Beyond the Tower of Babel …

Danube_at_Vienna_May19

A week after I return from China, I was off on my travels again; this time to Vienna for a workshop between molecular ecology specialists and ECOSTAT, the committee of Member State representatives who oversee ecological aspects of Water Framework Directive implementation.   As ever, I found some time to visit some art galleries around the meeting and, as Vienna has one of the most impressive collections of paintings by Pieter Brueghel, I could not resist spending some time in front of his “Tower of Babel”.  A few years ago I cheerfully included this picture in a talk on EU ecological assessment methods, as we tried to make sense of the myriad national approaches.   Three years after the Brexit vote, however, it seems to better reflect UK domestic politics where, ironically, language is one of the few things that all protagonists do have in common.

The River Danube seems to encapsulate the reasons why Europe needs collaborative thinking on the state of the environment.  It is the second longest river in Europe, after the Volga, and flows through ten countries, with tributaries extending into nine more.   Eight of the nine countries through which the river flows are members of the EU (the ninth, Serbia, is in the process of joining) so the river represents a case study, of sorts, on whether EU environmental policies actually work.   This is not just an academic question: ecologists are generally in favour of integrated management of entire catchments whilst the EU operates on a principle of “subsidiarity”, which means that decision-making is devolved to the lowest competent authority (individual Member States in the case of the environment).   Finding the right balance between these principles takes a lot of patient discussion and is one reason why EU decision-making can appear to be agonisingly slow.

Breughel_Tower_of_Babel

Pieter Bruegel’s “Tower of Babel” in the Kunsthistorisches Museum in Vienna.

And there are more problems: the Water Framework Directive evaluates the sustainability of water bodies by their naturalness yet very large rivers such as the Danube have been very heavily modified by human use for centuries.   The river has been broadened, deepened and impounded, and its banks have been straightened and strengthened in order to make it navigable, and there is a huge human population, with associated industry, living on its banks.  The stretch of the Danube along which I walked on my last morning in Vienna was also lined with embankments to protect the surrounding land from flooding but these, at the same time, cut the river off from the ecological benefits of the floodplain.

What hope for a large river such as the Danube in the face of all these challenges?   First of all, when dealing with rivers such as these we need to adjust our expectations, recognising that they are so central to the economic life of the regions through which they flow that there are limits to their capacity to ever resemble truly natural rivers.   Once we have done this, we can start to unpick the challenges that can be addressed by individual Member States.  In the case of water quality, in particular, the story for the Danube is encouraging and European environmental legislation has played its role in this process.  By the time the Danube reaches the borders with Romania, for example, nutrient concentrations are low enough for many of the benthic algal-communities to meet criteria for “good ecological status”.

You can see this in the graph below, from a paper that we’ve published recently.   The Romanian sites are largely clustered at the top left hand side of the graph, relative to data from other countries – indicating low phosphorus concentrations and good ecology (expressed as “ecological quality ratios”, EQRs).   Thanks to an extensive exercise that took place a few years before I started grappling with the Romanian data, we already had a consensus view of the EQR boundaries for high and good status, and most of the Romanian data fits into the band representing “good status”.  That’s encouraging and whilst these communities are just one element of a much more complex ecosystem, but it is a clear step in the right direction.

RO_VLR_intercalibration

The relationship between dissolved phosphorus and ecological stataus of the phytobenthos (expressed as the Ecological Quality Ratio, EQR, based on the intercalibration common metric (which gives a harmonised view of status between Member States).   Horizontal lines show the average position of “high” (blue) and “good” (green) status boundaries.   RO = Romanian data; XGIG = data from other Member States.   See Kelly et al. (2018) for more details.  

Romania is, of course, a long way downstream from where I was standing in Vienna.  Before the Danube gets there it has to cross Slovakia, Hungary and Serbia.  The river also forms the boundary between Romania and Bulgaria for about 300 kilometres, so it is important that there is joined-up thinking between those responsible for water quality on the two opposite banks.  That’s why the EU is so important for the environment on a pan-European scale.  It is easy for those of us crammed onto our insignificant archipelago in the north-west corner of the continent to overlook this, but the Danube is really a great success stories for European environmental collaboration and, indeed, a reason for staying with this ambitious project into the future.   Too late, I know, but it needs to be said.

Reference

Kelly, M.G., Chiriac, G., Soare-Minea, A., Hamchevici, C. & Birk, S. (2018).  Defining ecological status of phytobenthos in very large rivers: a case study of practical implementation of the Water Framework Directive in Romania.  Hydrobiologia 828: 353-367.

Vienna_sights_May19

Sightseeing in Vienna: Stefansdom, the historic cathedral in the city centre and the Ferris wheel at the Prater amusement park, which played a starring role in Graham Greene’s The Third Man.

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The complexities of measuring mass…

Benthotorch_in_action

Once upon a time, measuring the quantity of algae growing on the beds of streams and rivers was a painstaking, slow process that invariably revealed large amounts of spatial and temporal variation that, very often, obscured the ecological signals you were looking for. That has changed in the last decade thanks to the availability of field fluorimeters such as the BenthoTorch.  This makes it much quicker and easier to measure chlorophyll concentrations, the usual proxy for algal quantity.  Thanks to devices such as this it is now much easier to discover that your ecological signal is masked by spatial and temporal variation.

We’ve generated a lot of data about the fluxes of algae in the River Ehen using a BenthoTorch over the past five years and are in a position where we can start to make some generalisations about how the quantity of algae vary over the course of a year.  In broad terms, the results I showed in “The River Ehen in January” back in 2014 have not varied greatly over subsequent years, with peak biomass in mid-winter and low biomass in the summer (due, we presume, to intense grazing by invertebrates).  Curiously, we see a much less distinctive seasonal pattern in the nearby Croasdale Beck, but that is a story for another day….

The BenthoTorch uses an algorithm to partition the fluorescence signal between three major algal groups and, though this is not without issues (see below), I thought it might be interesting to see how these groups varied with biomass trends, and consider how this links to ecological theory.  The first group I’m considering are the green algae which, in this river, are mainly filamentous forms.   The general pattern, seen in the graph below, is for a gradual increase in the proportion of green algae, which fits with the current understanding of thicker biofilms having greater structural complexity with filamentous algae out-competing attached single celled algae to create a “canopy” of algae that are more effective at capturing light and other resources.  The relationship is, however, strongly wedge-shaped so, whilst many of the thickest biofilms have a lot of green algae, there are also thick biofilms where green algae are scarce or even non-existent.  Croasdale Beck shows a similar, but less pronounced, trend.

green_algae_in_Ehen

Relationship between the proportion of green algae and the total quantity of benthic algae (expressed as chlorophyll concentration) in the River Ehen (a.) and Croasdale Beck (b.).   The blue lines show quantile regression fits at p = 0.8, 0.5 and 0.2.   The image at the top of the post shows Ben Surridge using a BenthoTorch to measure algal biomass beside Croasdale Beck in Cumbria.

The second graph shows that this pattern of a gradual increase in proportion is also the case for diatoms and, once again, there is a broad wedge of points with an upward trend.  But, once again, there are also samples where biomass is high but diatoms are present in very low numbers or are even absent.   What is going on?

The problem is clear I think, if one looks at the final image in “The only way is up …” where the very patchy nature of algal communities in the River Ehen (and, indeed, many other rivers).   There are plenty of algae on this boulder, but not organised in a homogeneous manner: some zones on the boulder are almost pure diatom whilst others are almost pure green algae (and there are also zones that are almost pure Lemanea– I’ll come to that in a future post).   We try to sample the stones as randomly as possible so you can see the potential for getting very different numbers depending on where, on a stone, we point the BenthoTorch’s sensor.

diatoms_in_the_Ehen

Relationship between the proportion of diatoms and the total quantity of benthic algae (expressed as chlorophyll concentration) in the River Ehen (c.) and Croasdale Beck (d.).   The blue lines show quantile regression fits at p = 0.8, 0.5 and 0.2.  

With experience, you can make an educated guess about the types of algae present in a biofilm.  I’ve tried to capture this with my watercolours, using washes of raw sienna for the diatoms and a grass-green for the green algae, which roughly matches the colour of their respective growths in the photo in my earlier post.   The two groups of algae a are relatively distinct on that particular boulder.   The top row roughly matches the upper “edge” of the graph showing variation in diatoms, whilst the bottom row emulates the upper “edge” of the graph showing variation in green algae.  These are the two extreme situations; however, we also often see darker brown growths in the field, which can be recreated by mixing the raw sienna and grass-green together.  When I peer through a microscope I often see green algae smothered in diatoms: genera such as Oedogoniumare particularly prone as they have less mucilage than some of the others we find in the Ehen. Their filaments often host clusters of Fragilariacells as well as Achnanthidium minutissimum, whilst stalked Gomphonemaand chains of Tabellaria flocculosaoften grow through the tangle of green filaments.   The dark brown colour is deepened yet further by the colour of the underlying rock, so my effort on white watercolour paper is a little misleading.

colour_patches

A colour chart showing how different proportions of green algae and diatoms influence the colour of biofilms.

The final graph shows how, as the average biomass increases in the River Ehen, so the variability in biomass also increases.   The River Ehen is one of the cleanest rivers I know but I suspect that this pattern in benthic algal quantity could be reproduced in just about any river in the country. What I would not expect to see in any but the purest and most natural ecosystems is quite so much variation in the types of algae present.   Once there is a little enrichment, so I would expect the algae to become more of a monoculture of a dominant filamentous alga plus associated epiphytes.  Like much that happens in the microscopic world of rivers, it is easier to describe than it is to measure.

That, however, is only part of the story but I’ll come back to explain the patterns in the other main groups of algae in the Ehen and Croasdale Beck in my next post.

mean_biomass_by_stdev

The relationship between mean chlorophyll density and the standard deviation (based on measurements from five separate stones) for samples from the River Ehen and Croasdale Beck. 

 

Croasdale Beck in February

Ennerdale_Feb19

My latest trip to the west Cumbria coincided with the period of freakily warm weather that marked the end of February (in marked contrast to a year previously when we were in the midst of the “Beast from the East”).   It felt like spring had come early although the skeletal outlines of leafless trees were incongruous against the backdrop of blue skies and, despite feeling the warmth of the sun on our faces as we worked, the water still had a wintery chill when the time came to plunge in my arm.

There were thick growths of algae on the bed of Croasdale Beck: a quick check with my microscope later showed this to be mostly Odontidium mesodonand Gomphonema parvulumand this piqued my curiosity to see how different species responded to the fluctuations in biomass that we observe in the streams in this region. I’ve talked about this before (see “A tale of two diatoms …”), suggesting that Platessa oblongellatended to dominate when biofilms were thin whilst Odontidium mesodon preferred thicker biofilms.  That was almost two years ago and I now have more data with which to test that hypothesis, and also to see if any other common taxa had an equally strong preference for particular states.

Croasdale_cobble_Feb19

A cobble from the bed of Croasdale Beck in February 2019 showing a brown biofilm (approx. 1.7 micrograms per square centimetre) dominated by Gomphonema parvulumand Odontidium mesodon.   The photograph at the top of the post shows Ennerdale Water photographed on the same day.

I should also be clear that, in Croasdale Beck especially, diatoms are the main algal component of the biofilm, so they are not so much responding to a particular state of the biofilm as actively contributing biomass to create that state.  The other photosynthetic organism that is obvious to the naked eye in this part of Croasdale Beck is the cyanobacterium Chamaesiphon fuscus (see “A bigger splash …”) but this forms crusts on stone surfaces rather than contributing to the superstructure of the biofilm itself. We do find other filamentous algae, but intermittently and in smaller quantities.

We’ll look at Platessa oblongellafirst, bearing in mind that this was shown to be a mixture of two species about halfway through our study (see “Small details in the big picture …”).   The graph below, therefore, does not differentiate between these two species although, from my own observations, I have no reason to believe that they behave differently.   What I have done in these graphs is to divide the biomass measurements and the percent representation of these taxa in each sample into three categories: low, middle and high.   In each case, “low” represents the bottom 25 per cent of measurements, “high” represents the top 25 per cent of measurements and “middle” represents all the rest. The left-hand graph shows biomass (as chlorophyll a concentration) as a function of the relative abundance of the diatom whilst the right-hand graph shows the opposite: the relative abundance of the diatom as a function of the biomass.  These graphs bear out what I suggested in my earlier post: that Platessa oblongella(and P. saxonica) are species whose highest relative abundances occur when the biofilm is thin.  So far, so good.

P.oblongella

Relationship between relative abundance of Platessa oblongella (including P. saxonica) and biomass in Croasdale Beck, Cumbria.  a. shows biomass (as chlorophyll a) as a function of the relative abundance of the two species (Kruskal-Wallis test, p = 0.047) whilst b. shows the relative abundance as a function of biomass (p = 0.057).

My second prediction in my earlier post was that Odontidium mesodonpreferred moderate or thick biofilms; however, whilst there is a clear trend in the data, differences between low, middle and high values of neither biomass nor relative abundance are significant.   The explanation may lay in the strong seasonality that O. mesodondisplays, thriving in spring but less common at other times of year (see “More about Platessa oblongella and Odontidium mesodon”).  However, there are no strong seasonal patterns in biomass in Croasdale Beck, and this disjunction introduces enough noise into the relationship to render it not significant.

O.mesodon

Relationship between relative abundance of Odontidium mesodon and biomass in Croasdale Beck, Cumbria.  a. shows biomass (as chlorophyll a) as a function of the relative abundance of O. mesodon (Kruskal-Wallis test, p = 0.568) whilst b. shows the relative abundance as a function of biomass (p = 0.060).

I then tried looking at the relationship between relative abundance and biomass for a few other common taxa but with mixed results.   None of Achnanthidium minutissimum, Gomphonema parvulum complex or Fragilaria pectinalis showed any clear relationship; however, when I looked at Fragilaria gracilis, a different pattern emerged, with a significant relationship between the quantity of biomass and the proportion of this species in the sample.  That, too, is not a great surprise as I often see clusters of Fragilaria gracilis cells growing epiphytically on filamentous algae within the biofilm.  Whilst Platessa oblongella, which sits flat on the stone surface, seems to be a species that thrives when the biofilm is thin, so Fragilaira gracilisis favoured by a more complex three-dimensional structure, where it can piggy-back on other algae to exploit the light.   I suspect, however, that in a stream such as Croasdale Beck, where the substratum is very mobile, Fragilaira gracilis will also be one of the first casualties of a scouring spate which will, in turn, open up the canopy allowing Platessa oblongella back.   Even though my results for Odontidium mesodonare not significant, I still think it plays a part in this sequence, occupying the intermediate condition when some biomass has accumulated.  It looks to me as if it also likes cooler conditions which then complicates interpretation of my results.

Indeed, I am being rather selective in the results that I have included here.  Three of the six species I investigated showed no response and one of the three that I did include showed a trend rather than a statistically-convincing effect.  I suspect that the situation will rarely be as simple as I have shown for Platessa oblongella and Fragilaira gracilis.  Nonetheless, there is enough here to make me want to scratch a little more and try to understand this topic better.

F.gracilis

Relationship between relative abundance of Fragilaria gracilis and biomass in Croasdale Beck, Cumbria.  a. shows biomass (as chlorophyll a) as a function of the relative abundance of F.gracilis (Kruskal-Wallis test, p = 0.010) whilst b. shows the relative abundance as a function of biomass (p = 0.036).

Croasdale_Beck_Feb19

Croasdale Beck, photographed in February 2019. 

How Craticula got its name

Here is a puzzle for anyone who is learning to identify diatoms: how many species are shown in the plate below?   All share the same size and outline but they are very different in other respects, including several that we would normally regard as important for separating different species.   The left-hand image is an isolated girdle band, so let’s leave that to one side for the moment.  What about the two middle valves?   Both have a raphe in two parts, that runs along the midline, but the arrangement of their striae is very different.   And how do these relate to the pair on the right, which seem to have stout silica bars which traverse the cell?

The answer is that all belong to the same species: Craticula cuspidata.   Image b. is the way that it is most often seen (although it is not a particularly common species in the UK).   You should be able to see the raphe and fine striae which are more-or-less parallel to one another and perpendicular to the midline of the valve.   If you look with a scanning electron microscope, you’ll see that each of the striae is composed of a series of round or elliptical pores, equidistantly spaced so that the striae may appear to be running longitudinally as well as across the valve.

Craticula-cuspidata-all-bits-Pitsford-Jan2019

Craticula cuspidatafrom Pitsford Water, January 2019.   a. isolated girdle band; b.  “normal” valve; c. valve at “heribaudii stage”; d., e.: valves at “craticulae” stage. Scale bar: 10 micrometres (= 1/100thof a millimetre).  Photos: Chris Carter.

Although the genus Craticula was described in the 19thcentury by Grunow, it was considered to be part of the genus Naviculafor most of the 20thcentury.  We now regard the strictly parallel striae as one of the characteristics of Craticula but, if you think of it within in the broader realm of “Navicula” (basically, boat-shaped diatoms with a central raphe), many of which have radiate striae, then you might be happy to consider valve c. as being related to valve b.   In this case, it would have been called “Navicula cuspidata var. heribaudii”.   However, in 1979 Anne-Marie Schmid of the University of Salzburg, grew cultures of “normal” Craticula cuspidata in increasing salt concentrations and was able to show this (and the structures seen in images d. and e.) were responses to the stresses that this caused.

Under certain conditions, it seems, the normal process of cell division breaks down so that, rather than producing two daughter cells, each composed of two silica valves, just one “internal valve” is produced so that there are, in effect, three valves for two cells.  One of the cells then degenerates leaving a single functional cell albeit with one extra valve.   This phenomenon is not confined to Craticula but seems to be better understood for this genus than for others for reasons that I will come to shortly.   In this particular case, the internal valve has a similar outline to the parent, but a different arrangement of striae

Images d. and e. show another aspect of the same phenomenon: the formation of a “craticula” (from the Latin for “grid-iron”).  Schmid showed that this stage actually happens at lower salt concentrations than the “heribaudii” stage but that it, too, is related to the formation of these “internal valves”.   There is a thickening of silica along the central rib, after which transverse “buttresses” grow out and, finally, a silica band is laid down around the edges of the valve.  Schmid suggested that the resulting structures were resting stages, noting that she had found such structures in ponds in the Namib Desert that were only wetted for short periods every other year or so.  When they dried up, salinity increased very rapidly and these “resting spores” lay in the bottom muds protected by layers of “jelly” (i.e. extracellular polysaccharides).  About 11 days after she re-suspended them in distilled water, she observed viable cells gliding around again.

In the early 1990s, it became clear for other reasons that members of this genus were quite different from Naviculaso the original name was resurrected.  That leaves us with the unusual situation of a genus that is named after rarely-seen monstrosities.   It would be akin to naming Fragilaria “twisty diatoms” because, as we saw in “A twist in the tale …” a different form of stress causes a characteristic reaction in members of that genus.    Because Craticula is not a particularly common genus, and because “craticulae” valves are a relatively rare phenomenon within that genus, it is likely that most people have never seen the structure after which it was named.

References

Mann, D.G. & Stickle, A.J. (1991).  The genus Craticula. Diatom Research6: 79-107.

Schmid, A.-M. (1979).  Influence of environmental factors on the development of the valve in diatoms.  Protoplasma99: 99-115.

Life out of water …

Last time I wrote, I mentioned that those diatom genera that did not have to be permanently submerged in order to thrive (so-called “aerophilous diatoms”) often appeared together in samples.   Having seen some Luticola muticaearly in my analysis of the sample from Castle Eden Burn, it was no surprise to find Diadesmisand Simonsenialater in the same analysis.   If anything, the biggest surprise was that I did not also find Hantzschia amphioxys, another habitué of the damp fringes of diatom society.

A quick analysis of my database puts these thoughts into context.   There are 6500 samples in my database, so we can see, from the total number of records of each of the aerophilous genera that these are relatively scarce in the samples I encounter.  That is largely because my sampling approaches are biased against the habitats where these thrive (more about this below).   Aerophilous diatoms are more common than you might think; it is scientists with a yearning to learn more about them that is in short supply.

Hantzschiaand Simonseniaare both less frequent and less abundant than the other two genera, never occurring in numbers exceeding ten per cent of the total but, when they form more than one per cent of the total, there is a very high chance that you will also find other aerophilous taxa in the sample.   Humidophilaand Luticolaare sometimes found in higher numbers, and when this is the case, then the proportion of other aerophilous taxa is also often high: 75 per cent of samples where Humidophilais abundant, for example, have at least one other aerophilous taxon present at one per cent or more.

Frequency of other aerophilous genera in samples with Hantzschia, Humidophila, Luticolaand Simonsenia.    Each genus is represented by two rows: records where it formed 10 per cent or more of the total number of valves and records where it formed more than one per cent.   Similarly, records for other aerophilous genera are also stratified into those where they comprise more than 10 per cent of the total and those where they comprise more than one per cent.  

Genus number of records   other aerophilous genera
>10% >1%
Hantzschia 147 >10% n/a n/a
>1% 0.50 0.70
Humidophila 248 >10% 0.25 0.75
>1% 0.09 0.29
Luticola 630 >10% 0.09 0.35
>1% 0.05 0.16
Simonsenia 61 >10% n/a n/a
>1% 0.50 1.00

Over the years, I have come to use this information informally as a way of knowing whether the results of an analysis are likely to be giving me useful insights into ecological condition.   Many of the samples I analyse are collected by other people and sent to me.   These samplers should have been working to protocols that ensure that they check that the stones they choose were fully submerged for some time prior to their visit.  However, the person collecting the sample may have to make a judgement about river and lake level fluctuations in the period before their visit.  Finding lots of cells of aerophilous taxa in a sample is a good hint that something is awry.

The German method for ecological status assessment actually uses the proportion of aerophilous taxa as a check on the reliability of an assessment.    I suspect that they are not the only ones, but They have a list of 46 species that they regard as aerophilous taxa, and use a threshold of five per cent in a sample as a threshold.   The genera I’ve discussed all feature prominently, along with representatives of 19 other genera. Most of these are represented by only one or two species, although there are seven species of Nitzschia, five of Pinnulariaand six of Stauroneis.   I suspect that some species on this list are more tolerant of desiccation than others. We do not know enough of the physiological mechanisms behind this tolerance but it would seem that a few genera (Hantzschia, Humidophila, Luticiola) have definitely got this hard-wired into their genotypes, whilst other genera have members which are mostly aquatic in their habit but with a few exceptions able to survive out of water for some time.   I, personally, would trust the five per cent threshold if it was restricted to the hardcore aerophilous genera, with other taxa on the list providing supporting evidence. I would also add the proviso that there should be more than one aerophilous taxon contributing to that five per cent.  I would be happier, too, if there were a few experimental studies behind these lists and thresholds but, as ever with the world of diatoms, taxonomists are several steps ahead of the physiologists and so we are heavily dependent on anecdotal information when interpreting results.

List of taxa regarded as aerophilous in the German system for assessing ecological status in rivers. 

Name Authority
Achnanthes coarctata (Brébisson) Grunow in Cleve & Grunow 1880
Chamaepinnularia parsura (Hustedt) C.E.Wetzel & Ector in Wetzel et al. 2013
Cosmioneis incognita (Krasske) Lange-Bertalot in Werum & Lange-Bertalot 2004
Denticula creticola (Østrup) Lange-Bertalot & Krammer 1993
Diploneis minuta Petersen 1928
Eolimna subadnata  (Hustedt) G. Moser, Lange-Bertalot & Metzeltin 1998
Fallacia egregia (Hustedt) D.G. Mann 1990
Fallacia insociabilis (Krasske) D.G. Mann 1990
Fistulifera pelliculosa (Brébisson ex Kützing) Lange-Bertalot 1997
Halamphora montana (Krasske) Levkov 2009
Halamphora normanii (Rabenhorst) Levkov 2009
Hantzschia abundans Lange-Bertalot 1993
Hantzschia amphioxys (Ehrenberg) Grunow 1880
Hantzschia elongata (Hantzsch) Grunow 1877
Hantzschia graciosa Lange-Bertalot 1993
Hantzschia subrupestris Lange-Bertalot 1993
Hantzschia vivacior Lange-Bertalot 1993
Humidophila aerophila (Krasske) Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot & Kopalová, 2014
Humidophila brekkaensis (J.B.Petersen) D. Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot & Kopalová, 2014
Humidophila contenta (Grunow) Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot & Kopalová, 2014
Humidophila perpusilla (Grunow) Lowe, Kociolek, Johansen, Van de Vijver, Lange-Bertalot & Kopalová, 2014
Luticola cohnii (Hilse) D.G. Mann 1990
Luticola dismutica (Hustedt) D.G.Mann1990
Luticola mutica (Kützing) D.G. Mann 1990
Luticola nivalis (Ehrenberg) D.G. Mann 1990
Luticola nivaloides (W.Bock) J.Y.Li & Y.Z.Qi 2018
Luticola paramutica (W. Bock) D.G. Mann 1990
Luticola pseudonivalis (W.Bock) Levkov, Metzeltin & A.Pavlov 2013
Luticola saxophila (W.Bock ex Hustedt) D.G.Mann 1990
Mayamaea nolensoides (W. Bock) Lange-Bertalot 2001
Melosira dickiei (Thwaites) Kützing 1849
Muelleria gibbula (Cleve) Spaulding & Stoermer 1997
Neidium minutissimum Krasske 1932
Nitzschia aerophila Hustedt 1942
Nitzschia bacillarieformis Hustedt 1922
Nitzschia disputata J.R. Carater 1971
Nitzschia harderi Husedt 1949
Nitzschia modesta Hustedt 1950
Nitzschia terrestris (J.B. Petersen) Hustedt 1934
Nitzschia valdestriata Aleem & Hustedt 1951
Orthoseira dendroteres (Ehrenberg) Genkal & Kulikovskiy in Kulikovskiy et al. 2010
Orthoseira roseana (Rabenhorst) Pfitzer 1871
Pinnularia borealis Ehrenberg 1843
Pinnularia frauenbergiana E. Reichardt 1985
Pinnularia krookii (Grunow) Hustedt 1942
Pinnularia largerstedtii (Cleve) Cleve-Euler 1934
Pinnularia obscura Krasske 1932
Simonsenia delognei (Grunow) Lange-Bertalot 1979
Stauroneis agrestis J.B. Petersen 1915
Stauroneis borrichii (J.B.Petersen) J.W.G.Lund 1946
Stauroneis gracillima Hustedt 1943
Stauroneis lundii Hustedt 1959
Stauroneis muriella J.W.G. Lund 1946
Stauroneis obtusa Lagerstedt 1873
Surrirella terricola Lange-Bertalot & Alles 1996
Tryblionella debilis Arnott ex O’Meara 1873

Reference

Schaumburg, J., Schranz, C., Steizer, D., Hofmann, G., Gutowski, A. & Forester, J. (2006).  Instruction protocol for the ecological assessment of running waters for implementation of the EC Water Framework Directive: macrophytes and phytobenthos.  Bavarian Environment Agency

Tales from a dry river bed …

Two weeks ago I stood in a dry stream bed at Castle Eden Dene, wondering at the absence of water yet also conscious that many of the stones that littered the surface had a slipperiness that suggested not only that they had been wet relatively recently, but also that the surface biofilms (which impart this slipperiness) might still be intact.   A first look at a portion of this film under my microscope suggested that this might well be the case: I could certainly see some diatoms, and some green algae cells, but most were very small and that there was also a lot of particles, both inorganic and organic, that made viewing these algae quite difficult.   Since then, I’ve prepared a permanent slide from this material, so I can now take a closer look and get a better idea of what diatoms thrive in a dry stream bed in mid-winter in northern England.

A quick analysis of the sample found 34 species, of which four were abundant (comprising over 60% of the total) and the remainder were relatively infrequent.   The most abundant species was Amphora pediculus, which I’ve written about before, and which was not a surprise, as it is a species that thrives in the hard water that I would have expected in a stream draining a limestone catchment.  The other three common species wereHumidophila contenta, Luticola muticaand Simonsenia delognei, all of which are known to survive in habitats that are not permanently submerged.   These are relatively uncommon in the typical samples that I encounter but when they do occur in large numbers, they are often found together.   It is another facet of the “London Bus” paradigm that I described in the previous post, except this time it is a characteristic assemblage of species from different genera, rather than from a single genus or family.

Castle_Eden_diatoms_Jan19

Some of the diatoms from Castle Eden Burn, January 2019: a. Nitzschia nana; b. – g. Luticola mutica; h. – k. Humidophila contenta.   Scale bar: 10 micrometres (= 1/100thof a millimetre). 

Diatoms in the genus Humidophilahas changed names twice over the course of my career.   Back in the 1980s, species from this genus, as well as Luticolawere considered to be part of the Navicula which was regarded as a “dump for all bilaterally symmetrical [e.g. boat-shaped] raphid diatoms lacking particularly distinctive features” according to Frank Round, Dick Crawford and David Mann.    They split several groups of species away from Naviculato create new genera, one of which was Luticola.  In other cases, to resurrect old genera that had been subsumed into Naviculain the first half of the 20thcentury.  One of these resurrected genera was Diadesmiswhich differed from “true” Naviculain several respects, not least of which was a tendency to form ribbon-like colonies.   A more recent study suggested that Diadesmis, itself, needed to be split, with several species being moved to yet another new genus, Humidophila.   Unfortunately, the criteria on which this was based are not easily seen with the light microscope.  However, one by-product of this split was that all the species within the genera that are associated with damp, rather than fully-submerged habitats, ended up in the new genus rather than in Diadesmis.   That lends weight to the split, suggesting that there is more to the separation than just minor differences in the details of the cell wall.

The final species that was common in Castle Eden Burn was Simonsenia delognei.   This is another small diatom and, as I could not get good photographs from this sample, I have included photographs from another site to show what it looks like.  It is a very delicate diatom, easily overlooked when scanning a slide, particularly as it usually only occurs in small numbers.  That, again, might be because I usually look at samples from fully-submerged habitats.   Here, it formed about 12 per cent of the total number of valves, which is four times as many as I have previously found.

Simonsenia_delognei.jpg

Simonsenia delogneifrom Ballyfinboy River, Co. Tipperary, August 2014.   Scale bar: 10 micrometres (= 1/100thof a millimetre).  Photographs: Lydia King.

I’m quite intrigued, now, to see how the algal communities change over the course of the year. Are these diatoms that can tolerate drying ever-presents or will their proportions fluctuate over the course of the year as the stream comes and goes?   And what is it that makes some diatoms cope with these dry periods?   The ability to live out of water is associated with a few genera in particular, so what is it about their genetic make-up that lets them thrive.  What about Amphora pediculusand the other diatoms that I associate with submerged habitats? Am I looking at dormant but viable cells (I did not see many healthy chloroplasts when I made my initial observations) or are these diatom carcasses strewn across an arid desert?    At the risk of sliding into metaphor-overload, does this mean that Humidophila, Luticolaand Simonseniaare the cacti of the diatom world?

References

Lowe, R.L., Kociolek, P., Johannsen, J.R., van de Vijver, B., Lange-Bertalot, H. & Kopalová, K. (2014).  Humidophilagen. nov., a new genus for a group of diatoms (Bacillariophyta) formerly within the genus Diadesmis: species from Hawai’I, including one new species.  Diatom Research29: 351-360.

Round, F.E., Crowford, R.M. & Mann, D.G. (1990).  The Diatoms: Biology and Morphology of the Genera.   Cambridge University Press, Cambridge.

A twist in the tale …

hamsterley_forest_jan19\

After my sojourn in East Durham, described in the previous post, I have travelled back to the Pennines for this one, crossing the River Wear at Wolsingham before driving up onto the fells and finally dropping down to the woodlands that are Hamsterley Forest.  This is a large man-made plantation, dating from the 1930s and popular for recreation. In January, however, the forest is quiet, and I only have a few mountain bikers and a lone dog walker for company as I peer into the peaty waters of Euden Beck.   This stream rises on the open fells of Hamsterley Common, between Weardale and Teesdale, before flowing through the forest and joining Spurl’s Wood Beck just downstream from where I am standing, to become Hamsterley Beck.  This then joins the Wear a few kilometres downstream from Wolsingham.

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Euden Beck, just above the forest drive in Hamsterley Forest, January 2019.  The photograph at the top of the post shows a view towards Hamsterley Forest. 

There is a mixture of diatoms growing on the stones here but I am most interested in the genus Fragilaria today.   One of the curiosities of this genus is that we often find several representatives growing at the same site at the same time, reminiscent of the old adage about London buses (“you wait ages, and then three come along at once”).   I’ve written about this before (see “Baffled by the benthos (2)” and “When is a diatom like a London bus?”) and Euden Beck is another good example of this conundrum in practice.

Today, I could see quite a few cells of Fragilaria teneraand smaller numbers ofF. gracilisplus a newly-described species that I will talk more about later in the post.  Fragilaria teneraforms long, needle-like cells, often clustering together to form sea urchin-like masses growing out from either a filamentous alga or particle to which they are attached (see “Food for thought in the River Ehen” for an illustration).  Most of the ones that I saw in my samples from Euden Beck were either single cells or pairs of cells, presumably following a recent division. Note how the second cell from the left in the figure below is not as straight as the others.   This is something that I often see with Fragilaria populations in streams in the northern Pennines, and indicates that there may be heavy metal pollution in the water.  There are a lot of abandoned lead mines in the northern Pennines and, sure enough, when I looked at a large scale map, I found one that I had not previously noticed in the upper part of Euden Beck’s catchment.

fragilaria_tenera.euden_jan19

Live cells of Fragilaria tenera(a. – d.) and F. heatherae from Euden Beck, January 2019.   a., b. and e. are valve views; c. and d. are girdle views.  Scale bar: 10 micrometres (= 1/100thof a millimetre). 

The next image shows these valve abnormalities even more clearly, with almost all of the cells showing aberrations in their outline.   These images are from an older sample; the curiosity here is that whilst most of the Fragilaria tenera valves were twisted, fewer of the valves of Fragilaria gracilisare twisted, whilst few of the valves of the third Fragilaria species show any abnomality in their outline at all.   This species is very common in northern Pennine streams, and I have often seen distorted valves of this species in streams polluted by mine discharges.  This makes the discrepancy between the outlines of this and Fragilaria tenera in Euden Beck particularly intriguing.

fragilaria_tenera_rt#55

Fragilaria tenera from a sample collected from Euden Beck in June 2012.  Scale bar: 10 micrometres (= 1/100thof a millimetre).   Photographs: Lydia King.

I say “Fragilaria gracilis” with a modicum of trepidation as a recent study in which I have been involved, suggests that there may well be at least two species.  These are, as far as we can tell, indistinguishable using characteristics that can be seen with the light microscope though we know that they are genetically quite distinct, and both are widespread, turning up not just in the UK but in other parts of Europe too.

The third species, to the best of our knowledge, does not match the description of any other Fragilaria species, and we are in the process of publishing it as a new species, Fragilaria heatherae.   We have found it a number of samples, not just from the UK but also from sites elsewhere in Europe.   These, by comparison with the other two species, show very little distortion at all.   Whilst several authors have noted this phenomenon in the past, the physiological cause is still not understood. My guess is that the metal ions are displacing a metal co-factor in an enzyme that is involved in the process of laying down the silica cell wall.   Fragilaria seems to be particularly susceptible, but this may be because their long needle-like cells show the distortions more clearly than in some genera but, based on the evidence from Euden Beck, there are clearly differences in susceptibility between species.

Once again, I seem to be ending a post having asked more questions than I have answered. That is always frustrating but another way of looking at this is to realise that the frontiers of ecology are only ever a short drive away from where you are now.  It is very nice to cross oceans to visit rain forests and coral reefs, but there are adventures to be had closer to your doorstep.

fragilaria_gracilis_rt#55

Fragilaria gracilis from a sample collected from Euden Beck in June 2012.  Scale bar: 10 micrometres (= 1/100thof a millimetre).   Photographs: Lydia King.

fragilaria_heatherae_rt#55

Fragilaria heatherae” from a sample collected in Euden Beck in June 2012.  Scale bar: 10 micrometres (= 1/100thof a millimetre).   Photographs: Lydia King

References

Duong, T.T., Morin, S., Herlory, O. & Feurtet-Mazel, A. (2008). Seasonal effects of cadmium accumulation in periphytic diatom communities of freshwater biofilms.  Aquatic Toxicology90: 19-28.

Falasco, E., Bona, F., Ginepro, M., Hlúbiková, D., Hoffmann, L. & Ector, L. (2009). Morphological abnormalities of diatom silica walls in relation to heavy metal contamination and artificial growth conditions.  Water SA35: 595-606.

McFarland, B.H., Hill, B.H. & Willingham, W.T. (1996). Abnormal Fragilaria spp. (Bacillariophyceae) in Streams Impacted by Mine Drainage. Journal of Freshwater Ecology 12: 141-149.