Life in the colonies …

Another outcome of my visit to Ennerdale Water a couple of weeks ago in July (see “Life in the Deep Zone”) was some tiny green spheres in the sample I collected from one of the small streams flowing into the lake’s north-west corner.   The stream was very short, little more than a seepage arising from a wet rush-dominated area of a field just twenty metres or so from the lake margin and, at the point which I sampled, there was a tangle of filamentous algae (Stigeoclonium, Mirsrospora and Mougeotia) as well as a distinct diatom-dominated film on exposed stones.    The colonies looked like tiny peas in my sample tray but I suspect that they were attached to rocks or aquatic vegetation before I disrupted them. Under the microscope, these turned out to be colonies of the green alga Chaetophora pisiformis, a relative of Draparnaldia and Stigeoclonium, both of which I have written about before (see “The exception that proves the rule …” and “A day out in Weardale …”.  Like those, Chaetophora has branched filaments but they differ in forming well-defined colonies that are visible to the naked eye.

The pictures below show the form of colonies very clearly.  Chaetophora colonies are firm to the touch and cannot easily be squashed under a coverslip.   I overcame this by using a cavity slide, and taking one of the smallest colonies that I could find in order to photograph it with as little damage as possible.  Note how there is a very clear edge to the colony, whereas Draparnaldia and Stigeoclonium have a mass of filaments and mucilage but no obvious border between the “colony” and the surrounding environment.  Draparnaldia sometimes forms discrete colonies (see “The exception that proves the rule …”) but these are much softer and more easily squashed onto a slide.

Top: colonies of Chaetophora pisiformis from a small stream flowing into Ennerdale Water, with a one cent coin for scale; bottom left: lower power (x40) view of a colony.  The picture frame is about two millimetres across; bottom right: medium power (x100) view of the same colony.

Viewed at higher magnifications, the branches of the filaments are clear. They tend to be clustered towards the tops of the filaments and, in this case at least, end abruptly, rather than tapering to fine hairs.  I explained in the posts mentioned above how these fine hairs are used by the algae as means of capturing the nutrients that they need.  Chaetophora can form these hairs, but it does so less often, in my experience, than Draparnaldia and Stigeoclonium.   There will be dead and decaying vegetation in the rush-dominated swamp from which the stream originates, and the enzymes that these algae produce will be able to harvest any phosphorus from organic particles that result from this decay.  That’s the theory for Stigeoclonium at least, but I suspect that the colonies of Chaetophora are also highly efficient recycling units: the filaments are embedded in a firm mucilage that is far more than an inert polysaccharide gunk.   Any phosphorus that is released from a filament will be far more likely to be hoovered up by another filament than to drift downstream whilst the phosphatase enzymes will also be on hand at the colony surface to savenge any stray nutrients from the seepage.  These tight colonial forms are, in other words, fortresses of plenty in an otherwise inhospitable landscape: well adapted to nutrient-stressed situations and, as a paucity of nutrients is the natural condition of streams, the presence of these colonies is a good sign that this stream is in good condition.

Filaments of Chaetophora pisiformis from a small stream flowing into Ennerdale Water, July 2018.  Scale bar: 20 micrometres (= 1/50th of a millimetre). 

Reference

Whitton, B.A. (1988).  Hairs in eukaryotic algae.   pp. 446-480.  In: Algae and the Aquatic Environment (edited by F.E. Round).  Biopress, Bristol.

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Some like it hot …

My reflections on algae that thrive in hot weather continued recently when I visited a river in another part of the country.  As this is the subject of an ongoing investigation, I’ll have to be rather vague about where in the country this river flows; suffice it to say it is in one of those parts of the country where the sun was shining and your correspondent returned from a day in the field with browner (okay, redder) arms than when he started.   Does that narrow it down?

A feature of some of the tributaries, in particular, was brown, filamentous growths which, in close up, could be seen to be speckled with bubbles of oxygen: a sure sign that they were busy photosynthesising.  These were most abundant in well-lit situations at the edges of streams, away from the main flow.   Under the microscope, I could see that these were dominated by the diatom Melosira varians, but there were also several filaments of the cyanobacterium Oscillatoria limosa, chains of the diatom Fragilaria cf capucina and several other green algae and diatoms present.

Melosira varians is relatively unusual as it is a diatom that can be recognised with the naked eye – the fragile filaments are very characteristic as is its habitat – well lit, low-flow conditions seem to suit it well.   It does seem to prefer nutrient-rich conditions (see “Fertile speculations …”) but it can crop up when nutrient concentrations are quite low, so long as the other habitat requirements are right for it.  The long chains of Melosira (and some other diatoms such as Fragilaria capucina and Diatoma vulgare) help the cells to become entangled with the other algae.   I could see this at some sites where the Melosira seemed to grow around a green alga that had been completely smothered by diatoms and was, I presume, withering and dying.  In other cases, the Melosira filaments are much finer and seem to attach directly to the rocks.   Neither arrangement is robust enough for Melosira to resist any more than a gentle current which is why it is often most obvious at the edges of streams and in backwaters.   As is the case for Ulva flexuosa, described in the previous post, I suspect that the first decent rainfall will flush most of this growth downstream.   Another parallel with Ulva is that, despite this apparent lack of adaptation to the harsh running water environment, Melosira varians is more common in rivers and streams than it is in lakes.

Melosira varians-dominated filaments at the margins of a stream.  Top photograph shows the filaments smothering cobbles and pebbles in the stream margins (frame width: approximately one metre); bottom photograph shows a close-up (taken underwater) of filaments with oxygen bubbles (frame width: approximately one centimetre).

Algae from the filaments illustrated above: a. and b.: Melosira varians; c. Fragilaria cf capucina; d. Oscillatoria limosa.  Scale bar: 20 micrometres (= 1/50th of a millimetre).  

The graphs below support my comments about Melosira varians preferring nutrient rich conditions to some extent.  Many of our records are from locations that have relatively high nutrient concentrations; however, there are also a number of samples where M. varians is abundant despite lower nutrient concentrations.   How do we explain this?   About twenty years ago, Barry Biggs, Jan Stevenson and Rex Lowe envisaged the niche of freshwater algae in terms of two primary factors: disturbance and resources.   “Resources” encompasses everything that the organism needs to grow, particularly nutrients and light, whilst “disturbance” covers the factors such as grazing and scour that can remove biomass.   They used this framework to describe successions of algae, from the first cells colonising a bare stone through to a thick biofilm.   As the biofilm gets thicker, so the cells on the stone get denser and, gradually, they start to compete with each other for light, leading to shifts in composition favouring species adapted to growing above their rivals (see “Change is the only constant …”).

The relationship between Melosira varians and nitrate-nitrogen (left: “NO3-N”) and dissolved phosphorus (right: “PO4-P”).   The vertical lines show the average positions of concentrations likely to support high (red), good (green), moderate (orange) and poor (red) ecological status (see note at end of post for a more detailed explanation).

They suggested that filamentous green algae were one group well adapted to the later stages of these successions but these, in turn, create additional opportunities for diatoms such as M. varians which can become entangled amongst these filaments and access more light whilst being less likely to being washed away.   If there is a period without disturbance then the Melosira can overwhelm these green algal filaments.   Nutrients, in this particular case, do play a role but, in this case, are probably secondary to other factors such as low disturbance and high light.  Using the terminology I set out in “What does it all mean?”, I would place M. varians in the very broad group “b”, with the caveat that the actual nutrient threshold below which Melosira cannot survive in streams is probably relatively low.   Remember that phosphorus, the nutrient that usually limits growth in freshwater, comprises well under one per cent of total biomass, so a milligram of phosphorus could easily be converted to 100 milligrams of biomass in a warm, stable, well-lit backwater.

Schematic diagram showing the approximate position of Melosira varians on Biggs et al.’s conceptual habitat matrix.

The final graph shows samples in my dataset where Melosira varians was particularly abundant and this broadly supports all that has gone before: Melosira is strongly associated with late summer and early autumn, when the weather provides warm, well-lit conditions with relatively few spates.

The case of Meloisra varians is probably a good example of the problem I outlined in “Eutrophic or euphytic?”  I have seen similar growths of diatoms in other rivers recently, due to the prolonged period of warm, dry conditions.  It is easy to jump to the conclusion that these rivers have a nutrient problem.  They might have, but we also need to consider other possibilities.   Like Ulva flexuosa in the previous post, Melosira varians is an alga that is enjoying the heatwave.

Distribution of Melosira varians by season.   The line represents sampling effort (percent of all samples in the dataset) and vertical bars represent samples where M. varians forms >7% of all diatoms (90th percentile of samples, ranked by relative abundance). 

Reference

Biggs, B.J.F., Stevenson, R.J. & Lowe, R.L. (1991). A habitat matrix conceptual model for stream periphyton. Archiv für Hydrobiologie 143: 21-56.

Notes on species-environment plots

These are based on interrogation of a database of 6500 river samples collected as part of DARES project.  Phosphorus standards are based on the Environment Agency’s standard measure, which is unfiltered molybdate reactive phosphorus.  This approximates to “soluble reactive phosphorus” or “orthophosphate-phosphorus” in most circumstances but the reagents will react with phosphorus attached to particles that would have been removed by membrane filtration. The current UK phosphorus standards for rivers that are used here are site specific, using altitude and alkalinity as predictors.  This means that a range of thresholds applies, depending upon the geological preferences of the species in question.  The plots here show boundaries based on the average alkalinity (50 mg L-1 CaCO3) and altitude (75 m) in the whole dataset.

There are no UK standards for nitrate-nitrogen in rivers; thresholds in this report are based on values derived using the same principles as those used to derive the phosphrus standards and give an indication of the tolerance of the species to elevated nitrogen concentrations.  However, they have no regulatory significance.

 

 

Eutrophic or euphytic?

A paper has just been published that should be required reading for anyone interested in the management of nutrients in in ecology.   It is a follow-up of a 2006 paper with the catchy title “How green is my river” that set out to provide a conceptual framework for how rivers responded to enrichment by nutrients.   That original paper contained several good ideas but, crucially, not all of them were underpinned by evidence.  A decade on, several of the predictions and statements made in that original paper have been tested, and the time has come to re-examine and modify that original conceptual model.

My reaction to the 2006 paper was that it was very interesting but not fully reflective of the rivers in my part of Britain, whose rougher topography produced quite different responses to nutrient enrichment than that proposed in their original model.   That criticism has been addressed in the revised version, which places greater emphasis on the physical habitat template, which means that it is more broadly applicable than the original version.   But that, in turn, got me wondering about the continued relevance of a term such as “eutrophication” to rivers.

People have been using the term “eutrophic” to describe lakes with high concentrations of nutrients since early in the 20th century.   As the century progressed, evidence of a causal relationship between inorganic nutrients and algal biomass, and the consequences for other components of lake ecosystems grew.   With this foundation, it has then become possible to predict the benefits of reducing nutrients and there are plenty of case studies, particularly from deep lakes, that demonstrate real improvements as nutrient concentrations have declined.

Attempts to apply the same rationale to rivers have, however, met with far less success.   Legislation to reduce nutrients in rivers has been in force in Europe since 1991 (the Urban Wastewater Treatment Directive, followed by the Water Framework Directive) and whilst this has led to reductions in concentrations of phosphorus in rivers (see  “The state of things, part 2”), there has, in most cases, not been a corresponding improvement in ecology.   There are a number of reasons for this but, at the heart, there was a failure to understand that the tight coupling between nutrients and biology that was the case in deep lakes did not also occur in running waters.   What was needed was recognition of fundamental differences between lakes and rivers, and “How green is my river?” and, now, this new paper have both contributed to this.

However, one consequence of recognising the importance of the physical habitat template alongside nutrients is to challenge the relevance of the term “eutrophic” when describing rivers.   “Eutrophic” literally means “well-nourished” so is appropriate in situations where high nutrients cause high plant or algal biomass.   This high biomass (strictly speaking, the primary production arising from this biomass) then creates problems for the rest of the ecosystem (night-time anoxia caused by plants consuming oxygen being a good example).   If high biomass can arise due to, let’s say, removal of bankside shade or alteration to the flow regime, perhaps (but not always) in combination with nutrients, then perhaps we need a term that does not imply a naïve cause-effect relationship with a single pressure?

My suggestion is to shift the focus from nutrients to plant growth by using the term “euphytic” (“too many plants”) as this would shift the emphasis from simply driving down nutrient concentrations (expensive and not always successful) towards reducing secondary effects.  It is possible that strategies such as planting more bankside trees, for example, or altering the flow regime or channel morphology (see “An embarrassment of riches …”) may be just as beneficial, in some cases, as reducing nutrient concentrations.   That said, we also have to bear in mind that nutrients may have an effect well downstream, so focus on amelioration of effects within a particular stream segment will never be a complete solution.

I should emphasise that a lot of work has been done in recent years to understand the concentrations of nutrients that should be expected in undisturbed conditions, and also to understand the nutrient concentrations that lead to changes in community structure in both macrophytes and algae.   These show that many rivers around Europe do have elevated concentrations of nutrients and I am not trying to side-step these issues.  I do, however, think it is important that regulators can prioritise those rivers in greatest need of remediation and, in most cases, they do this without considering the risk of secondary effects.

It is, largely, a matter of semantics.   I have been involved in many conversations over the past couple of decades about how to improve the state of our rivers.  Many of those have centred on the importance of reducing nutrient concentrations (which would be, indisputably, a major step towards healthier rivers).  But there is more to it than that.  And Mattie O’Hare and colleagues are helping to open up some new vistas in this paper.

Note: the photograph at the top of this post shows the River Wear at Wolsingham.  This stretch of the river captures many of the challenges facing river ecologists: nutrient concentrations are relatively low and there is good bankside shade.  However, the flow of the river is highly altered due to impoundments upstream and a major water transfer scheme.  How do all these factors interact to create the often prolific algal growths that can be seen here, particularly in winter and spring?

References

Hilton, J., O’Hare, M., Bowes, M.J. & Jones, J.I. (2006).  How green is my river?  A new paradigm of eutrophication in rivers.   Science of the Total Environment 365: 66-83.

O’Hare, M.T., Baattrup-Pedersen, A., Baumgarte, I., Freeman, A., Gunn, I.D.M., Lázár, A.N., Wade, A.J. & Bowes, M.J. (2018).  Responses of aquatic plants to eutrophication in rivers: a revised conceptual model.   Frontiers in Plant Science.   9: 451

Unlikely neighbours …

One of the lessons I learned from writing “A tale of two diatoms …” is that we can often learn more about the ecology of a species by contrasting its behaviour with that of another species rather than by just relating the distribution of that species to features of its environment.  I came across another example of this when I was writing up the results of the latest “ring-test” that UK diatom analysts undertake to maintain their competence.

The sample came from a stream in east Devon (the one that had a walk-on part in “The challenging ecology of a freshwater diatom”).  This stream receives effluent from a small sewage works but our sample comes from just upstream of this works.   We know that the stream downstream of the sewage works is quite polluted but were also interested in the condition of the stream above the works.   This has proved to be challenging and, it seems, there are some pollution sources, including septic tanks and runoff from fields, that mean that the stream already shows signs of impact before it reaches the sewage works.   There are, however, mixed messages when we look at the aquatic flora, and some of the diatoms that are abundant are characteristic of low or only slight enrichment.

One feature of the stream that was quite unusual was a relatively large number of cells of Reimeria uniseriata, a relative of Reimeria sinuata which is quite common.  Both of these are illustrated below: note that R. uniseriata tends to be slightly larger and has distinctly punctate striae.  However, when I looked at the distribution of these species in response to water chemistry, I could see few differences, with most of the records suggesting a preference for water with low or slightly elevated phosphorus concentrations.   Reimeria sinuata is more common than R. uniseriata and when the latter is found, the former is usually present too.  They seem to be able to share their habitat quite comfortably.

Reimeria sinuata from Polly Brook, Devon, December 2016.   a. – f.: valve views; g.: girdle view focussed on ventral side.  Scale bar: 10 micrometres (= 1/100th of millimetre).  Photos: Lydia King.

Reimeria uniseriata from Polly Brook, December 2016.  h., i.: valve views; j.: girdle view focussed on dorsal side; k., l.: girdle views focussed on ventral side.   Scale bar: 10 µm (= 1/100th of millimetre).  Photos: Lydia King.

In other words, we cannot learn very much from looking at differences in the distribution of these two species of Reimeria, given our current state of knowledge.  There is, however, one other “compare and contrast” within the data that I collected from Polly Brook that is more intriguing.   If Reimeria sinuata, in particular, usually indicates a healthy stream, possibly with a little nutrient enrichment, Rhoicosphenia abbreviata is more often associated with enriched conditions.   We have met this diatom before (see “Cladophora and friends” amongst other posts) and I have explained that it is often found growing as an epiphyte on other algae.  We rarely see situations where both species are abundant at the same time, as the graph below shows.

The relative distribution of Reimeria sinuata and Rhoicosphenia abbreviata in the 6500 UK stream and river samples in the DARES dataset.   The horizontal and vertical lines indicate 10% relative abundance of each species.

When I started looking at stream algae there was a prevailing assumption that there were strong causal relationships between the species of diatom that were found at a site and the level of chemical pressures.  In the case of phosphorus, in particular, I am now not convinced that the evidence supports this whilst, at the same time, am more convinced that we should be able to, at the very least, describe what a healthy stream algal community looks like and give reasons.  I use the word “describe” because I think that many of us have been preoccupied with counting and measuring, often at the expense of a qualitative understanding.  These two species illustrate my point as when I look down a microscope and see Reimeria sinuata, I can usually assume that the stream where it was growing was reasonably healthy, even if the nutrients are a little higher than would be ideal.  On the other hand, seeing lots of cells of Rhoicosphenia makes me suspect that there has been a breakdown in the functioning of the healthy community.  These conclusions would be irrespective of what the chemistry or the values that biological indices told me.

Two species is barely enough to base a credible assessment upon but we could stir more into the mix: I often find Reimeria sinuata with Achnanthidium minutissimum, and that, in in summer especially, suggests strong top-down control by grazers, which means that pathways of energy flow have not been disrupted.   And Rhoicosphenia, as I have already mentioned, is associated with Cladophora which, in abundance, suggests a breakdown in these pathways, as shown by Michael Sturt and colleagues from University College, Cork, a few years ago.   That Polly Brook has both Reimeria and Rhoicosphenia in abundance suggests that it might just be at the tipping point between these two states.

The naïve answer to making sure that the upper stretches of Polly Brook do not cross this threshold would be to manage the nutrients.  However that is not quite as easy as it sounds in an agricultural catchment.   It could be that managing other aspects of the riparian environment are equally effective at keeping the stream in a healthy condition but that takes us into areas where the evidence is still accumulating.  It could be that the simplistic determinism that drove much of the development of biological assessment methods actually held back the gathering of that evidence for a long time.  Reimeria sinuata – and it’s cousin, R. uniseriata – stand as two reminders that there is more to the management of aquatic ecosystems than strong correlations.

Reference

Sturt, M.M., Jansen, M.A.K. & Harrison, S.A.C. (2011).  Invertebrate grazing and riparian shade as controllers of nuisance algae in a eutrophic river.  Freshwater Biology 56: 2580-2593.

The challenging ecology of a freshwater diatom?

amp_pedi_pollybrook

Amphora pediculus from Polly Brook, Devon, December 2016. Scale bar: 10 micrometres (= 1/100th of a millimetre).

The images above show one of the commonest diatoms that I find in UK waters.  It is a tiny organism, often less than 1/100th of a millimetre long, which means that it tests the limits of the camera on my microscope.  In recent months, however, it is not just the details on Amphora pediculus’ cell wall that I am struggling to resolve: I also find myself wondering how well we really understand its ecology.

The received wisdom is that Amphora pediculus favours hard water, does not like organic pollution and is relatively tolerant of elevated concentrations of inorganic nutrients.  This made it a very useful indicator species in a period of my career when we were using diatoms to identify sewage work s where investment in nutrient-removal technology might yield ecological benefits.  There were many nutrient-rich rivers, particularly in the lowlands, where any sample scraped from the upper surface of a stone was dominated by these tiny orange-segment-shaped diatom valves.   Unfortunately, twenty years on, many of those same rivers have much lower concentrations of nutrients (see “The state of things, part 2”) but still have plenty of Amphora pediculus.   Did I get the ecology of this species wrong?

The graph below shows some data from the early- and mid- 1990s showing how the abundance of Amphora pediculus was related to phosphorus.   The vertical lines on this graph show the average position of the boundaries between phosphorus classes based on current UK standards.   Records for A. pediculus are clustered in the “moderate” and “poor” classes, supporting my initial assertion that this species is a good indicator of nutrient-enriched conditions, but there are also samples outside this range where it is also abundant, so A. pediculus is only really useful when it is one of a number of strands of evidence.

aped_v_p

The relationship between Amphora pediculus and reactive phosphorus in UK rivers, based on data collected in the early-mid 1990s.  Vertical lines show the average boundaries between high and good (blue), good and moderate (green), moderate and poor (orange) and poor and bad (red) status classes based on current UK standards and the two arrows show the optima based on this dataset (right) and data collected in the mid-2000s (left).

If we weight each phosphorus measurement in the dataset by the proportion of Amphora pediculus at the same site (i.e. so that sites where A. pediculus is abundant are given greater weight), we get an idea of the point on the phosphorus gradient where A. pediculus is most abundant.   We can then infer that this is the point at which conditions are most suitable for the species to thrive.  In ecologist’s shorthand, this is called the “optimum” and, based on these data, we can conclude that the optimum for A. pediculus is 154 ug L-1 phosphorus.  The right hand arrow indicates this point on the graph below. However, I then repeated this exercise using another, larger, dataset, collected in the mid-2000s.   This yielded an optimum of 57 ug L-1 phosphorus (the left hand arrow on the graph), less than half of that suggested by the 1990s dataset.   There are, I think, two possible explanations:

First, the 1990s phosphorus gradient was based on single phosphorus samples collected at the same time that the diatom sample was collected (mostly spring, summer and autumn) whilst the mid-2000s phosphorus gradient was based (mostly) on the average of 12 monthly samples.  As phosphorus concentrations, particularly in lowland rivers, tend to be higher in summer than at other times of the year, it is possible that part of the difference between the two arrows is a result of different approaches.  (For context, in the 1990s, when I first started looking at the effect of nutrients in rivers, phosphorus was not routinely measured in many rivers, so we had no option but to do the analyses ourselves, and certainly did not have the budget or time to collect monthly samples).

However, another possibility is that the widespread introduction of phosphorus stripping in lowland rivers in the period between the mid-1990s and mid-2000s means that the average concentration of phosphorus in the rivers where conditions favour Amphora pediculus have fallen.   In other words, A. pediculus is tolerant of high nutrient conditions but is not that bothered about the actual concentration.   My guess is that it thrives under nutrient-rich conditions so long as the water is well-oxygenated and, as biochemical oxygen demand is generally falling, and dissolved oxygen concentrations rising (see “The state of things, part 1”), this criterion, too is widely fulfilled.   I suspect that both factors probably contribute to the change in optima.

But the second point in particular raises a different challenge:  We often slip into casual use of language that implies a causal relationship between a pressure such as phosphorus and biological variables whereas, in truth, we are looking at correlations between two variables.   Causal relationships are, in any case, quite hard to establish and the effect that we call “eutrophication” is really the result of interactions between a number of factors acting on the biology.   All of these simplifications mean that it is useful, from time to time, to look back to see if assumptions made in the past still hold.   In this case, I suspect that some of our indices might need a little fine-tuning.  There is no disgrace in this: the evidence we had in the 1990s led us to both to a conclusion about the relative sensitivity of Amphora pediculus to nutrients but also fed into a large-scale “natural experiment” in which nutrient levels in UK rivers were steadily reduced.   When we evaluate the results of that natural experiment we see we need to adjust our hypotheses.  That’s the nature of science.  As the sign on the door of a friend who is a parasitologist reads: “if we knew what we were doing, it wouldn’t be research”.

References

The 1990s dataset (89 records) is mostly based on data used in:

Kelly M.G. & Whitton B.A. (1995).   A new diatom index for monitoring eutrophication in rivers.   Journal of Applied Phycology 7: 433-444.

The mid-2000s dataset (1145 records) comes from:

Kelly, M.G., Juggins, S., Guthrie, R., Pritchard, S., Jamieson, B.J., Rippey, B, Hirst, H & Yallop, M.L. (2008).   Assessment of ecological status in UK rivers using diatoms.   Freshwater Biology 53: 403-422.

Hard science in hard water?

Having started to think about the ecology of small Fragilaroid diatoms in a recent post (see “When is a diatom like a London bus?”), I thought that it might pay to look in more detail at the habitats that these taxa do like, in the hope that this will help us to understand why they occur together so often.   I am just looking at two “species” in this post: “Staurosirella pinnata” (which we suspect to be a complex of several species) and “Staurosira construens” (which is also a complex, as the records in my database merge a number of varieties, most of which have subsequently been raised to the status of species in their own right).

One problem has to be confronted at the outset: these taxa also share a propensity to form chains which remain intact even after we’ve made slides.   This means that we often encounter aggregates of five or more cells, which violates the assumptions of random distributions of diatoms that underpin our statistical methods.   No-one, to my knowledge, has found a satisfactory means of dealing with this, but it should be borne in mind when considering the graphs which follow.

The first graph shows the distribution of records of these species in my database along an alkalinity gradient, and generally confirm the preference of both species for hard water.   I have highlighted two outliers on the chart for Staurosira construens. These samples are from the same location, the upper reaches of the River Wey (South) in Surrey, which receive a mixture of soft water, flowing off the Greensand, and harder water from the surrounding areas.   I have encountered anomalies between diatoms and water chemistry in this area before, which are probably the result of the complex hydrology of the area.

Staurosira_versus_alkalinity

The distribution of Staurosirella pinnata (left) and Staurosira construens (right) along an alkalinity gradient. Records from the “DARES” dataset.   Two outliers from the River Wey (South) are highlighted.

The next two graphs show the distribution of records along phosphorus and nitrogen gradients and these show opposite responses: both seem to be most abundant when phosphorus is low and nitrogen is high. Again, we have the problem of the two outliers from soft water sites confusing the view for Staurosira construens but we can generalise and say that neither species is likely to be abundant (meaning > 10 per cent of all valves) except when these conditions are met.

The horizontal red lines on these graphs show the range of phosphorus and nitrogen measured in a single river, the River Wylye, during a study in 2011-2012. I have included these lines to give a rough idea of the precision that we should expect when defining the preferences of a diatom.   The River Wylye is a chalk stream, which tend to have relatively stable hydrology, so the range of nutrient concentrations measured in these streams is probably lower than is the case for many rivers.

Staurosira_versus_PO4-P

The distribution of Staurosirella pinnata (left) and Staurosira construens (right) along an reactive phosphorus gradient. Records from the “DARES” dataset.   Vertical lines represent the approximate position of high (blue), good (green), moderate (orange) and poor (red) status boundaries.   The horizontal line shows the range of concentrations encountered in the River Wylye, Wiltshire in 2011-2012.

Staurosira_versus_nitrate

The distribution of Staurosirella pinnata (left) and Staurosira construens (right) along a nitrate-N gradient. Records from the “DARES” dataset.   Vertical lines represent the position of the (Irish) high (blue) and good (green) status boundaries.   The horizontal line shows the range of concentrations encountered in the River Wylye, Wiltshire in 2011-2012

Ecological assessment using diatoms is largely based on indices that calculate the relative position of a sample along a quality gradient based on a combination of the known ecology of the species and the representation of that species in the sample.   This means that the result is most strongly influenced by the most common species and anything that occurs below about five per cent has little influence. These charts suggest that Staurosirella pinnata and Staurosira construens will both be good indicators of a combination of low phosphorus and high nitrogen in hard water; however, there are a “tail” of records that extend into other types of water.   One valid question is whether the individuals responsible for these occurrences outside the “optimum” are the same species as those that are abundant at low P / high N / hard water.   Given what I wrote above about both of these taxa probably being complexes, this is a possibility.   However, the generally low numbers means that solving taxonomic riddles will be unlikely to lead to a great increase in precision in ecological assessments.

Personally, I lean towards the options I suggested in Baffled by the benthos (2) – that diversity within samples may be controlled by a wide range of factors unrelated to anthropogenic pressures and that interspecific diversity may give insights into ecological resilience. The problem is that this hypothesis is easier to propose than it is to test. It is not impossible to test; however, the hegemony of taxonomically-inclined diatomists over those with a genuine interest in functional ecology means that will probably remain no more than a theory for some time to come …

Michael McCarthy: Nature Studies

Here’s a link to an interesting article in today’s Independent about the state of England’s chalk streams.   They are not, says Environment Editor Michael McCarthy, in a very good condition, pointing to the problems of “diffuse pollution” (a catch-all phrase for all the pollutants that find their way into our rivers across the land, rather than via industrial or sewage effluents).   Principal amongst these is the run-off of agricultural chemicals, particularly nutrients.   This leads, says McCarthy, quoting members of the Salmon and Trout Association, to the growth of “blanket weed”, filamentous algae such as Cladophora glomerata.  Blanket weed smothers the gravel on the river bed which, in turn, makes life difficult for the bugs on which the trout and salmon feed.

Wylye_KD_110510_1_squeezed

The River Wylye at Kingston Deverill: a classic example of an English chalk stream. photographed in May 2010.

All this forms a narrative that has been repeated many times by river users and environmental scientists, including many in the Environment Agency.  The condition even has a name: Chalk Stream Malaise.   The problem is that there is no “smoking gun” to link the run-off to the decline of salmon and trout beyond reasonable doubt.   I had to research the literature on this as part of a study a few years back and was surprised about how little hard evidence I could find.  The circumstantial evidence is strong, so long as you cast your net wider than just English chalk streams, but I could not find one paper in a peer-reviewed literature that demonstrated an unambiguous link between nutrients and blanket weed, or between blanket weed and salmon and trout populations, specifically in chalk streams.   The reality is that there is much else happening in the surrounding catchments, including over-abstraction by water companies, which also affects stream ecology.   Nor are nutrients the sole consequence of increased agricultural production: silt and pesticides also enter the rivers and have their own effects, all of which are difficult to disentangle from one another. The result is a plot of interweaving motives and alibis that would make Hercule Poirot blanch.

See also streams of consciousness, my post from 21 August.