Analysis of the biological traits of diatoms in intermittent rivers ...

Analysis of the biological traits of diatoms in intermittent rivers

Alba Alemany Sancho

Degree of Biology

Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals de la Universitat de

Barcelona

Tutors: Núria Bonada & Joan Gomà

Presentation date: 30/06/2020

Abstract

Intermittent rivers predominate in Mediterranean areas; under these conditions, drying or no-

flowing phases are common. Aquatic taxa have resistance and resilience strategies to

withstand or recover after the drying period. In this research, we set the grounds for a rewetting

experiment that analyses the reestablishment of diatoms eliminating the effect of algal

recolonisation by drift. Furthermore, we also examined several diatom biological traits that may

potentially be beneficial as the degree of intermittency increases; the main analysed traits

were: “adnate”, “pad”, “colonial”, “ecological guilds”, “mobile”, “oxygen requirements”, “cell

size”, “pioneer” and “moisture tolerance”. These coded traits were compared amongst

ephemeral, intermittent and perennial rivers. Overall, the comparisons indicated that there

were no statistically significant differences neither in the composition of species nor in

biological traits. However, a higher variability of several of these metrics was observed for both

perennial and also intermittent rivers. Notwithstanding, some constant traits were detected in

intermittent rivers, indicating that lack of water flow may select pedunculated, mobile, non-

colonial, no-adnate, no-pad taxa. Further physiological traits, such as mucilage secretion and

other possible biofilm resistance mechanisms, should be analysed to predict the impact of

climate change scenarios on diatom communities in mediterranean rivers.

Sustainable development goals

Considering future climate change scenarios for freshwater ecosystems, the study of diatom

communities in intermittent rivers can be particularly significant in a long-term scale because

changes in flow patterns can have effects on diatom biodiversity. This project aims to shed

light into diatoms biological traits that may enable them to withstand or recover from the dry

period in intermittent rivers. Most of the research done on freshwater biodiversity and

intermittent rivers, has been done using aquatic macroinvertebrates, whereas diatoms have

been largely neglected. The results obtained from this project could be used for predicting

future diatom biodiversity patterns and their biological traits in Mediterranean river ecosystems.

Our results indicate that there are no significant differences between intermittent and perennial

rivers from Catalonia, nevertheless more research should be conducted to relate further

biological traits that may be suitable especially under climate change conditions.

Therefore, the sustainable development goals addressed in this study are included in

the issue of Planet. This impacts on Goal 13 “Take urgent action to combat climate change

and its impacts”, especially regarding 13.3: “Improve education, awareness-raising and human

and institutional capacity on climate change mitigation, adaptation, impact reduction and early

warning”. Furthermore, it affects Goal 15 “Protect, restore and promote sustainable use of

terrestrial ecosystems, sustainably manage forests, combat desertification, and halt and

reverse land degradation and halt biodiversity loss”, focusing on 15.1 “By 2020, ensure the

conservation, restoration and sustainable use of terrestrial and inland freshwater ecosystems

and their services, in particular forests, wetlands, mountains and drylands, in line with

obligations under international agreements”.

Adaptation of the TFG to the COVID-19 confinement

This project began in September 2019, starting with the setting of the rewetting experiment

and the laboratory procedures in order to obtain all diatom samples. Subsequently, microscope

observations, iconographies and identifications from all the samples were conducted, as well

as a photographic herbarium of the taxa found. The next step was to quantify each species on

each sample, nevertheless, that could not be accomplished due to the start of lockdown

measures in March 2020. Consequently, this project was modified by compiling information

from potential biological traits of diatom species found in the routine biomonitoring program of

Catalonian rivers from the Catalan Water Agency (“Agència Catalana de l’Aigua”) and

comparing the different river types.

INDEX

1. INTRODUCTION ....................................................................................................................................... 1

1.1 A brief introduction on Diatoms.................................................................................2

1.2 Diatoms biological traits to intermittence .................................................................4

2. HYPOTHESES ........................................................................................................................................... 7

3. OBJECTIVES............................................................................................................................................. 8

4. MATERIAL AND METHODS .................................................................................................................... 9

4.1 Methods for Objective 1 .............................................................................................9

The study area ..................................................................................................................9

Sampling ...........................................................................................................................9

Experimental design .......................................................................................................10

Laboratory protocol ........................................................................................................10

4.2 Methods for Objective 2 ...........................................................................................11

4.3 Methods for Objective 3 ...........................................................................................11

5. RESULTS ................................................................................................................................................. 12

5.1 Results for Objective 1 .............................................................................................12



6. DISCUSSION ........................................................................................................................................... 18

7. CONCLUSION ......................................................................................................................................... 20

8. BIBLIOGRAPHY ..................................................................................................................................... 21

1

1. INTRODUCTION

The mediterranean climate regions are located between temperate and desert climate regions

in most of the continents (Dallman,1998), and usually extending between 30° and 40° latitude.

The regions include areas around the Mediterranean Basin, coastal California, central Chile,

the Cape area in South Africa and the south and southwestern Australia (Aschmann, 1973).

These regions are characterised by summer droughts and moderate rainy winters (Bonada &

Resh, 2013) and are considered biodiversity hotspots with high levels of endemisms (Myers

et al., 2000; Smith & Darwall, 2006; Bonada et al., 2007). This high biodiversity level is the

result of combined causes: landscape heterogeneity (Koniak & Noy-Meir, 2009), high seasonal

and predictable precipitation and hydrological conditions (Bonada et al., 2007) and historical

effects (e.g. the use of these areas as refuges during the Pleistocene glaciations in the northern

hemisphere; Hewitt, 2004).

A large proportion of rivers in mediterranean climate regions are intermittent (also

known as temporary), which are defined as rivers that lack of superficial flow at some period

of time (Bonada & Resh, 2013). Ephemeral rivers are also frequent in these regions and are

characterised by brief water flow periods (just after rainfalls) followed by long dry periods

(Hancock, 2017). Both river types are characterised by having large variations on hydrological

connectivity as they alternate between an expansion phase (lotic or wet period) to a contraction

phase (lentic or dry period) (Sabater, 2008; Bernal et al., 2013). As a result, they are shifting

habitat mosaics between fully aquatic and fully terrestrial habitats (Datry et al., 2016). On the

contrary, perennial rivers are characterised for having a constant and continuous water flow,

most of them constantly connected to groundwater (Meinzer, 1923). Despite their high

occurrence in mediterranean climate regions, many of them are becoming intermittent due to

global change (Larned et al., 2010). Flow intermittence is, however, not a unique feature of

mediterranean climate regions. Intermittent rivers are present on every continent and account

for more than 50% of the global river network (Tooth, 2000; Datry et al, 2014). Most of the

global river network is, therefore, intermittent rather than perennial.

Intermittent (and ephemeral) rivers are considered natural disturbed ecosystems due

to the extreme flow variability along the year. As a result, organisms found in these rivers have

specific resistance and resilience strategies to withstand or recover from flow intermittence

(Bogan et al, 2017). Despite following these strategies, the full recovery of the pre-drought

community is unlikely to occur and generally it has a different species composition (Lawrence

et al., 2010). Resistance refers to the capacity of an ecological unit (e.g. a taxon) to endure the

harsh conditions in situ (Lake, 2000), summer drying in our case. It can include strategies

2

being able to tolerate dry river beds, such as desiccation-tolerant stages (summer diapause)

(Williams, 2006), or the ability to persist in a local perennial refuge like remnant pools (Boulton

et al, 1992). On the contrary, resilience refers to the capacity of an ecological unit that rebounds

after disturbance (Nimmo et al, 2015), and should include species with traits that allow them

to colonize passively by drift (i.e. the movement with flow) or by aerial dispersion, or actively

by means of their own motile structures (for example to the hyporheic zone; Del Rosario and

Resh, 2000). In general, the resilience strategy is meant to be more frequent than resistance

in Mediterranean rivers (Fox & Fox, 1986).

1.1 A brief introduction on Diatoms

Diatoms or Bacillariophyceae are unicellular or colonial microscopic algae characterised for

having chloroplasts with four membranes, thylakoids in stacks of three, fucoxanthin as a

pigment, and chrysolaminarin as a reserve product (Bhattacharya et al., 1992). The most

recognisable and unique feature is a cell wall composed of silica, constituting the frustule

(Pickett-Heaps et al., 1990). There is a high heterogeneity on frustule morphology, being the

most diverse protists, with estimations of 100000 taxa (Mann & Drop,1996). They can be found

on all kinds of moist habitats with access to light, from freshwater ecosystems to oceans, and

even in soils (Falkowski, et al, 1998). Generally, they are planktonic and live in suspension or

attached to a substrate, but can also be found forming filaments or colonial forms. Our study

focuses on benthic taxa, which mainly includes the periphyton attached on stones or sediment

surfaces in the river beds.

The frustule structure looks like an exoskeleton

made of hydrated silica (SiO2) and surrounded by

various organic compounds (Volcani, 1981) (Figure 1).

It consists of two halves, the epitheca and the

hypotheca; fitting inside one into the other, as the first

covers the second one (Round et al., 1990). The

frustule plays a role in gas exchange with the

environment, osmotic regulation, and mucilage release to

move or to adhere onto the substrate.

The thecae are made up of the valve, a more or less flattened plate, and the pleura on

the sides and at least one cingulum, which consists of siliceous linking bands between the

valve and the pleura; each band is called a copula or girdle band and is released after the

vegetative cell division that produced the valve. The valves can have various ornamentations

on their surface (Kroger et al., 1996). They have pores useful for nutrients intake and releasing

waste (Round et al., 1990).

Figure 1. Diagram showing the general

features of pennate diatoms (Taylor et al,

2006)

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Figure 2. Schematic representation of typical shapes of centric (A) and pennate (B) diatoms (Sorvari, 2001)

Considering their symmetry, they can be classified as centric (radial symmetry) or

pennate diatoms (bilateral symmetry) (Schmid et al, 1981; Mann, 1986) (Figure 2). The centric

ones do not have raphe and are typically planktonic, whereas pennate can have it and are

generally benthic or epipelic (living at the interface of water and sediment) (Round et al., 1990).

Consequently, the ability to move (in a forward and backward movement) is exclusive of the

pennate diatoms that have a raphe. This movement is useful to avoid sedimentation in benthic

diatoms and also facilitates sexual reproduction between genetically different individuals.

The life cycle of diatoms is diplontic, and includes a long vegetative stage during which

the cells divide mitotically, and a short stage in which sexual reproduction occurs, followed by

a complex developmental process leading to the formation of new vegetative cells

(auxospores) (Mann, 1993). During mitosis, the two thecae become the daughter’s epitheca,

so each of them will have to generate the hypotheca before separation (cytokinesis). As a

result, there is a gradual cell size reduction on the daughter cell that uses the mother hypotheca

as the epitheca, whereas the other one is the same size as the parent cell (MacDonald, 1969;

Pfitzer, 1971). Consequently, the auxosporulation process occurs, in order to re-establish the

cell size and also to achieve genetic recombination. The process of sexual reproduction in

centric diatoms is by means of oogamy, where gametes fuse, expand and form the auxospore

that contains new thecae, which eventually form the initial free cell (Von Stosch, 1950). On the

other hand, pennate diatoms carry out isogamy, forming a zygote on each parent cell, which

will become the auxospore (Round et al., 1990; Chepurnov and Mann, 2004). It has also been

studied apomixis, the production of auxospores without previous fecundation (Drebes, 1977).

If cells do not experience a sexual phase or auxosporulation process, they will divide mitotically

until they reach a critical small size threshold and die (Geitler, 1932)

4

Diatoms are used as bioindicators, that means that the presence or absence of a

species or group of them can reflect the quality of that environment. They are used to assess

the ecological quality of freshwater ecosystems, due to their specific ecological requirements

and responses to various biogeographical and physicochemical factors. Water flow, in

particular, is the factor that has a massive impact on all other physiochemical characteristics

(Stevenson, 1996).

1.2 Diatoms biological traits to intermittence

Several biological traits can potentially provide diatoms some ability to withstand changes in

water flow and intermittence, but also could be a solution to severe habitat conditions, even in

perennial streams (Vander Vorste et al., 2016). Biological traits are characters or observable

features that can be inherited or acquired in response to the environment. They include life

history, morphological, physiological and behavioural characteristics that are related to

ecological aspects (Naeem et al, 1999). This is based on the habitat template theory, which

considers the habitat as a templet for natural selection to select individuals/species with traits

adapted to the characteristics of the habitat (Southwood ,1977).

The analyses regarding diatoms traits can be conducted considering life-forms, cell sizes or

ecological guilds (Rimet et al., 2012):

I. Regarding life-forms, diatoms can be solitary cells or can form colonies. The

solitary ones can be not attached (if they are floating or free moving) or attached

to substrates in different ways: adnate (using their valve face), mucilage pad

(releasing mucilage on a pole) or mucilage stalk (the pore cells produce a stalk)

(Round et al., 1990). The colonial forms can be classified in: chain colonies,

ribbon colonies, zig-zag colonies, rosette colonies, star colonies, arbuscular

colonies, mucous tubule colonies. Moreover, each species can show different

life forms on every stage of its life-cycle.

II. The cell size is a property of each species but also varies in response to the

nutrient availability (Finkel et al., 2009) and current velocity (Tuji, 2000).

III. Ecological guilds regroup the varieties of life forms (Table 1) considering taxa

that coexist, but which may have adapted in different ways to abiotic factors

(Devito et al., 2004). The following groups are described:

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Table 1. Taxa assignment to ecological guilds, adapted from Passy (2007) and Rimet (2012)

Ecological guilds Taxa

Low-profile

Characterised by its resistance to

disturbances and not tolerating nutrient

enrichment. It includes species of short size

and slow moving following the r-strategy

(Biggs et al.,1998). This strategy refers to the

ability to reproduce and disperse rapidly and

exponentially, as a result they are successful

while colonising unpredictable changing

environments.

Achnanthidium, Achnanthes,

Amphora, Brachysira, Cymbella,

Cymbopleura, Cocconeis, Delicata,

Diploneis, Discostella, Encyonema,

Encyonopsis, Eucocconeis,

Fragilaria, Karayevia, Kolbesia,

Meridion, Nupela, Planothidium,

Platessa, Rhoicosphenia, Reimeria

High-profile

Sensitive to turbulence, but it is benefited from

nutrient enrichment. It includes large or

colonial species.

Aulacoseira, Achnanthidium

catenatum, Diadesmis, Diatoma,

Eunotia, Encyonema, Fragilaria,

Frustulia Gomphonema,

Gomphoneis, Gomphosphenia,

Melosira, Pleurosira,

Pseudostaurosira, Staurosira,

Staurosirella,

Tabularia, Ulnaria

Motile

Includes fast moving species that have the

ability to select their habitat, in spite of being

sensitive to high current velocities but tolerant

to high nutrients (Passy, 2007).

Adlafia, Bacillaria, Caloneis,

Craticula, Delicata, Denticula,

Eolimna, Epithemia, Fallacia,

Fistulifera, Mastogloia

Geissleria, Gyrosigma, Hippodonta,

Luticola, Mayamaea, Navicula,

Naviculadicta, Nitzschia, Nupela,

Sellaphora, Simonsenia, Stauroneis,

Surirella, Rhopalodia ,Tryblionella

Planktonic

Taxa with morphological adaptations for lentic

habitats and able to resist sedimentation

(Rimet et al, 2012).

Cyclotella, Cyclostephanos,

Nitzschia, Stephanodiscus

6

Diatoms can withstand the dry period through several resistance and resilience

strategies. Diapause can be one of the multiple resistance mechanisms they use to persist in

a local reach during the dry period. This is possible due to the fact that diatoms are part of the

periphyton, described by Sabater (2007), as a microecosystem composed of a complex

mucopolysaccharide matrix with embedded autotrophic and heterotrophic microorganisms that

has the natural ability to respond and recuperate from stress, such as drying. This is believed

to be feasible for regrowth when water starts flowing again, but no studies have tested it so

far. Furthermore, the diapause stage is not related to the auxospore formation; unlike other

algae, the auxospore is neither a resting stage nor a specialised cell for dispersal (Van den

Hoek et al., 1995). Another resistance mechanism could be the ability to remain active in local

remnant pools (Dodds et al.,1996; Robson et al., 2008). Similarly, other refuges such as dry

biofilm on stones, debris, dry sediments, organic matter and moist leaves can maintain a

certain humidity for them to endure the drying periods facilitating the recovery (Webb et al.

2012).

Active or passive movement from refuges or perennial pools to new flowing habitats

are considered resilience mechanisms. These are linked to species with a raphe that enables

them to actively migrate or to species more prone to be passively dispersed by drift, colonising

downstream areas appears such in perennial rivers (Peterson 1996). Other resilience

mechanisms would be related to passive dispersal through wind (Ehrenberg, 1849), waterbirds

(Figuerola et al., 2002) or on moist surfaces of several animals (Stoyneva, 2016). Despite all

these potential resistance and resilience mechanisms, Robson (2008) suggested that diatoms

mainly used dry biofilm on stones (resistance) and drift (resilience) to recolonise after the dry

period in intermittent rivers.

It has also been studied that only a week of dry period has a long-term effect on diatoms

metacommunity, and that it did not returned to the initial state after a 28-day rewetting (Barthès

et al, 2015). In addition, there are recognised differences on taxonomic composition of diatoms

meta-communities considering flow intermittence. Tornés et al. (2013) found that a perennial

flowing stream shows the highest proportion of nestedness in diatom communities of the

Iberian Peninsula, while on the contrary, idiosyncratic taxa and lower species richness seems

to be typical on intermittent rivers. Nestedness occurs when the taxa of a community with lower

numbers of species tends to be subsets of the taxa at richer sites (Wright et al, 1992); following

different patterns consisting of connecting local communities by dispersal of multiple

interacting species (Leibold et al., 2004). Idiosyncratic taxa do not follow this pattern, they are

characterised for being present in unexpected poor species sites or absent in species-rich sites

(Atmar & Patterson, 1993). Diatoms have efficient passive dispersal strategies (mainly drift)

7

which are meant to lower nested patterns; hence, unexpected presences and absences would

be common (Soininen, 2008; Ruhí et al., 2013).

2. HYPOTHESES

Following the biological traits of aquatic invertebrates analysed on “The biota of intermittent

rivers and ephemeral streams” by Stubbington et al (2017), we built up a table regarding the

possible diatom traits to endure the dry period in intermittent rivers, considering both resistance

and resilience traits. Contrary, perennial rivers may be characterised by the absence or less

presence of these studied traits. This table constitutes a summary of the hypotheses of this

study. Besides the listed traits, some others could be also considered, such as pH, salinity,

nitrogen uptake metabolism, trophic state, or the chlorophyll-a levels (Van Dam, 1994; Falasco

et al., 2016). However, more autoecological information is needed about all these traits and

therefore, we focused on more simple traits mostly related to morphology (Table 2).

Table 2. Potential diatom biological traits that may promote resistance and/or resilience strategies to intermittence.

Resistance traits Rationale

Large cell size Large diatoms tend to be epipelic, at the interface of sediments and

water (Round et al.,1990), therefore having more potential abilities to

survive in dry sediments.

Adnate The adhesion to the substrate by their valve face or girdle view,

promotes a moist microhabitat in which they may withstand the drying

period.

Pad The production of mucilage on a pole that sticks to substrate may be

useful to stay hydrated.

Colonial Promotes the adhesion to the substrate by creating a moist microhabitat

useful to withstand the drying period.

Planktonic guild

Taxa included in this guild are adapted to lentic environments, having

morphological adaptations that enable them resisting to sedimentation.

(Passy, 2007)

Low Oxygen

requirements

May be able to endure the anoxic conditions in remnant pools with

accumulated organic matter.

Mobile (raphe) They can move on the stone surface, remaining in contact with the

water, as the upper part of the river dries faster.

Motile guild

May be able to select their habitat and locally migrate to parts with

moisture under the stone.

8

Resilience traits Rationale

Small cell size Taxa with a fast-growing rate, may be competitive in recolonising.

Non-Colonial Free moving individual cells may be able to disperse easily.

Pioneer Able to endure harsh drying conditions, these species are the first to

colonise the disrupted environment. (Passy, 2007)

Low profile guild

Enhances resistance to disturbances, by reproducing and dispersing

exponentially, following the r strategy. (Passy, 2007)

Moisture

tolerance

May be able to survive on wet and moist or temporarily dry places. (Van

Daam, 1994)

3. OBJECTIVES

The main aim of this research is to assess the responses of flow intermittence on diatoms by

setting up and experiment and by compiling and analysing diatom community data in perennial

and intermittent rivers. The specific objectives are as follows:

OBJECTIVE 1: To assess the temporal dispersal ability of diatom species (resistance

strategies) in intermittent rivers by setting a rewetting experiment that avoids the

recolonization by resilience strategies.

OBJECTIVE 2: To create a list of biological resistance/resilience traits of diatom species

collected in several perennial and intermittent mediterranean rivers in Catalonia.

OBJECTIVE 3: To compare trait composition of diatom species in perennial and

intermittent mediterranean rivers in Catalonia.

9

4. MATERIAL AND METHODS

4.1 Methods for Objective 1

The study area

Three rivers from Sant Llorenç del Munt i l'Obac Natural Park, located in Vallés Occidental

county in central Catalonia, were selected: Sanana, Talamanca and Santa Creu1. They are

very close to each other but differ in the degree of flow intermittence, with Sanana being the

most permanent, Talamanca having intermediate values of flow intermittence, and Santa Creu

being the most ephemeral.

The area has a typical mediterranean climate, with an average annual rainfall of 682

mm, mainly falling in autumn, followed by spring and summer being the driest. We obtained

the following temperature data from sensors (HOBOs) placed several sites along the three

rivers (Figure 4); the mean temperature in all three rivers oscillates from 5-9 ºC in winter, while

reaching 20-26 ºC in summer. The year 2019 was extremely dry in winter and the precipitation

rate obtained by sensors oscillated around 0mm.

Figure 3. Mean temperature variations on the three rivers during 2019. Data collected from HOBOs located in

several sites along the three rivers.

Sampling

Field work was carried out during May 2019, as part of the MECODISPER project sampling

campaign. The diatom sampling started with the collection of 3 stones of each river. Then, the

upper surfaces of the stones were scrubbed with a toothbrush and samples were fixed in 4%

formaldehyde. A total of 9 (3 from each chosen river) samples of diatoms were collected.

1 Maps of the studied rivers and MECODISPER campaign sampling points are on the Appendix 1.

0,00

10,00

20,00

30,00

Jan Feb Mar Apr May June July Aug Sept Oct Nov Dec

Mea

n T

emp

erat

ure

(ºC

)

2019

Temperature in rivers 2019

Talamanca Sanana & Santacreu

10

Experimental design

Nine calcareous stones of different sizes were placed on each river on 28 th may 2019 to allow

the colonisation by diatoms. On 9-12 July 2019, stones and encrusted substrate were

extracted and left to dry out during the whole summer at the terrace of the Section of Ecology

at the University of Barcelona, in order to simulate the drying period in natural conditions.

Experimentally, all stones experienced drought for the same period of time, regardless of

belonging to rivers with a different degree of intermittency. Encrusted algae from bedrock was

also collected to ensure that colonisation of stones was completed. On 26th September 2019,

just after the summer period, the experimental rehydration of the substrates started. Firstly,

the substrates were placed in plastic tuppers and treated water without chlorine was added to

simulate the rewetting. A hole was made on each tupper lid to insert aerators, which would

provide oxygen into the water. The lid helped to isolate each sample from the outside, to

minimise the risk of contamination by propagules from the air. A total of 27 tuppers with one

stone each and 27 more with a sample of encrusted substrate (9+9 samples per river) were

set up and randomly placed at the terrace’s shelves, bearing in mind that there might be some

variations in weather conditions and light according to their location.

The growing in the community was monitored over time on both substrates. Three

stones and 3 pieces of encrusted substrate from each river were randomly extracted at three

different times. Nevertheless, the experimental design was different for each type of substrate,

due to the fact that different stones were taken on each extraction, whereas for encrusted

substrate the extractions were always collected from the same samples; thus, we also aimed

to study the succession of the exact same community over time.

For stones, the three extractions were conducted every 20-25 days, whereas for

encrusted substrates, the first extractions were done after 30 days, and the last one after 25

days. This difference in the extractions was due to the fact that the encrusted substrate seemed

to have more cyanobacteria after the rewetting.

Laboratory protocol

Each sample, stone or encrusted substrate, was scraped using a toothbrush. For the encrusted

substrate sample, the green part was detached from the original substrate using a blade, and

then the obtained periphyton were poured in flasks. Then, it was fixed using 4 % formaldehyde

and the sediments were allowed to settle for 24 hours. Part of each sample was poured on a

digestion tube and 110 volume hydrogen peroxide was added, in order to eliminate the organic

matter. The tubes were let to digest on a sand bath in a hot plate, at 100ºC. After 24h, the

hydrogen peroxide was removed while keeping the sediments, filling with distilled water and

adding drops of hydrochloric acid to acidify and remove calcium carbonate. After 24h, the

supernatant was removed and cleaned with distilled water again. A day later, the sediment

11

material was taken and resuspended on flasks, verifying with an optical microscope that the

density of frustules was the ideal for observation. Using a pipette, the sample was mixed and

a drop was put on a cover glass. Then it was let to dry for a day and after that permanent

samples were mounted using Naphrax resin.

The identification was based on the frustule shape, size and structure, mostly

considering the valve surface ornamentation. The observations were conducted using a Zeiss

microscope and its camera. After having identified the taxa present, all the species on each

sample should have been quantified. The ideal would have been that 400 valves were counted

for each sample using an optic microscope. After that, in order to infer whether the diatoms

found and identified were actually able to regrow from dry substrates, a viability study should

have been conducted, in which the proportion of alive cells (full frustules and chloroplasts)

between dead cells (empty frustules, no chloroplasts) had been determined. This procedure

can still be done at any time, due to the fact that all samples and extractions were first

preserved in formaldehyde, preserving both alive and dead cells.

%𝑣𝑖𝑎𝑏𝑖𝑙𝑖𝑡𝑦 =𝑎𝑙𝑖𝑣𝑒 𝑑𝑖𝑎𝑡𝑜𝑚𝑠

𝑑𝑒𝑎𝑑 𝑑𝑖𝑎𝑡𝑜𝑚𝑠𝑥100


This objective focused on analysing data from the Catalan Water Agency (“Agència Catalana

de l’Aigua”), consisting on diatom inventories from different types of rivers: 6 perennial

(Bruguent, Portella, Merlès, Major, Òsor and Guilla) 6 intermittent (Glorieta, Santa

Creu,Guanta, Avencó, Daró and Orlina) and 1 ephemeral (Riudaura). Samples from 1

ephemeral river were included for comparison with perennial and intermittent ones as an

extreme case of flow intermittence. No more data from other ephemeral rivers was available.

Perennial and intermittent rivers include 3 calcareous and 3 siliceous sites each, as diatoms

are very sensitive to geology and physico-chemical characteristics of the substrate (Cantonati

et al., 2009). These rivers were sampled in 2016-2018. For each taxon identified on each river,

biological traits were obtained from Rimet & Bouchez (2012) and Van Daam (1994) (Appendix

2).


In order to compare trait composition of diatom species in perennial and intermittent (and also

ephemeral) rivers, data from the Catalan Water Agency was divided and built in two different

excel matrixes. The first matrix (A) refers to taxa identified as columns and different rivers as

rows; it contains the abundance of each taxon for each river. The second matrix (B) includes

12

biological traits as columns and species as rows. Therefore, a third matrix (C)2 was obtained.

This matrix included the proportion of each biological trait in each river. Statistical analyses

were done first on the taxonomic matrix (A) by means of richness, Shannon diversity,

Multidimensional Scaling (MDS) and ANOSIM analysis. Richness refers to the total number of

unique species found at a site, whereas Shannon-Weiner diversity index represents species

weighted by abundance. While MDS allows to have a better view of diatoms communities

organization in a multidimensional space, ANOSIM tests for differences in community

composition among groups (perennial, intermittent and ephemeral rivers in our case). Kruskal-

Wallis tests were applied to test for significant differences between taxonomic metrics

(richness and Shannon diversity) and the proportion of each biological trait among the three

river types. All these statistical analyses were carried out in R (R Development Core Team,

2019) and using the packages vegan (v2.5-6; Oksanen & Blanchet, 2019), for richness and

diversity, and ade4 (v1.7-15; Dray & Dufour, 2018) to obtain the C matrix. In addition, a

Principal Components Analysis (PCA) was conducted using the Excel software XLSTAT in our

matrix B, regrouping taxa for each type of river.

5. RESULTS

5.1 Results for Objective 1

The taxa identified in all three studied rivers were the following:3

• Achnantes cf trinoidis

• Achnantidium minutissimum

• Brachisyra liliana

• Brachisyra vitrea

• Cyclostephanos dubius

• Cyclotella meneguiniana

• Cymbella cf compacta

• Cymbella cf laevis

• Cymbella cf lancettula

• Cymbella cymbiformes

• Cymbella excisa

• Cymbopleura amphicephala

2 This C Matrix is on the Appendix 3. 3 For further identification and taxonomic details, consult the photographic herbarium attached on the

Appendix 5.

13

• Denticula tenuis

• Diploneis elliptica

• Encyonema silesiacum

• Encyonopsis cesatii

• Encyonopsis minuta

• Eucocconeis flexella

• Eunotia bidens

• Eunotia denticulata

• Fragilaria cf radians

• Fragilaria dilatata

• Fragilaria ulna

• Gomphonema gracile

• Gomphonema lateripunctatum

• Gomphonema micropus

• Gomphonema parvulis

• Gomphonema vibrio

• Gyrosigma cf attenuatum

• Hantschia abundans

• Mastogloia lacustris

• Mastogloia smithii

• Navicula criptotenella

• Navicula vandamii

• Nitzschia denticula

• Pinnularia cf sinistra

• Pinnularia cf subcapita

• Pinnularia nobilis

• Rhopalodia parallela

• Stephanodiscus minutulus


The coded biological traits for each species are found on a table in the Appendix 2.

Most of the obtained values are coherent to each taxon, although there is a lack of data in

reference to “Oxygen requirements” and “Moisture tolerance” traits.

14


Kruskal-Wallis rank sum test richness by temp1 Kruskal-Wallis chi-squared = 1.4628, df = 2, p-value = 0.4812 Shannon by temp1 Kruskal-Wallis chi-squared = 0.91209, df = 2,

p-value = 0.6338

Species richness was visually higher in intermittent rivers, followed by perennial ones but no

significant differences were found (Figure 3). Moreover, perennial rivers appeared to have a

higher diversity than ephemeral rivers but overall, no significant differences were found. In both

graphics, it is noticeable that intermittent rivers have a higher variability in both taxonomic

metrics.

Figure 4. Dimension reduction via multidimensional scaling (MDS) showing the three types of rivers. I

(Intermittent), E (Ephemeral) and P (Perennial)

There seemed to be a high similarity between communities from the three river types (Figure 4) and no significant differences were observed with the ANOSIM analysis (Table 4).

Figure 3. Boxplot of the distribution of species

richness and diversity in the three river types. For

each type, the horizontal line represents the median

value of the distribution, and the whiskers reach the

highest and lowest value within 95% of the

distribution. (E)= Ephemeral, (I)=Intermittent and

(P)=Perennial.

15

Table 4. ANOSIM for communities and traits between the three types of rivers.

For communities: ANOSIM statistic R: -0.1201 Significance: 0.835

For traits: ANOSIM statistic R: -0.04028 Significance: 0.58

This indicates that there are no differences neither in communities nor in traits between

intermittent, ephemeral and perennial rivers. Moreover, the negative R values in both analyses

suggest more similarity between rivers than within rivers.

16

Figure 5. Boxplots of the distribution of each trait in the three river types. For each type, the horizontal line

represents the median value of the distribution, and the whiskers reach the highest and lowest value within 95% of

the distribution. (E)= Ephemeral, (I)=Intermittent and (P)=Perennial.

The comparison of each trait amongst river types resulted in a tendency of some traits (“size”,

“pad”, “adnate”, “colonial” and “high-profile guild”) to increase progressively as the degree of

intermittence decreases (Figure 5). However, it also seemed to be a high variability for

perennial rivers, including taxa with wide ranging values for “colonial”, “pad”, “mobile”, “stalk”

and “high-profile”.

On the other hand, while intermittent rivers also showed variability in some traits such

“pioneer”, it seemed that several are constant, including presence of “non-colonial”, “low-profile

guild”, “mobile”, “pedunculate”, “stalk” and absence of “pad” and “adnate” traits. In addition,

regarding the ephemeral river, the traits that appeared to predominate are: “pioneer”, “stalk”

and “planktonic guild”. Nevertheless, Kruskal-Wallis tests4 were conducted for each river type

per trait and all the obtained P-values showed non-significant differences.

4 The Kruskal-Wallis analyses and P-values are attached in the Appendix.

17

Figure 6. Principal Components Analysis (PCA) for species with biological traits as variables and considering

hydrological groups: “Perennial”, “Intermittent”, “Ephemeral” and “All”. Main Species codes5 : Achnanthes pyrenaica

(APYR), Amphora pediculus (APED), Brachysira neoexilis(BNEO) , Cocconeis placentula(CPPL), Cymbopleura

subaequalis (CSAQ ), Diploneis separanda(DSEP), Eucocconeis flexella (EUFL), Eunotia sp (EUNO), Eunotia

bilunaris (EBIL), Encyonema minutum (ENMI), Encyonopsis microcephala (ENCM), Fragilaria perminuta (FPEM),

Gomphonema utae (GUTA), Gomphonema parvulum (GPAR), ), Nitzschia sp. (NITZ), Pinnularia schoenfelderi

(PSHO), Reimeria uniseriata (RUMI), Tabellaria fenestrata (TFEN).

Our PCA (Figure 6), showed that the first component accounted for 29,96% of the information,

and the second component for 18,33%; together they contained 48,82%. The influential

variables are the following biological traits: Mobile, Non-colonial, Motile guild, Colonial,

Pedunculate and Stalk. The first component separated species with mobile, non-colonial traits

and motile guild on the right and colonial and pedunculate traits on the left (Table 5). For the

second component, the variables that weigh are: stalk, pedunculate, low profile guild and also

motile guild with a negative value. Thus, high values of this component indicate species that

have traits such as stalk, peduncle and belong to the low-profile guild, whereas low values of

this component refer to species without these traits that facilitate attachment to the substrate

and belong to the Motile-guild.

One of the centromeres from species belonging to ephemeral rivers is located

negatively on both axes, these taxa are colonial and belong to the high-profile guild or

5 All species Codes can be found on Appendix 2.

Mobile

Pedunculate

Stalk

Colonial

Non-Colonial

Motile guild

ACAF

ADMI

ADPYAPED

COCOCEUGCPLI

CCYM

CAEXCPARCSUT

DMON ESLE

EOMIFAUTFGRAFRCP

GANGGCBCGEXLGGRAGLAT

GOLC

GPUM

MVAR

MCIR

NCTENGRENRCHNVENNINC

PLFRPTLA

RSIN

RABB

SSEMSSTM

UACUULNAUULN

EUNO

EBILFPEM

GPAR

TFEN

APYRADGL

ADMS

BNEO

CPPLCPLE

CYMBCAFFCLAECPPV

CSAQ

DOBLDSEP

ENMI

ECKRENCMECPMESUM

EUFL

FRAG

FTEN

FULN

GACU

GANTGELGGMIN

GOLI

GROSGTER

GUTA

KCLE

MSMIMAPENANTNCARNRADNTPT

NITZ

NZABNIARNDENNDISNFICNFONNILANIMENPAENZSSNZSUNTABPMIC

RUNI

SEPA

ADCA

ADCTADRU

ADEUADEX

ACHD

ACLI

ADNM

AR…

ADCS

CEUOCPEDCOPL

CRAC

CEXFCSLP

CBAM

DDEL

DTEN

DIPL

ENVE

ECESECFAECRX

ESBTESBM

EMIC

FSBHFSAPFPECFVAU

GOMP

GMIC

GMIS

GPRI

MAATMPMINCINNCRYNCTONLANNIGR

NLIN

PPROPSHO

PLFTPLTD

UBICUDEL

-5

-4

-3

-2

-1

0

1

2

3

4

5

6

-6 -5 -4 -3 -2 -1 0 1 2 3 4 5

F2 (

18,3

3 %

)

F1 (29,96 %)

Biplot

All

18

planktonic guild. The other cluster is located below the first component and above the second

component.

Regarding species belonging to intermittent rivers, it appears to be a high variability,

with multiple centromeres: two below both axis, these include colonial species belonging to the

high-profile guild. Another centromere is positive for the first component and negative for the

second component, characterised for being non-colonial and belonging to the motile guild.

There are two more clusters located positive on both axes, includes taxa with traits to attach

to the substrate and belonging to the low-profile guild and non-colonial. In addition, Moreover,

there are also 3 clusters located on a negative first component and a positive second

component, composed of colonial species belonging to low profile guild and having structures

such as a stalk or peduncle.

In reference to species belonging to perennial rivers, there seems to be a high variability

on the traits found and consequently their distribution is wide along all axes.

Table 5. Factor loadings for each component.

F1 F2 F3 F4 F5

Size class -0,072 0,022 -0,164 0,759 -0,339

Mobile 0,758 0,287 -0,326 0,004 0,109

Pioneer 0,106 0,264 0,219 -0,219 0,667

Adnate 0,334 0,219 0,526 0,584 0,129

Pedunculate -0,602 0,647 -0,018 -0,338 -0,276

Pad -0,657 -0,150 0,456 -0,120 -0,350

Stalk -0,099 0,814 -0,395 -0,261 -0,005

Colonial -0,844 -0,177 -0,050 0,223 0,360 Non-Colonial 0,844 0,177 0,050 -0,223 -0,360 High-Profile guild -0,622 0,211 -0,605 0,229 0,037 Low-Profile guild 0,272 0,610 0,628 0,153 -0,003

Motile guild 0,543 -0,730 -0,239 -0,226 -0,014 Planktonic guild -0,457 -0,257 0,471 -0,376 -0,050

6. DISCUSSION

According to predictions on climate change and human alterations, low-flow and drying periods

are expected to increase in Mediterranean regions, with intermittent and ephemeral rivers

being even more frequent (Datry et al., 2014). We found that diatoms did not show statistically

significant differences on taxonomic or trait patterns amongst different river types, most

probably due to the fact that diatoms have short life cycles and are able to quickly respond to

environmental changes.

19

While resilience is studied as the leading cause for algal recolonization by means of

drift, there is a lack of knowledge especially regarding diatom physiological traits that may

enable them with resistant strategies. Our original experiment aimed to be the stepping stone

to further analyses of biofilm resistance to drying periods, shedding light into the pattern of

recolonisation considering both the degree of intermittence and habitat heterogeneity,

experimentally eliminating the effect from drift. Nevertheless, our final study results are mostly

based on morphological traits and ecological guilds, which cannot consider all the possible

stresses that may have an impact on the diatom communities and distribution patterns. The

ecological guilds defined by Passy (2007) are an interesting mechanism to regroup other

biological traits; in our study, the low-profile guild appears to increase following the degree of

intermittency, contrary to the high-profile guild. However, this approach does not seem

convenient when analysing several biological traits and hence, analysing traits independently

is essential. Furthermore, Van Dam (1994) and Falasco (2016) defined several physiological

traits: pH, salinity, Nitrogen uptake metabolism, trophic state, or the Chlorophyll-a levels, but

there is a lack of data for the majority of species sampled by Catalan Water Agency and for

this reason, our results comparing oxygen and moisture traits between different types of rivers

do not seem significant. Further research should also incorporate traits considering mucilage

production as potential and essential strategy to withstand drying conditions. Other valuable

biological traits that should be addressed may be aerial dispersal by animals or wind and the

occurrence of taxa in drift

While comparing traits between river types, one of the major traits that appears to

respond to flow intermittence is diatom size, which increased as the degree of intermittence

decreases, suggesting resistance. However, Round et al. (1990) stated that large diatoms tend

to be epipelic, growing and adhering at the interference of sediments and water, while Leira et

al. (2009) suggested that small cell sizes thrive on flowing rivers with low nutrients levels. Our

different approach is based on the fact that a small cell size implies a fast-growing and

recolonisation rate, usually being pioneer taxa, predominant in changing habitats like

intermittent rivers. On the contrary, a large cell size seems better adapted to stable conditions

like perennial streams. For this river type, most of the analysed traits seemed to have a wide

variability, hinting that other variables may be more influential in shaping diatoms communities

in perennial rivers.

Our results also found variability in intermittent rivers, regarding richness, diversity and

the pioneer trait, as well as in the dispersion on the MDS, and multiple centromeres on the

“Principal Components Analysis”. This is likely to be due to the different timings since the drying

periods started for each river. Unlike macroinvertebrates or other aquatic taxa, diatoms

complete their life cycle rapidly, recolonising and re-establishing the community in short

20

periods of time; therefore, the information about the perturbation is lost in a short time and for

this reason, the analysed rivers should be on the same desiccation stage for a more objective

insight into changes in diatom communities.

However, some other traits appeared to be constant in most intermittent rivers, such as

mobile, pedunculate, non-colonial, no-adnate, no-pad; indicating potential resistance and

resilience strategies. The traits mobile and non-colonial could possibly be correlated, free

individual cells may be able to move locally on a stone, reaching remaining water. In reference

to the peduncle structure, it is thought to be merely structural as a main response to reach light

(Round et al., 1990), but may also be considered as a resistance strategy in intermittent rivers,

whereas the assumed structures to stay attached (adnate and pad) are mostly absent,

suggesting that these traits are not probably resistance mechanisms. Despite the fact that

these traits show no statistically significant differences, the tendency to predominate in the

majority of intermittent rivers seems to evoke that the temporary lack of water flow can

potentially act as an environmental filtering, shaping the community composition, selecting

taxa with these apparent constant traits.

Regarding ephemeral streams, the tendency in traits correlates to intermittent rivers,

but includes pioneer taxa. Nevertheless, the analysed data only corresponds to a single river;

hence, a complete study should have included the same number of samples for each river type

classification.

7. CONCLUSION

In this research, we were not able to test for resistance traits of diatoms from the initial rewetting

experiment; however, our obtained laboratory samples can still be analysed. Our main study

tried to assess potential biological traits that may influence resistance and also resilience

strategies in intermittent rivers. This was carried out by coding traits and assigning them to real

data of diatom communities belonging to several rivers differing in the degree of flow

intermittence.

To sum up, we found no statistically significant differences on traits or community

composition between river types. Although both perennial and intermittent rivers showed a

wide variability, some constant tendencies were observed especially regarding intermittent

rivers. Our approach identifies traits such as mobile, pedunculate, non-colonial, no-adnate and

no-pad to be favourably selected in this river type. In reference to our hypotheses, it seems

that resistance traits predominate, but resilience strategies are also implied. In addition, we

suggest that conducting more species inventories on each river type would be likely to have

21

statistically significant results. Further research should also aim to study rivers that are on a

similar stage in the drying period, in order to reduce the variability.

Acknowledgements

We thank all the people involved in FEHM-lab research group, especially my tutors, Núria

Bonada and Joan Gomà. Also, to Miguel Cañedo-Argüelles for the sampling and data

collected and for helping on the setting of the experimental rewetting. Special thanks to

Carolina Solà and ACA for sending us the data collected from different rivers.

8. BIBLIOGRAPHY

-Barthès, A., Leflaive, J., Coulon, S., Peres, F., Rols J.L. & Ten-Hage,L. (2014). Impact of

Drought on Diatom Communities and the Consequences for the use of Diatom Index Values

in the River Maureillas (Pyrénées‐Orientales, France). River Research and Applications, (31),

993-1002.

-Berthon, B., Bouchez, A. & Rimet, F. (2011). Using diatom life-forms and ecological guilds to

assess organic pollution and trophic level in rivers: a case study of rivers in south-eastern

France. Hydrobiologia, (673), 259–271.

-Bogan, M. T., Chester, E. T., Datry, T., Murphy, A. L., Robson, B. J., Ruhi, A., Whitney, J. E.

(2017). Resistance, Resilience, and Community Recovery in Intermittent Rivers and

Ephemeral Streams. Intermittent Rivers and Ephemeral Streams, 349–376.

-Boix, D., García, E., Gascón, S., Benejam, L., Tornés,E., Sala,J., Benito,J., Munné, A., Solà,

C. & Sabater,S. (2010). Response of community structure to sustained drought in

Mediterranean rivers. Journal of Hidrology, (383), 135-146.

-Bonada, N. & Resh, V.H. (2013). Mediterranean-climate streams and rivers: geographically

separated but ecologically comparable freshwater systems. Hydrobiologia, (719), 1–29

-Chepurnov, V. A., Mann, D. G., Sabbe, K., & Vyverman, W. (2004). Experimental Studies on

Sexual Reproduction in Diatoms. International Review of Cytology, (23), 91–154.

-Datry, T., Larned, S. T., & Tockner, K. (2014). Intermittent Rivers: A Challenge for Freshwater

Ecology. BioScience, 64(3), 229–235.

-Datry, T., Pella, H., Leigh, C., Bonada, N., & Hugueny, B. (2016). A landscape approach to

advance intermittent river ecology. Freshwater Biology, 61(8), 1200–1213.

-De Tommasi, E., Gielis, J., & Rogato, A. (2017). Diatom Frustule Morphogenesis and

Function: A Multidisciplinary Survey. Marine Genomics, (35), 1–18.

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-Falasco, E., Piano, E., & Bona, F. (2016). Suggestions for diatom-based monitoring in

intermittent streams. Knowledge & Management of Aquatic Ecosystems, (417), 38, 1-12

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European rivers. Knowledge and Management of Aquatic Ecosystems, (406), 1-10.

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in seasonally-flowing streams. Freshwater Biology, (53), 2385-2401.

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9. APPENDIX

A.1 Maps of the three studied rivers

Figure 7. Topographic map of the three rivers location. a)Talamanca, b) Santa Creu and C) San Ana

https://www.google.es/maps

Figure 8. Maps of the three studied rivers. FEHM-lab

A.2 Table of Results for Objective 2

Table 6. Analysis of biological traits for each species (Data from ACA)

CO

DE

Hyd

rolo

gySize

classM

ob

ileP

ion

ee

rA

dn

ateP

ed

un

culate

Pad

StalkC

olo

nial

No

n-C

olo

nialH

igh-P

rofile

guild

Low

-Pro

file gu

ildM

otile

guild

Plan

kton

ic guildO

xygen

Mo

isture

Ach

nan

the

s pyre

naica H

uste

dt

AP

YRIn

term

iten

t2

10

01

01

01

01

00

Ach

nan

thid

ium

affine

(Gru

no

w) C

zarne

cki A

CA

FA

ll1

10

01

01

10

10

00

Ach

nan

thid

ium

caled

on

icum

(Lange

-Be

rtalot)Lan

ge-B

ertalo

t A

DC

AP

erm

ane

nt

11

00

10

10

10

10

0

Ach

nan

thid

ium

caten

atum

(Bily &

Marvan

) Lange

-Be

rtalot

AD

CT

Pe

rman

en

t1

10

01

01

10

10

00

Ach

nan

thid

ium

dru

artii Rim

et &

Co

uté

in R

ime

t & al.

AD

RU

Pe

rman

en

t1

10

01

01

10

10

00

Ach

nan

thid

ium

eu

trop

hilu

m (Lan

ge-B

ertalo

t)Lange

-Be

rtalot

AD

EUP

erm

ane

nt

11

00

10

10

10

10

0

Ach

nan

thid

ium

exile

(Kü

tzing) H

eib

erg

AD

EXP

erm

ane

nt

21

00

10

10

10

10

0

AC

HN

AN

THID

IUM

F.T. Kü

tzing

AC

HD

Pe

rman

en

t1

11

01

01

01

01

00

Ach

nan

thid

ium

gracillimu

m (M

eiste

r)Lange

-Be

rtalot

AD

GL

Inte

rmite

nt

11

00

10

10

10

10

0

Ach

nan

thid

ium

line

are W

.Smith

A

CLI

Pe

rman

en

t1

10

01

01

01

01

00

Ach

nan

thid

ium

min

utissim

um

(Kü

tzing) C

zarne

cki A

DM

IA

ll1

11

01

01

01

01

00

Ach

nan

thid

ium

ne

om

icroce

ph

alum

Lange

-Be

rtalot&

F.Staab

AD

NM

Pe

rman

en

t1

10

01

01

10

10

00

Ach

nan

thid

ium

pyre

naicu

m (H

uste

dt) K

ob

ayasi A

DP

YA

ll2

10

01

01

01

01

00

Ach

nan

thid

ium

rostro

pyre

naicu

m Jü

ttne

r & C

ox

AR

PY

Pe

rman

en

t1

10

01

01

10

10

00

Ach

nan

thid

ium

sp.

AD

CS

Pe

rman

en

t1

10

01

01

01

01

00

Ad

lafia min

uscu

la (Gru

no

w) Lan

ge-B

ertalo

t A

DM

SIn

term

iten

t1

10

00

00

01

00

10

Am

ph

ora p

ed

iculu

s (Kü

tzing) G

run

ow

A

PED

All

11

11

00

00

10

10

02

3

Brach

ysira ne

oe

xilis Lange

-Be

rtalot

BN

EOIn

term

iten

t2

10

00

00

01

01

00

CO

CC

ON

EIS C.G

. Ehre

nb

erg

CO

CO

All

51

01

00

00

10

10

03

2

Co

ccon

eis e

uglyp

ta Ehre

nb

erg e

me

nd

Ro

me

ro &

Jahn

C

EUG

All

51

01

00

00

10

10

0

Co

ccon

eis e

uglyp

toid

es (G

eitle

r) Lange

-Be

rtalot

CEU

OP

erm

ane

nt

51

01

00

00

10

10

0

Co

ccon

eis p

ed

iculu

s Ehre

nb

erg

CP

EDP

erm

ane

nt

51

01

00

00

10

10

02

1

Co

ccon

eis p

lacen

tula Eh

ren

be

rg var. pse

ud

olin

eata G

eitle

r C

PP

LIn

term

iten

t4

10

10

00

01

01

00

32

Co

ccon

eis p

lacen

tula Eh

ren

be

rg var.eu

glypta(Eh

r.)Gru

no

w

CP

LEIn

term

iten

t5

10

10

00

01

01

00

32

Co

ccon

eis p

lacen

tula Eh

ren

be

rg var.line

ata (Ehr.)V

an H

eu

rck C

PLI

All

51

01

00

00

10

10

03

2

Co

ccon

eis p

seu

do

line

ata (Ge

itler) Lan

ge-B

ertalo

t C

OP

LP

erm

ane

nt

41

01

00

00

10

10

03

2

Craticu

la accom

od

a (Hu

sted

t) Man

n

CR

AC

Pe

rman

en

t3

10

00

00

01

00

10

CYM

BELLA

C.A

gardh

C

YMB

Inte

rmite

nt

31

00

10

10

10

10

0

Cym

be

lla affinis K

ützin

g var.affinis

CA

FFIn

term

iten

t3

10

01

01

01

01

00

12

Cym

be

lla cymb

iform

is Agard

h

CC

YMA

ll5

10

01

01

01

10

00

12

Cym

be

lla excisa K

ützin

g var. excisa

CA

EXA

ll4

10

01

01

01

01

00

Cym

be

lla excisifo

rmis K

ramm

er var.e

xcisiform

is C

EXF

Pe

rman

en

t4

10

01

01

01

01

00

Cym

be

lla laevis N

aege

li in K

utzin

g var.laevis

CLA

EIn

term

iten

t2

10

01

01

01

01

00

Cym

be

lla parva (W

.Sm.) K

irchn

er in

Co

hn

C

PA

RA

ll2

10

01

01

01

01

00

Cym

be

lla pe

rparva K

ramm

er

CP

PV

Inte

rmite

nt

21

00

10

10

10

10

0

Cym

be

lla sub

lep

toce

ros K

ramm

er

CSLP

Perm

an

ent

41

00

10

10

10

10

0

Cym

be

lla sub

trun

cata Kram

me

r var.sub

trun

cata C

SUT

All

31

00

10

10

10

10

0

Cym

bo

ple

ura am

ph

icep

hala K

ramm

er

CB

AM

Pe

rman

en

t4

10

00

00

10

01

00

Cym

bo

ple

ura su

bae

qu

alis (Gru

no

w) K

ramm

er var. su

bae

qu

alis C

SAQ

Inte

rmite

nt

41

00

00

01

00

10

0

De

licata de

licatula (K

ützin

g) Kram

me

r var. de

licatula

DD

ELP

erm

ane

nt

31

00

00

01

01

00

0

De

nticu

la ten

uis K

ützin

g D

TENP

erm

ane

nt

31

00

00

00

10

01

01

1

Diato

ma m

on

iliform

is Kü

tzing ssp

.mo

nilifo

rmis (m

on

iliform

e?)

DM

ON

All

30

00

11

01

01

00

0

DIP

LON

EIS C.G

. Ehre

nb

erg e

x P.T. C

leve

D

IPL

Pe

rman

en

t2

10

10

00

01

01

00

Dip

lon

eis o

blo

nge

lla (Nae

geli) C

leve

-Eule

r D

OB

LIn

term

iten

t2

10

10

00

01

01

00

14

Dip

lon

eis se

paran

da Lan

ge-B

ertalo

t D

SEPIn

term

iten

t2

10

10

00

01

01

00

Encyo

ne

ma m

inu

tum

(Hilse

in R

abh

.) D.G

. Man

n in

Ro

un

d C

rawfo

rd &

Man

n

ENM

IIn

term

iten

t2

10

00

00

10

10

00

Encyo

ne

ma sile

siacum

(Ble

isch in

Rab

h.) D

.G. M

ann

ESLE

All

31

00

00

01

01

00

0

Encyo

ne

ma ve

ntrico

sum

(Agard

h) G

run

ow

in Sch

mid

t & al.

ENV

EP

erm

ane

nt

21

00

00

01

01

00

0

Encyo

no

psis ce

satii (Rab

en

ho

rst) Kram

me

r EC

ESP

erm

ane

nt

41

00

11

00

10

10

0

Encyo

no

psis falaise

nsis (G

run

ow

) Kram

me

r EC

FAP

erm

ane

nt

21

00

11

00

10

10

0

Encyo

no

psis kram

me

ri Re

ichard

t EC

KR

Inte

rmite

nt

11

00

11

00

10

10

0

Encyo

no

psis m

icroce

ph

ala (Gru

no

w) K

ramm

er

ENC

MIn

term

iten

t3

10

01

10

01

01

00

Encyo

no

psis m

inu

ta Kram

me

r & R

eich

ardt

ECP

MIn

term

iten

t1

10

01

10

01

01

00

Encyo

no

psis su

bm

inu

ta Kram

me

r & R

eich

ardt

ESUM

Inte

rmite

nt

11

00

11

00

10

10

0

Eolim

na crassu

lexigu

a (E.Re

ichard

t) Re

ichard

t EC

RX

Pe

rman

en

t1

10

01

10

01

01

00

Eolim

na m

inim

a(Gru

no

w) Lan

ge-B

ertalo

t EO

MI

All

11

00

00

00

10

01

0

Eolim

na su

bm

inu

scula (M

ang.) M

ose

r Lange

-Be

rtalot &

Me

tzeltin

f. ano

rESB

TP

erm

ane

nt

21

00

00

00

10

01

0

Eolim

na su

bm

inu

scula (M

angu

in) M

ose

r Lange

-Be

rtalot &

Me

tzeltin

ESB

MP

erm

ane

nt

21

00

00

00

10

01

0

Euco

ccon

eis fle

xella (K

ützin

g) Me

ister

EUFL

Inte

rmite

nt

51

00

11

00

11

00

0

EUN

OTIA

C.G

. Ehre

nb

erg

EUN

OEp

he

me

ral3

10

01

10

01

10

00

Eun

otia b

ilun

aris (Ehr.) M

ills var. bilu

naris

EBIL

Eph

em

eral

41

00

00

01

01

00

02

3

Eun

otia m

icroce

ph

ala Krasske

EM

ICP

erm

ane

nt

41

00

00

01

01

00

01

4

Fallacia sub

ham

ulata (G

run

ow

in V

. He

urck) D

.G. M

ann

FSB

HP

erm

ane

nt

21

00

00

00

10

01

0

Fistulife

ra sapro

ph

ila (Lange

-Be

rtalot &

Bo

nik) Lan

ge-B

ertalo

t FSA

PP

erm

ane

nt

11

00

00

00

10

01

0

FRA

GILA

RIA

H.C

. Lyngb

ye

FRA

GIn

term

iten

t2

00

01

10

10

10

00

Fragilaria austriaca (G

run

ow

) Lange

-Be

rtalot

FAU

TA

ll1

00

01

10

10

00

01

1

Fragilaria gracilis Østru

p

FGR

AA

ll1

00

01

10

10

00

01

1

Fragilaria pe

ctinalis (O

.F. Mü

ller) G

ray FP

ECP

erm

ane

nt

10

00

11

01

00

00

11

Fragilaria pe

rmin

uta (G

run

ow

) Lange

-Be

rtalot

FPEM

Eph

em

eral

10

00

11

01

01

00

0

Fragilaria recap

itellata Lan

ge-B

ertalo

t & M

etze

ltin

FRC

PA

ll1

00

01

10

10

00

01

1

Fragilaria ten

era (W

.Smith

) Lange

-Be

rtalot

FTENIn

term

iten

t2

00

01

10

01

00

01

12

Fragilaria uln

a (Nitzsch

.) Lange

-Be

rtalot var. u

lna

FULN

Inte

rmite

nt

50

00

11

01

01

00

03

2

Fragilaria vauch

eriae

(Kü

tzing) P

ete

rsen

FV

AU

Pe

rman

en

t1

00

01

10

10

00

01

1

GO

MP

HO

NEM

A C

.G. Eh

ren

be

rg G

OM

PP

erm

ane

nt

31

00

10

10

11

00

0

Go

mp

ho

ne

ma acu

min

atum

Ehre

nb

erg var.acu

min

atum

G

AC

UIn

term

iten

t5

10

01

01

10

10

00

22

Go

mp

ho

ne

ma an

gustatu

m (K

ützin

g) Rab

en

ho

rst G

AN

GA

ll3

10

01

01

01

10

00

1

Go

mp

ho

ne

ma an

gustu

m A

gardh

G

AN

TIn

term

iten

t4

10

01

01

01

10

00

1

Go

mp

ho

ne

ma cym

be

lliclinu

m R

eich

ardt &

Lange

-Be

rtalot

GC

BC

All

31

00

10

10

11

00

0

Go

mp

ho

ne

ma e

legan

tissimu

m R

eich

ardt &

Lange

-Be

rtalot in

Ho

fman

n &

al.G

ELGIn

term

iten

t3

10

01

01

01

10

00

Go

mp

ho

ne

ma e

xilissimu

m(G

run

.) Lange

-Be

rtalot &

Re

ichard

t G

EXL

All

31

00

10

10

11

00

0

Go

mp

ho

ne

ma gracile

Ehre

nb

erg

GG

RA

All

41

00

10

10

11

00

01

3

Go

mp

ho

ne

ma late

ripu

nctatu

m R

eich

ardt &

Lange

-Be

rtalot

GLA

TA

ll4

10

01

01

01

10

00

13

Go

mp

ho

ne

ma m

icrop

us K

ützin

g var. micro

pu

s G

MIC

Pe

rman

en

t4

10

01

01

01

10

00

23

Go

mp

ho

ne

ma m

inu

sculu

m K

rasske

GM

ISP

erm

ane

nt

21

00

10

10

11

00

0

Go

mp

ho

ne

ma m

inu

tum

(Ag.)A

gardh

f. min

utu

m

GM

INIn

term

iten

t3

10

01

01

01

10

00

Go

mp

ho

ne

ma o

livaceu

m (H

orn

em

ann

) Bré

bisso

n var. o

livaceu

m

GO

LIIn

term

iten

t3

10

01

01

10

10

00

21

Go

mp

ho

ne

ma o

livaceu

m var.calcare

a (Cle

ve) C

leve

in V

an H

eu

rck G

OLC

All

31

00

10

11

01

00

02

1

Go

mp

ho

ne

ma p

arvulu

m (K

ützin

g) Kü

tzing var. p

arvulu

m f. p

arvulu

m

GP

AR

Eph

em

eral

31

00

10

10

11

00

04

3

Go

mp

ho

ne

ma p

um

ilum

(Gru

no

w) R

eich

ardt &

Lange

-Be

rtalot

GP

UM

All

21

00

10

10

11

00

0

Go

mp

ho

ne

ma p

um

ilum

var. rigidu

m R

eich

ardt &

Lange

-Be

rtalot

GP

RI

Pe

rman

en

t2

10

01

01

01

10

00

Go

mp

ho

ne

ma ro

sen

stockian

um

Lange

-Be

rtalot &

Re

ichard

t G

RO

SIn

term

iten

t4

10

01

01

01

10

00

Go

mp

ho

ne

ma te

rgestin

um

(Gru

no

w in

Van

He

urck) Sch

mid

t in Sch

mid

t & al.

GTER

Inte

rmite

nt

41

00

10

10

11

00

01

3

Go

mp

ho

ne

ma u

tae Lan

ge-B

ertalo

t & R

eich

ardt

GU

TAIn

term

iten

t4

10

01

01

01

10

00

Karaye

via cleve

i (Gru

no

w) B

ukh

tiyarova var.cle

vei

KC

LEIn

term

iten

t3

10

01

10

01

01

00

Masto

gloia sm

ithii Th

waite

s M

SMI

Inte

rmite

nt

41

00

00

00

10

01

03

Mayam

aea ato

mu

s (Kü

tzing) Lan

ge-B

ertalo

t var.atom

us

MA

AT

Pe

rman

en

t2

10

00

00

01

00

10

Mayam

aea ato

mu

s var. pe

rmitis (H

uste

dt) Lan

ge-B

ertalo

t M

AP

EIn

term

iten

t1

10

00

00

01

00

10

Mayam

aea p

erm

itis (Hu

sted

t) Bru

de

r & M

ed

lin

MP

MI

Pe

rman

en

t1

10

00

00

01

00

10

Me

losira varian

s Agard

h

MV

AR

All

50

00

00

01

01

00

03

2

Me

ridio

n circu

lare (G

reville

) C.A

.Agard

h var. circu

lare

MC

IRA

ll4

00

01

10

10

01

00

21

Navicu

la anto

nii Lan

ge-B

ertalo

t N

AN

TIn

term

iten

t3

10

00

00

01

00

10

Navicu

la cari Ehre

nb

erg

NC

AR

Inte

rmite

nt

41

00

00

00

10

01

0

Navicu

la cincta (Eh

r.) Ralfs in

Pritch

ard

NC

INP

erm

ane

nt

31

00

00

00

10

01

03

4

Navicu

la crypto

cep

hala K

ützin

g N

CR

YP

erm

ane

nt

31

00

00

00

10

01

03

2

Navicu

la crypto

ten

ella Lan

ge-B

ertalo

t N

CTE

All

31

00

00

00

10

01

03

2

Navicu

la crypto

ten

ello

ide

s Lange

-Be

rtalot

NC

TOP

erm

ane

nt

21

00

00

00

10

01

0

Navicu

la gregaria D

on

kin

NG

RE

All

31

00

00

00

10

01

04

3

Navicu

la lance

olata (A

gardh

) Ehre

nb

erg

NLA

NP

erm

ane

nt

41

00

00

00

10

01

03

3

Navicu

la radio

sa Kü

tzing

NR

AD

Inte

rmite

nt

51

00

00

00

10

01

02

3

Navicu

la reich

ardtian

a Lange

-Be

rtalot var. re

ichard

tiana

NR

CH

All

21

00

00

00

10

01

0

Navicu

la tripu

nctata (O

.F.Mü

ller) B

ory

NTP

TIn

term

iten

t4

10

00

00

01

00

10

23

Navicu

la ven

eta K

ützin

g N

VEN

All

21

00

00

00

10

01

0

NITZSC

HIA

A.H

. Hassall

NITZ

Inte

rmite

nt

30

00

00

00

10

01

0

Nitzsch

ia abb

reviata H

uste

dt in

Schm

idt &

al. N

ZAB

Inte

rmite

nt

11

00

00

00

10

01

0

Nitzsch

ia archib

aldii Lan

ge-B

ertalo

t N

IAR

Inte

rmite

nt

21

00

00

00

10

01

02

Nitzsch

ia de

nticu

la Gru

no

w

ND

ENIn

term

iten

t2

10

00

00

01

00

10

Nitzsch

ia dissip

ata (Kü

tzing) G

run

ow

ssp.d

issipata

ND

ISIn

term

iten

t4

10

00

00

01

00

10

23

Nitzsch

ia filiform

is var.con

ferta (R

ichte

r) Lange

-Be

rtalot

NFIC

Inte

rmite

nt

21

00

00

00

10

01

03

3

Nitzsch

ia fon

ticola G

run

ow

in V

an H

eu

rck N

FON

Inte

rmite

nt

31

00

00

00

10

01

02

1

Nitzsch

ia gracilis Han

tzsch

NIG

RP

erm

ane

nt

21

00

00

00

10

01

02

1

Nitzsch

ia inco

nsp

icua G

run

ow

N

INC

All

11

00

00

00

10

01

03

3

Nitzsch

ia lacuu

m Lan

ge-B

ertalo

t N

ILAIn

term

iten

t1

10

00

00

01

00

10

1

Nitzsch

ia line

aris(Agard

h) W

.M.Sm

ith var.lin

earis

NLIN

Pe

rman

en

t5

10

00

00

01

00

10

23

Nitzsch

ia me

dia H

antzsch

. N

IME

Inte

rmite

nt

41

00

00

00

10

01

0

Nitzsch

ia pale

acea (G

run

ow

) Gru

no

w in

van H

eu

rck N

PA

EIn

term

iten

t2

10

00

00

01

00

10

32

Nitzsch

ia spe

cies

NZSS

Inte

rmite

nt

31

00

00

00

10

01

0

Nitzsch

ia sup

ralitore

a Lange

-Be

rtalot

NZSU

Inte

rmite

nt

11

00

00

00

10

01

0

Nitzsch

ia tabe

llaria (Gru

no

w) G

run

ow

in C

l. & G

run

ow

N

TAB

Inte

rmite

nt

21

00

00

00

10

01

0

Parlib

ellu

s pro

tractus (G

run

ow

) Witko

wski Lan

ge-B

ertalo

t & M

etze

ltin

PP

RO

Pe

rman

en

t4

10

00

00

10

10

00

Pin

nu

laria micro

stauro

n (Eh

r.) Cle

ve var. m

icrostau

ron

P

MIC

Inte

rmite

nt

51

00

00

00

10

01

03

3

Pin

nu

laria scho

en

feld

eri K

ramm

er

PSH

OP

erm

ane

nt

51

00

00

00

10

01

0

Plan

oth

idiu

m fre

qu

en

tissimu

m(Lan

ge-B

ertalo

t)Lange

-Be

rtalot

PLFR

All

21

01

00

00

10

10

0

Plan

oth

idiu

m fre

qu

en

tissimu

m(Lan

ge-B

ertalo

t)Lange

-Be

rtalot f. an

orm

al

PLFT

Pe

rman

en

t3

10

10

00

01

01

00

Plan

oth

idiu

m lan

ceo

latum

(Bre

bisso

n e

x Kü

tzing) Lan

ge-B

ertalo

t P

TLAA

ll3

10

10

00

01

01

00

PLA

NO

THID

IUM

Ro

un

d &

Bu

khtiyaro

va P

LTDP

erm

ane

nt

31

01

00

00

10

10

0

Re

ime

ria sinu

ata (Gre

gory) K

ocio

lek &

Stoe

rme

r R

SINA

ll3

10

11

01

01

01

00

Re

ime

ria un

iseriata Sala G

ue

rrero

& Fe

rrario

RU

NI

Inte

rmite

nt

31

01

10

10

10

10

0

Rh

oico

sph

en

ia abb

reviata (C

.Agard

h) Lan

ge-B

ertalo

t R

AB

BA

ll3

00

01

01

10

01

00

22

Sellap

ho

ra parap

up

ula Lan

ge-B

ertalo

t SEP

AIn

term

iten

t1

10

00

00

01

00

10

Sellap

ho

ra sem

inu

lum

(Gru

no

w) D

.G. M

ann

SSEM

All

11

00

00

00

10

01

0

Sellap

ho

ra stroe

mii (H

uste

dt) K

ob

ayasi in M

ayama Id

ei O

sada &

Nagu

mo

SSTM

All

11

00

00

00

10

01

0

Tabe

llaria fen

estrata (Lyn

gbye

) Kü

tzing

TFENEp

he

me

ral3

00

01

10

01

10

00

1

Uln

aria acus (K

ützin

g) Ab

oal

UA

CU

All

50

00

11

01

01

00

0

Uln

aria bice

ps (K

ützin

g) Co

mp

ère

U

BIC

Pe

rman

en

t5

00

01

10

10

10

00

ULN

AR

IA C

om

pè

re

ULN

AA

ll5

00

01

10

10

10

00

Uln

aria de

licatissima (W

.Smith

) Ab

oal &

Silva U

DEL

Pe

rman

en

t5

00

01

10

10

10

00

Uln

aria uln

a (Nitzsch

) Co

mp

ère

U

ULN

All

50

00

11

01

01

00

0

Table key

-Size class: one of these 5 values can be given to the taxon: 1: biovolume between 0-99 µm3,

2: 100-299 µm3, 3: 300-599 µm3, 4: 600-1499 µm3, 5: over 1500 µm3. (Rimet, 2012)

-Mobile: one of these 2 values can be given to the taxon: 0: immobile taxon, 1: mobile taxon

(has a raphe). (Rimet, 2012)

-Pioneer: one of these 2 values can be given to the taxon: 0: non-pioneer taxon, 1: taxon

considered as pioneer. (Rimet, 2012)

-Adnate: one of these 2 values can be given to the taxon: 0: non adnate taxon, 1: adnate

taxon. (Rimet, 2012)

-Pedunculate (stalk or pad attached to substrate): one of these 2 values can be given to the

taxon: 0: non pedunculate, 1: pedunculate. (Rimet, 2012)

-Pad (attached to substrate):one of these 2 values can be given to the taxon: 0: not producing

a pad, 1: producing a pad. (Rimet, 2012)

-Stalk (attached to substrate): one of these 2 values can be given to the taxon: 0: not producing

a stalk, 1: producing a stalk. (Rimet, 2012)

-Colonial: one of these 2 values can be given to the taxon: 0: not forming colonies, 1: can form

colonies. (Rimet, 2012)

CODE

HydrologySize class

Mobile

PioneerAdnate

PedunculatePad

StalkColonial

Non-ColonialHigh-Profile guild

Low-Profile guild

Motile guild

Planktonic guildO

xygenM

oisture

Achnanthes pyrenaica Hustedt APYR

Intermitent

21

00

10

10

10

10

0

Achnanthidium affine (Grunow

) Czarnecki ACAF

All1

10

01

01

10

10

00

Achnanthidium caledonicum

(Lange-Bertalot)Lange-Bertalot ADCA

Permanent

11

00

10

10

10

10

0

Achnanthidium catenatum

(Bily & M

arvan) Lange-Bertalot ADCT

Permanent

11

00

10

11

01

00

0

Achnanthidium druartii Rim

et & Couté in Rim

et & al.

ADRUPerm

anent1

10

01

01

10

10

00

Achnanthidium eutrophilum

(Lange-Bertalot)Lange-Bertalot ADEU

Permanent

11

00

10

10

10

10

0

Achnanthidium exile (Kützing) Heiberg

ADEXPerm

anent2

10

01

01

01

01

00

ACHNAN

THIDIUM F.T. Kützing

ACHDPerm

anent1

11

01

01

01

01

00

Achnanthidium gracillim

um (M

eister)Lange-Bertalot ADGL

Intermitent

11

00

10

10

10

10

0

Achnanthidium lineare W

.Smith

ACLIPerm

anent1

10

01

01

01

01

00

Achnanthidium m

inutissimum

(Kützing) Czarnecki ADM

IAll

11

10

10

10

10

10

0

Achnanthidium neom

icrocephalum Lange-Bertalot&

F.Staab ADN

MPerm

anent1

10

01

01

10

10

00

Achnanthidium pyrenaicum

(Hustedt) Kobayasi ADPY

All2

10

01

01

01

01

00

Achnanthidium rostropyrenaicum

Jüttner & Cox

ARPYPerm

anent1

10

01

01

10

10

00

Achnanthidium sp. ADCS

Permanent

11

00

10

10

10

10

0

Adlafia minuscula (Grunow

) Lange-Bertalot ADM

SInterm

itent1

10

00

00

01

00

10

Amphora pediculus (Kützing) Grunow

APED

All1

11

10

00

01

01

00

23

Brachysira neoexilis Lange-Bertalot BN

EOInterm

itent2

10

00

00

01

01

00

COCCO

NEIS C.G. Ehrenberg

COCO

All5

10

10

00

01

01

00

32

Cocconeis euglypta Ehrenberg emend Rom

ero & Jahn

CEUGAll

51

01

00

00

10

10

0

Cocconeis euglyptoides (Geitler) Lange-Bertalot CEUO

Permanent

51

01

00

00

10

10

0

Cocconeis pediculus Ehrenberg CPED

Permanent

51

01

00

00

10

10

02

1

Cocconeis placentula Ehrenberg var. pseudolineata Geitler CPPL

Intermitent

41

01

00

00

10

10

03

2

Cocconeis placentula Ehrenberg var.euglypta(Ehr.)Grunow

CPLEInterm

itent5

10

10

00

01

01

00

32

Cocconeis placentula Ehrenberg var.lineata (Ehr.)Van Heurck CPLI

All5

10

10

00

01

01

00

32

Cocconeis pseudolineata (Geitler) Lange-Bertalot CO

PLPerm

anent4

10

10

00

01

01

00

32

Craticula accomoda (Hustedt) M

ann CRAC

Permanent

31

00

00

00

10

01

0

CYMBELLA C.Agardh

CYMB

Intermitent

31

00

10

10

10

10

0

Cymbella affinis Kützing var.affinis

CAFFInterm

itent3

10

01

01

01

01

00

12

Cymbella cym

biformis Agardh

CCYMAll

51

00

10

10

11

00

01

2

Cymbella excisa Kützing var. excisa

CAEXAll

41

00

10

10

10

10

0

Cymbella excisiform

is Kramm

er var.excisiformis

CEXFPerm

anent4

10

01

01

01

01

00

Cymbella laevis N

aegeli in Kutzing var.laevis CLAE

Intermitent

21

00

10

10

10

10

0

Cymbella parva (W

.Sm.) Kirchner in Cohn

CPARAll

21

00

10

10

10

10

0

Cymbella perparva Kram

mer

CPPVInterm

itent2

10

01

01

01

01

00

Cymbella subleptoceros Kram

mer

CSLPPerm

anent4

10

01

01

01

01

00

Cymbella subtruncata Kram

mer var.subtruncata

CSUTAll

31

00

10

10

10

10

0

Cymbopleura am

phicephala Kramm

er CBAM

Permanent

41

00

00

01

00

10

0

Cymbopleura subaequalis (Grunow

) Kramm

er var. subaequalis CSAQ

Intermitent

41

00

00

01

00

10

0

Delicata delicatula (Kützing) Kramm

er var. delicatula DDEL

Permanent

31

00

00

01

01

00

0

Denticula tenuis Kützing DTEN

Permanent

31

00

00

00

10

01

01

1

Diatoma m

oniliformis Kützing ssp.m

oniliformis (m

oniliforme?)

DMO

NAll

30

00

11

01

01

00

0

DIPLON

EIS C.G. Ehrenberg ex P.T. Cleve DIPL

Permanent

21

01

00

00

10

10

0

Diploneis oblongella (Naegeli) Cleve-Euler

DOBL

Intermitent

21

01

00

00

10

10

01

4

Diploneis separanda Lange-Bertalot DSEP

Intermitent

21

01

00

00

10

10

0

Encyonema m

inutum (Hilse in Rabh.) D.G. M

ann in Round Crawford &

Mann

ENM

IInterm

itent2

10

00

00

10

10

00

Encyonema silesiacum

(Bleisch in Rabh.) D.G. Mann

ESLEAll

31

00

00

01

01

00

0

Encyonema ventricosum

(Agardh) Grunow in Schm

idt & al.

ENVE

Permanent

21

00

00

01

01

00

0

Encyonopsis cesatii (Rabenhorst) Kramm

er ECES

Permanent

41

00

11

00

10

10

0

Encyonopsis falaisensis (Grunow) Kram

mer

ECFAPerm

anent2

10

01

10

01

01

00

Encyonopsis kramm

eri Reichardt ECKR

Intermitent

11

00

11

00

10

10

0

Encyonopsis microcephala (Grunow

) Kramm

er EN

CMInterm

itent3

10

01

10

01

01

00

Encyonopsis minuta Kram

mer &

Reichardt ECPM

Intermitent

11

00

11

00

10

10

0

Encyonopsis subminuta Kram

mer &

Reichardt ESUM

Intermitent

11

00

11

00

10

10

0

Eolimna crassulexigua (E.Reichardt) Reichardt

ECRXPerm

anent1

10

01

10

01

01

00

Eolimna m

inima(Grunow

) Lange-Bertalot EO

MI

All1

10

00

00

01

00

10

Eolimna subm

inuscula (Mang.) M

oser Lange-Bertalot & M

etzeltin f. anorESBT

Permanent

21

00

00

00

10

01

0

Eolimna subm

inuscula (Manguin) M

oser Lange-Bertalot & M

etzeltin ESBM

Permanent

21

00

00

00

10

01

0

Eucocconeis flexella (Kützing) Meister

EUFLInterm

itent5

10

01

10

01

10

00

EUNO

TIA C.G. Ehrenberg EUN

OEphem

eral3

10

01

10

01

10

00

Eunotia bilunaris (Ehr.) Mills var. bilunaris

EBILEphem

eral4

10

00

00

10

10

00

23

Eunotia microcephala Krasske

EMIC

Permanent

41

00

00

01

01

00

01

4

Fallacia subhamulata (Grunow

in V. Heurck) D.G. Mann

FSBHPerm

anent2

10

00

00

01

00

10

Fistulifera saprophila (Lange-Bertalot & Bonik) Lange-Bertalot

FSAPPerm

anent1

10

00

00

01

00

10

FRAGILARIA H.C. Lyngbye FRAG

Intermitent

20

00

11

01

01

00

0

Fragilaria austriaca (Grunow) Lange-Bertalot

FAUTAll

10

00

11

01

00

00

11

Fragilaria gracilis Østrup

FGRAAll

10

00

11

01

00

00

11

Fragilaria pectinalis (O.F. M

üller) Gray FPEC

Permanent

10

00

11

01

00

00

11

Fragilaria perminuta (Grunow

) Lange-Bertalot FPEM

Ephemeral

10

00

11

01

01

00

0

Fragilaria recapitellata Lange-Bertalot & M

etzeltin FRCP

All1

00

01

10

10

00

01

1

Fragilaria tenera (W.Sm

ith) Lange-Bertalot FTEN

Intermitent

20

00

11

00

10

00

11

2

Fragilaria ulna (Nitzsch.) Lange-Bertalot var. ulna

FULNInterm

itent5

00

01

10

10

10

00

32

Fragilaria vaucheriae (Kützing) Petersen FVAU

Permanent

10

00

11

01

00

00

11

GOM

PHON

EMA C.G. Ehrenberg

GOM

PPerm

anent3

10

01

01

01

10

00

Gomphonem

a acuminatum

Ehrenberg var.acuminatum

GACU

Intermitent

51

00

10

11

01

00

02

2

Gomphonem

a angustatum (Kützing) Rabenhorst

GANG

All3

10

01

01

01

10

00

1

Gomphonem

a angustum Agardh

GANT

Intermitent

41

00

10

10

11

00

01

Gomphonem

a cymbelliclinum

Reichardt & Lange-Bertalot

GCBCAll

31

00

10

10

11

00

0

Gomphonem

a elegantissimum

Reichardt & Lange-Bertalot in Hofm

ann & al.

GELGInterm

itent3

10

01

01

01

10

00

Gomphonem

a exilissimum

(Grun.) Lange-Bertalot & Reichardt

GEXLAll

31

00

10

10

11

00

0

Gomphonem

a gracile Ehrenberg GGRA

All4

10

01

01

01

10

00

13

Gomphonem

a lateripunctatum Reichardt &

Lange-Bertalot GLAT

All4

10

01

01

01

10

00

13

Gomphonem

a micropus Kützing var. m

icropus GM

ICPerm

anent4

10

01

01

01

10

00

23

Gomphonem

a minusculum

Krasske GM

ISPerm

anent2

10

01

01

01

10

00

Gomphonem

a minutum

(Ag.)Agardh f. minutum

GM

INInterm

itent3

10

01

01

01

10

00

Gomphonem

a olivaceum (Hornem

ann) Brébisson var. olivaceum

GOLI

Intermitent

31

00

10

11

01

00

02

1

Gomphonem

a olivaceum var.calcarea (Cleve) Cleve in Van Heurck

GOLC

All3

10

01

01

10

10

00

21

Gomphonem

a parvulum (Kützing) Kützing var. parvulum

f. parvulum

GPAREphem

eral3

10

01

01

01

10

00

43

Gomphonem

a pumilum

(Grunow) Reichardt &

Lange-Bertalot GPUM

All2

10

01

01

01

10

00

Gomphonem

a pumilum

var. rigidum Reichardt &

Lange-Bertalot GPRI

Permanent

21

00

10

10

11

00

0

Gomphonem

a rosenstockianum Lange-Bertalot &

Reichardt GRO

SInterm

itent4

10

01

01

01

10

00

Gomphonem

a tergestinum (Grunow

in Van Heurck) Schmidt in Schm

idt & al.

GTERInterm

itent4

10

01

01

01

10

00

13

Gomphonem

a utae Lange-Bertalot & Reichardt

GUTAInterm

itent4

10

01

01

01

10

00

Karayevia clevei (Grunow) Bukhtiyarova var.clevei

KCLEInterm

itent3

10

01

10

01

01

00

Mastogloia sm

ithii Thwaites

MSM

IInterm

itent4

10

00

00

01

00

10

3

Mayam

aea atomus (Kützing) Lange-Bertalot var.atom

us M

AATPerm

anent2

10

00

00

01

00

10

Mayam

aea atomus var. perm

itis (Hustedt) Lange-Bertalot M

APEInterm

itent1

10

00

00

01

00

10

Mayam

aea permitis (Hustedt) Bruder &

Medlin

MPM

IPerm

anent1

10

00

00

01

00

10

Melosira varians Agardh

MVAR

All5

00

00

00

10

10

00

32

Meridion circulare (Greville) C.A.Agardh var. circulare

MCIR

All4

00

01

10

10

01

00

21

Navicula antonii Lange-Bertalot

NAN

TInterm

itent3

10

00

00

01

00

10

Navicula cari Ehrenberg

NCAR

Intermitent

41

00

00

00

10

01

0

Navicula cincta (Ehr.) Ralfs in Pritchard

NCIN

Permanent

31

00

00

00

10

01

03

4

Navicula cryptocephala Kützing

NCRY

Permanent

31

00

00

00

10

01

03

2

Navicula cryptotenella Lange-Bertalot

NCTE

All3

10

00

00

01

00

10

32

Navicula cryptotenelloides Lange-Bertalot

NCTO

Permanent

21

00

00

00

10

01

0

Navicula gregaria Donkin

NGRE

All3

10

00

00

01

00

10

43

Navicula lanceolata (Agardh) Ehrenberg

NLAN

Permanent

41

00

00

00

10

01

03

3

Navicula radiosa Kützing

NRAD

Intermitent

51

00

00

00

10

01

02

3

Navicula reichardtiana Lange-Bertalot var. reichardtiana

NRCH

All2

10

00

00

01

00

10

Navicula tripunctata (O

.F.Müller) Bory

NTPT

Intermitent

41

00

00

00

10

01

02

3

Navicula veneta Kützing

NVEN

All2

10

00

00

01

00

10

NITZSCHIA A.H. Hassall

NITZ

Intermitent

30

00

00

00

10

01

0

Nitzschia abbreviata Hustedt in Schm

idt & al.

NZAB

Intermitent

11

00

00

00

10

01

0

Nitzschia archibaldii Lange-Bertalot

NIAR

Intermitent

21

00

00

00

10

01

02

Nitzschia denticula Grunow

N

DENInterm

itent2

10

00

00

01

00

10

Nitzschia dissipata (Kützing) Grunow

ssp.dissipata N

DISInterm

itent4

10

00

00

01

00

10

23

Nitzschia filiform

is var.conferta (Richter) Lange-Bertalot N

FICInterm

itent2

10

00

00

01

00

10

33

Nitzschia fonticola Grunow

in Van Heurck N

FON

Intermitent

31

00

00

00

10

01

02

1

Nitzschia gracilis Hantzsch

NIGR

Permanent

21

00

00

00

10

01

02

1

Nitzschia inconspicua Grunow

N

INC

All1

10

00

00

01

00

10

33

Nitzschia lacuum

Lange-Bertalot N

ILAInterm

itent1

10

00

00

01

00

10

1

Nitzschia linearis(Agardh) W

.M.Sm

ith var.linearis N

LINPerm

anent5

10

00

00

01

00

10

23

Nitzschia m

edia Hantzsch. N

IME

Intermitent

41

00

00

00

10

01

0

Nitzschia paleacea (Grunow

) Grunow in van Heurck

NPAE

Intermitent

21

00

00

00

10

01

03

2

Nitzschia species

NZSS

Intermitent

31

00

00

00

10

01

0

Nitzschia supralitorea Lange-Bertalot

NZSU

Intermitent

11

00

00

00

10

01

0

Nitzschia tabellaria (Grunow

) Grunow in Cl. &

Grunow

NTAB

Intermitent

21

00

00

00

10

01

0

Parlibellus protractus (Grunow) W

itkowski Lange-Bertalot &

Metzeltin

PPROPerm

anent4

10

00

00

10

10

00

Pinnularia microstauron (Ehr.) Cleve var. m

icrostauron PM

ICInterm

itent5

10

00

00

01

00

10

33

Pinnularia schoenfelderi Kramm

er PSHO

Permanent

51

00

00

00

10

01

0

Planothidium frequentissim

um(Lange-Bertalot)Lange-Bertalot

PLFRAll

21

01

00

00

10

10

0

Planothidium frequentissim

um(Lange-Bertalot)Lange-Bertalot f. anorm

al

PLFT

Permanent

31

01

00

00

10

10

0

Planothidium lanceolatum

(Brebisson ex Kützing) Lange-Bertalot PTLA

All3

10

10

00

01

01

00

PLANO

THIDIUM Round &

Bukhtiyarova PLTD

Permanent

31

01

00

00

10

10

0

Reimeria sinuata (Gregory) Kociolek &

Stoermer

RSINAll

31

01

10

10

10

10

0

Reimeria uniseriata Sala Guerrero &

Ferrario RUN

IInterm

itent3

10

11

01

01

01

00

Rhoicosphenia abbreviata (C.Agardh) Lange-Bertalot RABB

All3

00

01

01

10

01

00

22

Sellaphora parapupula Lange-Bertalot SEPA

Intermitent

11

00

00

00

10

01

0

Sellaphora seminulum

(Grunow) D.G. M

ann SSEM

All1

10

00

00

01

00

10

Sellaphora stroemii (Hustedt) Kobayasi in M

ayama Idei O

sada & N

agumo

SSTMAll

11

00

00

00

10

01

0

Tabellaria fenestrata (Lyngbye) Kützing TFEN

Ephemeral

30

00

11

00

11

00

01

Ulnaria acus (Kützing) Aboal UACU

All5

00

01

10

10

10

00

Ulnaria biceps (Kützing) Compère

UBICPerm

anent5

00

01

10

10

10

00

ULNARIA Com

père ULN

AAll

50

00

11

01

01

00

0

Ulnaria delicatissima (W

.Smith) Aboal &

Silva UDEL

Permanent

50

00

11

01

01

00

0

Ulnaria ulna (Nitzsch) Com

père UULN

All5

00

01

10

10

10

00

-Non-colonial: one of these 2 values can be given to the taxon: 0: can form colonies, 1: not

forming colonies. (Rimet, 2012)

-Guilds(high profile, low profile, motile, planktonic): one of these 2 values can be given to

the taxon: 0: do not belong to this guild, 1: belong to this guild. (Rimet, 2012)

-Oxygen requirements:1:continuously high (about 100% saturation),2:fairly high (above 75%

saturation), 3:moderate (above 50% saturation), 4:low (above 30% saturation), 5:very low

(about 10% saturation). (Van Dam, 1994)

-Moisture: 1:never, or only very rarely, occurring outside water bodies, 2: mainly occurring in

water bodies, sometimes on wet places, 3: mainly occurring in water bodies, also rather

regularly on wetand moist places, 4: mainly occurring on wet and moist or temporarily dry

places, 5 : nearly exclusively occurring outside water bodies. (Van Dam, 1994)

A.3 Matrix (C)

This C Matrix includes the proportion of each trait per studied river Size Mobile Pioneer Adnate Pedunculate Brugent 2.266173 0.5430457 0.09649035 0.00000000 0.88811119 Portella 3.098310 0.5043504 0.07810781 0.00000000 0.93679368 Merles 1.544109 0.8578716 0.54550910 0.00000000 0.93168634 Major 2.448145 0.9347935 0.20802080 0.05360536 0.97219722 Osor 1.974892 0.9352806 0.00470141 0.26647994 0.09582875 Guilla 3.880148 0.9994002 0.03948421 0.94092363 0.05787685 Glorieta 1.329900 0.9666000 0.76820000 0.00000000 0.95420000 SantaCreu 1.396540 0.9903990 0.80168017 0.00240024 0.96859686 Guanta 2.376162 0.9800020 0.14168583 0.00739926 0.94280572 Avenco 2.137900 0.9779000 0.00250000 0.05420000 0.94580000 Daro 1.542717 0.9729892 0.69947979 0.02711084 0.86984794 Orlina 2.850570 0.8522705 0.06041208 0.46029206 0.25175035 Ridaura 1.377200 0.9448000 0.75740000 0.00000000 0.96880000 Pad Stalk Colonial Non.Colonial Brugent 0.3934606539 0.49465053 0.46945305 0.4618538 Portella 0.5052505251 0.43154315 0.71747175 0.2825283 Merles 0.1517303461 0.77995599 0.26305261 0.7346469 Major 0.0513051305 0.92089209 0.05590559 0.9347935 Osor 0.0144043213 0.08142443 0.05741723 0.9282785 Guilla 0.0005997601 0.05727709 0.00019992 0.9994002 Glorieta 0.0334000000 0.92080000 0.07940000 0.9081000 SantaCreu 0.0362036204 0.93239324 0.00720072 0.9927993 Guanta 0.2366763324 0.70612939 0.02999700 0.9700030 Avenco 0.0148000000 0.93100000 0.02210000 0.9779000 Daro 0.0123049220 0.85754302 0.03211285 0.9555822 Orlina 0.0145029006 0.23724745 0.14772955 0.8522705 Ridaura 0.0456000000 0.92320000 0.04320000 0.9376000 High.Profile_guild Low.Profile_guild Motile_guild Brugent 0.439256074 0.4566543 0.00000000 Portella 0.734473447 0.2510251 0.01450145 Merles 0.170434087 0.6757351 0.02580516 Major 0.673167317 0.3152315 0.00230023 Osor 0.009302791 0.3528058 0.62358708

Guilla 0.017493003 0.9807077 0.00139944 Glorieta 0.092000000 0.8622000 0.02910000 SantaCreu 0.075007501 0.8935894 0.02660266 Guanta 0.174082592 0.7885211 0.03739626 Avenco 0.913800000 0.0813000 0.00490000 Daro 0.155662265 0.7537015 0.07833133 Orlina 0.019303861 0.6249250 0.35577115 Ridaura 0.180200000 0.7814000 0.00240000 Planktonic_guild Oxygen Moisture Brugent 0.03539646 0.04559544 0.01809819 Portella 0.00000000 0.03880388 0.11421142 Merles 0.12572515 0.21074215 0.07791558 Major 0.00000000 0.20252025 0.17931793 Osor 0.00000000 0.23847154 0.22636791 Guilla 0.00000000 2.79438225 1.88404638 Glorieta 0.00420000 0.12090000 0.07090000 SantaCreu 0.00480048 0.08470847 0.12341234 Guanta 0.00000000 0.14198580 0.19388061 Avenco 0.00000000 0.06900000 0.04190000 Daro 0.00000000 0.27581032 0.13735494 Orlina 0.00000000 1.34926985 1.23274655 Ridaura 0.01680000 0.06480000 0.04320000

A.4 Kruskal-Wallis Tests for the three river types per each trait

Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 3.7692, df = 2, p-value = 0.1519 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 2.6593, df = 2, p-value = 0.2646 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 2.2198, df = 2, p-value = 0.3296 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 1.3023, df = 2, p-value = 0.5214 NULL

Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 2.4505, df = 2, p-value = 0.2937 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 1, df = 2, p-value = 0.6065 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 3.8022, df = 2, p-value = 0.1494 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 2.0549, df = 2, p-value = 0.3579 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 2.0549, df = 2, p-value = 0.3579 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 0.37363, df = 2, p-value = 0.8296 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 1.3626, df = 2, p-value = 0.5059 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 3.5495, df = 2, p-value = 0.1695

NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 1.7429, df = 2, p-value = 0.4184 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 1.1648, df = 2, p-value = 0.5585 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp1 Kruskal-Wallis chi-squared = 1.1648, df = 2, p-value = 0.5585 no hi ha diferències en lloc per 3 categories NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 2.9388, df = 1, p-value = 0.08648 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 2.4694, df = 1, p-value = 0.1161 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 1.6531, df = 1, p-value = 0.1985 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 0, df = 1, p-value = 1 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2

Kruskal-Wallis chi-squared = 1.3061, df = 1, p-value = 0.2531 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 0.73469, df = 1, p-value = 0.3914 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 3.449, df = 1, p-value = 0.06329 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 2.0408, df = 1, p-value = 0.1531 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 2.0408, df = 1, p-value = 0.1531 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 0.020408, df = 1, p-value = 0.8864 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 1.3061, df = 1, p-value = 0.2531 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 2.0408, df = 1, p-value = 0.1531 NULL Kruskal-Wallis rank sum test

data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 0.026531, df = 1, p-value = 0.8706 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 0.020408, df = 1, p-value = 0.8864 NULL Kruskal-Wallis rank sum test data: matriuC[, i] and temp2 Kruskal-Wallis chi-squared = 0.18367, df = 1, p-value = 0.6682 A.5 Herbarium for the identified species (Objective 1)

Làmina 1

X1500

Figura 1: Fragilaria cf radians

Figura 2: Fragilaria dilatata

Figura 3-4: Fragilaria ulna

Figura 5-7: Rhopalodia parallela

Figura 8: Gyrosigma cf attenuatum

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Figura 1-4: Cymbopleura amphicephala

Figura 5-7: Eucocconeis flexella

Figura 8: Diploneis elliptica

Figura 9-10: Eunotia bidens

Figura 11-13 Eunotia denticulata

Làmina 3

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Figura 1-7: Mastogloia smithii

Figura 8-10: Mastogloia lacustris

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Làmina 4

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Figura 1: Cyclotella meneguiniana

Figura 2: Stephanodiscus minutulus

Figura 3: Cyclostephanos dubius

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Figura 1-3: Cymbella excisa

Figura 4-5: Cymbella cf compacta

Figura 6: Cymbella cf laevis

Figura 7-8: Encyonema silesiacum

Figura 9-10: Cymbella cymbiformes

Figura 11: Cymbella cf lancettula

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Làmina 6

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Figura 1-6: Brachisyra vitrea

Figura 7: Brachisyra liliana

Figura 8-9: Navicula vandamii

Figura 10: Navicula criptotenella

Figura 11-16: Encyonopsis cesatii

Figura 17-18: Encyonopsis minuta

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Figura 1: Pinnularia cf subcapita

Figura 2-3: Pinnularia cf sinistra

Figura 4: Pinnularia nobilis

Figura 5: Denticula tenuis

Figura 6-7: Nitzschia denticula

Figura 8: Hantschia abundans

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Figura 1-7: Gomphonema lateripunctatum

Figura 8-11: Gomphonema micropus

Figura 12-13 : Gomphonema gracile

Figura 14: Gomphonema vibrio

Figura 15-20: Gomphonema parvulis

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Figura 1-7: Achnantidium minutissimum

Figura 8: Achnantes cf trinoidis

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Analysis of the biological traits of diatoms in intermittent rivers ...

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