Carlos

Vila-Viçosa who provided input on sampling points. To them I am grateful also.

To those above and also to Ana Jesus, Cristiana Vieira, Ginevra Coradeschi , Helena

Hespanhol, Joana Marques, Luís Carlos Seabra, Maria Martín-Seijo, Valentina Bellavia

and all others who contributed to a good work environment and provided fond

memories of the last two years. No matter where we are we will always be the group

from lab 1.36.

Ana Luísa Ramos, you were a great companion and a valuable friend throughout all of

this. Even if the vicissitudes of life meant that we weren’t together as much in these last

months, you never ceased to be supportive and present whenever needed and I shall

never forget that.

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Cláudia Oliveira, your input and advice contributed a good deal to this thesis and our

chats about various and diverse subjects, ranging from the taxonomy of “higher” plants

to XX century history, or computer memory hierarchy always managed to cheer up

even the gloomiest of days. I am truly thankful for everything.

To my mother Conceição and father Carlos, for raising me and helping me these long

years, and providing me with an environment where I was free to explore and inquire,

and for supporting my choice to become a scientist. I love you dearly.

To my brother Rodrigo and my sister Priscila, for putting up with me and my (often)

incoherent babbling about some scientifical subject or other. You are both extremely

annoying. I love you.

To Patrícia Martins, missing in action in faraway Britain, but never more distant than a

keyboard stroke. Although in all likelihood you will not be able to attend this thesis

defence, know that I consider you to be there in spirit. I hope we can be together again

soon.

Ana Maria de Melo and Pedro Emanuel da Silva. What can I say. You two are the best

Friends I could have ever had and your companionship over the last 24 years has been

a blessing. Thank you for your friendship, support, patience, affection and mostly, for

being the wonderful people that you are. This thesis is dedicated to you both. Let’s

spend many more decades exploring this wonderful world!


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Abstract

The wood of the Maloideae has long frustrated those that attempted to study it

anatomically. Previous literature has been inconsistent in its treatment of the group,

particularly as to the level of identification possible. In this work we perform a wide

survey of the wood of three common Portuguese Maloideae: Crataegus monogyna,

Pyrus cordata and Pyrus bourgaeana. They are compared in terms of commonly used

wood anatomical characters in order to determine if an identification using these

characters is indeed possible. We conclude that using only these traditional characters,

it is not possible to distinguish the wood of these three species from one another.

Furthermore, we compared the obtained anatomical characters with the growing

environment of our specimens. A few expected and unexpected trends appeared.

Notably, we report the appearance of scalariform perforation plates on some

individuals of Pyrus cordata, a character that to our knowledge was previously

undescribed for this species. We also conclude that enough data has been collected to

suggest that ray-size might be a good indicator of environmental conditions, at least in

this group.

Keywords: Wood anatomy, Maloideae, Ecological gradients

Resumo

A anatomia da madeira das Maloideae há muito que é um tema de estudo frustrante. A

literatura existente trata a anatomia das Maloideae de forma algo inconsistente no que

diz respeito ao nível de detalhe taxonómico que é passível de ser obtido. Neste

trabalho executamos uma pesquisa abrangente das madeiras de três espécies de

Maloideae: Crataegus monogyna, Pyrus cordata e Pyrus bourgaeana. Comparamos as

suas madeiras usando os caracteres anatómicos mais comummente usados de forma

a determinar a possibilidade de uma identificação com base neste critério. Concluímos

que com base nestes caracteres não é possível distinguir a madeira destas três

espécies. Comparámos também os caracteres analisados com as condições

ambientais. Assinalámos a presença de placas de perfuração escalariformes em

indivíduos de Pyrus cordata, um carácter previamente inédito nesta espécie.

Concluímos também que os dados recolhidos são suficientes para sugerir que o

tamanho dos raios poderá ser um bom indicador de condições ambientais, pelo menos

neste grupo.

Palavras chave: Anatomia de madeiras, Maloideae, Gradientes ecológicos


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Table of contents

Aknowledgements ···································································································· 4

Abstract ················································································································· 7

Table of contents ····································································································· 9

State of the art ······································································································· 11

Materials and methods ···························································································· 16

Field Sampling ····································································································································· 16

Laboratory work – preparation of specimens and slides ······························································· 18

Laboratory work – observation and analysis of slides ··································································· 20

Data analyses ······································································································································ 25

Results ················································································································ 27

Sampling sites – environmental characteristics ·············································································· 27

Anatomical characteristics of the species ························································································ 35

Anatomical characteristics and environmental factors··································································· 46

Discussion ············································································································ 57

Anatomical characters of note ··········································································································· 57

Suitability of wood anatomical characters for differentiation between Crataegus monogyna,

Pyrus cordata and Pyrus bourgaeana.····························································································· 62

Closing remarks ··································································································································· 65

References ··········································································································· 67

Annex 01 ·············································································································· 72

Annex 02 ·············································································································· 79

Annex 03 ·············································································································· 81

Annex 04 ·············································································································· 83


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State of the art

Wood is a biological material, produced by higher plants as a complex interaction of

molecular building blocks and microstructural organization. It is a fibrous substance

whose main building blocks are lignin polymers, as well as cellulose, hemicellulose and

residual amounts of other materials, arranged in the cell walls of the xylem of woody

plants (mostly Gymnosperms and Dicotyledons) (Forest products laboratory 1987).

These cells are arranged in turn in complex patterns which are mainly responsible for

the wide variety of properties exhibited by different types of wood. Wood, taken as a

whole, is therefore an extremely versatile raw material, finding use as a construction

material (both in structures and in ships), tools, furniture, musical and sports material,

as well as fuel both processed and not. Despite having been used as a raw material by

humanity since prehistoric times, wood continues to be in high demand in the 21st

century, in applications where by local absence of technological infrastructure, lack of

suitable cost-effective alternatives, or merely by the status it confers see its continued

use.

Apart from its uses in manufacture and industry, wood can also be used as a window to

the past. While the principle of using tree rings to assess the age of a given tree has

been well known since at least the Renaissance, one can also combine this information

with other characters present in wood in order to extract past environmental information

from a given tree (Schweingruber 1996). One can, for example, track the occurrence of

drought years, or unusually harsh winters, or even episodic events such as avalanches

by the signatures they leave on tree rings. By cross dating several trees, it is even

possible to create a continuous history of a regions climate (Douglass 1941). In recent

decades, tree ring research has also aided studies in climate change (Dittmar, Zech &

Elling 2003; Di Filippo et al. 2010), and the study of preserved wood remains

(waterlogged, charred, desiccated or fossil) has been successfully used in archaeology

in order to help reconstruct past customs, practices and environments (Lev-Yadun

2007; Marguerie & Hunot 2007; Martín Seijo et al. 2011).

Tree rings, however, are at their most visible in regions with temperate climates. In

tropical climates, tree rings become fainter due to the lesser degree of seasonal

variability, whilst in regions with more extreme climates, tree rings become too narrow

to be distinguished without the aid of a microscope (Schweingruber 1996). This means

that the main thrust of dendroecological research has so far been primarily on the wood


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of temperate regions. Traditionally, studies have focused on sites with either stable

precipitation (where most variability is therefore attributable to temperature) or stable

temperatures (with variability attributed to precipitation), with studies then proceeding to

temperate regions, where growth is affected by both temperature and precipitation

(Campbell 1949; Von Jazewitsch 1961; Schweingruber 1996). Wood growth also varies

from species to species within the same region (Büntgen et al. 2007). As a result, in

cases where wood cannot be primarily ascribed to a species, it is essential to be able

to reliably identify the wood to the lowest taxonomical level possible.

In the case of many European woods, existing literature allows a reliable identification

down to genus, or even species level (Schweingruber 1990; Vernet et al. 2001;

Akkemik & Yaman 2012). Multimedia tools are also available that can assist in the

identification of many taxa, such as Delta Intkey (Dallwitz, Paine & Zurcher 1993) or the

Inside Wood database (Wheeler 2011), provided that proper diagnosis characters are

on-hand. The science of identifying wood down to a specific taxon relies on the

microscopic observation of three diagnostic sections. These rely on the axial-radial

alignment of the xylemic structures in order to allow a clear visualization of wood

microstructure. The three sections are: Transverse, Longitudinal Radial and

Longitudinal Tangential. If correctly obtained, these allow one to visualize such

characters as: size and arrangement of pores, disposition of parenchyma, ray size and

composition, presence of helical thickenings and/or crystals, among several others.

Fig. 01 – The three diagnostic sections of wood. From left to right: Transverse (Ficus carica, large pores), Tangential

(Pistacia terebinthus, wide rays from head-on), Radial (Pistacia terebinthus, heterogeneous rays in profile).

The significance given to precise numerical measurements of these characters varies:

Both Vernet et al. (2001) and Akkemik & Yaman (2012) make note of pore average

diameter in their descriptions, while Schweingruber (1990) mentions only proportions or

relative size classes. Nevertheless, despite the large number of characters available,


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certain plant groups cannot be reliably identified down to a fine level. Family Fabaceae

and subfamily Maloideae are two such examples. In these cases, wood variability is so

large that a large sampling of individuals would be required in order to reliably ascribe

diagnosis characters to species (Schweingruber 1990). This is particularly unfortunate

in the case of the Maloideae.

Subfamily Maloideae, of the Family Rosacea, includes a series of economically

relevant species. These include: Pyrus communis L. (Pear), Malus domestica (Borkh.)

Borkh. (Apple), Cydonia oblonga Mill. (Quince) and Eriobotrya japonica (Thunb.) Lindl.

(Loquat). Most of these species have been cultivated since ancient times (4000-3000

BCE) (Janick 2005). Nearly all members of this subfamily are edible, either raw or

cooked (e.g. Pyrus, Cydonia), although some are unpalatable and considered famine

foods (e.g. Crataegus). Additionally, a wide range of its members are cultivated as

ornamentals and fragrants (e.g. Sorbus, Crataegus), and a few for their timber (e.g.

Malus) (Hummer & Janick 2009). Several species play an important ecological role.

Therefore, it is considerably unfortunate that the wood of these species is generally

considered undistinguishable according to the current state-of-the-art.

The taxon has undergone a number of revisions in recent times, thanks to the

widespread availability of molecular typing methods that has allowed a more

phylogenetic approach to this group’s classification (Potter et al. 2007; Hummer &

Janick 2009). Most revisions group members of subfamily Spiraeoideae closely with

members of this taxon. In the work by Potter et al. (2007), subtribe Pyrinae is made

part of subfamily Spiraeoideae (Amygdaloideae as of 2011), and now corresponds to

former subfamily Maloideae, now also including the former Prunoideae. However,

since wood science still mostly follows the Maloideae as a subfamily, in part due to it

mapping nicely to economically important species, we shall be using the subfamily

such as defined by Schulze-Menz (Engler & Melchior 1967) in this work.


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In a preliminary exploration of which characters would be most helpful in a putative

attempt to distinguish between wood of members of subfamily Maloideae, as well as of

those that would be most affected by environmental factors, a large sampling was

taken of three members of this taxon: Crataegus monogyna, Pyrus cordata and Pyrus

bourgaeana. These species were selected on the basis of their ease of identification in

the field, and their widespread nature, both in geography and in the environment.

According to Akkemik & Yaman (2012) members of eastern mediterranean Maloideae

can be distinguished by their simple perforation plates, possessing fibres with

distinctively bordered pits, exclusively solitary pores, and rays between 1-3 cells wide.

They further suggest that Crataegus can be distinguished from Pyrus by the presence

of helical thickenings in the narrow vessels of the former, and libriform fibers in the

latter. Vernet et al. (2001) provide a key where Pyrus is distinguished from Crataegus.

Pyrus is characterized as having isolated pores in the latewood 10 to 40 micrometers

wide, as well as narrow heterogeneous rays 1-2 cells wide and up to 20-25 cells high.

Its ray-vessel pits are un-bordered. Crataegus is characterized as having latewood

pores isolated or in groups of two, 15-60 micrometers wide, with homogeneous or

heterogeneous rays 1-4 cells wide and up to 35 cells high. They do however, add the

caveat that distinguishing between Maloideae is difficult. Schweingruber (1999) notes

the Maloideae as being distinguished by their relatively small, isolated and regularly

distributed pores, thick walled fibres, and homogeneous to slightly heterogeneous rays

1-4 cells wide and averaging 15 cells in height. Both Vernet et al. and Schweingruber

note no libriform fibers in their studied species of Maloideae. It should be noted that the

studied species of Crataegus and Pyrus varied between authors and may in part justify

discrepancies in these works (Table 01).


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Table 01 – Maloideae species mentioned in the anatomical studies cited. *Malus communis in the original source.

Anatomical atlas: Species of Maloideae mentioned:

Akkemik & Yaman (2012) Crataegus aronia (L.) Bosc. ex. DC.; Crataegus

monogyna Jacq.; Pyrus serikensis Güner & Duman;

Pyrus syriaca Boiss.; Sorbus torminalis (L.) Crantz.

Vernet et al. (2001) Amelanchier ovalis Medik. ; Crataegus sp.; Cotoneaster

nebrodensis (Guss.) C.Koch; Cotoneaster tomentosus

Lindl.; Cotoneaster integerrimus Medik.; Cotoneaster

vulgaris Lindl.; Pyrus amygdalus Vill.; Pyrus communis

L.; Sorbus aria (L.) Crantz; S.domestica L.; S.torminalis

(L.) Crantz; S.aucuparia L.; Malus domestica Borkh*

Schweingruber (1999) Amelanchier ovalis Medik.; Cotoneaster granatensis

Boiss; Cotoneaster integerrimus Medik. ; Cotoneaster

nebrodensis (Guss.) C.Koch; Cotoneaster nummularia

Fischer & C.A. Meyer; Crataegus calycina Peterm.;

Crataegus laciniata Ucria ;Crataegus monogyna Jacq.;

Crataegus pycnoloba Boiss. & Heldr. ; Cydonia oblonga

Miller; Eriobotrya japonica (Thunb.) Lindley; Malus

domestica Borkh.; Malus sylvestris Miller; Mespilus

germanica L.; Pyracantha coccinea M.J. Roemer;

Pyrus amygdaliformis Vill. ; Pyrus communis L.; Pyrus

pyraster Burgsd.; Sorbus aria (L.) Crantz; S. aucuparia

L.; S. chamaemespilus (L.) Crantz; S. domestica L.


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It is the objective of this work to attempt to exhaustively characterise the wood anatomy

of three Portuguese Maloideae: Crataegus monogyna, Pyrus cordata, and Pyrus

bourgaeana. It will then attempt to discern which differences (if any) between

anatomical characters can be ascribed to the plants taxon, and which are mainly

ascribed to environmental factors. It is hoped such a study will provide a solid basis for

discussion on the status of wood identification and interpretation in subfamily

Maloideae.

Fig. 02 – From left to right: Crataegus monogyna, Pyrus cordata, Pyrus bourgaeana.

Materials and methods

Field Sampling

In order to retrieve sufficient material to permit a study of the different ecological trends,

it was decided to do the widest sampling possible within the temporal and budgetary

constraints available. The species in this study were selected partially due to their

presence in a wide array of environments as well as for their general ease of field

identification and their widespread occurrence. C. monogyna, P. bourgaeana and P.

cordata are all found in forest margins, in a wide variety of soils (Castroviejo 1986-

2012), simplifying their search in the field. As a starting point, the herbarium of the

University of Porto (PO) and the herbarium of the University of Trás-os-montes e Alto

Douro (HVR) were parsed in search of previously collected specimens to which

geographical data could be ascribed. After eliminating those specimens whose


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geographical data was exceedingly poor, 22 usable points were found for C. monogyna,

12 for P. bourgaeana and 9 for P. cordata. In addition to these, two points were added

for P. bourgaeana and one for P. cordata by referencing the Biodiversity4all database.

The coordinates for 6 additional P. bourgaeana points were courtesy of Carlos Vila-

Viçosa, who obtained them in the course of his post-graduate work. In all, 52 points

were identified for sampling. Additional areas of interest, which might yield additional

specimens, were identified based on the Flora-on and Anthos databases.

Sampling was carried out in the course of 8 field trips occurring in the period between

August 7 and November 7, 2015. Trees identified in the field were assigned a

specimen code, had their GPS coordinates marked, some basic local environmental

data noted, and core and wood samples extracted. A total of 120 points were sampled,

spread across continental Portugal. 32 of these points fell within the Atlantic

biogeographical region and 88 in the Mediterranean region. (European Environment

Agency 2016). There was a northernly bias to the sampling effort, due both to the

proximity to the University of Porto and the available geographical information in

planning field excursions. The precise location of each point can be found in Fig. 06

and annex 01.

Some environmental data was retrieved in the field by empirical observation

concurrently with the sampling effort: Sampling point altitude, Shade density (Three

classes), Light direction (Four classes), Soil abundance (Three classes) and Terrain

slope (Three classes). Estimated tree height was also recorded.

Table 02 – Environmental characters registered at the sampling location.

Variable Classes

Altitude a.s.l. (meters) Quantitative

Shade Full sun, partial shade, full shade

Light direction North, East, South, West

Soil abundance Residual, Sparse, Abundant

Terrain slope Negligible, moderate, high.

Height (meters) Quantitative

Core samples were taken using a Haglöf 4.3x400mm two-threaded Pressler increment

borer, with the sample extracted at chest height whenever possible, and at the highest

practical accessible point whenever not. The drill bit was removed once the borer


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reached the midway point of the trees diameter and the extracted core transferred to a

polypropylene straight drinking straw for safekeeping. For larger core samples, two

straws were taped end-to-end.

Wood samples were removed by selecting an adequately sized (more than 5 cm in

diameter) main branch, and extracting an approximately 50mm long section from the

branch. The specimen code was recorded on the sample and a botanical sample was

taken from the plant for addition to the Herbarium of the University of Porto. These

were taken to serve as vouchers for the specimens so that their identification could be

authenticated by later researchers.

As a result of these field sampling expeditions, 120 specimens were retrieved for this

dissertation. These 120 specimens comprised: 58 C. monogyna, 24 P. bourgaeana, 21

P. cordata, 7 P. communis and 10 Pyrus sp.. The decision to take samples from P.

communis was made in the field due to its relative abundance and the contrast it would

provide with non-domesticated Pyrus. All samples of P. communis were either wild or

escaped specimens, with the exception of 4 cultivated individuals in Alentejo. The 10

Pyrus sp. samples correspond to individuals who presented neither fruiting structures

nor their remains, and whose leaves were too intermediate in character to reliably

ascribe to any of the Portuguese Pyrus species. A list of sampled individuals and their

geographical coordinates is available in annex 01.

Laboratory work – preparation of specimens and slides

The extracted wood samples were left to air-dry until they reached moisture equilibrium

with the atmosphere. This method was preferred to kiln drying, both due to its allowing

of a slower drying (minimizing radial deformation), and to the lack of a kiln at the

storage and analysis area, which would severely complicate the logistics of one’s use.

Larger and moister samples, where it was felt air drying would be too slow or leave

them at risk of fungal attack, were instead immersed in 96% alcohol and left to dry.

Alcohol replaces the water inside xylem cells and cell walls, and upon evaporation

causes less deformation upon the wood samples, due to its weaker polarity, and

therefore swell ability, when compared to water (Stamm 1935). The immersion and

saturation of the wood using PEG 1500 (Polyethyleneglycol) was considered (Mitchell

1972), both to prevent deformation and to stabilize the wood during later microtome


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sectioning, but was ultimately rejected, due to the large number of steps required,

increasing both time and effort required for each sample. Linseed oil was applied to the

exposed surfaces of the dried wood samples to reduce re-hydration while they waited

further processing.

Once adequately dried, each sample was processed from its raw state into a

rectangular cuboid with dimensions 20x20x40mm. The faces of these cuboids were

oriented such as to coincide with the three diagnostic sections of wood anatomy. The

cuboid dimensions were sufficient to obtain adequately sized slices from a microtome,

with a healthy margin of error, while being small enough to unobtrusively store in the

space available in the herbarium of the University of Porto. Unprocessed samples were

mounted on a 75mm forged steel vise, and cutting planes were carefully marked before

being cut close to the reference size with a hacksaw. The cut surfaces were then

adjusted by abrasion using aluminium oxide sandpaper with increasingly finer grits,

these being in order: p220, p320, p400, p500 and p600. In practice, it was quickly

ascertained that only the two coarser grits were necessary to obtain an adequate

surface for the microtome, since the microtome’s blade by its own action would polish

the cutting surface. However, the surfaces polished up to the finest grit (p600) did

present a very high contrast with sufficient detail to visualize anatomical structures

when viewed under a stereomicroscope, so it might be worthwhile to pursue finer

abrasion when producing show- and educational pieces.

It is intended for these samples to enhance and modernize the wood reference

collection (xylarium), at the University of Porto. The size of the cuboids is also sufficient

to allow standardised physical tests to be carried out if desired, i.e., specific gravity

(UNE 56531), Hygroscopicity (UNE 56532) and volumetric shrinkage (UNE 56533).

Non-standardised tests other than bending stress tests could also be carried out on

these cuboids (Kasal & Anthony 2004).

Once the cuboids were obtained, they were mounted on an Ulbrecht-Reichert sliding

sledge microtome to obtain 20 m thick wood slices of the tree diagnostic sections. An

initial slice was made to expose fresh wood, and then 3-4 slices were obtained from

one of the surfaces, and gently brushed onto a duly identified petri dish. The cuboid

was then repositioned on the jig, and the process was repeated. This continued until all

three planes were sampled. These then proceeded directly to staining.


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The detailed staining protocol can be found in annex 02. In short, samples were

stained in Alcian blue and Safranine, then dehydrated and transferred to Xylene before

permanent mounting in Canada balsam. After staining, sections were observed under

a Nikon SMZ 2 stereomicroscope to evaluate if they had properly taken up the stain

and if they had any remaining water. In some cases, quality justifying, duplicates or

triplicates were retained. Slides were identified with specimen codes and anatomical

section, and stored pending analysis.

Laboratory work – observation and analysis of slides

Slides were imaged under a NIKON Eclipse 50i compound microscope using

transmitted light and no special filters. Images were acquired using a Nikon DS-Fi1

digital camera and associated NIS-Elements F image capture software. Images were

recorded from the three diagnosis sections. When recording images from the tangential

section, care was taken to image the areas where the rays appeared head-on, as

opposed to the edges of the slice, where due to the geometry of the wood, they could

appear slightly edge-on, distorting measurements.

The images were analyzed using the public domain ImageJ 1.45 (Rasband 1997-2016)

software, with select quantitative and qualitative characteristics noted, as presented on

table 03. Areas were calculated by the use of threshold functions to select only those

areas of interest. Counting of objects was done using the particle analysis function.

Distances were measured using the line tool. Aspect ratios, averages, maximums and

minimums were obtained by using the inbuilt tools in the measurement command. For

all quantitative measurements, a 10x objective was used, corresponding to a 100x total

magnification. When taking into account the cameras narrow aperture, this resulted in a

field of view featuring an area of 750000 m2. Qualitative measurements were obtained

by direct observation of the sample.


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Table 03 – Wood anatomical characteristics selected for imaging, and corresponding diagnosis section.* Homogeneous

rays are those composed solely of procumbent cells. Heterogeneous type I rays have one row of upright cells in their

periphery, Heterogeneous type II rays have interspersed rows of upright and procumbent cells.

Character Description Diagnosis section

evaluated

Number of pores Total number of pores

visible in field.

Transverse

Pore grouping Whether pores are

exclusively solitary, or

appear in clusters.

Transverse

Maximum number of pores

per cluster

The maximum number

of pores observed in a

single cluster.

Transverse

Pore cluster orientation Whether pore clusters

are oriented according

to the tangential

direction, radial

direction, or neither.

Transverse

Total number of pore clusters Total number of

clusters visible in field.

Transverse

Total pore area (m2) The sum total of the

area of all pores.

Transverse

Area of smallest pore (m2) Area of the smallest

pore visible

Transverse


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Area of largest pore (m2) Area of the largest pore

visible

Transverse

Average pore area (m2) Average area of all

pores.

Transverse

Average pore aspect ratio The average of the ratio

between the long axis

and the short axis.

Transverse

Direction of semi-major axis Whether the semi-

major axis has a radial

or a tangential

orientation.

Transverse

Height of tallest ray (m) The height from top cell

to bottom cell of the

tallest ray visible.

Tangential

Height of shortest ray (m) The height from top cell

to bottom cell of the

shortest ray visible.

Tangential

Average ray height (m) The average height of

all rays visible in field

Tangential

Number of rays/mm2 The total number or

rays visible in the field,

divided by field area.

Tangential

Width of widest ray ( No.

cells)

The number of cells at

the widest point of the

widest ray visible.

Tangential


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Width of narrowest ray (No.

cells)

The number of cells at

the widest point of the

narrowest ray visible.

Tangential

Height of tallest ray (No. of

cells)

The number of cells

from bottommost to

topmost, in the tallest

ray visible. When

different cell columns

would yield different

results, the one with the

greater number of cells

was counted.

Tangential

Height of shortest ray (No. of

cells)

The number of cells

from bottommost to

topmost, in the shortest

ray visible. When

different cell columns

would yield different

results, the one with the

greater number of cells

was counted.

Tangential

Presence of two distinct ray

sizes

Whether the rays

visible form two distinct

populations or not.

Tangential

Type of vessel perforations Whether the vessel

perforations observed

were simple,

scalariform, or both.

Radial


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Ray heterogeneity Whether the rays were

homogeneous or

heterogeneous.

Radial

Presence of crystals Whether crystals of any

type were present.

Radial

Presence of helical

thickenings

Whether helical

thickenings were

present in the xylem

(Excluding primary

xylem)

Radial

The selection of these characters for analysis was based mostly on their use in wood

anatomy atlases to distinguish between different taxa (Schweingruber 1990; Vernet et

al. 2001; Akkemik & Yaman 2012). A few were added due to the ease of their recording

(The command that returns the average pore area also allows the return of its

maximum and minimum values.) to evaluate their potential to provide relevant

information. Although not generally considered a diagnostic character, pore eccentricity

data was included since its collection did not add any significant extra effort to the

analysis, and a measurement tool was readily available.


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Fig. 03 – Crataegus monogyna. From left to right: Transverse section, Tangential section, Radial section.

Fig. 04 – Pyrus cordata. From left to right: Transverse section, Tangential section, Radial section.

Fig. 05 – Pyrus bourgaeana. From left to right: Transverse section, Tangential section, Radial section.

Data analyses

After anatomical data was collected, efforts turned to geospatial analysis. Field

coordinates (Taken from GPS using ETRF 89 reference data) were entered into

ArcMap 10.2. These were then superposed over environmental data shapefiles

obtained from Atlas do Ambiente (Agência Portuguesa do Ambiente 2011). The list of

shapefiles used can be found on table 04. For each point, the polygon in each

shapefile to which it belonged was obtained, and the data registered.


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Table 04 – Shapefiles used for spatial analysis. *Time period information unavailable.

Shapefile Description

AtAmb_1001111_Insolacao_Cont Average No. of hours of sunlight

per annum. 1931-1960.

AtAmb_1002111_Temperatura_Cont Annual average of daily mean air

temperature. 1931-1960.

AtAmb_1003111_RadiacaoSolar_Cont Solar irradiance. 1938-1970

AtAmb_1009111_EvapotranspiracaoReal_Cont Real evapotranspiration.*

AtAmb_1013111_CLitologica_Cont Lithological map.

AtAmb_1041111_Precipitacao_NrDiasAno_Cont Average No. of rain days per

annum. 1931-1960.

AtAmb_1042111_Precipitacao_QuantTotal_Cont Average total annual rainfall. 1931-

1961.

AtAmb_1052111_GeadaNrDiasAno_Cont Average No. of frost days per

annum. 1941-1960.

AtAmb_3001111_CSolos_Cont Soil map.

Having collected all the data, it was then organized into an Excel spreadsheet

(Microsoft Office Excel 14.0) that collated the specimen code, species, Geographical

coordinates, anatomical data, site environmental data, and GIS-derived environmental


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data. In order to prepare the data for statistical analysis, non-binary quantitative

information was transformed using equation 01. The transformed data had values

between 0 and 1.

𝑿𝒏 =𝑿 − 𝑿𝒎𝒊𝒏

𝑿𝒎𝒂𝒙 − 𝑿𝒎𝒊𝒏

Equation 01 – Where X is the value of the individual entries, Xmin and Xmax are the minimum and maximum of the

entire value range and Xn is the normalized output of the equation.

In order to search for correlations between the gathered characters, statistical analysis

was performed using the past 3.0 free software program (Hammer, Harper & Ryan

2001). Species and sites were entered into the program as group type data, all others

as generic data types. Spearman correlation analysis was performed in an attempt to

find which characters exhibited a significant taxonomical or environmental signal.

Cluster analysis was performed using the neighbor-joining algorithm with a bootstrap

value of 10000 to try to understand if the species and/or sites clustered naturally

according to the studied characters. A Betula pendula sample analyzed in the same

manner as the other species was used to root the dendrogram. Principal component

analysis was performed to attempt to visualize any explainable trends in the variance of

the data.

Results

Sampling sites – environmental characteristics

A nested percentage table (table 05) summarizes the environmental variables collected

in situ. Not all possible combinations appear in this table since some of these (e.g.

abundant soil in high terrain slope) are far less likely to occur than others. As for the

various classes; 62.5% of the points sampled belonged to the “abundant” soil

abundance class, 60.8% to the “negligible” terrain slope class and 60% to the “full sun”

shade class. This is hardly surprising, reflecting better odds of finding the target


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vegetation when the site is favorable to plant growth. Somewhat surprisingly, of the

trees under some degree of shade, 42.6% belonged to the “North” light direction class,

in spite of Portugal’s location in the Northern hemisphere ensuring that southerly facing

surfaces receive, a priori, more sunlight. The collected environmental data can be

found in annex 03.

Typically, most sampled sites possessed only one of each species under study,

although the sites near Portel and Montemor-o-Novo (Évora district, Baixo Alentejo)

yielded both Crataegus monogyna and Pyrus bourgaeana, and the site near Miradouro

da Boneca (Braga district, Minho) yielded Crataegus monogyna and Pyrus cordata.

Over half the sampled C. monogyna came from north of the river Douro, with 26

individuals proceeding from the Minho region, and 6 from the Trás-os-montes region.

Of the remainder, 8 individuals came from the Beira Litoral, 5 from the Beira Alta and

13 were sampled in the Alentejo. P. cordata and P. bourgaeana were rather more

restricted in their sampling distribution. P. cordata was sampled only once south of the

Douro, in Alentejo. P. bourgaeana was sampled in Alentejo almost exclusively, again

with a single exception, an individual sampled in Minho. This mutually exclusive

distribution was already evident from the distribution data available from various

sources (Flora-on: Flora de Portugal Interactiva. 2014; Anthos. Information System of

the plants of Spain. 2011; Castroviejo 1986-2012). It was initially hoped that individuals

could be collected along the boundaries of these two species’ distribution area, which

would have allowed a more precise understanding of the presence of a gradient in their

anatomical characters, however poor data on sampling points in this area, together

with time constraints prevented this.

Pyrus communis was found almost always in close proximity to P. cordata or P.

bourgaeana, and it often exhibited signs of grafting. In the Alentejo region at least,

exchanges with local inhabitants revealed that such grafting is apparently a cultural

practice, performed almost as if to “tame” wild pear trees, with the result that the

ancestry of some of the trees is potentially problematic. These were nevertheless still

included in the final analysis, under the assumption that they would present either

environmental or anatomical differences that would make them useful as a control

group. These individuals are marked with a † in the table in annex 01.


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Fig. 06 – Location of the sites sampled in the course of this dissertation. Location codes are referenced in annex 01.


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Table 05 – Nested percentages of qualitative environmental data

Full shade 19,17%

Abundant 14,17%

Moderate 5,83%

Negligible 8,33%

Residual 1,67%

High 1,67%

Sparse 3,33%

High 0,83%

Moderate 0,83%

Negligible 1,67%

Full sun 60,00%

Abundant 30,83%

High 0,83%

Moderate 2,50%

Negligible 27,50%

Residual 10,00%

High 1,67%

Moderate 2,50%

Negligible 5,83%

Sparse 19,17%

High 5,83%

Moderate 5,00%

Negligible 8,33%

Partial shade 20,83%

Abundant 17,50%

High 2,50%

Moderate 6,67%

Negligible 8,33%

Sparse 3,33%

High 1,67%

Moderate 0,83%

Negligible 0,83%

Total 100,00%


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A summary of the spatial analysis is provided in figures 07-14. The analysis revealed

that 40% of the samples fell within the cambisol soil class, with 77.5% of these being

humic cambisols. This could be due to the northernly bias in the sampling effort,

Northern Portugal being particularly rich in Humic Cambisols. As an exercise, removing

all sites to the north of the Douro river results in 50% of the remaining sites falling into

the Lithosol category. As for soil lithology, it was mostly granitic or schistose. Together

these two lithologies represented 66.66% of all sites.

The sampling sites fell within three different rainfall categories: 50-75, 75-100 and >100

days, being roughly well represented in all three categories (37.50%, 22.50% and

40.00% respectively), but when taking into account rainfall amount the largest class

was actually 600 to 700 mm (with 25%), a rather moderate amount.

The abundance of sampling sites at the extreme values of evapotranspiration

decreased sharply, as it likewise did for increasing duration of frost. Again, this is likely

a result of the greater abundance of vegetation at the more favorable conditions. A

local maximum at frost levels of 20-30 days corresponds to sampling performed in

moderately well-conserved mountainous regions, the more frost-free lowlands being

under greater anthropic pressure.

Solar radiation was significantly more well represented at the >140 Kcal/cm2 class.

Since northern Portugal is more homogenous in terms of received solar radiation than

the south (Agência Portuguesa do Ambiente 2011) this may reflect a sampling bias.

Amount of sunlight hours was much more evenly distributed, northern Portugal’s

geography allowing a greater amount of variation than the relatively flat Alentejo.


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Fig. 07 – Percentage of sites sampled in each Soil class.

Fig. 08 – Percentage of sites sampled in each Rainfall class.


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Fig. 09 – Percentage of sites sampled in each Total Rainfall class.

Fig. 10 – Percentage of sites sampled in each Evapotranspiration class.


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Fig. 11 – Percentage of sites sampled in each Frost class.

Fig. 12 – Percentage of sites sampled in each Solar radiation class.


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Fig. 13 – Percentage of sites sampled in each Sunlight class.

Fig. 14 – Percentage of sites sampled in each Temperature class.

Anatomical characteristics of the species

The anatomical characters observed showed some interesting trends. C. monogyna

showed less variation than other species for a number of characters, despite having

the greatest number of specimens and being sampled across a greater range of sites.

Despite being typical of regions with higher levels of rainfall, P. cordata showed a mean

average pore area lower than that of P. bourgaeana, a species more typical of areas

0.00%

5.00%

10.00%

15.00%

20.00%

25.00%

30.00%

35.00%

7.5-10 10-12.5 12.5-15 15-16 16-17.5 >17.5

Pe

rce

nt

tota

l

Average temperature; ºC

Temperature


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were water stress is a greater concern. This is the opposite of what was expected and

might be connected to frost resistance strategies as discussed in the next chapter. A

greater occurrence of pore clusters in P. bourgaeana was also noted.

Pore aspect ratio measurements were notably clustered, with the semi major axis

tending to be approximately twice the length of the semi minor axis. Outliers could be

explained by sections in which the pores were at a degree to the normal. Ray density

in P. communis was notably higher than any of the other species. Despite the small

sample number, the magnitude of the difference was still notable.

Most of the specimens retrieved showed two distinct types of ray sizes, one tall and

narrow, the other short and narrow. This was expected since it is a commonly

described anatomical feature of the Maloideae (Schweingruber 1990). Those that did

not show two clearly distinct classes did so by having more intermediate sizes of rays

that blurred the line between classes, and not due to a lack of short rays.

Much more surprising was the presence of scalariform perforation plates in the vessels

of a number of P. cordata specimens (figure 45). Although a small sample, this was

completely unexpected since one of the anatomical features of the Maloideae are their

simple perforation plates (Schweingruber 1990), and to our knowledge, scalariform

plates have not been previously described in a member of this group. The specimens

with this character were all sampled from the same site, (Corno de Bico, Viana do

Castelo district, Minho) and it might represent an environmental adaption to frost,

discussed in the next chapter.

Type I heterogeneous rays were the most common ray type. Type II and homogenous

rays were less frequent but nonetheless present, and Type III heterogeneous were

completely absent (as expected). Prismatic crystals were present in about a quarter of

the samples, but relative proportions varied greatly, in C. monogyna being present in

one-fifth of the samples and in P. bourgaeana in three quarters. When present, crystals

were located in the axial elements i.e. no crystals were found in the ray cells. Helical

thickenings were imaged in 4 of the specimens, an unexpectedly low amount.

Anatomical atlases indicate that the presence of this character in Maloideae wood is

variable (Schweingruber 1990; Akkemik & Yaman 2012), but personal experience

indicated that its presence was substantially more frequent than 4/121.


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Except for one individual (MI1 5.7), all specimens presented their pores with the semi

major axis in the radial direction. Direction of semi major axis was thus dropped as a

variable from further analysis.

Fig. 15 – Boxplot showing the distribution of the Number of pores variable, grouped by taxon.

Fig. 16 –Boxplot showing the distribution of the Total number of pore clusters variable, grouped by taxon.


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Fig. 17 – Boxplot showing the distribution of the Maximum number of pores per cluster variable, grouped by taxon. Note

that only individuals who had pore clusters present were accounted for in this graph.

Fig. 18 – Bar graph showing Pore cluster orientation, grouped by taxon.


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Fig. 19– Boxplot showing the distribution of the Total pore area variable, grouped by taxon.

Fig. 20 – Boxplot showing the distribution of the Area of smallest pore variable, grouped by taxon.


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Fig. 21 – Boxplot showing the distribution of the Area of largest pore variable, grouped by taxon.

Fig. 22 – Boxplot showing the distribution of the Average pore area variable, grouped by taxon.


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Fig. 23 – Boxplot showing the distribution of the Average pore aspect ratios variable, grouped by taxon.

Fig. 24 – Boxplot showing the Distribution of the Height of tallest ray in micrometers variable, grouped by taxon.


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Fig. 25 – Boxplot showing the Distribution of the Height of shortest ray in micrometers variable, grouped by taxon.

Fig. 26 – Boxplot showing the distribution of the Height of tallest ray variable, grouped by taxon.


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Fig. 27 – Boxplot showing the distribution of the Number of rays per square millimeter variable, grouped by taxon.

Fig. 28 – Boxplot showing the distribution of the Width of widest ray variable, grouped by taxon.


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Fig. 29 – Boxplot showing the distribution of the Height of tallest ray in number of cells variable, grouped by taxon.

Fig. 30 – Boxplot showing the distribution of the Height of shortest ray in number of cells variable, grouped by taxon.


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Fig. 31 – Bar graph showing the Presence of two distinct ray sizes, grouped by taxon.

Fig. 32– Bar graph showing the Vessel perforation type, percent of each taxon’s abundance.

Fig. 33– Bar graph showing the Ray heterogeneity, grouped by taxon.


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Fig. 34– Bar graph showing the Presence of crystals, grouped by taxon.

Fig. 35 – Bar graph showing the Presence of helical thickenings, grouped by taxon.

Anatomical characteristics and environmental factors

Principal component analysis (PCA) on the anatomical characters failed to reveal any

significant clustering of data. The first component accounted for merely 16.15% of the

total variance of the dataset, while the second accounted for 13.4%. The first four

components when taken together accounted for 49.1%. An alternate approach using

log transformation of the original data had little effect on either the explained variance,

or the distribution of the samples on the plot. It thus proved impossible to use the PCA

to produce clusters or to reduce the number of variables under analysis.


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Analysis of similarities (ANOSIM) performed using the 5 species as groups showed

high similarity between all groups for all anatomical characteristics. The highest R

value was that obtained for the Rays per square mm character, with a value of 0.138.

Six characters (Total pore area, Height of tallest ray (m), Average ray height, Width of

widest ray, Height of tallest ray (cells) and Presence of helical thickenings) displayed a

negative R value, suggesting that differences were larger within groups than between

them.

The dendrogram produced by the neighbor-joining algorithm reinforced this notion, with

no discernible clusters emerging from the tree, and with several of the nodes showing

0% support after 10000 bootstrap replicates.

ANOSIM was then performed against the environmental variables, again using the 5

species as groups. Overall R values were again low, however pairwise comparison of

the groups did show some trends. P. cordata and P. bourgaeana showed marked

dissimilarity on the following variables: Altitude (R:0.8574 ; P:0.0001), Soil class

(R:0.5706 ; P: 0.0001), No. days rainfall (R:0.6580 ; P:0.0001), Lithology (R:0.5948 ;

P:0.0001), Evapotranspiration (R:0.5639 ; P:0.0001), Solar radiation (R:0.7070 ;

P:0.0001), Mean temperature (R:0.7519 ; P:0.0001), Hours sunshine (R:0.7734 ;

P:0.0001) and Total rainfall (R:0.7130 ; P:0.0001). These differences are quite possibly

the result of the North Portugal/South Portugal divide in the distribution of these two

species. Significant differences were also returned for the Crataegus monogyna/Pyrus

cordata combination for the following variables: Mean temperature (R:0.7519 ; P:

0.0001) and Total rainfall (R:0.6319 ; P:0.0001).

When comparing variables among themselves using Spearman correlation, several

showed a p value under 0.01. These included several that correlated with latitude

and/or longitude, which might go a long way to explain pairwise correlation detected

between several of the environmental variables, since in most cases these have a

stronger correlation with geodata than with each other. When looking at correlations

between environmental and anatomical variables, the following correlations show up as

highly significant: Pore numbers correlated with seven soil classes, but otherwise

showed only a weaker (P=0.0266) correlation with solar radiation; Of the three Pore

cluster orientation classes, Random cluster orientation showed a strong correlation with

Full shade (0.00891); Both the Area of smallest pore and the Average pore area

characters similarly showed high correlation with Full shade (0.004231 and 0.00334


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respectively); The height of shortest ray in m correlated with No. days rainfall

(0.00574) and Evapotranspiration (0.00728); Average ray height correlated with No.

days rainfall (0.000696), Evapotranspiration (0.000933), Solar radiation (0.00225),

Mean temperature (0.00674), Hours sunshine (0.00232) and Total rainfall (0.00446);

the Number of rays per square mm correlated with the Residual soil abundance class

(0.00575) and with Solar radiation (0.00725); Height of tallest ray correlated with Full

shade (0.000601);The Type of vessel perforations correlated with full shade (3.18x10-5)

and with Evapotranspiration (0.000921); Heterogeneous Type I rays were correlated

with partial shade (0.00791) while Heterogeneous type II were correlated with the

Residual soil abundance class (3.53x10-5), No. days rainfall (0.00467),

Evapotranspiration (0.00276) and Solar radiation (0.00360). Homogeneous rays also

correlated with No. days rainfall and Solar radiation (0.00734 and 3.23x10-5

respectively) but also showed correlation with Mean temperature (0.00116), Hours

sunshine (0.000531) and Total rainfall (0.00674). Fifty-four other variable combinations

showed a lesser degree of correlation (0.01 < p < 0.05). These can be visualized in the

table in annex 04.

Several soil and lithology classes presented strong correlations with one or more

anatomical characters. However two classes dominated the sampling for both variables

(Humic cambisols for soils and Granites for lithology), which as discussed earlier is

most likely an artifact of the sampling effort. Furthermore since these variables were

broken up into multiple classes (19 soil classes and 11 lithology classes) many of these

ended up having only a handful of samples. On that basis they are presented here

separately. Of the soils: Humic cambisols correlated with Number of pores (P =

0.00267), Average ray height in m (0.00203), Type of vessel perforation (0.00241)

and Heterogeneous type II rays (0.00421). Schist associated cambisols correlated with

total pore area (0.00193), Area of largest pore (0.00500) and Average pore area

(0.00314). Chromic cambisols correlated with width of widest ray (0.00402) and

presence of helical thickenings (0.000222). Eutric lithosols correlated with

homogeneous rays (2.44x10-7) and Heterogeneous type I rays (0.000715). These exact

same correlations repeated themselves in the Luvisol associated eutric lithosols soil

class. Eutric fluvisols correlated with the number of rays per square mm. Orthic luvisols

correlated with pore grouping (4.92x10-6), Maximum number of pores per cluster

(0.00258), Total number of clusters (0.00325) and Average ray height in m (0.000225).

Albic luvisols correlated with the Maximum number of pores per cluster (0.00523),

Random cluster orientation (0.00712), Width of widest ray (0.00174) and Presence of


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crystals (0.00488). Chromic luvisols correlated with the Tangential cluster orientation

(0.00440) and with Average pore area (0.00195). Distric regosols correlated with Total

pore area (0.00293), Area of smallest pore (0.00879), Area of largest pore (0.0100)

and average pore area (0.00410). Gleyic solonchaks correlated with pore cluster

orientation (0.00406). Orthic podzols correlated with width of narrowest ray (1.31x10-5)

and Homogeneous rays (0.000180).

As for lithology: Granite correlated with average Ray height (0.00401), Homogeneous

rays (0.00677) and Heterogeneous type II rays (0.00341). Schist and greywacke

correlated with Pore grouping (0.00893), Area of smallest pore (0.00320), Area of

largest pore (0.00570) and Average pore area (0.000481). Schist, amphibolites, mica

schist, greywackes, quartzites and gneisses correlated with Height of tallest ray in m

(0.00189), Height of tallest ray in cells (0.000705) and Presence of helical thickenings

(2.88x10-5). Aeolian sands correlated with Total pore area (0.00293), Area of smallest

pore (0.00880), Area of largest pore (0.00985) and Average pore area (0.00410).

Sands, pebble s, clays and weakly consolidated sandstone correlated with height of

shortest ray (0.00581) and width of widest ray (0.00400). Sands, gravels, limestones

and clays correlated with width of narrowest ray (0.000183), Homogeneous rays

(0.000160) and Heterogeneous type I rays (0.00389). Finally, Shale, graywacke and

sandstone correlated with average ray height (0.00682) and homogeneous rays

(2.33x10-5). The full list of correlations can be found in annex 04.


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Fig.36 – PC1/PC2 scatter plot. Dots coloured as per species: Red, Crataegus monogyna; Blue, Pyrus cordata; Yellow,

Pyrus bourgaeana; Violet, Pyrus communis; Black, Pyrus sp.


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Fig. 37 – PC1/PC3 scatter plot. Dots coloured as per species: Red, Crataegus monogyna; Blue, Pyrus cordata; Yellow,



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Fig. 42 – Dendrogram constructed using neighbor-joining algorithm with 10000 bootstrap replicates. The numbers at

each branch represent the percentage of replicates for which the node was still supported. Betula pendula was included

to serve as a rooting point.


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Discussion

Anatomical characters of note

The analysis performed on the wood anatomical characters of C. monogyna, P.

cordata and P. bourgaeana revealed a number of noteworthy facts that merit a more in-

depth discussion.

Fig. 43 – Transverse section of Pyrus cordata Mi1 5.12 (Left) and of Pyrus bourgaeana Aj4 4.1 (Right). Both images

taken at 100x magnification.

As mentioned previously, despite being found mostly in moist northwestern Portugal,

our samples of Pyrus cordata proved to have lower average pore areas than Pyrus

bourgaeana, which were sampled mostly in the drier Alentejo and Trás-os-montes

regions. This is somewhat peculiar, since as a general rule, plants tend to have

narrower pores in more arid conditions (Lovisolo & Schubert 1998; Carlquist 1966). In

terms of conductivity, larger pore diameters are more efficient than narrower ones, as a

result of the Hagen-Poiseuille law:

𝚽 = 𝚷. 𝐑𝟒.∆ 𝐩

8L

Equation 02 – Where is the flow rate (volume), P is the pressure gradient between the ends of the vessel, is the

viscosity of the fluid in the vessel and L is the length of the vessel.

A notable result of this equation is that if a plant were to shrink its pores to half their

diameter (all other things being equal) it would now need 16 times as many pores to

equal the same flow rate, even though these new pores would have a combined pore


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area 4 times larger than the original. So long as a plant is capable of generating

enough negative pressure in its vessels so as to maintain cohesion-tension, it would

benefit from having the widest pores it can support. However wider pores have

disadvantages. Larger pores are at greater risk of embolization (Zimmermann 1983;

Sperry et al. 1994), and their presence is therefore a tradeoff between conductive

efficiency and risk of hydraulic failure. Plants that live in conditions of water stress in

general show a tendency to have smaller diameter pores than plants in more mesic

conditions, and this is often interpreted as a response to increased embolism risk. The

higher negative pressure maintained in the xylem under conditions of drought

increases the probability of embolisms spreading via the air-seeding mechanism,

where air from adjoining air-filled vessels gets aspirated into functional vessels (Sperry

& Tyree 1988). However despite the smaller pores observed in P. cordata, this should

not be taken to mean that the pores of P. bourgaeana are uncharacteristic of arid

climates. The mean pore area for this species was 792.9 m2 which for an ideal

circular pore translates into a diameter of 31.77m. This falls within the size range

typical of other published descriptions of Pyrus in Mediterranean habitats (Akkemik &

Yaman 2012). What is peculiar is rather that P. cordata consistently shows smaller

values than its southerly counterpart (685.5m2 and 29.54m respectively). Since high

correlations were found between average pore area and full shade, one hypothesis is

that the reduction in photosynthetic activity under conditions of shade has an even

more drastic effect on pore size than water stress. The relative frequencies of

occurrence of shade conditions (47.6% P. cordata under conditions of full shade versus

16.6% for P. bourgaeana) would seem to bear this out. The fact that there is only a

weak inverse relationship between Number of pores and Maximum pore size, serves to

reinforce the notion that reduced conductivity and not safety from embolism is the

driving force behind pore size reduction.


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Fig. 44 – Plot of Area of largest pore vs. Number of pores for Pyrus bourgaeana and Pyrus cordata.

In contrast, the greater occurrence of pore clusters observed in P. bourgaeana meshes

correctly with the established literature, where greater occurrence of pore clustering

has been likewise connected to greater water stress, the clusters being interpreted as

offering greater hydraulic safety by creating “bypasses” around embolised vessel

elements, and serving as a reservoir from where embolised pores can be refilled

(Lindorf 1994; Lens et al. 2010; Rita et al. 2015). The fact that the relationship between

greater clustering and aridity is maintained but that of average pore area is not

reinforces the notion that and additional environmental variable is at work where pore

area is concerned. A more directed study focusing solely on these characters might be

warranted in order to better understand the relative response of Pyrus to light and

water stress.

Most significant of all was the presence of scalariform perforation plates in the vessels

of Pyrus cordata sampled at Corno de Bico, since according to the existing literature

(Schweingruber 1990; Vernet et al. 2001; Akkemik & Yaman 2012) one of the

diagnostic characters of the Maloideae is their simple perforation plates. In fact

scalariform perforation plates are rare in the Rosaceae (Eyde 1975). In the sampled

individuals, the perforation plates were exclusively scalariform (i.e. no simple

perforation plates were observed), formed an angle of 10-23 degrees with the

longitudinal axis and possessed 20-24 bars, at least some of which were forked.


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Scalariform perforation plates (particularly those with large numbers of bars and low

angles to the longitudinal) have been described as a primitive character, lost in favor of

the derived simple perforation plates. They mostly persist in species characteristic of

mesic habitats, the number of taxa possessing them increasing with latitude and

altitude (Carlquist 1975; Baas 1976; Lens et al. 2016). Scalariform perforation plates

have been described as having a resistivity-increasing effect on vessels, possibly to as

high as double the value of the unobstructed vessel lumen. Furthermore, their

resistivity seems to place an upper constraint on the diameter displayed by pores

(Christman & Sperry 2010). It is thus argued that they are retained by taxa that are not

faced with high selective pressure for conductive efficiency (Lens et al. 2016).

Fig. 45 – To the left: Scalariform perforation plate of a Pyrus cordata specimen sampled at Corno de Bico, Mi1 5.12. To

the right: Simple perforation plate of another Pyrus cordata, Mi1 5.14, for comparison. Both images are at 500x

magnification.

Although Corno de Bico’s location in the Atlantic bioclimatic region, as well as its

location between the Lima and Coura river valleys, place it as one of the most rainy

sites in continental Portugal (Beja 2008), this alone does not explain the re-appearance

of a presumed ancestral character in a species characterized by the presence of its

more derived counterpart. Zimmerman (1983) suggested that the retention of


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scalariform plates confers some sort of advantage to plants that must endure frost

conditions (Zimmermann 1983). Low temperatures are known to be a risk factor for

vessel embolism, with outgassing caused by the freezing of sap followed by gas bubble

fusion following the re-establishment of negative xylem pressures once sap-flow is

restored leading to vessel occlusion by vapor. It is theorized that scalariform plates

provide plants with a mechanism to combat this phenomenon, either by trapping and/or

splitting gas bubbles as they ascend, limiting their surface area and thus making it

easier to redissolve them in the liquid (Zimmermann 1983), or by forcing subdivisions in

gas bubbles, restricting them to the length a vessel element, simplifying vessel refilling.

(Sperry 1986). Although the plants were in a relatively low temperature area (monthly

averages 8.6 – 21.4 ºC, reaching a minimum of -5ºC in the winter (Beja 2008)), with

moderate amounts of frost in winter (30-40 days (Agência Portuguesa do Ambiente

2011)), they were not the only individuals in such conditions and in fact several others

were sampled in more extreme temperature and frost conditions without exhibiting

scalariform perforation plates. However, they were the only ones that combined these

factors with a high evapotranspiration rate (>800mm (Agência Portuguesa do Ambiente

2011)) and full shade.

Depending on which factor is considered dominant, one can be tempted to offer two

potential explanations for the presence of scalariform plates. If evapotranspiration is

considered the most significant factor, one could expect that the plant’s conductive

fluids would be under a greater amount of negative xylem pressure, due to stomatal

opening and decreased water potential at the plant’s leaves. This would then leave the

plant at a greater risk of embolism, since the decreased pressure would make it easier

for dissolved gasses to come out of solution, and conversely, more difficult for gas

bubbles to dissolve back into the sap (Zimmermann 1983). This combined greater risk

of embolism might be serious enough for the plant to sacrifice conductive efficiency in

favor of greater safety from embolism. However we find this to be the less likely

scenario: although the amount of plant cover means that transpiration is almost

certainly an important contribution to total evapotranspiration, the sampled plants were

all part of the understory, experiencing greater humidity and lower temperature

conditions as well as greater protection from wind that would make transpiration less

intense than the total average for the area would suggest.

If we consider shade conditions to be the driving factor, it can instead be argued that

the reduced levels of photosynthesis results in a higher leaf water potential that in turn


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would lead to a reduced need for high vessel conductivity. The presence of the

scalariform plates may then represent the uptake of a feature whose benefits (greater

protection from embolism) now offset the opportunity cost (upper limits on conductivity).

Previous studies performed on Fagus sylvatica indicate that trees can adjust their

vulnerability to embolisms according to light levels (Cochard, Lemoine & Dreyer 1999).

If we then consider that most of the overstory is composed of deciduous plants such as

oak, whose loss of foliage in the winter months leaves the understory exposed, likely

leading to enhanced transpiration precisely at the time where increased negative xylem

pressures would be more likely to lead to embolism, the case for scalariform

perforation plates appearing in conditions where Pyrus cordata is simultaneously under

severe freeze/thaw embolism risk and low photosynthesis seems to be stronger.

Ultimately, a larger, more directed sampling would be required, comparing different

sites under similar conditions to verify if scalariform perforation plates were not peculiar

to the population at Corno de Bico, and to confirm that the individuals sampled in

Corno de Bico where not a fluke. In situ measurements of transpiration rates might also

help comprehend the role that evapotranspiration plays in the plant’s physiology.

Suitability of wood anatomical characters for differentiation between Crataegus

monogyna, Pyrus cordata and Pyrus bourgaeana.

One of the ultimate objectives of this work was to establish if it was possible to

distinguish between the woods of Crataegus monogyna, Pyrus cordata and Pyrus

bourgaeana using wood anatomical characters. Despite using several characters

frequently used in wood identification we were unable to ascertain any consistent and

significant difference between the three species. No categorical character was

consistently present (or absent) in these species, and no scalar character presented a

variance that enabled us to definitely ascribe a certain value range to a particular

species. Our conclusion is thus that, at least while using the characters analyzed in this

work, which included all of the major characters generally used in anatomical wood

identification (Schweingruber 1990), it is not possible to reliably tell apart C. monogyna,

P. cordata, and P. bourgaeana.

However, in comparing our results with the available literature, some discrepancies

appeared (besides the aforementioned presence of scalariform perforation plates).

Specifically, comparing our obtained values we noted that for pore size, Ray height and


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Ray width the differences were as presented in table 06. Note that although the Pyrus

species presented by each author varies, they are included here since they are

sometimes used to ascribe a wood sample to a genus, since the provided keys do not

refer to this or other supraspecifical taxons.

Table 06 – Comparison of values for selected characters in literature and in the present study *Diameter calculated

indirectly from maximum pore area. Reported values are those of the median of the maximum, while values inside

parenthesis are those of the maximum value measured (ignoring outliers).

Crataegus

monogyna

Pyrus sp.

Vernet et al. (2001) Pore diameter 15-40 m

Ray height up to 35 cells

Ray width 1-4 cells

Pore diameter 10-40 m

Ray height 20 (25)

Ray width 1-2(3)

Akkemik & Yaman

(2012)

Pore diameter <50 m


Ray width 1-4 cells

Pyrus serikensis Pyrus syriaca

Pore diameter <50 m


Ray width 1-3 cells

<50 m

Up to 46 cells

1-3 cells

Present study* Pore diameter 45(64) m

Ray height up to 26(51)

cells

Ray width 1-2(6) cells

Pyrus cordata Pyrus bourgaeana

Pore diameter 42 (60) m

Ray height up to 22 (48)

Ray width 1-2 (3) cells

45 (59)m

30 (46) cells

1-2 (4) cells

The differences observed in these characters are likely the result of different sampling

environments and populations. However we would be remiss not to point out that

extreme caution must be taken when using such precise characters as an identification

aide when dealing with species in far-ranging areas. One would note that we did not


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mention the work by Schweingruber (1990) in the above table. In fact, our observations

did not diverge from the treatment in this atlas. This is chiefly for two reasons: Firstly,

the Maloideae are only considered in their whole, rather than attempting a species-

level identification. Secondly, non-categorical characters are only used sparingly (e.g.

vessel size is only described as three comparative classes.) which though having the

effect of reducing resolving capability, also makes this guide more tolerant of variability.

We thus recommend that when identifying a botanical specimen solely on the features

of its wood anatomy, extreme care be given when accounting for wood character

variability. In particular, we must emphasize that when attempting to identify Maloideae

wood, identification down to the species level is not possible where P. cordata, P.

bourgaeana or C. monogyna are concerned, unless some other element is available to

aid identification. Furthermore, although the number of samples was much lower than

for the other species, the sampled Pyrus communis also did not show significant

differences in its wood anatomy, despite being sampled in much the same sites as the

wild species. Although grafting between P. communis and P. bourgaeana was

apparently commonplace in Alentejo, the similarity was preserved even in those

individuals that showed no evidences of grafting. We therefore feel that attempting to

resolve P. communis wood from that of the remaining species is also ill-advised.

Ultimately, in conditions where wood identification is necessary on the basis solely on

wood anatomy (e.g. in archaeobotanical studies) we recommend that any identification

resulting in any of these species be reduced to Crataegus/Pyrus.

The remaining objective, identifying environmental variables that had the most

influence on these species, was partly successful. As would be expected, P. cordata

and P. bourgaeana showed dissimilarity on several environmental characters

associated with their distribution area. Continental Portugal is progressively warmer

and drier as one advances towards the south, and there is also an aridity gradient

towards its interior that is particularly marked in the northern region (Agência

Portuguesa do Ambiente 2011), where a majority of the samples were taken. P.

cordata thus occurred mostly in the moister and cooler regions of Portugal, with P.

bourgaeana showing the opposite trend. The other dissimilarities registered were

strongly correlated with the geographical coordinates, and it is probable that they

correlated simply due to their geographical distribution in Portugal, and not because

they are constraining the species distribution. C. monogyna also showed dissimilarity

with P. cordata, again in temperature and rainfall. Since C. monogyna has a wider

distribution than either species of Pyrus (Flora-on: Flora de Portugal Interactiva 2014),


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and it was sampled in places where neither Pyrus appeared, but where temperature

and humidity were closer to that of the preferential environment of P. bourgaeana, this

dissimilarity is also within expectations.

Environmental influence on individual anatomical characters was likewise observed. As

discussed above, vessel size showed a correlation with rainfall, albeit the opposite of

that which was expected (the potential reasons for which have already been discussed),

but increasing levels of total rainfall also resulted in shorter rays, and an increase in

heterogeneous type II Rays. Plants that were in full shade had a greater occurrence of

vessels in radial multiples, smaller minimum and maximum vessel size and shorter

rays. Taller rays conversely, appeared when evapotranspiration was high, as again did

heterogeneous type II rays. Homogeneous rays were more frequent in areas with more

hours of sunlight. There was also a slight trend towards shorter rays in sites with higher

mean temperatures.

In short, it seems that ray size, one of the most plastic anatomical characters, is also

the most responsive to environmental pressures, responding to shade,

evapotranspiration and rainfall. These factors have an impact on the water potential

(either directly or indirectly via effects on photosynthetic activity), and the plasticity of

the xylem rays might reflect their adaption to different flow rates. Through more

controlled and aimed studies, it might yet be possible to elucidate these relationships,

at least within well-defined groups.

Closing remarks

This study analysed the wood of a large number of specimens from three specific

species of the Maloideae, taken from the widest practically possible sampling in

continental Portugal. Their anatomy was studied systematically, with the main

characters used in wood identification being noted and statistical analysis being

performed both to search for significant differences between the species and to search

for those characters that correlated strongly with the environment of the sampling sites.

The amount and extent of our studies, and their results, has left us confident in our

assertion that the wood from Pyrus cordata, Pyrus bourgaeana and Crataegus

monogyna is anatomically indistinguishable from each other. It is likely that other

species in the same genera are also not distinguishable on this basis (e.g. Pyrus

communis). In studies where it is necessary to rely solely on wood anatomy, it is highly


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recommended that no identification going down to the species-level is attempted. If

interpretation to a fine level is necessary, we recommend using the identification

Crataegus/Pyrus. Given the economical relevance of these species, we understand

that authors would be anxious to achieve a more clear-cut identification, however, we

cannot in good faith support a greater precision on the basis of wood anatomy alone.

Although the scope of our work did not allow for a comprehensive study of the

Maloideae, it did however reinforce a fact admitted by most wood anatomists: That the

wood of the Maloideae is a difficult subject of study, and that exercise of caution is

advised in their identification. A more comprehensive study of the Maloideae might yet

result in the consensus that any identification below subfamiliar level is under most

circunstances not possible.

It is also possible that less commonly used characters, such as vessel pit morphology

and arrangement or cell wall thickness, might yet be used to allow the differentiation of

problematic species. However, since these characters tend to be smaller and more

easily damaged, their potential usefulness is limited to studies where wood is well-

preserved.

The correlation between environment and anatomical characters proved interesting,

and revealed some relationships that merit further research. Environmental data is

most frequently derived from characters in the transverse section, and this study would

seem to support that variations of ray anatomy in Crataegus/Pyrus can be ascribed to

environmental factors. More detailed studies, with greater control for environmental

variables are necessary before anything definite can be said, and the scope should be

enlarged to other angiosperms to verify if this trend is observed more widely.


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Annex 01 List of sampling points and their coordinates

Site Taxa Latitude (º,','' N)* Longitude (º,','' W)* Xylarium code

P 2 Pyrus cordata 41°58'35.28'' 08°13'43.97'' PO-XI00002



MI1 5.3 Pyrus cordata 41°53'28.93'' 08°30'10.89'' PO-XI00021








VR1 1.5 Pyrus cordata 41°30'45.21'' 08°04'10.02'' PO-XI00038


VR1 2.5 Crataegus monogyna 41°29'49.49" 07°47'23.76" PO-XI00047



MI2 1.12 Pyrus communis 41°44'47.10'' 08°09'15.57'' PO-XI00072






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MI2 2.1 Crataegus monogyna 41°43'38.95" 08°11'05.70" PO-XI00080








MI2 6.2 Pyrus bourgaeana 41°52'01.22'' 08°24'08.04'' PO-XI00098

VR2 2.1 Pyrus sp. 41°11'29.72'' 07°58'44.86'' PO-XI00103

VR2 2.4 Pyrus sp. 41°11'29.30'' 07°48'44.71'' PO-XI00106

VR2 2.5 Pyrus sp. 41°11'29.30'' 07°48'44.71'' PO-XI00107

VR2 6.1 Pyrus sp. 41°11'03.69'' 07°50'08.08'' PO-XI00117

VR2 8.5 Pyrus sp. 40°59'25.94'' 07°55'56.38'' PO-XI00125

AV1 3.1 Crataegus monogyna 40°34’29.79'' 08°30’37.86'' PO-XI00137







MI3 4.1 Crataegus monogyna 41°48'21.79'' 08°51'22.24'' PO-XI00171





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JT 1.7 Crataegus monogyna 39°27'46.59'' 08°58'27.71'' PO-XI00215

AJ1 1.1 Crataegus monogyna 39°03'04.05'' 08°51'05.45'' PO-XI00228

AJ1 1.4 Crataegus monogyna 39°02’56.34'' 08°51’03.63'' PO-XI00231

AJ1 1.7 Crataegus monogyna 39°02’48.53'' 08°50’57.69'' PO-XI00234

AJ1 1.8 Crataegus monogyna 39°02'48.53'' 08°50'57.69'' PO-XI00235

AJ1 3.2 Crataegus monogyna 38°40'15.82" 08°22'22.27" PO-XI00238


AJ1 5.1 Pyrus bourgaeana 38°37'00.70" 08°13'10.50" PO-XI00242




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AJ2 1.2 Pyrus bourgaeana 38º11'45.09'' 07°30’03.07'' PO-XI00254






AJ3 2.3 Craetaegus monogyna 38°17'00.84" 07°43'06.93" PO-XI00263



AJ3 3.1 Pyrus bourgaeana 38°17'56.65'' 07°43'11.10'' PO-XI00266

AJ3 3.2 Pyrus communis 38°17'56.65'' 07°43'11.10'' PO-XI00267

AJ3 3.3 Pyrus communis 38°17'56.65'' 07°43'11.10'' PO-XI00268

AJ3 4.1 Pyrus sp. 38°16'17.10" 07°42'05.46" PO-XI00269

AJ3 4.2 Crataegus monogyna 38°16'17.10'' 07°42'05.46" PO-XI00270

AJ3 4.3 Pyrus sp. 38°16'17.10" 07°42'05.46" PO-XI00271

AJ3 4.4 Crataegus monogyna 38°16'17.10'' 07°42'05.46" PO-XI00272

AJ3 5.1 Pyrus communis 37°31'05.88" 08°36'35.39" PO-XI00273

AJ3 5.2 Pyrus communis 37°31'05.88" 08°36'35.39" PO-XI00274


AJ3 6.2 Pyrus bourgaeana 37°30'51.54'' 08°35'25.44" PO-XI00279


AJ4 2.1 Pyrus bourgaeana† 38°08'07.67" 08°34'54.50" PO-XI00282



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AJ4 3.1 Pyrus cordata 38°27'06.91" 08°30'39.27'' PO-XI00285







TM1 2.2 Crataegus monogyna 40°52'17.18" 06°58'36.92" PO-XI00296

TM1 2.3 Crataegus monogyna 40°52'17.18" 06°58'36.92" PO-XI00297

TM1 3.4 Crataegus monogyna 40º52’48.54" 06°50’11.32'' PO-XI00301

TM1 3.5 Crataegus monogyna 40º52’48.54" 06°50’11.32'' PO-XI00302

TM1 3.6 Crataegus monogyna 40°52'44.71" 06°50’14.46" PO-XI00303

TM2 3.1 Pyrus sp. 41°24'46.79" 06°50'06.93" PO-XI00314




TM3 1.5 Pyrus sp. 41°24'00.76" 06°20'57.19" PO-XI00332

TM3 1.7 Pyrus sp. 41°24'00.76" 06°20'57.19" PO-XI00334

TM3 2.2 Crataegus monogyna 41°48’47.37" 06°50’06.41" PO-XI00339

TM3 2.4 Crataegus monogyna 41°48’47.37" 06°50’06.41" PO-XI00341

*Coordinates expressed in WGS84 Geographic coordinate system.

†These specimens showed evidence of grafting, most likely with Pyrus communis


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Annex 02 Staining protocol for histological wood sections

Reagents:

- Safranine, general purpose grade (Fisher scientific)

- Alcian Blue 8GX, certified. (Acros organics)

- Javel water

- Ethanol

-Xylene

- Canada balsam (Sigma-Aldrich)

- Desionized water

Stock solutions:

1. Dilute ethanol in water to obtain 96%, 70% & 50% grades.

2. Safranine stock solution is obtained by dissolving 0.1g in 10ml 70% ethanol. Staining

grade Safranine is obtained by diluting this stock solution 10X.

3. Alcian blue staining solution is obtained by dissolving 0.1g in 10 ml water. This

solution can be used directly for staining.

Procedure:

1. Soak each wood slice in Javel water for 3 minutes

2. Transfer to Alcian blue staining solution and leave to stain for 3 minutes.

3. Transfer the slices to an initial dehydration series of alcohol: 50, then 70%, letting it

bathe 1 minute in each.

4. Transfer to Safranine staining solution and leave to stain for 3 minutes.

5. Remove the slice and immerse it in 96% and 100% (absolute) alcohol, 1 minute

each. Observe under a stereomicroscope until the level of safranine in the tissue has

dropped to an acceptable level. If necessary, extend the time spent in alcohol.

6. After dehydration, transfer the slice to xylene for 1 minute. If the Xylene turns cloudy,

then return the slice to absolute alcohol for additional dehydration, and replace the

xylene.


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7. If the xylene remains clear, the stained section can be then transferred to a glass

slide and if necessary, carefully uncurled. Mount with a drop of Canada balsam. Leave

the slides in a heated environment to eliminate air bubbles.

FCUP

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Annex 03 Collected environmental data

Sites whose soil classes are marked with an * are in anthropized environments.

Site Species Plant height (m) Soil abundance Terrain slope Shade Light direction Altitude Soil class Nr. days frost Nr. days rainfall Lithology Evapotranspiration (mm) Solar radiation (Kcal/Cm2) Mean Temperature (ºC) Hours Sunshine Rainfall (mm)

P 2 Pyrus cordata n/d Residual High Full sun S 955 Humic cambisol 20-30 >100 Granites 700-800 <140 10-12.5 1800-1900 2400-2800

P 3 Pyrus cordata n/d Abundant Negligible Full sun n/a 961 Rankers 20-30 >100 Granites 700-800 <140 10-12.5 <1800 2400-2800

P 4 Pyrus cordata 5 Sparse High Partial shade E 813 Humic cambisol 20-30 >100 Granites 700-800 <140 10-12.5 <1800 2400-2800

MI1 5.3 Pyrus cordata 2.5 Abundant Moderate Full shade n/a 627 Humic cambisol 20-30 >100 Granites >800 <140 10-12.5 2100-2200 2000-2400

MI1 5.4 Pyrus cordata 2.5 Abundant Moderate Full shade n/a 645 Humic cambisol 20-30 >100 Granites >800 <140 10-12.5 2100-2200 2000-2400

MI1 5.6 Pyrus cordata 2 Abundant Negligible Full shade n/a 637 Humic cambisol 20-30 >100 Granites >800 <140 10-12.5 2100-2200 2000-2400




MI1 5.13 Pyrus cordata 2 Sparse Negligible Full shade N 628 Humic cambisol 20-30 >100 Granites >800 <140 10-12.5 2100-2200 2000-2400

MI1 5.14 Pyrus cordata 4 Sparse Moderate Full sun n/a 614 Humic cambisol 20-30 >100 Granites >800 <140 10-12.5 2100-2200 2000-2400

VR1 1.5 Pyrus cordata 2 Sparse High Full shade n/a 523 Humic cambisol 40-50 75-100 Granites 600-700 <140 10-12.5 2300-2400 1200-1400

VR1 1.9 Pyrus cordata 6 Abundant Negligible Full shade n/a 535 Humic cambisol 40-50 75-100 Granites 600-700 <140 10-12.5 2300-2400 1200-1400

VR1 2.5 Crataegus monogyna 2 Residual High Full shade S 651 Schist associated cambisol 30-40 75-100 Schist and Greywacke 600-700 140-145 10-12.5 2300-2400 1400-1600

VR1 3.4 Pyrus cordata 1 Sparse Negligible Full sun n/a 902 Schist associated cambisol >80 75-100 Schist and Greywacke 600-700 140-145 10-12.5 2300-2400 1600-2000

VR1 4.3 Pyrus cordata 4 Residual High Full shade N 835 Rankers 60-70 75-100 Granites 600-700 140-145 7.5-10 2300-2400 1400-1600

MI2 1.12 Pyrus communis 4 Abundant Negligible Partial shade W 605 Rankers 10-20 >100 Granites 600-700 <140 10-12.5 1800-1900 >2800

MI2 1.13 Pyrus communis 3 Abundant Negligible Partial shade W 605 Rankers 10-20 >100 Granites 600-700 <140 10-12.5 1800-1900 >2800

MI2 1.14 Pyrus cordata 3.5 Abundant Negligible Partial shade W 605 Rankers 10-20 >100 Granites 600-700 <140 10-12.5 1800-1900 >2800

MI2 1.15 Pyrus cordata 3.5 Abundant Negligible Partial shade W 605 Rankers 10-20 >100 Granites 600-700 <140 10-12.5 1800-1900 >2800

MI2 1.17 Pyrus cordata 2 Abundant Moderate Partial shade n/a 636 Rankers 10-20 >100 Granites 600-700 <140 10-12.5 1800-1900 >2800

MI2 1.19 Pyrus communis 4 Abundant Moderate Full sun W 597 Rankers 10-20 >100 Granites 600-700 <140 10-12.5 1800-1900 >2800

MI2 2.1 Crataegus monogyna 2.5 Abundant Negligible Full sun n/a 817 Humic cambisol 10-20 >100 Granites 700-800 <140 10-12.5 1800-1900 2400-2800



MI2 2.4 Crataegus monogyna 2 Abundant Negligible Full sun n/a 817 Humic cambisol 10-20 >100 Granites 700-800 <140 10-12.5 1800-1900 2400-2800

MI2 2.5 Pyrus cordata 2 Abundant Negligible Full sun n/a 817 Humic cambisol 10-20 >100 Granites 700-800 <140 10-12.5 1800-1900 2400-2800



MI2 2.8 Pyrus cordata 2 Abundant Negligible Full sun n/a 817 Humic cambisol 10-20 >100 Granites 700-800 <140 10-12.5 1800-1900 2400-2800

MI2 6.2 Pyrus bourgaeana 1 Sparse High Partial shade N 75 Humic cambisol 30-40 >100 Granites >800 <140 10-12.5 2200-2300 2000-2400

VR2 2.1 Pyrus sp. 1 Abundant Moderate Full shade n/a 664 Humic cambisol 40-50 75-100 Granites 600-700 140-145 10-12.5 2300-2400 1400-1600

VR2 2.4 Pyrus sp. 2 Abundant Moderate Full shade n/a 664 Eutric lithosol (luvisol associated) 30-40 75-100 Schist and Greywacke 500-600 140-145 15-16 2500-2600 800-1000

VR2 2.5 Pyrus sp. 2 Abundant Moderate Full shade n/a 664 Eutric lithosol (luvisol associated) 30-40 75-100 Schist and Greywacke 500-600 140-145 10-12.5 2500-2600 800-1000

VR2 6.1 Pyrus sp. n/d Abundant Negligible Full shade E 416 Eutric lithosol (luvisol associated) 30-40 75-100 Schist and Greywacke 500-600 140-145 15-16 2400-2500 800-1000

VR2 8.5 Pyrus sp. 1 Sparse High Full sun n/a 1107 Rankers 40-50 >100 Granites 600-700 145-150 7.5-10 2200-2300 1600-2000

AV1 3.1 Crataegus monogyna 0.5 Sparse Negligible Full sun S 953 Eutric fluvisol 5-10 75-100 Alluvium 600-700 140-145 12.5-15 2500-2600 800-1000



AV1 3.6 Crataegus monogyna 1.5 Abundant Negligible Full sun n/a 953 Eutric fluvisol 5-10 75-100 Alluvium 600-700 140-145 12.5-15 2500-2600 800-1000

AV1 5.1 Crataegus monogyna 1 Sparse Negligible Full shade n/a 150 Schist associated cambisol 1-5 75-100 Schist and Greywacke 600-700 140-145 12.5-15 2300-2400 1400-1600

AV1 5.2 Crataegus monogyna 0.5 Sparse Moderate Full shade n/a 150 Schist associated cambisol 1-5 75-100 Schist and Greywacke 600-700 140-145 12.5-15 2300-2400 1400-1600

AV1 7.2 Crataegus monogyna 4 Sparse Negligible Partial shade N 641 Luvisol associated cambisol 10-20 >100 Schist and Greywacke 600-700 145-150 7.5-10 2200-2300 2000-2400

MI3 4.1 Crataegus monogyna 3.5 Abundant Moderate Partial shade W 19 Distric regosol 1-5 >100 Aeolian sands* 700-800 <140 12.5-15 2400-2500 1200-1400

MI3 4.2 Crataegus monogyna 3.5 Abundant Moderate Partial shade W 19 Distric regosol 1-5 >100 Aeolian sands* 700-800 <140 12.5-15 2400-2500 1200-1400

MI3 4.3 Crataegus monogyna 3 Abundant High Partial shade W 19 Distric regosol 1-5 >100 Aeolian sands* 700-800 <140 12.5-15 2400-2500 1200-1400



MI3 6.1 Crataegus monogyna 1.5 Residual Negligible Full sun N 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.2 Crataegus monogyna 1.5 Sparse Negligible Full sun N 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.3 Crataegus monogyna 1 Residual Negligible Full sun N 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.4 Crataegus monogyna 1 Residual Negligible Full sun N 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.7 Crataegus monogyna 1.5 Sparse Negligible Full sun N 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.8 Crataegus monogyna 1 Sparse Negligible Full sun N 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.9 Crataegus monogyna 1.5 Residual Negligible Full sun W 125 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.10 Crataegus monogyna 1 Residual Negligible Full sun W 129 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.11 Crataegus monogyna 1 Residual Negligible Full sun S 129 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.12 Crataegus monogyna 1.5 Residual Negligible Full sun S 129 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.13 Crataegus monogyna 2 Residual Moderate Full sun n/a 131 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.14 Crataegus monogyna 2 Residual Moderate Full sun n/a 131 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200


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Site Species Plant height (m) Soil abundance Terrain slope Shade Light direction Altitude Soil class Nr. days frost Nr. days rainfall Lithology Evapotranspiration (mm) Solar radiation (Kcal/Cm2) Mean Temperature (ºC) Hours Sunshine Rainfall (mm)

MI3 6.15 Crataegus monogyna 1 Residual High Full sun n/a 131 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.16 Crataegus monogyna 0.5 Sparse High Full sun W 131 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

MI3 6.17 Crataegus monogyna 1 Sparse High Full sun W 131 Humic cambisol < 1 >100 Granites 700-800 <140 12.5-15 2400-2500 1000-1200

JT 1.7 Crataegus monogyna 1.5 Abundant Negligible Full sun n/a 169 Calcic luvisol 20-30 75-100 Conglomerate,Sandstone,Limestone,Marlstone 600-700 140-145 15-16 2300-2400 800-1000

AJ1 1.1 Crataegus monogyna 2 Abundant Negligible Full sun n/a 5 Gleyic solonchaks 20-30 50-75 Alluvium 500-600 145-150 15-16 2800-2900 500-600

AJ1 1.4 Crataegus monogyna 2 Abundant Negligible Full sun n/a 5 Gleyic solonchaks 20-30 50-75 Alluvium 500-600 145-150 15-16 2800-2900 500-600

AJ1 1.7 Crataegus monogyna 2.5 Abundant Negligible Full sun N 5 Gleyic solonchaks 20-30 50-75 Alluvium 500-600 145-150 15-16 2800-2900 500-600

AJ1 1.8 Crataegus monogyna 2.5 Abundant Moderate Full sun N 5 Gleyic solonchaks 20-30 50-75 Alluvium 500-600 145-150 15-16 2800-2900 500-600

AJ1 3.2 Crataegus monogyna 1.5 Abundant Moderate Partial shade S 120 Orthic luvisol 40-50 50-75 Xistos argilosos, Grauvaques, Arenitos 450-500 145-150 15-16 2800-2900 600-700

AJ1 3.4 Crataegus monogyna 3 Abundant Moderate Full sun S 120 Orthic luvisol 40-50 50-75 Xistos argilosos, Grauvaques, Arenitos 450-500 145-150 15-16 2800-2900 600-700

AJ1 5.1 Pyrus bourgaeana 3 Abundant Negligible Partial shade S 250 Orthic luvisol 30-40 50-75 Schist and Greywacke 450-500 145-150 15-16 2800-2900 600-700

AJ1 5.2 Pyrus bourgaeana 3 Abundant Negligible Full shade n/a 250 Orthic luvisol 30-40 50-75 Schist and Greywacke 450-500 145-150 15-16 2800-2900 600-700

AJ1 5.4 Pyrus bourgaeana 3 Abundant Negligible Full shade n/a 250 Orthic luvisol 30-40 50-75 Schist and Greywacke 450-500 145-150 15-16 2800-2900 600-700

AJ1 6.2 Crataegus monogyna 2.5 Sparse High Full sun E 200 Orthic luvisol 40-50 50-75 Schist and Greywacke 450-500 145-150 15-16 2800-2900 600-700

AJ1 6.3 Crataegus monogyna 2.5 Sparse High Full sun n/a 200 Orthic luvisol 40-50 50-75 Schist and Greywacke 450-500 145-150 15-16 2800-2900 600-700

AJ1 6.4 Pyrus bourgaeana 2 Abundant High Full sun E 200 Orthic luvisol 40-50 50-75 Schist and Greywacke 450-500 145-150 15-16 2800-2900 600-700

AJ2 1.2 Pyrus bourgaeana 2 Sparse Moderate Partial shade N 168 Ferric luvisol 10-20 50-75 Metavulcanite 400-450 160-165 16-17.5 3000-3100 500-600

AJ2 2.2 Pyrus bourgaeana 3 Abundant Negligible Full sun n/a 200 Albic Luvisol 20-30 50-75 Sands, pebbles, clays, weakly consolidated sandstone* <400 160-165 >17.5 3000-3100 400-500




AJ3 1.1 Pyrus bourgaeana 7 Abundant Negligible Full sun n/a 327 Chromic luvisol 20-30 50-75 Metavulcanite* 450-500 150-155 16-17.5 3000-3100 600-700

AJ3 2.3 Crataegus monogyna 1.5 Abundant Negligible Partial shade n/a 321 Chromic luvisol 20-30 50-75 Rochas carbonatadas 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 2.4 Crataegus monogyna 1.5 Abundant Negligible Partial shade n/a 321 Chromic luvisol 20-30 50-75 Rochas carbonatadas 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 2.5 Pyrus bourgaeana 1 Abundant Negligible Full sun E 321 Chromic luvisol 20-30 50-75 Rochas carbonatadas 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 3.1 Pyrus bourgaeana 3 Abundant Negligible Full sun n/a 325 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 150-155 16-17.5 3000-3100 600-700

AJ3 3.2 Pyrus communis 3 Abundant Negligible Full sun n/a 325 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 150-155 16-17.5 3000-3100 600-700

AJ3 3.3 Pyrus communis 4.5 Abundant Negligible Full sun n/a 325 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 150-155 16-17.5 3000-3100 600-700

AJ3 4.1 Pyrus sp. 1.5 Abundant Moderate Partial shade N 323 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 4.2 Crataegus monogyna 1.5 Abundant Moderate Partial shade N 323 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 4.3 Pyrus sp. 3 Abundant Moderate Partial shade N 323 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 4.4 Crataegus monogyna 1.5 Abundant Moderate Partial shade N 323 Chromic luvisol 20-30 50-75 Metavulcanite 450-500 155-160 16-17.5 3000-3100 600-700

AJ3 5.1 Pyrus communis 2 Sparse Negligible Full sun N 150 Eutric lithosol 1-5 75-100 Xistos argilosos, Grauvaques, Arenitos 450-500 155-160 15-16 2800-2900 700-800

AJ3 5.2 Pyrus communis 1.5 Sparse Negligible Full sun N 150 Eutric lithosol 1-5 75-100 Xistos argilosos, Grauvaques, Arenitos 450-500 155-160 15-16 2800-2900 700-800

AJ3 6.1 Pyrus bourgaeana 6 Abundant Moderate Full shade n/a 75 Eutric lithosol 1-5 75-100 Xistos argilosos, Grauvaques, Arenitos 450-500 155-160 15-16 2800-2900 700-800

AJ3 6.2 Pyrus bourgaeana 1.5 Abundant Moderate Full shade n/a 75 Eutric lithosol 1-5 75-100 Xistos argilosos, Grauvaques, Arenitos 450-500 155-160 15-16 2800-2900 700-800

AJ4 1.1 Crataegus monogyna 1.5 Sparse High Full sun N 75 Plinthic luvisol 5-10 50-75 Arenitos, calcários, areias, cascalheiras, argilas 450-500 155-160 15-16 2900-3000 600-700

AJ4 2.1 Pyrus bourgaeana † 2 Abundant Negligible Full sun n/a 172 Eutric lithosol 5-10 50-75 Xistos argilosos, Grauvaques, Arenitos 500-600 150-155 16-17.5 2900-3000 700-800



AJ4 3.1 Pyrus cordata 2.5 Sparse Moderate Full sun n/a 25 Albic Luvisol 30-40 50-75 Arenitos, calcários, areias, cascalheiras, argilas 450-500 150-155 16-17.5 2800-2900 600-700

AJ4 4.1 Pyrus bourgaeana 2 Sparse Moderate Full sun n/a 78 Orthic podzol 30-40 50-75 Arenitos, calcários, areias, cascalheiras, argilas 500-600 150-155 15-16 2800-2900 600-700

AJ4 4.2 Pyrus bourgaeana 1.5 Sparse Negligible Full sun n/a 78 Orthic podzol 30-40 50-75 Arenitos, calcários, areias, cascalheiras, argilas 500-600 150-155 15-16 2800-2900 600-700

AJ4 5.1 Pyrus bourgaeana 1.5 Residual Moderate Full sun n/a 93 Orthic podzol 40-50 50-75 Arenitos, calcários, areias, cascalheiras, argilas 500-600 150-155 15-16 2800-2900 600-700

AJ4 6.1 Pyrus bourgaeana † 3 Sparse Moderate Full sun n/a 50 Orthic podzol 40-50 50-75 Arenitos, calcários, areias, cascalheiras, argilas 500-600 150-155 15-16 2800-2900 600-700



TM1 2.2 Crataegus monogyna 3 Abundant Negligible Full shade n/a 726 Schist and quartzite associated cambisol 60-70 75-100 Sands, pebbles, clays, weakly consolidated sandstone* <400 145-150 12.5-15 2800-2900 500-600

TM1 2.3 Crataegus monogyna 3 Abundant Negligible Full shade n/a 726 Schist and quartzite associated cambisol 60-70 75-100 Sands, pebbles, clays, weakly consolidated sandstone* <400 145-150 12.5-15 2800-2900 500-600

TM1 3.4 Crataegus monogyna 2 Abundant Negligible Full sun n/a 590 Orthic luvisol 50-60 75-100 Schist and Greywacke 400-450 150-155 12.5-15 2800-2900 500-600



TM2 3.1 Pyrus sp. 5 Sparse High Full sun n/a 455 Chromic luvisol 30-40 50-75 Xistos, anfibolitos, micaxistos, grauvaques, quartzitos, gnaisses 450-500 145-150 15-16 2700-2800 600-700

TM2 5.1 Crataegus monogyna 3 Abundant Negligible Partial shade N 554 Eutric lithosol (luvisol associated) 40-50 50-75 Xistos, anfibolitos, micaxistos, grauvaques, quartzitos, gnaisses 450-500 145-150 12.5-15 2600-2700 800-1000



TM3 1.5 Pyrus sp. n/d Abundant Negligible Full sun n/a 652 Dystric cambisol 10-20 50-75 Schist and Greywacke 400-450 150-155 12.5-15 2500-2600 600-700

TM3 1.7 Pyrus sp. n/d Abundant Negligible Full sun n/a 652 Dystric cambisol 10-20 50-75 Schist and Greywacke 400-450 150-155 12.5-15 2500-2600 600-700

TM3 2.2 Crataegus monogyna 2 Abundant Negligible Full sun W 848 Chromic cambisol 70-80 75-100 Xistos, anfibolitos, micaxistos, grauvaques, quartzitos, gnaisses 500-600 145-150 7.5-10 2600-2700 1000-1200

TM3 2.4 Crataegus monogyna 3 Abundant Negligible Full sun W 848 Chromic cambisol 70-80 75-100 Xistos, anfibolitos, micaxistos, grauvaques, quartzitos, gnaisses 500-600 145-150 7.5-10 2600-2700 1000-1200


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Annex 04 Correlation matrix (Spearman’s D) for studied variables.

p values for Spearman’s rank correlation for studied variables. p values under 0.05 are marked out in grey. p values under 0.01 are marked out in black.

No. of vessels Vessel grouping Max. No. Vessels/cluster Radial Cluster Tangential Cluster Random Cluster No. of clusters Total vessel area Area, smallest vessel Area, largest vessel Average vessel area Vessel aspect ratio Height of tallest ray (m) Height of shortest ray (m) Average ray height

Number of vessels xx 0.88781 0.019918 0.5113 0.72878 0.49223 3.20E-04 6.62E-07 0.00022453 0.031931 0.005994 0.25651 0.36687 0.12969 0.45126

Vessel grouping 0.88781 xx 4.54E-15 0.0079083 0.0079083 4.03E-06 3.08E-14 0.00076351 0.0097361 0.12468 0.0017469 0.052085 0.82855 0.012177 0.53301

Maximum number of vessels per cluster 0.019918 4.54E-15 xx 0.094914 0.080795 0.00028031 4.64E-21 1.01E-05 0.05542 0.038085 0.003495 0.080547 0.42113 7.74E-05 0.051328

Radial Cluster orientation 0.5113 0.0079083 0.094914 xx 0.0096928 7.14E-06 0.37111 0.055258 0.18456 0.63352 0.21855 4.69E-04 0.5506 0.19115 0.79223

Tangential Cluster orientation 0.72878 0.0079083 0.080795 0.0096928 xx 7.14E-06 0.10712 0.090998 0.20521 0.16852 0.035379 0.9867 0.10299 0.21727 0.10875

No Cluster orientation 0.49223 4.03E-06 0.00028031 7.14E-06 7.14E-06 xx 3.16E-05 0.0035943 0.031929 0.81479 0.065446 0.21873 0.51597 0.99575 0.32463

Total number of clusters 0.00031979 3.08E-14 4.64E-21 0.37111 0.10712 3.16E-05 xx 2.40E-10 0.021558 0.10088 0.00028082 0.41226 0.24698 0.020937 0.22238

Total vessel area (m2) 6.62E-07 0.00076351 1.01E-05 0.055258 0.090998 0.0035943 2.40E-10 xx 0.00024871 3.22E-05 1.37E-11 0.019309 0.94934 0.29265 0.82102

Area of smallest vessel (m2) 0.00022453 0.0097361 0.05542 0.18456 0.20521 0.031929 0.021558 0.00024871 xx 1.92E-05 2.83E-14 0.060383 0.87113 0.69667 0.82836

area of largest vessel (m2) 0.031931 0.12468 0.038085 0.63352 0.16852 0.81479 0.10088 3.22E-05 1.92E-05 xx 7.97E-13 0.33576 0.42101 0.39952 0.90445

Average vessel area (m2) 0.005994 0.0017469 0.003495 0.21855 0.035379 0.065446 0.00028082 1.37E-11 2.83E-14 7.97E-13 xx 0.2025 0.37574 0.71162 0.21825

Average vessel aspect ratio 0.25651 0.052085 0.080547 0.00046918 0.9867 0.21873 0.41226 0.019309 0.060383 0.33576 0.2025 xx 0.53036 0.71112 0.2861

Height of tallest ray (m) 0.36687 0.82855 0.42113 0.5506 0.10299 0.51597 0.24698 0.94934 0.87113 0.42101 0.37574 0.53036 xx 3.35E-02 1.70E-17

Height of shortest ray (m) 0.12969 0.012177 7.74E-05 0.19115 0.21727 0.99575 0.020937 0.29265 0.69667 0.39952 0.71162 0.71112 0.033456 xx 2.04E-08

Average ray height (m) 0.45126 0.53301 0.051328 0.79223 0.10875 0.32463 0.22238 0.82102 0.82836 0.90445 0.21825 0.2861 1.70E-17 2.04E-08 xx

No. of rays/mm2 0.66255 0.45409 0.96794 0.61204 0.24694 0.052783 0.20482 0.65892 0.090178 0.85608 0.47804 0.008629 0.063391 0.030802 0.00084547

Width of widest ray (cells) 0.0004472 0.45625 0.073427 0.4682 0.11016 0.92687 0.034487 0.0033442 0.7086 0.69156 0.36832 0.95803 0.097081 0.00957 0.0019019

Width of narrowest ray (cells) 0.13704 0.61708 0.11596 0.6263 0.6263 0.23672 0.12777 0.20919 0.93098 0.59325 0.85112 0.80612 0.25405 0.13697 0.14479

Height of tallest ray (cells) 0.86576 0.58109 0.92967 0.038819 0.069407 0.51727 0.34324 0.27232 0.059272 0.92111 0.0061377 0.81998 2.14E-14 0.80216 8.84E-07

Height of shortest ray (cells) 0.76857 0.70564 0.79572 0.8007 0.28372 0.33666 0.61698 0.7329 0.63091 0.19821 0.33795 0.72582 0.0029828 0.00017888 0.00020577

Presence of two distinct ray sizes 0.54714 0.22489 0.072319 0.25825 0.25825 0.4119 0.32268 0.73565 0.69994 0.010434 0.47942 0.44452 0.27312 3.14E-06 0.024503

Scalariform Vessels 0.31655 0.31113 0.00085009 3.18E-05 0.32394 0.086892 0.0033019 1 0.35701 0.1982 0.26018 0.0077821 0.018557 0.016469 0.26018

Homogeneous Rays 0.035797 0.22489 0.30579 0.094433 0.090809 0.33217 0.13838 0.91882 0.78659 0.83348 0.25143 0.23598 0.41475 0.049695 0.026603

Heterogeneous I Rays 0.89307 0.11342 0.0013057 0.15692 0.15692 0.32913 0.0086498 0.3454 0.37356 0.38518 0.85282 0.00028065 0.99785 0.28331 0.98498

Heterogeneous II Rays 0.035434 0.47292 0.0025324 0.00028151 0.98788 0.021162 0.077347 0.27063 0.39557 0.37474 0.31483 0.0007207 0.37842 0.00045456 0.015172

Presence of crystals 0.92119 0.096623 0.57784 0.61929 0.97545 0.08541 0.78168 0.60403 0.39957 0.052406 0.47361 0.69447 0.58733 0.00093136 0.014538

Presence of vessel thickenings 0.38832 0.31113 0.098369 0.11255 0.76402 0.49346 0.25185 0.15193 0.59866 0.0041591 0.023417 0.79239 0.12468 0.040622 0.015831

Plant height (m) 0.7046 0.43886 0.47522 0.90326 0.017942 0.17427 0.49444 0.093148 0.29675 0.32556 0.6661 0.6006 0.61728 0.59844 0.1096

Abundant soil 0.57841 0.060416 0.029989 0.43821 0.25721 0.21577 0.4925 0.24603 0.97621 0.68429 0.60649 0.39463 0.88361 0.54185 0.90505

Sparse soil 0.98549 0.29386 0.63732 0.14244 0.89392 0.055334 0.87179 0.20636 0.1824 0.44322 0.1359 0.65373 0.96615 0.18731 0.45224

Residual soil 0.60364 0.20256 0.014103 0.34351 0.34351 0.63218 0.28005 0.85086 0.12836 0.16705 0.28965 0.84762 0.83483 0.011919 0.28227

High slope 0.84844 0.79904 0.86982 0.77101 0.34832 0.43858 0.53259 0.88894 0.83121 0.070571 0.58898 0.86285 0.075844 0.97361 0.36784

Moderate slope 0.55343 0.82982 0.9692 0.45551 0.45551 0.30904 0.99005 0.60656 0.6218 0.010757 0.34393 0.76815 0.97772 0.48697 0.84987

Negligible slope 0.88245 0.77997 0.97331 0.34199 0.34199 0.19682 0.53619 0.61897 0.6671 0.47618 0.75716 0.95069 0.19591 0.48447 0.67685

Full sun 0.2214 0.579 0.7755 0.44982 0.44982 0.097546 0.79264 0.44996 0.29656 0.93404 0.7649 0.61944 0.095676 0.1562 0.70457

Partial Shade 0.90739 1 0.21625 0.30825 0.90569 0.47221 0.91983 0.01141 0.18525 0.083844 0.01913 0.68149 0.28475 0.79099 0.27328

Full Shade 0.091596 0.4189 0.44068 0.035081 0.72847 0.0089082 0.51795 0.13973 0.0042314 0.035396 0.0033435 0.51772 0.31075 0.058216 0.57536

N Direction 0.8676 0.13331 0.2145 0.72847 0.72847 0.50691 0.36398 0.15654 0.65267 0.11403 0.20994 0.99468 0.61696 0.83621 0.2951

W Direction 0.88032 0.14134 0.60274 0.52601 0.96387 0.46953 0.50716 0.30635 0.21387 0.17174 0.3808 0.17783 0.856 0.17905 0.25151

S Direction 0.67954 0.10006 0.069217 0.44369 0.94448 0.43637 0.079977 0.58183 0.37976 0.71112 0.16126 0.039767 0.72173 0.35938 0.80129

E Direction 0.21452 1 0.2228 0.96159 0.26803 0.39763 0.35905 0.64106 0.38604 0.12275 0.85925 0.61304 0.10758 0.90068 0.27271

Altitude 0.64206 0.013953 0.028935 0.42533 0.065585 0.24559 0.14634 0.016556 0.21513 0.19577 0.06888 0.41534 0.055534 0.77064 0.011167

x(dec º N) 0.004041 0.38096 0.02819 0.17472 0.4313 0.79827 0.15337 0.2541 0.67526 0.35123 0.58889 0.034521 0.47823 0.00090126 0.00057498

y (dec º W) 0.70547 0.035439 0.013801 0.81026 0.23127 0.57611 0.5261 0.59217 0.16213 0.097249 0.74233 0.90275 0.30756 0.17848 0.71073

Humic cambisol (Soil 1) 0.002667 0.43105 0.03774 0.96344 0.96344 0.57163 0.24754 0.40351 0.96827 0.94562 0.43951 0.81573 0.31988 0.038267 0.0020259

Rankers (Soil 2) 0.02836 0.49002 0.54892 0.26358 0.13039 0.38153 0.26002 0.092211 0.22796 0.71235 0.71233 0.19513 0.34887 0.51067 0.92062

Schist associated cambisol (Soil 3) 0.039241 0.31113 0.3958 0.11255 0.11255 0.086892 0.14584 0.0019392 0.010961 0.0049951 0.0031385 0.16927 0.27921 0.44698 0.46469

Eutric lithosol (luvisol associated) (Soil 4) 0.011934 0.69811 0.72318 0.73636 0.51651 0.94773 0.92381 0.01355 0.53429 0.53425 0.2286 0.53422 0.28988 0.11047 0.027754

Eutric fluvisol (Soil 5) 0.67155 0.31113 0.3958 0.32394 0.76402 0.17081 0.71342 0.84926 0.61398 0.76996 0.95336 0.17385 0.48274 0.36457 0.34934

Luvisol associated cambisol (Soil 6) 0.91951 0.61708 0.51973 0.6263 0.6263 0.23672 0.42502 0.2309 0.10286 0.46158 0.21975 0.78385 0.19881 0.10277 0.12239

Distric regosol (Soil 7) 0.68871 0.25541 0.84876 0.96159 0.96159 0.39763 0.75145 0.0029334 0.0087893 0.0098523 0.0041021 0.63161 0.055968 0.64567 0.99476

Calcic luvisol (Soil 8) 0.22527 0.61708 0.51973 0.6263 0.040012 0.39802 0.35326 0.41065 0.41063 0.34807 0.53472 0.085786 0.87382 0.51586 0.67545

Gleyic solonchaks (Soil 9) 0.38431 0.31113 0.75591 0.0040605 0.32394 0.49346 0.25185 0.4559 0.84353 0.42978 0.96501 0.60879 0.24206 0.98833 0.38028

Orthic luvisol (Soil 10) 0.056116 4.92E-06 0.0025798 0.091514 0.091514 0.31165 0.0032535 0.56359 0.048424 0.095099 0.028687 0.026757 0.070964 0.19173 0.00022538

Ferric luvisol (Soil 11) 0.78386 0.61708 0.67508 0.6263 0.6263 0.23672 0.67403 0.98848 0.55396 0.26632 0.76175 0.11556 0.31916 0.18888 0.12239

Albic Luvisol (Soil 12) 0.91631 0.25541 0.0052321 0.26803 0.26803 0.0071159 0.012625 0.32787 0.90071 0.26705 0.42677 0.45006 0.011885 0.041073 0.011885

Chromic luvisol (Soil 13) 0.061171 0.76187 0.39373 0.31693 0.0043937 0.21897 0.66348 0.24821 0.14403 0.17788 0.0019524 0.97208 0.98604 0.14635 0.18067

Eutric lithosol (Soil 14) 0.011934 0.69811 0.72318 0.73636 0.51651 0.94773 0.92381 0.01355 0.53429 0.53425 0.2286 0.53422 0.28988 0.11047 0.027754

Plinthic luvisol (Soil 15) 0.17478 0.61708 0.67508 0.040012 0.6263 0.39802 0.42502 0.29202 0.15718 0.89663 0.57339 0.98848 0.44416 0.3053 0.15292

Orthic podzol (Soil 16) 0.23554 0.21082 0.50403 0.87371 0.87371 0.20442 0.28425 0.53122 0.42326 0.34756 0.23788 0.35373 0.47734 0.25252 0.1975

Schist and quartzite associated cambisol (Soil 17) 0.78983 0.47764 0.55162 0.48922 0.26592 0.81029 1 1 0.83757 0.85361 0.95095 0.48572 0.91834 0.24248 0.56589

Dystric cambisol (Soil 18) 0.022255 0.47764 0.41303 0.26592 0.48922 0.81029 0.15824 0.48581 0.75846 0.43593 0.57985 0.69685 0.96729 0.26816 0.62265

Chromic cambisol (Soil 19) 0.045613 0.28681 0.80684 0.26592 0.48922 0.23003 0.94253 0.18271 0.98364 0.65196 0.69685 0.064985 0.02165 0.88586 0.019416

nº days frost 0.53654 0.027339 0.075777 0.013559 0.018577 0.059831 0.23271 0.11576 0.029877 0.061442 0.094814 0.40469 0.12191 0.67751 0.01978

nº days rainfall 0.5768 0.67334 0.11039 0.91471 0.67617 0.55518 0.86791 0.97612 0.20916 0.28825 0.69995 0.058458 0.99136 0.0057414 0.0006962

Granites (Litho 1) 0.095836 0.70844 0.099206 0.57404 0.38817 0.9496 0.62545 0.90547 0.53832 0.89264 0.35031 0.3559 0.66189 0.020105 0.0040074

Schist and Greywacke (Litho 2) 0.15045 0.00893 0.041423 0.82066 0.29899 0.14069 0.028875 0.027567 0.0032029 0.0057014 0.00048127 0.92267 0.43107 0.55301 0.11123

Alluvium (Litho 3) 0.72847 0.17461 0.25506 0.10223 0.73636 0.94773 0.2308 0.59482 0.50883 0.344 0.68269 0.18819 0.4564 0.91526 0.28989

Aeolian sands (Litho 4) 0.68871 0.25541 0.84876 0.96159 0.96159 0.39763 0.75145 0.0029334 0.0087893 0.0098523 0.0041021 0.63161 0.055968 0.64567 0.99476

Conglomerate,Sandstone,Limestone,Marlstone (Litho 5) 0.91951 0.61708 0.51973 0.6263 0.6263 0.23672 0.42502 0.2309 0.10286 0.46158 0.21975 0.78385 0.19881 0.10277 0.12239

Metavulcanite (Litho 6) 0.27894 0.61708 0.67508 0.6263 0.6263 0.23672 0.47726 0.34813 0.64414 0.73987 0.69668 0.76175 0.89662 0.82854 0.59321

Sands, pebbles, clays, weakly consolidated sandstone (Litho 7) 0.62148 0.21082 0.13559 0.22298 0.87371 0.034421 0.082511 0.27849 0.4338 0.21931 0.60454 1 0.11462 0.005811 0.036115

Carbonate rocks (Litho 8) 0.51206 0.56039 0.93054 0.395 0.52955 0.76779 0.51006 0.29346 0.055325 0.087944 0.017359 0.013142 0.66812 0.46453 0.71774

Shale, Graywacke, Sandstone (Litho 9) 0.1457 0.30048 0.49848 0.52496 0.80945 0.59955 0.38101 0.053861 0.26019 0.28857 0.10114 0.14985 0.23369 0.11191 0.0068158

Sands, gravels, sandstones, limestones, clays (Litho 10) 0.081614 0.14491 0.28985 0.66569 0.62145 0.21794 0.26021 1 0.090284 0.33831 0.16482 0.28786 0.75224 0.12186 0.24278

Schist, Amphibolites, Mica schist, Greywackes, Quartzites, Gneiss (Litho 11) 0.62148 0.8348 0.91051 0.87371 0.87371 0.67228 0.18925 0.31759 0.80036 0.80035 0.51545 0.36002 0.0018878 0.28373 0.40596

Evapotranspiration (mm) 0.41126 0.35181 0.050117 0.12816 0.90787 0.5812 0.55819 0.6476 0.28308 0.15871 0.91332 0.17057 0.96227 0.0072829 0.00093329

Solar radiation (Kcal/Cm2) 0.026561 0.7098 0.71764 0.36888 0.57173 0.57015 0.52767 0.73105 0.76943 0.68001 0.17619 0.33747 0.99286 0.020215 0.0022536

Mean Temperature (ºC) 0.48585 0.28811 0.83471 0.068437 0.052083 0.44379 0.40382 0.30415 0.87211 0.85112 0.071363 0.090767 0.941 0.13253 0.0067412

Hours Sunshine 0.26913 0.47133 0.88941 0.15753 0.15361 0.5662 0.30861 0.34432 0.99387 0.93093 0.04581 0.049604 0.88653 0.087608 0.0023176

Rainfall (mm) 0.73348 0.49268 0.94525 0.21518 0.50056 0.31327 0.36664 0.2 0.62834 0.78822 0.16998 0.083818 0.85015 0.16921 0.0044574


84

Título

No. of rays/mm2 Width of widest ray Width of narrowest ray Height of tallest ray (cells) Height of shortest ray (cells) Two ray sizes Scalariform Vessels Homogeneous Heterogeneous I Heterogeneous II Crystals Thickenings Height Abundant Sparse Residual High Moderate Negligible

Number of vessels 0.66255 0.0004472 0.13704 0.86576 0.76857 0.54714 0.31655 0.035797 0.89307 0.035434 0.92119 0.38832 0.7046 0.57841 0.98549 0.60364 0.84844 0.55343 0.88245

Vessel grouping 0.45409 0.45625 0.61708 0.58109 0.70564 0.22489 0.31113 0.22489 0.11342 0.47292 0.096623 0.31113 0.4389 0.060416 0.29386 0.20256 0.79904 0.82982 0.77997

Maximum number of vessels per cluster 0.96794 0.073427 0.11596 0.92967 0.79572 0.072319 8.50E-04 0.30579 0.0013057 0.0025324 0.57784 0.098369 0.4752 0.029989 0.63732 0.014103 0.86982 0.9692 0.97331

Radial Cluster orientation 0.61204 0.4682 0.6263 0.038819 0.8007 0.25825 3.18E-05 0.094433 0.15692 0.00028151 0.61929 0.11255 0.9033 0.43821 0.14244 0.34351 0.77101 0.45551 0.34199

Tangential Cluster orientation 0.24694 0.11016 0.6263 0.069407 0.28372 0.25825 0.32394 0.090809 0.15692 0.98788 0.97545 0.76402 0.0179 0.25721 0.89392 0.34351 0.34832 0.45551 0.34199

No Cluster orientation 0.052783 0.92687 0.23672 0.51727 0.33666 0.4119 0.086892 0.33217 0.32913 0.021162 0.08541 0.49346 0.1743 0.21577 0.05533 0.63218 0.43858 0.30904 0.19682

Total number of clusters 0.20482 0.034487 0.12777 0.34324 0.61698 0.32268 3.30E-03 0.13838 0.0086498 0.077347 0.78168 0.25185 0.4944 0.4925 0.87179 0.28005 0.53259 0.99005 0.53619

Total vessel area (m2) 0.65892 0.0033442 0.20919 0.27232 0.7329 0.73565 1 0.91882 0.3454 0.27063 0.60403 0.15193 0.0931 0.24603 0.20636 0.85086 0.88894 0.60656 0.61897

Area of smallest vessel (m2) 0.090178 0.7086 0.93098 0.059272 0.63091 0.69994 0.35701 0.78659 0.37356 0.39557 0.39957 0.59866 0.2968 0.97621 0.1824 0.12836 0.83121 0.6218 0.6671

area of largest vessel (m2) 0.85608 0.69156 0.59325 0.92111 0.19821 0.010434 0.1982 0.83348 0.38518 0.37474 0.052406 0.0041591 0.3256 0.68429 0.44322 0.16705 0.070571 0.010757 0.47618

Average vessel area (m2) 0.47804 0.36832 0.85112 0.0061377 0.33795 0.47942 0.26018 0.25143 0.85282 0.31483 0.47361 0.023417 0.6661 0.60649 0.1359 0.28965 0.58898 0.34393 0.75716

Average vessel aspect ratio 0.008629 0.95803 0.80612 0.81998 0.72582 0.44452 0.0077821 0.23598 0.00028065 7.21E-04 0.69447 0.79239 0.6006 0.39463 0.65373 0.84762 0.86285 0.76815 0.95069

Height of tallest ray (m) 0.063391 0.097081 0.25405 2.14E-14 0.0029828 0.27312 0.018557 0.41475 0.99785 0.37842 0.58733 0.12468 0.6173 0.88361 0.96615 0.83483 0.075844 0.97772 0.19591

Height of shortest ray (m) 0.030802 0.00957 0.13697 0.80216 1.79E-04 3.14E-06 0.016469 0.049695 0.28331 0.00045456 0.000931 0.040622 0.5984 0.54185 0.18731 0.011919 0.97361 0.48697 0.48447

Average ray height (m) 0.00084547 0.0019019 0.14479 8.84E-07 0.00020577 0.024503 0.26018 0.026603 0.98498 0.015172 0.014538 0.015831 0.1096 0.90505 0.45224 0.28227 0.36784 0.84987 0.67685

No. of rays/mm2 xx 0.67493 0.14452 0.95893 0.023241 0.056188 0.24768 0.0076963 0.21885 0.1902 0.38752 0.18299 0.0172 0.010118 0.45926 0.0057535 0.90048 0.55105 0.59608

Width of widest ray (cells) 0.67493 xx 0.10299 0.18319 0.070709 0.00064398 0.97226 0.94263 0.92614 0.96725 0.053801 0.88939 0.8532 0.92297 0.84572 0.91099 0.36804 0.58255 0.85292

Width of narrowest ray (cells) 0.14452 0.10299 xx 0.1743 0.18267 0.59894 0.85268 0.057247 0.21266 0.64511 0.55507 0.85268 0.1398 0.19671 0.08327 0.71629 0.67442 0.069885 0.21266

Height of tallest ray (cells) 0.95893 0.18319 0.1743 xx 0.2712 0.95422 0.011329 0.95168 0.38171 0.23427 0.13111 0.13163 0.8696 0.30626 0.78014 0.28916 0.084415 0.68635 0.092529

Height of shortest ray (cells) 0.023241 0.070709 0.18267 0.2712 xx 9.17E-09 0.10305 0.89747 0.46674 0.42647 0.030443 0.80828 0.627 0.88376 0.76044 0.78938 0.14961 0.30919 0.82347

Presence of two distinct ray sizes 0.056188 0.00064398 0.59894 0.95422 9.17E-09 xx 0.86981 0.8443 0.93395 0.74942 0.09789 0.86981 0.6013 0.13852 0.44464 0.17643 0.19438 0.10962 0.59269

Scalariform Vessels 0.24768 0.97226 0.85268 0.011329 0.10305 0.86981 xx 0.28671 0.137 0.0021733 0.23188 0.7068 0.83 0.60095 1 0.46162 0.39479 0.26377 0.10408

Homogeneous Rays 0.0076963 0.94263 0.057247 0.95168 0.89747 0.8443 0.28671 xx 8.68E-13 0.0082336 0.71789 0.28671 0.1231 0.73247 0.44464 0.16222 0.073115 0.010059 0.41151

Heterogeneous I Rays 0.21885 0.92614 0.21266 0.38171 0.46674 0.93395 0.137 8.68E-13 xx 3.81E-10 0.078043 0.65301 0.7071 0.19418 0.74702 0.041337 0.11166 0.026671 0.32276

Heterogeneous II Rays 0.1902 0.96725 0.64511 0.23427 0.42647 0.74942 0.0021733 0.0082336 3.81E-10 xx 0.061203 0.083157 0.0312 0.041498 0.21381 3.53E-05 0.91994 0.95489 0.70405

Presence of crystals 0.38752 0.053801 0.55507 0.13111 0.030443 0.09789 0.23188 0.71789 0.078043 0.061203 xx 0.96924 0.9952 0.26013 0.71906 0.29562 0.70539 0.38568 0.71497

Presence of vessel thickenings 0.18299 0.88939 0.85268 0.13163 0.80828 0.86981 0.7068 0.28671 0.65301 0.083157 0.96924 xx 0.4326 0.11663 0.24219 0.46162 0.5705 0.26377 0.55659

Plant height (m) 0.017209 0.85318 0.1398 0.86964 0.62695 0.60126 0.83 0.12308 0.70713 0.0312 0.99516 0.43259 xx 7.81E-05 0.03773 0.0034702 0.73204 0.31015 0.63375

Abundant soil 0.010118 0.92297 0.19671 0.30626 0.88376 0.13852 0.60095 0.73247 0.19418 0.041498 0.26013 0.11663 ###### xx ####### 3.09E-07 1.38E-04 0.82433 0.004553

Sparse soil 0.45926 0.84572 0.083265 0.78014 0.76044 0.44464 1 0.44464 0.74702 0.21381 0.71906 0.24219 0.0377 4.26E-16 xx 0.022086 0.001222 1 0.023946

Residual soil 0.0057535 0.91099 0.71629 0.28916 0.78938 0.17643 0.46162 0.16222 0.041337 3.53E-05 0.29562 0.46162 0.0035 3.09E-07 0.02209 xx 0.13186 0.8583 0.37893

High slope 0.90048 0.36804 0.67442 0.084415 0.14961 0.19438 0.39479 0.073115 0.11166 0.91994 0.70539 0.5705 0.732 0.00013752 0.00122 0.13186 xx 0.011468 1.12E-08

Moderate slope 0.55105 0.58255 0.069885 0.68635 0.30919 0.10962 0.26377 0.010059 0.026671 0.95489 0.38568 0.26377 0.3102 0.82433 1 0.8583 0.011468 xx 6.37E-14

Negligible slope 0.59608 0.85292 0.21266 0.092529 0.82347 0.59269 0.10408 0.41151 0.32276 0.70405 0.71497 0.55659 0.6338 0.004553 0.02395 0.37893 1.12E-08 6.37E-14 xx

Full sun 0.45437 0.14687 0.40612 0.020922 0.90741 0.46798 0.014752 0.24012 0.65162 0.48793 0.57079 0.70559 0.0161 0.004841 0.06949 0.032255 0.73637 0.014926 0.0098905

Partial Shade 0.20375 0.89607 0.60796 0.53878 0.30509 0.44146 0.29873 0.18916 0.0079091 0.046785 0.78178 0.83536 0.01 0.012947 0.24478 0.041979 0.43327 0.093755 0.053626

Full Shade 0.77926 0.094365 0.6263 0.00060079 0.20564 0.99255 3.18E-05 0.99255 0.05894 0.015688 0.57657 0.76402 0.5914 0.21047 0.35062 0.623 0.77101 0.15045 0.34601

N Direction 0.46699 0.79056 0.6263 0.70357 0.66807 0.56871 0.76402 0.58146 0.34199 0.53327 0.12064 0.32394 0.2725 0.010342 0.0234 0.34351 0.77101 0.72947 0.99685

W Direction 0.70784 0.16994 0.69489 0.36785 0.30346 0.76195 0.42688 0.02443 0.21431 0.39835 0.25411 0.028889 0.0397 0.26926 0.21677 0.91156 0.051508 0.64289 0.34231

S Direction 0.015781 0.9415 0.76302 0.52725 0.63637 0.50585 0.54135 0.083663 0.53679 0.27924 0.75421 0.54135 0.0131 0.027242 0.7041 0.0036967 0.64511 0.79547 0.95522

E Direction 0.3242 0.10773 0.83483 0.61268 0.7283 0.92668 0.67274 0.92668 0.96902 0.88103 0.76179 0.034692 0.6226 0.90649 0.43072 0.40844 0.004149 0.20953 0.33164

Altitude 0.76555 0.68537 0.15277 0.6828 0.5061 0.90361 0.11925 0.22345 0.78175 0.33502 0.14491 0.59339 0.9844 0.01556 0.20178 0.0759 0.40184 0.0058975 0.0019444

x(dec º N) 0.056561 0.14076 0.34799 0.55043 0.79192 0.71407 0.0029908 4.12E-05 0.25089 0.0029707 0.084132 0.40041 0.2965 0.071825 0.53634 0.0002225 0.064996 0.12299 0.93571

y (dec º W) 0.011605 0.2493 0.47927 0.20936 0.68671 0.86591 0.38026 0.73799 0.041531 0.024092 0.16413 0.14974 0.0042 2.82E-07 0.01928 4.12E-05 0.22937 0.33761 0.042344

Humic cambisol (Soil 1) 0.036592 0.067572 0.50434 0.73067 0.44173 0.63846 0.0024135 0.054954 0.543 0.0042054 0.84256 0.17622 0.0193 0.0037664 0.73314 4.16E-05 0.42408 0.22035 0.54747

Rankers (Soil 2) 0.73081 0.63135 0.77584 0.19296 0.9341 0.37905 0.56407 0.10233 0.28091 0.69947 0.18659 0.56407 0.0939 0.32696 0.31909 0.95713 0.52982 0.93496 0.73698

Schist associated cambisol (Soil 3) 0.50087 0.97226 0.85268 0.013925 0.027881 0.16357 0.7068 0.28671 0.10408 0.35084 0.2634 0.7068 0.0318 0.0089185 0.01933 0.40015 0.5705 0.93638 0.65301

Eutric lithosol (luvisol associated) (Soil 4) 0.010611 0.48454 0.80344 0.67418 0.93982 0.62666 0.61414 2.44E-07 0.00071508 0.21113 0.10907 0.61414 0.8206 0.61656 0.82281 0.32378 0.25405 0.73668 0.55563

Eutric fluvisol (Soil 5) 0.0067876 0.97226 0.85268 0.47774 0.18704 0.28671 0.7068 0.86981 0.65301 0.68927 0.2634 0.7068 0.006 0.11663 0.01933 0.46162 0.39479 0.26377 0.10408

Luvisol associated cambisol (Soil 6) 0.096626 0.98631 0.92696 0.46992 0.16378 0.057247 0.85268 0.59894 0.42233 0.64511 0.55507 0.85268 0.1398 0.19671 0.08327 0.71629 0.67442 0.58117 0.42233

Distric regosol (Soil 7) 0.070697 0.01837 0.83483 0.818 0.11249 0.23159 0.67274 0.23159 0.067979 0.29482 0.075817 0.034692 0.0149 0.078084 0.1891 0.40844 0.004149 0.37008 0.0045855

Calcic luvisol (Soil 8) 0.67524 0.98631 0.92696 0.7071 0.78754 0.59894 0.85268 0.59894 0.21266 0.029913 0.55507 0.85268 0.413 0.19671 0.5637 0.71629 0.67442 0.58117 0.21266

Gleyic solonchaks (Soil 9) 0.37207 0.4239 0.85268 0.31601 0.34322 0.16357 0.7068 0.28671 0.10408 0.35084 0.23188 0.7068 0.8127 0.11663 0.24219 0.46162 0.39479 0.93638 0.55659

Orthic luvisol (Soil 10) 0.43105 0.2795 0.75073 0.6753 0.20448 0.29013 0.51989 0.76942 0.13627 0.11048 0.54467 0.51989 0.1344 0.46412 0.85567 0.20788 0.038138 0.67296 0.27491

Ferric luvisol (Soil 11) 0.31187 0.98631 0.92696 0.46992 0.18267 0.59894 0.85268 0.59894 0.42233 0.64511 0.55507 0.85268 0.8722 0.19671 0.08327 0.71629 0.67442 0.069885 0.21266

Albic Luvisol (Soil 12) 0.39287 0.0017417 0.83483 0.020294 0.049795 0.034303 0.67274 0.92668 0.37177 0.29482 0.004881 0.67274 0.0664 0.41095 0.79282 0.40844 0.33931 0.85773 0.37177

Chromic luvisol (Soil 13) 0.027973 0.15116 0.73888 0.037509 0.62735 0.23925 0.49953 0.077517 0.85226 0.093988 0.44636 0.49953 0.6292 0.0048532 0.03578 0.18634 0.12663 0.38994 0.66389

Eutric lithosol (Soil 14) 0.010611 0.48454 0.80344 0.67418 0.93982 0.62666 0.61414 2.44E-07 0.00071508 0.21113 0.10907 0.61414 0.8206 0.61656 0.82281 0.32378 0.25405 0.73668 0.55563

Plinthic luvisol (Soil 15) 0.7837 0.10299 0.92696 0.32573 0.18267 0.59894 0.85268 0.59894 0.21266 0.029913 0.55507 0.85268 0.413 0.19671 0.08327 0.71629 0.01729 0.58117 0.21266

Orthic podzol (Soil 16) 0.77701 0.043503 1.31E-05 0.89926 0.65296 0.4785 0.64213 0.00017961 0.023547 0.24906 0.13967 0.64213 0.2859 0.017821 0.01603 0.69669 0.29311 0.010346 0.15856

Schist and quartzite associated cambisol (Soil 17) 0.60799 0.21833 0.89642 0.60786 0.71773 0.45512 0.79199 0.3287 0.75261 0.51305 0.40193 0.79199 0.5965 0.2713 0.41225 0.60576 0.55077 0.43334 0.25447

Dystric cambisol (Soil 18) 0.22216 0.95141 0.89642 0.78171 0.25527 0.45512 0.79199 0.45512 0.25447 0.51305 0.43298 0.79199 0.1909 0.2713 0.41225 0.60576 0.55077 0.43334 0.25447

Chromic cambisol (Soil 19) 0.11413 0.0040232 0.89642 0.024609 0.25527 0.45512 0.79199 0.45512 0.75261 0.22446 0.40193 0.00022239 0.198 0.2713 0.41225 0.60576 0.55077 0.43334 0.25447

nº days frost 0.039704 0.15955 0.83779 0.2013 0.75747 0.61698 0.46348 0.69146 0.65909 0.31873 0.54764 0.37819 0.0206 2.38E-05 0.04566 0.0004058 0.70968 0.73665 0.51162

nº days rainfall 0.10213 0.03109 0.24631 0.11589 0.53544 0.55284 0.024162 0.0073401 0.95179 0.0046668 0.052889 0.94383 0.2682 0.021023 0.77769 0.0018565 0.10835 0.2517 0.86286

Granites (Litho 1) 0.072013 0.13989 0.43045 0.30196 0.43806 0.3561 0.010191 0.00677 0.99491 0.0034131 0.36592 0.11024 0.1886 0.026371 0.82907 0.0001118 0.2715 0.22693 0.69696

Schist and Greywacke (Litho 2) 0.68171 0.4192 0.66449 0.027882 0.37087 0.59338 0.37962 0.20074 0.2129 0.83133 0.53456 0.61099 0.6283 0.65265 0.19568 0.34528 0.13377 0.39872 0.77577

Alluvium (Litho 3) 0.029689 0.2181 0.80344 0.83569 0.52294 0.64907 0.61414 0.15331 0.16637 0.81834 0.86514 0.61414 0.1597 0.76384 0.26285 0.32378 0.25405 0.13417 0.029364

Aeolian sands (Litho 4) 0.070697 0.01837 0.83483 0.818 0.11249 0.23159 0.67274 0.23159 0.067979 0.29482 0.075817 0.034692 0.0149 0.078084 0.1891 0.40844 0.004149 0.37008 0.0045855

Conglomerate,Sandstone,Limestone,Marlstone (Litho 5) 0.096626 0.98631 0.92696 0.46992 0.16378 0.057247 0.85268 0.59894 0.42233 0.64511 0.55507 0.85268 0.1398 0.19671 0.08327 0.71629 0.67442 0.58117 0.42233

Metavulcanite (Litho 6) 0.13296 0.10299 0.92696 0.44369 0.78754 0.59894 0.85268 0.59894 0.42233 0.64511 0.090191 0.85268 0.0819 0.43858 0.5637 0.71629 0.67442 0.58117 0.42233

Sands, pebbles, clays, weakly consolidated sandstone (Litho 7) 0.36605 0.0040028 0.81855 0.2188 0.033721 0.085219 0.64213 0.76133 0.24867 0.24906 0.019563 0.64213 0.0393 0.052559 0.14849 0.36308 0.29311 0.16739 0.044633

Carbonate rocks (Litho 8) 0.48505 0.35267 0.87278 0.53902 0.087495 0.35826 0.74566 0.35826 0.16103 0.4211 0.30258 0.74566 0.0903 0.17604 0.31321 0.52554 0.46307 0.33521 0.16103

Shale, Graywacke, Sandstone (Litho 9) 0.069523 0.54176 0.77584 0.73072 0.90539 0.4261 0.56407 2.33E-05 0.014008 0.15254 0.066767 0.56407 0.8638 0.32696 0.84206 0.25895 0.19193 0.12104 0.73698

Sands, gravels, sandstones, limestones, clays (Litho 10) 0.84967 0.083159 0.00018281 0.78418 0.84854 0.81351 0.58824 0.00016044 0.003891 0.70124 0.085312 0.58824 0.565 0.0026031 0.00076 0.93967 0.83826 0.0069378 0.032333

Schist, Amphibolites, Mica schist, Greywackes, Quartzites, Gneiss (Litho 11) 0.8002 0.096734 0.81855 0.00070455 0.11698 0.18811 0.64213 0.18811 0.24867 0.95623 0.13967 2.88E-05 0.2344 0.052559 0.14849 0.36308 0.29311 0.16739 0.044633

Evapotranspiration (mm) 0.013307 0.11322 0.76819 0.12079 0.55092 0.4313 0.0009207 0.012857 0.81767 0.0027586 0.27187 0.78818 0.0594 0.0032944 0.4503 0.0008548 0.1486 0.63019 0.48916

Solar radiation (Kcal/Cm2) 0.0072501 0.094882 0.31419 0.16966 0.72673 0.85534 0.028454 3.23E-05 0.21418 0.0036013 0.16221 0.81947 0.311 0.080313 0.67029 0.0015289 0.071655 0.16701 0.88213

Mean Temperature (ºC) 0.07681 0.52279 0.43014 0.018982 0.84373 0.4821 0.01867 0.0011597 0.33885 0.021816 0.1995 0.53647 0.9766 0.25879 0.664 0.18009 0.070061 0.10274 0.81371

Hours Sunshine 0.04638 0.15236 0.44754 0.025217 0.23463 0.52969 0.024068 0.00053084 0.28261 0.017491 0.045936 0.90579 0.4628 0.11863 0.74748 0.090711 0.15984 0.21036 0.92631

Rainfall (mm) 0.16604 0.43611 0.35823 0.067544 0.43484 0.84186 0.02476 0.0067387 0.6211 0.021294 0.097585 0.70095 0.6291 0.27337 0.88075 0.1393 0.16497 0.41331 0.77343


85

Título

Full sun Partial Shade Full Shade N Direction W Direction S Direction E Direction Altitude x(dec º N) y (dec º W) Soil 1 Soil 2 Soil 3 Soil 4 Soil 5 Soil 6 Soil 7 Soil 8 Soil 9 Soil 10 Soil 11 Soil 12 Soil 13 Soil 14 Soil 15 Soil 16 Soil 17 Soil 18 Soil 19

Number of vessels 0.2214 0.90739 0.091596 0.8676 0.88032 0.67954 0.21452 0.64206 0.004041 0.70547 0.002667 0.02836 0.039241 0.011934 0.67155 0.91951 0.68871 0.22527 0.38431 0.056116 0.78386 0.91631 0.061171 0.011934 0.17478 0.23554 0.78983 0.022255 0.045613

Vessel grouping 0.579 1 0.4189 0.13331 0.14134 0.10006 1 0.013953 0.38096 0.035439 0.43105 0.49002 0.31113 0.69811 0.31113 0.61708 0.25541 0.61708 0.31113 4.92E-06 0.61708 0.25541 0.76187 0.69811 0.61708 0.21082 0.47764 0.47764 0.28681

Maximum number of vessels per cluster 0.7755 0.21625 0.44068 0.2145 0.60274 0.069217 0.2228 0.028935 0.02819 0.013801 0.03774 0.54892 0.3958 0.72318 0.3958 0.51973 0.84876 0.51973 0.75591 0.00258 0.67508 0.005232 0.39373 0.72318 0.67508 0.50403 0.55162 0.41303 0.80684

Radial Cluster orientation 0.44982 0.30825 0.035081 0.72847 0.52601 0.44369 0.96159 0.42533 0.17472 0.81026 0.96344 0.26358 0.11255 0.73636 0.32394 0.6263 0.96159 0.6263 0.004061 0.091514 0.6263 0.26803 0.31693 0.73636 0.040012 0.87371 0.48922 0.26592 0.26592

Tangential Cluster orientation 0.44982 0.90569 0.72847 0.72847 0.96387 0.94448 0.26803 0.065585 0.4313 0.23127 0.96344 0.13039 0.11255 0.51651 0.76402 0.6263 0.96159 0.040012 0.32394 0.091514 0.6263 0.26803 0.004394 0.51651 0.6263 0.87371 0.26592 0.48922 0.48922

No Cluster orientation 0.097546 0.47221 0.0089082 0.50691 0.46953 0.43637 0.39763 0.24559 0.79827 0.57611 0.57163 0.38153 0.086892 0.94773 0.17081 0.23672 0.39763 0.39802 0.49346 0.31165 0.23672 0.007116 0.21897 0.94773 0.39802 0.20442 0.81029 0.81029 0.23003

Total number of clusters 0.79264 0.91983 0.51795 0.36398 0.50716 0.079977 0.35905 0.14634 0.15337 0.5261 0.24754 0.26002 0.14584 0.92381 0.71342 0.42502 0.75145 0.35326 0.25185 0.003254 0.67403 0.012625 0.66348 0.92381 0.42502 0.28425 1 0.15824 0.94253

Total vessel area (m2) 0.44996 0.01141 0.13973 0.15654 0.30635 0.58183 0.64106 0.016556 0.2541 0.59217 0.40351 0.092211 0.0019392 0.01355 0.84926 0.2309 0.002933 0.41065 0.4559 0.56359 0.98848 0.32787 0.24821 0.01355 0.29202 0.53122 1 0.48581 0.18271

Area of smallest vessel (m2) 0.29656 0.18525 0.0042314 0.65267 0.21387 0.37976 0.38604 0.21513 0.67526 0.16213 0.96827 0.22796 0.010961 0.53429 0.61398 0.10286 0.008789 0.41063 0.84353 0.048424 0.55396 0.90071 0.14403 0.53429 0.15718 0.42326 0.83757 0.75846 0.98364

area of largest vessel (m2) 0.93404 0.083844 0.035396 0.11403 0.17174 0.71112 0.12275 0.19577 0.35123 0.097249 0.94562 0.71235 0.0049951 0.53425 0.76996 0.46158 0.009852 0.34807 0.42978 0.095099 0.26632 0.26705 0.17788 0.53425 0.89663 0.34756 0.85361 0.43593 0.65196

Average vessel area (m2) 0.7649 0.01913 0.0033435 0.20994 0.3808 0.16126 0.85925 0.06888 0.58889 0.74233 0.43951 0.71233 0.0031385 0.2286 0.95336 0.21975 0.004102 0.53472 0.96501 0.028687 0.76175 0.42677 0.001952 0.2286 0.57339 0.23788 0.95095 0.57985 0.69685

Average vessel aspect ratio 0.61944 0.68149 0.51772 0.99468 0.17783 0.039767 0.61304 0.41534 0.034521 0.90275 0.81573 0.19513 0.16927 0.53422 0.17385 0.78385 0.63161 0.085786 0.60879 0.026757 0.11556 0.45006 0.97208 0.53422 0.98848 0.35373 0.48572 0.69685 0.064985

Height of tallest ray (m) 0.095676 0.28475 0.31075 0.61696 0.856 0.72173 0.10758 0.055534 0.47823 0.30756 0.31988 0.34887 0.27921 0.28988 0.48274 0.19881 0.055968 0.87382 0.24206 0.070964 0.31916 1.19E-02 0.98604 0.28988 0.44416 0.47734 0.91834 0.96729 2.17E-02

Height of shortest ray (m) 0.1562 0.79099 0.058216 0.83621 0.17905 0.35938 0.90068 0.77064 0.00090126 0.17848 0.038267 0.51067 0.44698 0.11047 0.36457 0.10277 0.64567 0.51586 0.98833 0.19173 0.18888 4.11E-02 0.14635 0.11047 0.3053 0.25252 0.24248 0.26816 0.88586

Average ray height (m) 0.70457 0.27328 0.57536 0.2951 0.25151 0.80129 0.27271 0.011167 0.00057498 0.71073 0.0020259 0.92062 0.46469 0.027754 0.34934 0.12239 0.99476 0.67545 0.38028 0.000225 0.12239 1.19E-02 0.18067 0.027754 0.15292 0.1975 0.56589 0.62265 1.94E-02

No. of rays/mm2 0.45437 0.20375 0.77926 0.46699 0.70784 0.015781 0.3242 0.76555 0.056561 0.011605 0.036592 0.73081 0.50087 0.010611 0.006788 0.096626 0.070697 0.67524 0.37207 0.43105 0.31187 0.39287 2.80E-02 0.010611 0.7837 0.77701 0.60799 0.22216 0.11413

Width of widest ray (cells) 0.14687 0.89607 0.094365 0.79056 0.16994 0.9415 0.10773 0.68537 0.14076 0.2493 0.067572 0.63135 0.97226 0.48454 0.97226 0.98631 0.01837 0.98631 0.4239 0.2795 0.98631 0.001742 0.15116 0.48454 0.10299 0.043503 0.21833 0.95141 4.02E-03

Width of narrowest ray (cells) 0.40612 0.60796 0.6263 0.6263 0.69489 0.76302 0.83483 0.15277 0.34799 0.47927 0.50434 0.77584 0.85268 0.80344 0.85268 0.92696 0.83483 0.92696 0.85268 0.75073 0.92696 0.83483 0.73888 0.80344 0.92696 1.31E-05 0.89642 0.89642 0.89642

Height of tallest ray (cells) 0.020922 0.53878 0.00060079 0.70357 0.36785 0.52725 0.61268 0.6828 0.55043 0.20936 0.73067 0.19296 0.013925 0.67418 0.47774 0.46992 0.818 0.7071 0.31601 0.6753 0.46992 2.03E-02 0.037509 0.67418 0.32573 0.89926 0.60786 0.78171 0.024609

Height of shortest ray (cells) 0.90741 0.30509 0.20564 0.66807 0.30346 0.63637 0.7283 0.5061 0.79192 0.68671 0.44173 0.9341 0.027881 0.93982 0.18704 0.16378 0.11249 0.78754 0.34322 0.20448 0.18267 0.049795 0.62735 0.93982 0.18267 0.65296 0.71773 0.25527 0.25527

Presence of two distinct ray sizes 0.46798 0.44146 0.99255 0.56871 0.76195 0.50585 0.92668 0.90361 0.71407 0.86591 0.63846 0.37905 0.16357 0.62666 0.28671 0.057247 0.23159 0.59894 0.16357 0.29013 0.59894 0.034303 0.23925 0.62666 0.59894 0.4785 0.45512 0.45512 0.45512

Scalariform Vessels 0.014752 0.29873 3.18E-05 0.76402 0.42688 0.54135 0.67274 0.11925 0.0029908 0.38026 0.0024135 0.56407 0.7068 0.61414 0.7068 0.85268 0.67274 0.85268 0.7068 0.51989 0.85268 0.67274 0.49953 0.61414 0.85268 0.64213 0.79199 0.79199 0.79199

Homogeneous Rays 0.24012 0.18916 0.99255 0.58146 0.02443 0.083663 0.92668 0.22345 4.12E-05 0.73799 0.054954 0.10233 0.28671 2.44E-07 0.86981 0.59894 0.23159 0.59894 0.28671 0.76942 0.59894 0.92668 0.077517 2.44E-07 0.59894 0.00018 0.3287 0.45512 0.45512

Heterogeneous I Rays 0.65162 0.0079091 0.05894 0.34199 0.21431 0.53679 0.96902 0.78175 0.25089 0.041531 0.543 0.28091 0.10408 0.000715 0.65301 0.42233 0.067979 0.21266 0.10408 0.13627 0.42233 0.37177 0.85226 0.000715 0.21266 0.023547 0.75261 0.25447 0.75261

Heterogeneous II Rays 0.48793 0.046785 0.015688 0.53327 0.39835 0.27924 0.88103 0.33502 0.0029707 0.024092 0.0042054 0.69947 0.35084 0.21113 0.68927 0.64511 0.29482 0.029913 0.35084 0.11048 0.64511 0.29482 0.093988 0.21113 0.029913 0.24906 0.51305 0.51305 0.22446

Presence of crystals 0.57079 0.78178 0.57657 0.12064 0.25411 0.75421 0.76179 0.14491 0.084132 0.16413 0.84256 0.18659 0.2634 0.10907 0.2634 0.55507 0.075817 0.55507 0.23188 0.54467 0.55507 0.004881 0.44636 0.10907 0.55507 0.13967 0.40193 0.43298 0.40193

Presence of vessel thickenings 0.70559 0.83536 0.76402 0.32394 0.028889 0.54135 0.034692 0.59339 0.40041 0.14974 0.17622 0.56407 0.7068 0.61414 0.7068 0.85268 0.034692 0.85268 0.7068 0.51989 0.85268 0.67274 0.49953 0.61414 0.85268 0.64213 0.79199 0.79199 0.000222

Plant height (m) 0.016113 0.010014 0.59138 0.27251 0.039746 0.013097 0.62262 0.98444 0.29652 0.0041932 0.019303 0.093908 0.031793 0.82057 0.00602 0.1398 0.014928 0.41297 0.81273 0.13444 0.87224 0.066422 0.6292 0.82057 0.41297 0.28593 0.59651 0.19087 0.19801

Abundant soil 0.004841 0.012947 0.21047 0.010342 0.26926 0.027242 0.90649 0.01556 0.071825 2.82E-07 0.0037664 0.32696 0.0089185 0.61656 0.11663 0.19671 0.078084 0.19671 0.11663 0.46412 0.19671 0.41095 0.004853 0.61656 0.19671 0.017821 0.2713 0.2713 0.2713

Sparse soil 0.069487 0.24478 0.35062 0.023403 0.21677 0.7041 0.43072 0.20178 0.53634 0.019282 0.73314 0.31909 0.019332 0.82281 0.019332 0.083265 0.1891 0.5637 0.24219 0.85567 0.083265 0.79282 0.035783 0.82281 0.083265 0.016033 0.41225 0.41225 0.41225

Residual soil 0.032255 0.041979 0.623 0.34351 0.91156 0.0036967 0.40844 0.0759 0.00022246 4.12E-05 4.16E-05 0.95713 0.40015 0.32378 0.46162 0.71629 0.40844 0.71629 0.46162 0.20788 0.71629 0.40844 0.18634 0.32378 0.71629 0.69669 0.60576 0.60576 0.60576

High slope 0.73637 0.43327 0.77101 0.77101 0.051508 0.64511 0.0041491 0.40184 0.064996 0.22937 0.42408 0.52982 0.5705 0.25405 0.39479 0.67442 0.004149 0.67442 0.39479 0.038138 0.67442 0.33931 0.12663 0.25405 0.01729 0.29311 0.55077 0.55077 0.55077

Moderate slope 0.014926 0.093755 0.15045 0.72947 0.64289 0.79547 0.20953 0.0058975 0.12299 0.33761 0.22035 0.93496 0.93638 0.73668 0.26377 0.58117 0.37008 0.58117 0.93638 0.67296 0.069885 0.85773 0.38994 0.73668 0.58117 0.010346 0.43334 0.43334 0.43334

Negligible slope 0.009891 0.053626 0.34601 0.99685 0.34231 0.95522 0.33164 0.0019444 0.93571 0.042344 0.54747 0.73698 0.65301 0.55563 0.10408 0.42233 0.004586 0.21266 0.55659 0.27491 0.21266 0.37177 0.66389 0.55563 0.21266 0.15856 0.25447 0.25447 0.25447

Full sun xx 1.63E-11 1.61E-10 0.22036 0.17959 0.4685 0.96924 0.51628 0.42617 0.065402 0.21319 0.10256 0.1591 0.4982 0.092406 0.22869 0.006181 0.22869 0.092406 0.75266 0.22869 0.058807 0.19545 0.4982 0.40612 0.21852 0.79112 0.23807 0.23807

Partial Shade 1.63E-11 xx 0.006431 0.0030574 0.0001904 0.94619 0.96277 0.23544 0.95876 0.060863 0.0056593 0.0079128 0.29873 0.16368 0.29873 0.051253 9.25E-06 0.60796 0.29873 0.821 0.051253 0.24327 0.000786 0.16368 0.60796 0.1992 0.46628 0.46628 0.46628

Full Shade 1.61E-10 0.006431 xx 0.15763 0.037205 0.44369 0.96159 0.030918 0.29048 0.4451 0.14581 0.52496 0.0040605 0.51651 0.32394 0.6263 0.26803 0.6263 0.32394 0.9309 0.6263 0.26803 0.076621 0.51651 0.6263 0.87371 0.26592 0.48922 0.48922

N Direction 0.22036 0.0030574 0.15763 xx 0.037205 0.10924 0.26803 0.019723 0.4693 0.47754 0.64964 0.52496 0.32394 0.51651 0.32394 0.040012 0.26803 0.6263 0.11255 0.091514 0.040012 0.26803 0.037656 0.51651 0.040012 0.22298 0.48922 0.48922 0.48922

W Direction 0.17959 0.0001904 0.037205 0.037205 xx 0.19702 0.3723 0.13398 0.00016938 0.074452 0.58887 1.14E-04 0.42688 0.2869 0.42688 0.69489 6.66E-09 0.69489 0.42688 0.17407 0.69489 0.3723 0.1538 0.2869 0.69489 0.32629 0.5775 0.12555 0.12555

S Direction 0.4685 0.94619 0.44369 0.10924 0.19702 xx 0.49282 0.31162 0.75035 0.13592 0.95267 0.34898 0.22188 0.413 1.03E-06 0.76302 0.49282 0.76302 0.54135 0.017564 0.76302 0.49282 0.27292 0.413 0.76302 0.45051 0.6685 0.6685 0.6685

E Direction 0.96924 0.96277 0.96159 0.26803 0.3723 0.49282 xx 0.79776 0.63159 0.43443 0.59362 0.51718 0.67274 0.5713 0.67274 0.83483 0.63529 0.83483 0.67274 0.015077 0.83483 0.63529 0.44833 0.5713 0.83483 0.60179 0.76713 0.76713 0.76713

Altitude 0.51628 0.23544 0.030918 0.019723 0.13398 0.31162 0.79776 xx 9.90E-03 3.42E-10 0.27746 0.0048571 0.55846 0.037682 0.00129 0.41037 0.00044 0.59308 0.000689 0.38233 0.57325 0.24219 0.88171 0.037682 0.18394 0.018803 0.28615 0.086766 0.038297

x(dec º N) 0.42617 0.95876 0.29048 0.4693 0.00016938 0.75035 0.63159 0.0099024 xx 0.0034906 4.39E-14 0.061662 0.83778 1.10E-05 0.38025 0.76173 0.070868 0.61331 0.24203 0.044376 0.13697 0.002408 9.18E-05 1.10E-05 0.10901 0.041794 0.7584 0.49212 0.19641

y (dec º W) 0.065402 0.060863 0.4451 0.47754 0.074452 0.13592 0.43443 3.42E-10 0.0034906 xx 1.94E-05 0.12855 0.53909 0.013524 0.2193 0.9195 0.006414 0.08577 0.030441 0.15322 0.23077 0.029208 0.000118 0.013524 0.42714 0.44797 0.038358 0.017869 0.0205

Humic cambisol (Soil 1) 0.21319 0.0056593 0.14581 0.64964 0.58887 0.95267 0.59362 0.27746 4.39E-14 1.94E-05 xx 0.038085 0.17622 0.069866 0.17622 0.50434 0.12884 0.50434 0.17622 0.020681 0.50434 0.12884 0.01519 0.069866 0.50434 0.094735 0.34302 0.34302 0.34302

Rankers (Soil 2) 0.10256 0.0079128 0.52496 0.52496 0.00011425 0.34898 0.51718 0.0048571 0.061662 0.12855 0.038085 xx 0.56407 0.43946 0.56407 0.77584 0.51718 0.77584 0.56407 0.32375 0.77584 0.51718 0.30048 0.43946 0.77584 0.47608 0.68592 0.68592 0.68592

Schist associated cambisol (Soil 3) 0.1591 0.29873 0.0040605 0.32394 0.42688 0.22188 0.67274 0.55846 0.83778 0.53909 0.17622 0.56407 xx 0.61414 0.7068 0.85268 0.67274 0.85268 0.7068 0.51989 0.85268 0.67274 0.49953 0.61414 0.85268 0.64213 0.79199 0.79199 0.79199

Eutric lithosol (luvisol associated) (Soil 4) 0.4982 0.16368 0.51651 0.51651 0.2869 0.413 0.5713 0.037682 1.10E-05 0.013524 0.069866 0.43946 0.61414 xx 0.61414 0.80344 0.5713 0.80344 0.61414 0.3884 0.80344 0.5713 0.36545 1.05E-27 0.80344 0.53336 0.72373 0.72373 0.72373

Eutric fluvisol (Soil 5) 0.092406 0.29873 0.32394 0.32394 0.42688 1.03E-06 0.67274 0.0012899 0.38025 0.2193 0.17622 0.56407 0.7068 0.61414 xx 0.85268 0.67274 0.85268 0.7068 0.51989 0.85268 0.67274 0.49953 0.61414 0.85268 0.64213 0.79199 0.79199 0.79199

Luvisol associated cambisol (Soil 6) 0.22869 0.051253 0.6263 0.040012 0.69489 0.76302 0.83483 0.41037 0.76173 0.9195 0.50434 0.77584 0.85268 0.80344 0.85268 xx 0.83483 0.92696 0.85268 0.75073 0.92696 0.83483 0.73888 0.80344 0.92696 0.81855 0.89642 0.89642 0.89642

Distric regosol (Soil 7) 0.006181 9.25E-06 0.26803 0.26803 6.66E-09 0.49282 0.63529 0.00043952 0.070868 0.0064138 0.12884 0.51718 0.67274 0.5713 0.67274 0.83483 xx 0.83483 0.67274 0.46993 0.83483 0.63529 0.44833 0.5713 0.83483 0.60179 0.76713 0.76713 0.76713

Calcic luvisol (Soil 8) 0.22869 0.60796 0.6263 0.6263 0.69489 0.76302 0.83483 0.59308 0.61331 0.08577 0.50434 0.77584 0.85268 0.80344 0.85268 0.92696 0.83483 xx 0.85268 0.75073 0.92696 0.83483 0.73888 0.80344 0.92696 0.81855 0.89642 0.89642 0.89642

Gleyic solonchaks (Soil 9) 0.092406 0.29873 0.32394 0.11255 0.42688 0.54135 0.67274 0.00068938 0.24203 0.030441 0.17622 0.56407 0.7068 0.61414 0.7068 0.85268 0.67274 0.85268 xx 0.51989 0.85268 0.67274 0.49953 0.61414 0.85268 0.64213 0.79199 0.79199 0.79199

Orthic luvisol (Soil 10) 0.75266 0.821 0.9309 0.091514 0.17407 0.017564 0.015077 0.38233 0.044376 0.15322 0.020681 0.32375 0.51989 0.3884 0.51989 0.75073 0.46993 0.75073 0.51989 xx 0.75073 0.46993 0.24803 0.3884 0.75073 0.4266 0.65187 0.65187 0.65187

Ferric luvisol (Soil 11) 0.22869 0.051253 0.6263 0.040012 0.69489 0.76302 0.83483 0.57325 0.13697 0.23077 0.50434 0.77584 0.85268 0.80344 0.85268 0.92696 0.83483 0.92696 0.85268 0.75073 xx 0.83483 0.73888 0.80344 0.92696 0.81855 0.89642 0.89642 0.89642

Albic Luvisol (Soil 12) 0.058807 0.24327 0.26803 0.26803 0.3723 0.49282 0.63529 0.24219 0.0024084 0.029208 0.12884 0.51718 0.67274 0.5713 0.67274 0.83483 0.63529 0.83483 0.67274 0.46993 0.83483 xx 0.44833 0.5713 0.83483 0.60179 0.76713 0.76713 0.76713

Chromic luvisol (Soil 13) 0.19545 0.00078614 0.076621 0.037656 0.1538 0.27292 0.44833 0.88171 9.18E-05 0.00011809 0.01519 0.30048 0.49953 0.36545 0.49953 0.73888 0.44833 0.73888 0.49953 0.24803 0.73888 0.44833 xx 0.36545 0.73888 0.40416 0.63593 0.63593 0.63593

Eutric lithosol (Soil 14) 0.4982 0.16368 0.51651 0.51651 0.2869 0.413 0.5713 0.037682 1.10E-05 0.013524 0.069866 0.43946 0.61414 1.05E-27 0.61414 0.80344 0.5713 0.80344 0.61414 0.3884 0.80344 0.5713 0.36545 xx 0.80344 0.53336 0.72373 0.72373 0.72373

Plinthic luvisol (Soil 15) 0.40612 0.60796 0.6263 0.040012 0.69489 0.76302 0.83483 0.18394 0.10901 0.42714 0.50434 0.77584 0.85268 0.80344 0.85268 0.92696 0.83483 0.92696 0.85268 0.75073 0.92696 0.83483 0.73888 0.80344 xx 0.81855 0.89642 0.89642 0.89642

Orthic podzol (Soil 16) 0.21852 0.1992 0.87371 0.22298 0.32629 0.45051 0.60179 0.018803 0.041794 0.44797 0.094735 0.47608 0.64213 0.53336 0.64213 0.81855 0.60179 0.81855 0.64213 0.4266 0.81855 0.60179 0.40416 0.53336 0.81855 xx 0.74456 0.74456 0.74456

Schist and quartzite associated cambisol (Soil 17) 0.79112 0.46628 0.26592 0.48922 0.5775 0.6685 0.76713 0.28615 0.7584 0.038358 0.34302 0.68592 0.79199 0.72373 0.79199 0.89642 0.76713 0.89642 0.79199 0.65187 0.89642 0.76713 0.63593 0.72373 0.89642 0.74456 xx 0.85331 0.85331

Dystric cambisol (Soil 18) 0.23807 0.46628 0.48922 0.48922 0.12555 0.6685 0.76713 0.086766 0.49212 0.017869 0.34302 0.68592 0.79199 0.72373 0.79199 0.89642 0.76713 0.89642 0.79199 0.65187 0.89642 0.76713 0.63593 0.72373 0.89642 0.74456 0.85331 xx 0.85331

Chromic cambisol (Soil 19) 0.23807 0.46628 0.48922 0.48922 0.12555 0.6685 0.76713 0.038297 0.19641 0.0205 0.34302 0.68592 0.79199 0.72373 0.79199 0.89642 0.76713 0.89642 0.79199 0.65187 0.89642 0.76713 0.63593 0.72373 0.89642 0.74456 0.85331 0.85331 xx

nº days frost 0.026764 0.69932 0.013056 0.18 0.077139 0.2604 0.7596 0.00032239 0.0052219 2.71E-12 2.35E-05 0.21255 0.78409 0.74218 0.075521 0.24801 0.008597 0.61906 0.2572 0.002291 0.83779 0.45223 0.53503 0.74218 0.83779 0.36025 0.024905 0.13758 0.072435

nº days rainfall 0.43511 0.90087 0.39155 0.50667 0.00012083 0.91497 0.27263 0.02913 5.20E-19 7.77E-06 1.16E-13 0.0022355 0.92516 0.15532 0.92516 0.26572 0.011354 0.96301 0.01885 0.00302 0.24631 0.008362 2.49E-05 0.15532 0.24631 0.003715 0.94749 0.099661 0.94749

Granites (Litho 1) 0.76572 0.2343 0.29968 0.9306 0.11486 0.57293 0.39106 0.010549 2.78E-16 0.0012204 2.51E-20 8.16E-05 0.11024 0.032302 0.11024 0.43045 0.072912 0.43045 0.11024 0.00629 0.43045 0.072912 0.004145 0.032302 0.43045 0.048479 0.26283 0.26283 0.26283

Schist and Greywacke (Litho 2) 0.25608 0.22983 0.0058276 0.094669 0.26133 0.70735 0.0059233 0.12461 0.21614 0.0040693 0.001583 0.1779 3.01E-06 0.23895 0.37962 0.021133 0.32386 0.66449 0.37962 3.75E-10 0.66449 0.32386 0.11476 0.23895 0.66449 0.27772 0.53791 0.001059 0.53791

Alluvium (Litho 3) 0.14253 0.16368 0.73636 0.73636 0.2869 0.00069521 0.5713 0.74525 0.20564 0.15328 0.069866 0.43946 0.61414 0.49919 2.26E-09 0.80344 0.5713 0.80344 2.26E-09 0.3884 0.80344 0.5713 0.36545 0.49919 0.80344 0.24736 0.72373 0.72373 0.72373

Aeolian sands (Litho 4) 0.006181 9.25E-06 0.26803 0.26803 6.66E-09 0.49282 0.63529 0.00043952 0.070868 0.0064138 0.12884 0.51718 0.67274 0.5713 0.67274 0.83483 1.05E-27 0.83483 0.67274 0.46993 0.83483 0.63529 0.44833 0.5713 0.83483 0.60179 0.76713 0.76713 0.76713

Conglomerate,Sandstone,Limestone,Marlstone (Litho 5) 0.22869 0.051253 0.6263 0.040012 0.69489 0.76302 0.83483 0.41037 0.76173 0.9195 0.50434 0.77584 0.85268 0.80344 0.85268 1.05E-27 0.83483 0.92696 0.85268 0.75073 0.92696 0.83483 0.73888 0.80344 0.92696 0.81855 0.89642 0.89642 0.89642

Metavulcanite (Litho 6) 0.40612 0.60796 0.6263 0.6263 0.69489 0.76302 0.83483 0.87377 0.29188 0.29189 0.50434 0.77584 0.85268 0.80344 0.85268 0.92696 0.83483 0.92696 0.85268 0.75073 0.92696 0.83483 0.0027 0.80344 0.92696 0.81855 0.89642 0.89642 0.89642

Sands, pebbles, clays, weakly consolidated sandstone (Litho 7) 0.21852 0.1992 0.87371 0.22298 0.32629 0.45051 0.60179 0.94237 0.01104 0.00084895 0.094735 0.47608 0.64213 0.53336 0.64213 0.81855 0.60179 0.81855 0.64213 0.4266 0.81855 4.98E-15 0.40416 0.53336 0.81855 0.56587 6.00E-10 0.74456 0.74456

Carbonate rocks (Litho 8) 0.35858 0.04868 0.395 0.395 0.49325 0.59842 0.010784 0.85982 0.036318 0.091071 0.2435 0.61891 0.74566 0.66374 0.74566 0.87278 0.71569 0.87278 0.74566 0.57895 0.87278 0.71569 1.60E-07 0.66374 0.87278 0.68861 0.8201 0.8201 0.8201

Shale, Graywacke, Sandstone (Litho 9) 0.63552 0.45711 0.80945 0.80945 0.22309 0.11854 0.51718 0.0081237 1.24E-05 0.021606 0.038085 0.37643 0.56407 1.50E-21 0.56407 0.77584 0.51718 0.77584 0.56407 0.1599 0.77584 0.51718 0.30048 1.50E-21 0.77584 0.47608 0.68592 0.68592 0.68592

Sands, gravels, sandstones, limestones, clays (Litho 10) 0.092843 0.13476 0.62145 0.62145 0.25281 0.37937 0.54324 0.0027932 0.0063157 0.29023 0.051586 0.40644 0.58824 0.46806 0.58824 0.78927 0.54324 0.78927 0.58824 0.35436 0.78927 0.22405 0.33114 0.46806 0.000183 7.68E-21 0.70427 0.70427 0.70427

Schist, Amphibolites, Mica schist, Greywackes, Quartzites, Gneiss (Litho 11) 0.6407 0.07228 0.22298 0.049947 0.80614 0.45051 0.60179 0.042343 0.26009 0.00011327 0.094735 0.47608 0.64213 0.53336 0.64213 0.81855 0.60179 0.81855 0.64213 0.4266 0.81855 0.60179 0.57812 0.53336 0.81855 0.56587 0.74456 0.74456 6.00E-10

Evapotranspiration (mm) 0.44699 0.53514 0.1273 0.66803 0.015868 0.6697 0.31458 0.31808 2.89E-18 1.45E-10 2.16E-16 0.25669 0.57057 0.060808 0.57057 0.77947 0.017311 0.77947 0.55047 9.26E-05 0.13279 0.000454 0.00033 0.060808 0.32345 0.46074 0.017033 0.032775 0.67551

Solar radiation (Kcal/Cm2) 0.26405 0.98659 0.17938 0.19804 0.00023714 0.37914 0.64214 0.066781 6.76E-22 1.64E-06 1.42E-14 0.011403 0.7609 0.00055 0.7609 0.69611 0.013886 0.88059 0.42884 0.052916 0.084071 0.000316 8.16E-06 0.00055 0.14098 0.011774 0.57911 0.15289 0.57911

Mean Temperature (ºC) 0.12516 0.73898 0.014649 0.2537 0.016022 0.818 0.23067 3.01E-09 4.76E-17 0.81923 4.55E-08 6.70E-05 0.18458 0.003891 0.763 0.086918 0.73491 0.43014 0.11 0.064642 0.15299 0.000149 5.66E-07 0.003891 0.43014 0.048329 0.83257 0.83257 0.015049

Hours Sunshine 0.065429 0.98696 0.033012 0.12987 0.005617 0.85515 0.66581 9.21E-06 3.39E-18 0.01411 8.79E-11 1.96E-05 0.10367 0.00916 0.90579 0.32061 0.40621 0.42178 0.12391 0.008487 0.12875 0.001873 2.40E-07 0.00916 0.21986 0.05733 0.28074 0.93387 0.72436

Rainfall (mm) 0.01555 0.23863 0.067423 0.30908 0.0010978 0.94646 0.51132 1.91E-05 1.20E-14 0.012808 1.24E-09 5.63E-06 0.082645 0.30326 0.83618 0.26772 0.13551 0.91869 0.002585 0.000198 0.13689 0.000472 0.000835 0.30326 0.70457 0.02149 0.034642 0.19196 0.66361


86

Título

Nr. days frost Nr. days rainfall Litho 1 Litho 2 Litho 3 Litho 4 Litho 5 Litho 6 Litho 7 Litho 8 Litho 9 Litho 10 Litho 11 Evapotranspiration Solar radiation Mean Temperature Hours Sunshine Rainfall (mm)

Number of vessels 0.53654 0.5768 0.095836 0.15045 0.72847 0.68871 0.91951 0.27894 0.62148 0.51206 0.1457 0.081614 0.62148 0.41126 0.026561 0.48585 0.26913 0.73348

Vessel grouping 0.027339 0.67334 0.70844 0.00893 0.17461 0.25541 0.61708 0.61708 0.21082 0.56039 0.30048 0.14491 0.8348 0.35181 0.7098 0.28811 0.47133 0.49268

Maximum number of vessels per cluster 0.075777 0.11039 0.099206 0.041423 0.25506 0.84876 0.51973 0.67508 0.13559 0.93054 0.49848 0.28985 0.91051 0.050117 0.71764 0.83471 0.88941 0.94525

Radial Cluster orientation 0.013559 0.91471 0.57404 0.82066 0.10223 0.96159 0.6263 0.6263 0.22298 0.395 0.52496 0.66569 0.87371 0.12816 0.36888 0.068437 0.15753 0.21518

Tangential Cluster orientation 0.018577 0.67617 0.38817 0.29899 0.73636 0.96159 0.6263 0.6263 0.87371 0.52955 0.80945 0.62145 0.87371 0.90787 0.57173 0.052083 0.15361 0.50056

No Cluster orientation 0.059831 0.55518 0.9496 0.14069 0.94773 0.39763 0.23672 0.23672 0.034421 0.76779 0.59955 0.21794 0.67228 0.5812 0.57015 0.44379 0.5662 0.31327

Total number of clusters 0.23271 0.86791 0.62545 0.028875 0.2308 0.75145 0.42502 0.47726 0.082511 0.51006 0.38101 0.26021 0.18925 0.55819 0.52767 0.40382 0.30861 0.36664

Total vessel area (m2) 0.11576 0.97612 0.90547 0.027567 0.59482 0.002933 0.2309 0.34813 0.27849 0.29346 0.053861 1 0.31759 0.6476 0.73105 0.30415 0.34432 0.2

Area of smallest vessel (m2) 0.029877 0.20916 0.53832 0.003203 0.50883 0.008789 0.10286 0.64414 0.4338 0.055325 0.26019 0.090284 0.80036 0.28308 0.76943 0.87211 0.99387 0.62834

area of largest vessel (m2) 0.061442 0.28825 0.89264 0.005701 0.344 0.009852 0.46158 0.73987 0.21931 0.087944 0.28857 0.33831 0.80035 0.15871 0.68001 0.85112 0.93093 0.78822

Average vessel area (m2) 0.094814 0.69995 0.35031 0.000481 0.68269 0.004102 0.21975 0.69668 0.60454 0.017359 0.10114 0.16482 0.51545 0.91332 0.17619 0.071363 0.04581 0.16998

Average vessel aspect ratio 0.40469 0.058458 0.3559 0.92267 0.18819 0.63161 0.78385 0.76175 1 0.013142 0.14985 0.28786 0.36002 0.17057 0.33747 0.090767 0.049604 0.083818

Height of tallest ray (m) 0.12191 0.99136 0.66189 0.43107 0.4564 0.055968 0.19881 0.89662 1.15E-01 0.66812 0.23369 0.75224 0.0018878 0.96227 0.99286 0.941 0.88653 0.85015

Height of shortest ray (m) 0.67751 0.0057414 0.020105 0.55301 0.91526 0.64567 0.10277 0.82854 5.81E-03 0.46453 0.11191 0.12186 0.28373 0.0072829 0.020215 0.13253 0.087608 0.16921

Average ray height (m) 0.01978 0.0006962 0.0040074 0.11123 0.28989 0.99476 0.12239 0.59321 3.61E-02 0.71774 0.006816 0.24278 0.40596 0.00093329 0.0022536 0.0067412 0.0023176 0.0044574

No. of rays/mm2 0.039704 0.10213 0.072013 0.68171 0.029689 0.070697 0.096626 0.13296 0.36605 0.48505 0.069523 0.84967 0.8002 0.013307 0.0072501 0.07681 0.04638 0.16604

Width of widest ray (cells) 0.15955 0.03109 0.13989 0.4192 0.2181 0.01837 0.98631 0.10299 0.0040028 0.35267 0.54176 0.083159 9.67E-02 0.11322 0.094882 0.52279 0.15236 0.43611

Width of narrowest ray (cells) 0.83779 0.24631 0.43045 0.66449 0.80344 0.83483 0.92696 0.92696 0.81855 0.87278 0.77584 0.00018281 0.81855 0.76819 0.31419 0.43014 0.44754 0.35823

Height of tallest ray (cells) 0.2013 0.11589 0.30196 0.027882 0.83569 0.818 0.46992 0.44369 2.19E-01 0.53902 0.73072 0.78418 0.00070455 0.12079 0.16966 0.018982 0.025217 0.067544

Height of shortest ray (cells) 0.75747 0.53544 0.43806 0.37087 0.52294 0.11249 0.16378 0.78754 0.033721 0.087495 0.90539 0.84854 0.11698 0.55092 0.72673 0.84373 0.23463 0.43484

Presence of two distinct ray sizes 0.61698 0.55284 0.3561 0.59338 0.64907 0.23159 0.057247 0.59894 0.085219 0.35826 0.4261 0.81351 0.18811 0.4313 0.85534 0.4821 0.52969 0.84186

Scalariform Vessels 0.46348 0.024162 0.010191 0.37962 0.61414 0.67274 0.85268 0.85268 0.64213 0.74566 0.56407 0.58824 0.64213 0.0009207 0.028454 0.01867 0.024068 0.02476

Homogeneous Rays 0.69146 0.0073401 0.00677 0.20074 0.15331 0.23159 0.59894 0.59894 0.76133 0.35826 2.33E-05 0.00016044 0.18811 0.012857 3.23E-05 0.0011597 0.00053084 0.0067387

Heterogeneous I Rays 0.65909 0.95179 0.99491 0.2129 0.16637 0.067979 0.42233 0.42233 0.24867 0.16103 0.014008 0.003891 0.24867 0.81767 0.21418 0.33885 0.28261 0.6211

Heterogeneous II Rays 0.31873 0.0046668 0.0034131 0.83133 0.81834 0.29482 0.64511 0.64511 0.24906 0.4211 0.15254 0.70124 0.95623 0.0027586 0.0036013 0.021816 0.017491 0.021294

Presence of crystals 0.54764 0.052889 0.36592 0.53456 0.86514 0.075817 0.55507 0.090191 0.019563 0.30258 0.066767 0.085312 0.13967 0.27187 0.16221 0.1995 0.045936 0.097585

Presence of vessel thickenings 0.37819 0.94383 0.11024 0.61099 0.61414 0.034692 0.85268 0.85268 0.64213 0.74566 0.56407 0.58824 2.88E-05 0.78818 0.81947 0.53647 0.90579 0.70095

Plant height (m) 0.02059 0.26822 0.18859 0.62826 0.15965 0.014928 0.1398 8.19E-02 0.039301 0.09028 0.86379 0.56503 0.23442 0.059393 0.31097 0.97664 0.46279 0.62905

Abundant soil 2.38E-05 0.021023 0.026371 0.65265 0.76384 0.078084 0.19671 0.43858 0.052559 0.17604 0.32696 0.0026031 0.052559 0.0032944 0.080313 0.25879 0.11863 0.27337

Sparse soil 0.045662 0.77769 0.82907 0.19568 0.26285 0.1891 0.083265 0.5637 0.14849 0.31321 0.84206 0.00076128 0.14849 0.4503 0.67029 0.664 0.74748 0.88075

Residual soil 0.0004058 0.0018565 1.12E-04 0.34528 0.32378 0.40844 0.71629 0.71629 0.36308 0.52554 0.25895 0.93967 0.36308 0.0008548 0.0015289 0.18009 0.090711 0.1393

High slope 0.70968 0.10835 0.2715 0.13377 0.25405 0.004149 0.67442 0.67442 0.29311 0.46307 0.19193 0.83826 0.29311 0.1486 0.071655 0.070061 0.15984 0.16497

Moderate slope 0.73665 0.2517 0.22693 0.39872 0.13417 0.37008 0.58117 0.58117 0.16739 0.33521 0.12104 0.0069378 0.16739 0.63019 0.16701 0.10274 0.21036 0.41331

Negligible slope 0.51162 0.86286 0.69696 0.77577 0.029364 0.004586 0.42233 0.42233 0.044633 0.16103 0.73698 0.032333 0.044633 0.48916 0.88213 0.81371 0.92631 0.77343

Full sun 0.026764 0.43511 0.76572 0.25608 0.14253 0.006181 0.22869 0.40612 0.21852 0.35858 0.63552 0.092843 0.6407 0.44699 0.26405 0.12516 0.065429 0.01555

Partial Shade 0.69932 0.90087 0.2343 0.22983 0.16368 9.25E-06 0.051253 0.60796 0.1992 0.04868 0.45711 0.13476 0.07228 0.53514 0.98659 0.73898 0.98696 0.23863

Full Shade 0.013056 0.39155 0.29968 0.005828 0.73636 0.26803 0.6263 0.6263 0.87371 0.395 0.80945 0.62145 0.22298 0.1273 0.17938 0.014649 0.033012 0.067423

N Direction 0.18 0.50667 0.9306 0.094669 0.73636 0.26803 0.040012 0.6263 0.22298 0.395 0.80945 0.62145 0.049947 0.66803 0.19804 0.2537 0.12987 0.30908

W Direction 0.077139 1.21E-04 0.11486 0.26133 0.2869 6.66E-09 0.69489 0.69489 0.32629 0.49325 0.22309 0.25281 0.80614 0.015868 0.00023714 0.016022 0.005617 0.0010978

S Direction 0.2604 0.91497 0.57293 0.70735 0.000695 0.49282 0.76302 0.76302 0.45051 0.59842 0.11854 0.37937 0.45051 0.6697 0.37914 0.818 0.85515 0.94646

E Direction 0.7596 0.27263 0.39106 0.005923 0.5713 0.63529 0.83483 0.83483 0.60179 0.010784 0.51718 0.54324 0.60179 0.31458 0.64214 0.23067 0.66581 0.51132

Altitude 0.00032239 0.02913 0.010549 0.12461 0.74525 0.00044 0.41037 0.87377 0.94237 0.85982 0.008124 0.0027932 0.042343 0.31808 0.066781 3.01E-09 9.21E-06 1.91E-05

x(dec º N) 0.0052219 5.20E-19 2.78E-16 0.21614 0.20564 0.070868 0.76173 0.29188 0.01104 0.036318 1.24E-05 0.0063157 0.26009 2.89E-18 6.76E-22 4.76E-17 3.39E-18 1.20E-14

y (dec º W) 2.71E-12 7.77E-06 1.22E-03 0.004069 0.15328 0.006414 0.9195 0.29189 0.00084895 0.091071 0.021606 0.29023 1.13E-04 1.45E-10 1.64E-06 0.81923 0.01411 0.012808

Humic cambisol (Soil 1) 2.35E-05 1.16E-13 2.51E-20 0.001583 0.069866 0.12884 0.50434 0.50434 0.094735 0.2435 0.038085 0.051586 0.094735 2.16E-16 1.42E-14 4.55E-08 8.79E-11 1.24E-09

Rankers (Soil 2) 0.21255 0.0022355 8.16E-05 0.1779 0.43946 0.51718 0.77584 0.77584 0.47608 0.61891 0.37643 0.40644 0.47608 0.25669 0.011403 6.70E-05 1.96E-05 5.63E-06

Schist associated cambisol (Soil 3) 0.78409 0.92516 0.11024 3.01E-06 0.61414 0.67274 0.85268 0.85268 0.64213 0.74566 0.56407 0.58824 0.64213 0.57057 0.7609 0.18458 0.10367 0.082645

Eutric lithosol (luvisol associated) (Soil 4) 0.74218 0.15532 0.032302 0.23895 0.49919 0.5713 0.80344 0.80344 0.53336 0.66374 1.50E-21 0.46806 0.53336 0.060808 0.00054995 0.0038905 0.0091603 0.30326

Eutric fluvisol (Soil 5) 0.075521 0.92516 0.11024 0.37962 2.26E-09 0.67274 0.85268 0.85268 0.64213 0.74566 0.56407 0.58824 0.64213 0.57057 0.7609 0.763 0.90579 0.83618

Luvisol associated cambisol (Soil 6) 0.24801 0.26572 0.43045 0.021133 0.80344 0.83483 1.05E-27 0.92696 0.81855 0.87278 0.77584 0.78927 0.81855 0.77947 0.69611 0.086918 0.32061 0.26772

Distric regosol (Soil 7) 0.0085974 0.011354 0.072912 0.32386 0.5713 1.05E-27 0.83483 0.83483 0.60179 0.71569 0.51718 0.54324 0.60179 0.017311 0.013886 0.73491 0.40621 0.13551

Calcic luvisol (Soil 8) 0.61906 0.96301 0.43045 0.66449 0.80344 0.83483 0.92696 0.92696 0.81855 0.87278 0.77584 0.78927 0.81855 0.77947 0.88059 0.43014 0.42178 0.91869

Gleyic solonchaks (Soil 9) 0.2572 0.01885 0.11024 0.37962 2.26E-09 0.67274 0.85268 0.85268 0.64213 0.74566 0.56407 0.58824 0.64213 0.55047 0.42884 0.11 0.12391 0.0025851

Orthic luvisol (Soil 10) 0.0022908 0.0030195 0.0062904 3.75E-10 0.3884 0.46993 0.75073 0.75073 0.4266 0.57895 0.1599 0.35436 0.4266 9.26E-05 0.052916 0.064642 0.0084873 0.00019815

Ferric luvisol (Soil 11) 0.83779 0.24631 0.43045 0.66449 0.80344 0.83483 0.92696 0.92696 0.81855 0.87278 0.77584 0.78927 0.81855 0.13279 0.084071 0.15299 0.12875 0.13689

Albic Luvisol (Soil 12) 0.45223 0.0083623 0.072912 0.32386 0.5713 0.63529 0.83483 0.83483 4.98E-15 0.71569 0.51718 0.22405 0.60179 0.00045427 3.16E-04 1.49E-04 0.0018728 0.00047184

Chromic luvisol (Soil 13) 0.53503 2.49E-05 0.0041449 0.11476 0.36545 0.44833 0.73888 0.0026998 0.40416 1.60E-07 0.30048 0.33114 0.57812 0.00033017 8.16E-06 5.66E-07 2.40E-07 0.00083549

Eutric lithosol (Soil 14) 0.74218 0.15532 0.032302 0.23895 0.49919 0.5713 0.80344 0.80344 0.53336 0.66374 1.50E-21 0.46806 0.53336 0.060808 0.00054995 0.0038905 0.0091603 0.30326

Plinthic luvisol (Soil 15) 0.83779 0.24631 0.43045 0.66449 0.80344 0.83483 0.92696 0.92696 0.81855 0.87278 0.77584 0.00018281 0.81855 0.32345 0.14098 0.43014 0.21986 0.70457

Orthic podzol (Soil 16) 0.36025 0.0037148 0.048479 0.27772 0.24736 0.60179 0.81855 0.81855 0.56587 0.68861 0.47608 7.68E-21 0.56587 0.46074 0.011774 0.048329 0.05733 0.02149

Schist and quartzite associated cambisol (Soil 17) 0.024905 0.94749 0.26283 0.53791 0.72373 0.76713 0.89642 0.89642 6.00E-10 0.8201 0.68592 0.70427 0.74456 0.017033 0.57911 0.83257 0.28074 0.034642

Dystric cambisol (Soil 18) 0.13758 0.099661 0.26283 0.001059 0.72373 0.76713 0.89642 0.89642 0.74456 0.8201 0.68592 0.70427 0.74456 0.032775 0.15289 0.83257 0.93387 0.19196

Chromic cambisol (Soil 19) 0.072435 0.94749 0.26283 0.53791 0.72373 0.76713 0.89642 0.89642 0.74456 0.8201 0.68592 0.70427 6.00E-10 0.67551 0.57911 0.015049 0.72436 0.66361

nº days frost xx 6.04E-07 0.00083463 0.002117 0.67056 0.008597 0.24801 0.83779 0.097115 0.11327 0.66427 0.67376 0.0015808 5.93E-08 5.80E-05 0.89179 0.01951 0.0052512

nº days rainfall 6.04E-07 xx 3.17E-18 0.031718 0.063931 0.011354 0.26572 0.24631 0.0485 0.042853 0.038953 0.00072473 0.0485 2.77E-20 2.35E-21 4.88E-17 1.83E-20 8.64E-20

Granites (Litho 1) 0.00083463 3.17E-18 xx 0.000191 0.032302 0.072912 0.43045 0.43045 0.048479 0.16844 0.014324 0.021525 0.048479 3.92E-17 3.99E-18 1.93E-13 2.37E-17 1.89E-16

Schist and Greywacke (Litho 2) 0.0021168 0.031718 0.00019119 xx 0.23895 0.32386 0.021133 0.66449 0.27772 0.44867 0.1779 0.20604 0.27772 0.0012887 0.094496 0.98521 0.53397 0.058974

Alluvium (Litho 3) 0.67056 0.063931 0.032302 0.23895 xx 0.5713 0.80344 0.80344 0.53336 0.66374 0.43946 0.40694 0.53336 0.8954 0.50282 0.29334 0.21263 0.027385

Aeolian sands (Litho 4) 0.0085974 0.011354 0.072912 0.32386 0.5713 xx 0.83483 0.83483 0.60179 0.71569 0.51718 0.54324 0.60179 0.017311 0.013886 0.73491 0.40621 0.13551

Conglomerate,Sandstone,Limestone,Marlstone (Litho 5) 0.24801 0.26572 0.43045 0.021133 0.80344 0.83483 xx 0.92696 0.81855 0.87278 0.77584 0.78927 0.81855 0.77947 0.69611 0.086918 0.32061 0.26772

Metavulcanite (Litho 6) 0.83779 0.24631 0.43045 0.66449 0.80344 0.83483 0.92696 xx 0.81855 0.87278 0.77584 0.78927 0.81855 0.32345 0.31419 0.15299 0.12875 0.35823

Sands, pebbles, clays, weakly consolidated sandstone (Litho 7) 0.097115 0.0485 0.048479 0.27772 0.53336 0.60179 0.81855 0.81855 xx 0.68861 0.47608 0.50359 0.56587 2.62E-05 1.34E-03 0.005837 0.001535 4.84E-05

Carbonate rocks (Litho 8) 0.11327 0.042853 0.16844 0.44867 0.66374 0.71569 0.87278 0.87278 0.68861 xx 0.61891 0.64061 0.68861 0.084568 0.010124 0.012551 0.0079678 0.10853

Shale, Graywacke, Sandstone (Litho 9) 0.66427 0.038953 0.014324 0.1779 0.43946 0.51718 0.77584 0.77584 0.47608 0.61891 xx 0.40644 0.47608 0.018779 0.00082548 0.0018492 0.0044667 0.12109

Sands, gravels, sandstones, limestones, clays (Litho 10) 0.67376 0.00072473 0.021525 0.20604 0.40694 0.54324 0.78927 0.78927 0.50359 0.64061 0.40644 xx 0.50359 0.17251 0.0019088 0.011294 0.017094 0.013067

Schist, Amphibolites, Mica schist, Greywackes, Quartzites, Gneiss (Litho 11) 0.0015808 0.0485 0.048479 0.27772 0.53336 0.60179 0.81855 0.81855 0.56587 0.68861 0.47608 0.50359 xx 0.058322 0.32834 0.1987 0.51056 0.79836

Evapotranspiration (mm) 5.93E-08 2.77E-20 3.92E-17 0.001289 0.8954 0.017311 0.77947 0.32345 2.62E-05 0.084568 0.018779 0.17251 0.058322 xx 2.26E-22 1.98E-13 1.02E-18 2.73E-19

Solar radiation (Kcal/Cm2) 5.80E-05 2.35E-21 3.99E-18 0.094496 0.50282 0.013886 0.69611 0.31419 0.0013363 0.010124 0.000825 0.0019088 0.32834 2.26E-22 xx 1.76E-16 7.61E-22 2.34E-19

Mean Temperature (ºC) 0.89179 4.88E-17 1.93E-13 0.98521 0.29334 0.73491 0.086918 0.15299 0.005837 0.012551 0.001849 0.011294 0.1987 1.98E-13 1.76E-16 xx 2.47E-22 9.49E-20

Hours Sunshine 0.01951 1.83E-20 2.37E-17 0.53397 0.21263 0.40621 0.32061 0.12875 0.001535 0.007968 0.004467 0.017094 0.51056 1.02E-18 7.61E-22 2.47E-22 xx 4.33E-24

Rainfall (mm) 0.0052512 8.64E-20 1.89E-16 0.058974 0.027385 0.13551 0.26772 0.35823 4.84E-05 0.10853 0.12109 0.013067 0.79836 2.73E-19 2.34E-19 9.49E-20 4.33E-24 xx

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