Two-faced diatoms …

Back in March I reflected on the challenges involved in discriminating species of Gomphonema (see “Baffling biodiversity …”).   That there were several species in the sample which prompted the article was indisputable; that some of those species were, individually, quite variable was also clear.  The former issue I resolved, to some extent, by reference back to Hutchinson’s “Paradox of the Plankton” but the latter was harder to explain.

Part of the problem stems, I suspect, from the reliance on morphology to characterise species.  We assume that, because a group of organisms share a set of visible characteristics, then they must also share genes which determine those characteristics and that, in turn, implies a common ancestry.   Turning that assumption on its head, we assume that groups of microscopic algae that appear different to each other belong to different species.   However, a dog lover might point out that Chihuahuas and Great Danes look very different but are, in fact, the same species.   One of the challenges of those of us who study algae is deciding just how much variation in form is typical within a species, and at what point differences are such that they represent more than one species.

Gomphonema sarcophagus from Pitsford Water, Northamptonshire, showing Janus cells.  Photographs by Ingrid Jüttner.  Scale bar: 10 micrometres (= 1/100th of a millimetre).

So what should we make of the diatom valves in the image above?   The valve outlines and breadths are similar but the striae densities are so different that we might think that they belong to two separate species.   However, I recently stumbled, by chance, on a 1998 paper by Stacy McBride and Robert Edgar which discussed the topic of “Janus cells”.  Janus, you may remember, is the Roman god of time and is depicted with two faces, one looking back to the past and the looking to the future. His name has been appropriated, in this context, to describe diatoms that have frustules comprising two valves with different characteristics.   A few genera show consistent differences between the two valves – in Cocconeis and Planothidium, for example, one valve has a raphe whilst the other does not – and there are also differences in striae densities between the raphe and rapheless valves.   The term “Janus cell” is applied to diatoms where there are marked differences between the two valves but this is not a fundamental characteristic of the species or genus.   So, in the example above, we see some forms with much denser striae (11-13 in 10 mm) than others (7-8 in 10 mm).

We don’t know, from just looking at variability in populations, that this is not polymorphism within the species, in much the same way that some humans have attached ear lobes and others do not.   But, as diatom populations grow in number by repeated divisions of single cells, we can assume that most are clones of a small number of genotypes and, therefore, that the differences are due to ontogenetic variation.   What is interesting here is that this variation seems to create two distinct outcomes – coarsely or finely striated valves.  Some have suggested that such variation may be determined by differences in environmental conditions; however, the co-existence in a single population argues against this.

Gomphonema, as I have mentioned in earlier posts, is a genus that challenges taxonomists.  And, because ecologists depend upon taxonomists to give them a means of sorting diatom valves and frustules into meaningful categories, the environmental signals we get from Gomphonema species are often quite confused too.   The possibility of encountering Janus cells just throws one more curve ball into the mix.

Reference

McBride, S.A. & Edgar, B.K. (1998).   Janus cells unveiled: frustular morphometric variability in Gomphonema angustatum.   Diatom Research 13:293-310.

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

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Summertime blues …

My reflections on the effects of the heatwave on freshwater algae continued with the latest of my regular visits to the River Wear at Wolsingham.  A comparison of the picture above with that at the head of “Spring comes slowly up this way …” says it all: the sun was shining and the gravel berms that I usually use to enter the river were occupied by families with barbeques whilst their children splashed around in the water.   At times such as this, a grown man picking up stones and then vigorously brushing their tops with a toothbrush would have invited too many questions, so I slunk off 100 metres or so downstream and found a quieter spot to explore.

The biofilm in the main channel of the River Wear at Wolsingham, July 2018. 

The first thing I noticed was that the biofilm coating the submerged stones at the bottom of the river had a greenish tinge, rather than its usual chocolate brown appearance.  It also was more crusty and less slimy to the touch than I usually see in this river.  When I got a specimen under the microscope, I could see that the composition was completely different to that which I had observed in previous months.   Most samples from this location that I’ve looked at in the past have been dominated by diatoms, with occasional spring flourishes of filamentous green algae.  Today, however, the sample was dominated by small green algae – a group that I am not very confident at identifying.   My rough estimate is that these formed about three quarters of all the algae that I could see, with diatoms and cyanobacteria each accounting for about half of the remainder.   The most abundant greens were a tiny single-celled alga that I tentatively identified as Keratococcus bicaudatus, along with a species of Scenedesmus and Desmococcus communis.   There were also a number of cells of Monoraphidium arcuatum and some of Ankistrodesmus sp.

Two views of biofilms from the River Wear, Wolsingham in January 2018.   Left: from the main channel; right: from pools at the edge of the channel.

Green algae from the River Wear at Wolsingham, July 2018: a. Desmococcus communis; b. Monoraphidium arcuatum; c. Scenedesmus sp.; d. unidentified, possibly Keratococcus bicaudatus.  Scale bar: 10 micrometres (= 1/100th of a millimetre).

However, there were also pools at the side of the channel, away from the main current but not so cut off that they were isolated from the river itself.   These were dominated by dense, brown filamentous growths, very similar in appearance to the Melosira varians flocs I described in “Some like it hot …”.  The filaments, however, felt coarser to the touch and, in close-up, could be seen to be branched, even without recourse to a microscope.   Once I got these under the microscope, I could see that they were filaments of Cladophora glomerata, another green alga, but so smothered with epiphytic diatoms (mostly Cocconeis pediculus) that they appeared brown in colour.

This combination of Cladophora glomerata and Cocconeis pediculus in the backwaters were as much of a surprise as the green-algae-dominated biofilms in the main channel.   These are species usually associated with enriched rivers (see “Cladophora and friends”) and, whilst I have seen Cladophora in the upper Wear before, it is an unusual occurrence.   Just as for the prolific growths of Melosira varians described in “Some like it hot …” it is tempting to leap to the conclusion that this must be a sign that the river is nutrient-rich.  However, the same conditions will apply here as there: “nutrients” are not the only resource that can limit plant growth and a steady trickle of phosphorus combined with warm, sunny conditions is just as likely to lead to prolific growths as a more conventionally “polluted” river.

Cladophora filaments smothered by the diatom Cocconeis pediculus in a pool beside the River Wear at Wolsingham, July 2017.   The frame width of the upper image is about 1 cm; the scale bar on the lower images is 20 micrometres (= 1/50th of a millimetre).

Another way to think of these situations is that, just as even healthy people are occasionally ill, so healthy streams can go through short periods when, based on a quick examination of plants and animals present, they exhibit symptoms associated with polluted conditions or simply (as for the first sample I described) different to what we usually expect to find.   A pulse of pollution might have passed downstream or, as seems to be happening at the moment, an unusual set of conditions lad to different organisms thriving.   Just as the ability to fight off infection forms part of a doctor’s understanding of “health”, so I expect that the River Wear will, in a few weeks time, be back to its usual state.   Healthy ecosystems, just like healthy humans, show “resilience”.   The irony is that, in this case, the “symptoms” are most obvious at a time when we are enjoying a summer better than any we’ve had in recent years.

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.

 

 

A question of scale …

It has taken some time to convert the observations from my last visit to the River Wear (see “Spring comes slowly up this way …”) into a picture.  Then, if you remember, the river was balanced between its “spring” and “summer” guises, the cool, wet weather that we experienced in March seems to have held the plants and animals that I usually see at this time of year back.   The result was a patchiness that was easy to see with the naked eye, but harder to visualise at the microscopic level.

First there were quite a few diatoms, Achnanthidium minutissimum in particular, – that suggested a thin biofilm subject to grazing by invertebrates (and I could see some chironomid larvae moving amongst the biofilm as I was sampling).   However, there were also diatoms such as Ulnaria ulna and Gomphonema olivaceum that suggested a thicker biofilm.    And finally there were filaments of the green alga Stigeoclonium tenue, mostly in discrete patches.   I never see healthy filaments of Stigeoclonium tenue smothered in epiphytes, which I have always assumed to be due to the copious mucilage that surrounds the plants.  However, I wondered if, nonetheless, Stigeoclonium contributes to overall habitat patchiness for the diatoms, as they subtly alter the way that water flows across the stone, reducing drag and shear stress in a way that favours Gomphonema and Ulnaria.   This is just speculation, of course…

That brings me back to a familiar theme: the problems of understanding the structure of the microscopic world (see “The River Wear in January” and “Baffled by the benthos (1)”) and, tangentially, to a paper on organisms’ responses to climate that was quoted at a scientific meeting I attended recently.   In this, Kristen Potter and colleagues demonstrated that there was typically a 1000 to 10,000 fold difference between the scale at which the distribution of organisms is studied and the size of those organisms.   That might be enough to draw out some coarse-scale patterns in distribution of species, but organisms actually live in microclimates, which may be patchy and which can often be quite different to the prevailing macroclimate (the difference between being exposed to full sun in open grassland and in the shade of a forest being a good example).   They suggested that the ideal spatial resolution is between one and ten times the organism’s length/height.

I see no reason why the same challenge should not also apply to the pressures faced by organisms in rivers where, again, we can get a certain amount of useful information from a coarse analysis of distribution in relation to (let’s say) average nutrient concentrations in a reach, but cannot really understand the reasons behind the spatial and temporal variation that we see in our data.  This mismatch between the scale at which organisms respond and the scale at which we study them is, I suspect, an even bigger problem for those of us who study the microscopic world.

A second illustration came at the same meeting in a talk by Honor Prentice from the University of Lund in Sweden.  She was dabbling in molecular biology years before this became a fashionable pastime for ecologists and has, over her career, developed some fascinating insights into how the structure of both plant communities and populations of individuals vary over short distances.  Her work has focussed on the island of Öland in Sweden which has the largest extent of alvar (limestone pavement) in Europe.   The system of grikes (the slabs) and clints (the fissures which separate the grikes) create quite different microclimates – the cool, moist conditions in the latter can create bog-like conditions with much lower pH than the limestone clints.   These differences influence not just the composition of the community but the genetic structure of species within these communities too.  I left thinking that if she could detect such differences at a scale barely more than one order of magnitude greater than the organisms, then how much more variation am I missing, with perhaps a five order of magnitude difference between organism size and sampling scale?

Based on these two studies, we would need to sample biofilms at a scale of about 1 mm2 in order to get a meaningful understanding of habitat patchiness in stream benthic algae.  That might just be possible with Next Generation Sequencing technologies, though I am not sure how one would go about collecting environmental data at that scale needed to explain what is going on.  Meanwhile, I am left with the coarse approach to sampling that is inevitable when you are five orders of magnitude bigger than the organism that you want to collect, and my imagination.

References

Potter, K.A., Woods, H.A. & Pincebourde, S. (2013).  Microclimatic changes in global change biology.   Global Change Biology 19: 2932-2939.

Prentice, H.C., Lonn, M., Lefkovitch, L.P. & Runyeon, H. (1995).   Associations between allele frequencies in Festuca ovina and habitat variation in the alvar grasslands on the Baltic island of OlandJournal of Ecology 83: 391-402.

 

Return to Cyprus …

A few weeks ago I described some of the algae that I found during a visit to the Avgás Gorge (pictured above) in Cyprus, including a chain-forming Ulnaria (see “Cypriot delights …”).   I’ve now had a chance to prepare cleaned valves from this material so we can take a closer look.

The chain-forming habit had already led David Williams to suggest Ulnaria ungeriana (Grunow) Compère 2001 and more detailed observations have confirmed this.  This is a species that was actually first described from Cyprus (actually Northern Cyprus) and it was also recorded quite extensively during a survey of the island’s diatoms a few years ago.   Unfortunately, some of the key diagnostic characters – such as small marginal spines and striae composed of single rows of pores – cannot be seen with light microscopy but the former, at least, can be inferred from the chain-forming habit.   Note, too, how the long chains that dominated the population in the live state, fell apart when the sample was cleaned with oxidising agents and I did not see more than three cells joined together in the new preparation.

Ulnaria ungeriana from Avgás Gorge, Cyprus, April 2018.   Scale bar: 10 micrometres (= 1/100th of a millimetre).

The Ulnaria ungeriana cells are mostly about 100 – 150 mm long and 7-8 mm wide, with a striae density of 9-10 / 10 mm.   They have parallel sides, narrowing to rostrate to slightly sub-capitate ends, and central areas that reach to the valve margin and which are slightly longer than they are broad.   Unfortunately, most of these characteristics overlap with those of Ulnaria ulna in all but most recent identification guides.   This species was first described by Nitzsch in 1817; it would have been one of the more conspicuous diatoms visible with the relatively basic equipment available at the time, with a magnification of about 150x.  His drawings are of live cells, mostly in girdle view, which means that many of the details which modern diatomists use to discriminate species are not apparent.   Moreover, the material on which these drawings are based is no longer available so we cannot go back to this in order to ascertain the characteristics of the original Ulnaria ulna and, to increase the confusion yet further, it is possible that Nitzsch has illustrated more than one species (see the reference by Lange-Bertalot and Ulrich below).

It would be, in short, very easy to look at a population of Ulnaria ungeriana in the cleaned state and match it to the descriptions of Ulnaria ulna which, under various names, have appeared in the identification literature over the past 100 years or so.   You might just detect the small marginal spines if you have a good microscope and know what you are looking for.  In the live state, however, the ribbon-like colonies are a very distinctive feature yet these do not survive preparation, putting anyone who only encounters this species on a permanent slide at a distinct disadvantage.   It is a good example of how examination of live material can add valuable information to an understanding of a diatom species yet, inevitably, many diatomists make little time for examination of their samples before dropping them into their bubbling cauldrons of oxidising agents.

High magnification views of the ends and central portions of Ulnaria ungeriana valves.   Scale bar: 10 micrometres (= 1/100th of a millimetre). 

What do we know about the ecology of Ulnaria ungeriana?   Our survey of Cypriot streams a few years ago yielded 11 records, forming up to four percent of all diatoms in the sample.  This means it is both less widespread and less dominant in samples than some other Ulnaria species.   It was often found along with other Ulnaria species, in particular U. mondii and, though generally not associated with reference sites (one out of the 11 records), it was mostly found in relatively clean conditions.   It was also associated with sites with high conductivity, which corresponds with the limestone geology that we saw in the Avgás Gorge.   On the whole, these environmental preferences are similar to those of other Ulnaria species from Cyprus that we’ve studied (see reference in earlier post).

The last question is perhaps the hardest to answer.  What benefit does the chain-forming habit confer upon Ulnaria ungeriana?   Ulnaria often forms tufts of upright cells sharing a common pad of mucilage at the base, and it is often (but not exclusively) found as an epiphyte on other plants.   We can’t rule out the possibility that the Ulnaria ungeriana chains are not also attached at one end, but it is also possible that the chain-forming habit means that they are easily entangled with the Chara and filamentous green algae that I described in the earlier post.   Both mucilage pads and entangled chains fulfil the same role of keeping the alga in the same spot in the stream, particularly where there are other plants and filamentous algae to offer extra protection from the current.

There is some speculation in the final couple of sentences but that’s never a bad thing for an ecologist.  If nothing else, it provides me with a reason to return one day …

Ecological preferences of Ulnaria ungeriana at running water sites in Cyprus.  a. pH; b. conductivity; c. total nitrogen (TN) and d. total phosphorus (TP).  Arrows indicate the mean value for each variable, weighted by the relative abundance of Ulnaria ungeriana in the sample.

Reference

Krammer, K. & Lange-Bertalot, H. (1991).   Süsswasserflora von Mitteleuropa 2 Bacillariophyceae, 3 teil: Centrales, Fragilariaceae, Eunotiaceae.   Spektrum Akademischer Verlag, Heidelberg, Berlin.

Lange-Bertalot, H. & Ulrich, S. (2014).  Contributions to the taxonomy of needle-shaped Fragilaria and Ulnaria species.   Lauterbornia 78: 1-73.

Spring comes slowly up this way …*

I took a few minutes out on my trip to Upper Teesdale to stop at Wolsingham and collect one of my regular samples from the River Wear.  Back in March, I commented on the absence of Ulothrix zonata, which is a common feature of the upper reaches of rivers such as the Wear in early Spring (see “The mystery of the alga that wasn’t there …”).   I put this down to the unusually wet and cold weather that we had been experiencing and this was, to some extent, confirmed by finding prolific growths of Ulothrix zonata in late April in Croasdale Beck (see “That’s funny …”).   Everything seems to be happening a little later than usual this year.   So I should not have been that surprised to find lush growths of green algae growing on the bed of the river when I waded out to find some stones from which to sample.

These growths, however, turned out to be Stigeoclonium tenue, not Ulothrix zonata (see “A day out in Weardale”): it is often hard to be absolutely sure about the identity of an alga in the field and, in this case, both can form conspicuous bright green growths that are slimy to the touch.   Did I miss the Ulothrix zonata bloom in the River Wear this year?   Maybe.   Looking back at my records from May 2009 I see that I recorded quite a lot of narrow Phormidium filaments then but none were apparent in this sample.   That taxon thrived throughout the summer, so perhaps, again, its absence is also a consequence of the unusual weather.

Growths of Stigeoclonium tenue on a cobble in the River Wear at Wolsingham, May 2018.  

The photograph illustrates some of the problems that ecologists face: the distribution of algae such as Ulothrix zonata and Stigeoclonium zonata is often very patchy: there is rarely a homogeneous cover and, often, these growths are most prolific on the larger, more stable stones.   I talked about this in Our Patchwork Heritage; the difference now is that the patchiness is exhibited by different groups of algae, rather than variation within a single group.   Ironically, the patchiness is easier to record with the naked eye than by our usual method of sampling attached algae using toothbrushes.   That’s partly because we tend to sample from smaller substrata (the ones that we can pick up and move!) but also because of the complications involved in getting a representative sample.   We have experimented with stratified sampling approaches – including some stones with green algae, for example, in proportion to their representation on the stream bed – but that still means that we have to make an initial survey to estimate the proportions of different types of growth.

Under the microscope, therefore, the algal community looks very different.   There are fewer green cells and more yellow-brown diatom cells, these dominated by Achnanthidium minutissimum, elegant curved cells of Hannaea arcus and some Navicula lanceolata, still hanging on from its winter peak.   The patterns I described in The mystery of the alga that wasn’t there … are still apparent although the timings are all slightly adrift.

A view of the biofilm from the River Wear, Wolsingham in May 2018.

The schematic view below tries to capture this spatial heterogeneity.  On the left hand side I have depicted the edge of one of the patches of Stigeoclonium.   Healthy populations of Stigeoclonium do no support large populations of epiphytes, probably as a result of the mucilage that this alga produces.  My diagram also speculates that the populations of Gomphonema olivaceum-type cells and Ulnaria ulna may be living in the shadow of these larger algal growths, as neither is well adapted to the fast current speeds on more exposed rock surfaces.  Finally, on the right of the image, there are cells of Achnanthidium minutissimum, small fast-growing cells that can cope with both fast currents and grazing.   I have not included all of the taxa I could see under the microscope, partly because of the space available.  There is no Hannaea arcus or Navicula lanceolata and I have also left out the chain of Diatoma cells that you can see on the right hand side of the view down the microscope.

The speckled background in the image of the view down the microscope is, by the way, a mass of tiny bacteria, all jigging around due to Brownian motion.  The sample had sat around in the warm boot of the car for a few hours after collection so I cannot be sure that these were quite as abundant at the time of collection as they were when I came to examine it.  However, some people have commented on the absence of bacteria – known to be very abundant in stream biofilms – from my pictures, so these serve as a salutary reminder of an extra dimension that really needs to be incorporated into my next images.

Schematic view of the biofilm from the River Wear at Wolsingham, May 2018.  a. Stigeoclonium tenue; b. Gomphonema olivaceum complex; c. Ulnaria ulna; d. Meridion circulare; e. Achnanthidium minutissimum.   Scale bar: 10 micrometres (= 1/100th of a millimetre).

* Samuel Taylor Coleridge, Christabel (1816)

 

That’s funny …

The most exciting phrase to hear in science, the one that heralds new discoveries, is not “Eureka!” but “That’s funny”
Attributed to Issac Asimov

I have visited Croasdale Beck, in western Cumbria, twenty-eight times since 2015 and I thought I was beginning to understand it’s character (see “A tale of two diatoms” and “What a difference a storm makes”).   It is the unruly sibling of the River Ehen which, usually, offers a far less amenable environment for freshwater algae.  Last week, however, as we walked down the track towards the stream, we were confronted with the unexpected sight of a river bed that was bright green.  Our measurements, too, showed that not only was there a lot of algae in absolute terms, but there was far more here than we had measured in the River Ehen.  Usually, the situation is reversed, with the Ehen having more than Croasdale Beck.

Croasdale Beck at NY 087 170 looking upstream in April 2018.   The position of the gravel bar has shifted over the time that we have visited, with the wetted channel originally being at the right hand side, rather than being split into two.

It was hard to capture the extent of the algae growing on the river bed in a photograph, but the macroscopic image below captures the colour of the growths well, and you’ll have to use your imagination to scale this up to cover half of the stream bed.  Under the microscope, these growths turned out to be virtual monocultures of the green alga Draparnaldia glomerata.  This is common in clean rivers in spring time, and I often find it in the nearby River Ehen (see “The River Ehen in February”).  What my images do not show is the mucilage that surrounds the filaments.   In some cases, the growths can be almost jelly-like, so prolific is this mucilage.   One of the roles of this mucilage plays is to serve a matrix within which enzymes released by the fine hairs at the end of the filaments can act to release nutrients bound into tiny organic particles (see “A day out in Weardale …”).

Growths of Draparnaldia glomerata in Croasdale Beck (NY 087 170) in April 2018.  The upper image shows the filaments growing on submerged stones and the lower image shows the bushy side-branches growing from a central filament.  Scale bar: 100 micrometres (= 1/10th of a millimetre).

We also sample a site a couple of kilometres downstream on Croasdale Beck and, here again, the river bed was smothered in green growths.  I assumed that this, too, was Draparnaldia glomerata but, when I examined the filaments under the microscope, it turned out to be a different alga altogether: Ulothrix zonata (see “Bollihope Bhavacakra” and links therein).   There is little difference between the two sites that might explain this: the latter is slightly lower and is surrounded by rough pasture whilst the other is closer to the fells.   However, I have seen both Ulothrix zonata and Draparnaldia glomerata at several other sites in the vicinity, and a simplistic interpretation based on agricultural enrichment does not really work.

There were also a few obvious differences in the diatoms that I saw in the two samples.   In both cases, we sampled stones lacking green algae but, instead, having a thick brown biofilm.  Several taxa were common to both sites – Odontidium mesodon, for example (broadly confirming the hypothesis in “A tale of two diatoms …”) and Meridion circulare was conspicuous in both.   However, the lower site had many more cells of “Ulnaria ulna” than the upper site.   Again, there is no ready explanation but, at the same time, neither green algae or diatoms at either site suggests anything malign.

Filaments of Ulothrix zonata at Croasdale Beck (NY 072 161).   The upper filament is in a healthy vegetative state (although the cell walls are not as thickened as in many populations).  The lower filament is producing zoospores.   Scale bar: 25 micrometres (= 1/40th of a millimetre).

Diatoms in Croasdale Beck, April 2018.   a. upper site: note the abundance of Odontidium mesodon, plus cells of Gomphonema cf exilissimum, Achnanthidium minutissimum and Meridion circulare; b. lower site: note the presence of “Ulnaria ulna” as well as several of the taxa found at the upper site.   Scale bar: 25 micrometres (= 1/40th of a millimetre).  

So where does this take us?  I talked about the benefits of repeat visits to the same site in “A brief history of time wasting …” and I think that these data from Croasdale are making a similar point.  By necessity, most formal assessments of the state of ecology are based on very limited data, from which, at best, we get an estimate of the “average” condition of a water body over a period of time.  Repeat visits might lead to a more precise assessment of the “average” state but also give us a better idea of the whole range of conditions that might be encountered.  Here, I suspect, we chanced upon one of the extremes of the distribution of conditions.   Cold, wet weather in early spring delayed the growth of many plants – aquatic and terrestrial – as well as the invertebrates that graze them.   Then the period of warm, dry conditions that preceded our visit gave the algae an opportunity to thrive whilst their grazers are still playing “catch-up”.  I suspect that next time we visit Croasdale Beck will have its familiar appearance.   It is, nonetheless, sobering to think that this single visit could have formed fifty-percent of the evidence on which a formal assessment might have been made.