Mystery, wonder and joy

My Advent reading this year was Michael McCarthy’s The Moth Snowstorm: Nature and Joy (John Murray, 2015), a meditation on the reasons why humans love the natural world, and how engagement with nature can, in turn, be beneficial for our wellbeing.   His personal fascinations with butterflies, moths and birds provide most of the examples but, as I was reading this book at the same time as I was writing the previous post on “little round green things”.  As a result, I found myself reflecting on my own fascinations with the microscopic world.

A characteristic of ecologists, I have realised, is that there is almost always a tension between their scientific training and a primeval emotional response to nature.   This is not unique to ecology: geologists and astronomers certainly share it, but it is not a universal trait of scientists.  In those disciplines where it occurs, however, interactions with the natural world occasionally transcend strictly dispassionate objective observation and spill over into the language of joy and wonder.   “Joy” being, in McCarthy’s words, “concentrated happiness” whilst “wonder” is “a sort of astonished cherishing or veneration … often involving an element of mystery”.   We are straying away from the language of science and towards a religious and spiritual dimension that many ecologists would, I suspect, be reluctant to acknowledge.

“Mystery” is the word that ties together the disparate worlds of science and religion.   It implies “missing knowledge”, but much more than just an absence of necessary facts.   Every time I peer at samples from the River Ehen through my microscope I get the full gamut of joy-wonder-mystery-related emotions even though I have seen similar views many times before.   Part of this can be attributed to “missing knowledge” but not all.  I am acutely aware of my own shortcomings as I struggle to identify the organisms that I see, as well as the limitations of the taxonomic literature on which I depend.  I am, in addition, perpetually astonished that so much diversity can live on such a small scale and, even when I have done my best to name the algae present, I still struggle to explain why the communities differ over the space of a few metres and between our monthly visits.

Ehen_Mill_181212

Regular visits for five years have not diminished my wonder at the microscopic world of the River Ehen: this submerged boulder has obvious patches of brown diatoms and green algae, but also gaps where the algae are much less abundant. We can make coarse predictions about which species are likely to be found in particular locations, but the factors that determine their distribution on much finer scales are still shrouded in mystery.

The word “mystery” in short, carries an emotional heft that simply “not knowing” does not.  It rises above ignorance, partly because mystery, by definition, implies an awareness of this lack of knowledge.   The word “mystery”, in a modern, scientific context, also links to the concept of complexity, recognising that interactions between variables is often such that it is very difficult to predict outcomes.   That “astonished cherishing” that forms part of McCarthy’s definition of wonder needs to include an element of wariness.  We approach – or, at least, we should approach – ecosystems in the same cautious manner that Moses approached the burning bush.   Whether or not you believe in a higher power, recognition of both the complexity of nature and our limited understanding of this is humbling.   Humility, in turn, generates reverence, and we have completed the journey from the hard, dispassionate language of science to the fringes of spirituality and religion.

None of this precludes trying to improve our understanding of the natural world, nor of using this knowledge to inform decision-making.   What I have written above is no more than the Precautionary Principle, albeit expressed in quasi-mystical language.   Whilst the Precautionary Principle is an instrument of policy, my interpretation is more personal.   Each of us, individually, should be finding time to revel in the wonder of nature which, in turn, will fuel the sense of mystery and, in turn, temper any inclination to rush to intemperate conclusions.

Ehen_181212_diatoms

Some of the diatoms that are abundant in the River Ehen.  Top left: colonies of Gomphonema(see “Diatoms and dinosaurs” for more about this species); top right: colonies of Fragilaria tenera, which shares the habitat with at least two other similar representatives of the same genus; bottom left: Tabellaria flocculosa.  Genetic studies suggest that this, too, is probably a complex of morphologically-similar species.   Scale bar: 20 micrometres (= 1/50thof a millimetre).

We should, however, never assume that joy, wonder and a sense of mystery are ever-presents in the make-up of ecologists.   McCarthy makes the point that a love of nature is not a universal human attribute, although a propensity to love nature may be.   Just as that propensity can be nurtured through adolescence into an adult appreciation of the natural world, so a failure to exercise that appreciation as an adult can lead to it withering again.   I am acutely conscious that ecologists of middling seniority and above often spend more time staring at spreadsheets and in teleconferences than they do engaging directly with nature.  Within government agencies the reduction of time available for field ecology since the onset of austerity in the UK means that I often now deal with people who are unable to conjure visual images from the words and numbers that populate their datasets.  And, in my own work, I have to consciously make time to observe the natural world beyond the tight constraints of my professional life.

Above all, never forget that this love of nature exists in the first person, present tense or not at all. Natural history documentaries on the television and (dare I say) blogs such as mine are the herbs and spices that enliven your diet, but the naturalist’s basic sustenance needs a commitment that goes beyond staring at a spreadsheet or sitting on a couch.

 

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A day out in Wasdale

Irt_LundBridge_Nov18

A few days after my trip to Weardale I found myself beside the River Irt, a few hundred metres below the point where it flows out of Wastwater, in the western part of the Lake District.   Whereas the River Wear drains a catchment underlain by Carboniferous rocks, including a high proportion of limestone (see “Co. Durham’s secret Karst landscape”), the Irt’s catchment is largely underlain by ancient volcanic rocks, resulting in much softer water.   I was curious to see how different the algae were here compared to those in the Wear.

The river bed at this point is dominated by boulders of granite, which host a patchwork of mosses, filamentous algae and discrete growths of diatoms (visible on the right-hand side of the figure below).  Between these there were areas of pebbles and gravels, suggesting good habitat for freshwater mussels.   The patches of filamentous algae (mostly no more than a couple of centimetres in length) were a mixture of Mougeotiaand Zygnema, similar to the forms that I find in the River Ehen, a 30 minute drive to the north.   These two species differ in the form of their chloroplasts (Mougeotiahas a flat plate whilst Zygnemahas two star-shaped chloroplasts, attached by thin cytoplasmic strands to resemble an animal skin stretched on a frame) but are closely-related, both belonging to the family Zygnemtaceae.

Irt_substratum_Nov18

An underwater photograph of the substratum of the River Irt in November 2018 showing patches of filamentous green algae, mosses and (on the right-hand side) diatoms growing on granite boulders.

Irt_greens_Nov18

Filamentous green algae from the River Irt, November 2018.   The upper photograph shows cells from a filament of Mougeotiawhilst the lower image shows two filaments of Zygnema. Scale bar: 20 micrometres (= 1/50thof a millimetre).

In between the tufts of filamentous algae were apparently bare patches of rock (they almost certainly had a very thin biofilm that would be hard to sample in isolation from the lusher algal growths that shared their habitat) and some conspicuous orange-brown growths of colonial diatoms.  These turned out to be almost pure growths ofGomphonema hebridense, or a close relative (I can’t give a definitive answer until I have examined cleaned material), growing on long mucilaginous, sometimes branched, stalks to create a veritable “bush” of diatoms.  There were a few other species of diatom growing within this bush, most notably some cells of Achnanthidium (cf.) caledonicumthat seemed to be growing on short stalks attached to the Gomphonemastalks, but also a few cells of Gomphonema capitatum(which also grows on long stalks) and some chains of Tabellaria flocculosa.

Gomphonema hebridenseis a diatom that I have written about several times before, as it is also common in the River Ehen, and also presents a number of interesting challenges to taxonomists (see “Diatoms and dinosaurs”). Whatever future studies reveal, however, the presence of colonies of this (or these) species that are visible with the naked eye is something I associate with only the cleanest rivers in the country during the cooler times of year.  It should not have been a great surprise to me to find it flowing out of one of the most pristine lakes in England (see “The Power of Rock …”).

Gomphonema_Irt_x1000_Nov18

A close up of cells within a colony of Gomphonemacf hebridense.  Several mucilaginous stalks are also visible as well as (top left) a cell of Achnanthidiumcf caledonicum.   Scale bar: 10 micrometres (= 100th of a millimetre).

The predominance of boulders over smaller, more easily moved stones, suggests a river that has more energy than the River Ehen, one of my regular Lake District haunts.   Both flow out of lakes whose catchments include some of the wildest and most mountainous terrain in the country.   Lakes tend to act as shock absorbers in catchments, slowing down the water that pours off the fells after heavy rain.   Streams in this part of the world that have no such impediments to flow tend to have rocky, mobile beds and relatively sparse algal communities.   By contrast, the Irt and Ehen just below their respective lakes have relatively lush growths of algae.   The substrates of the two rivers, however, are very different: the Ehen having very few boulders in comparison to the Irt, due to the presence of a weir at the outfall. This allows Ennerdale Water to be used as a water supply for the towns of north west Cumbria but, at the same time, turns the lake into an even more effective hydrological shock absorber.  Yet more of the energy that should be washing smaller stones down the river is no longer available except after the most exceptional storms.

That’s my working hypothesis, then: the Irt is a river that is subject to just enough high energy events to move the rocky substrates around yet no so many that rich algal communities cannot develop between these.  The Ehen, by contrast, has fewer events, leading to fewer opportunities for the algae to be scoured away, whilst unregulated streams such as Croasdale Beck (see “What a difference a storm makes …”) have such regular scouring spates that the algal communities are usually sparse.   I might be wrong, of course and I might be back in a years time with a better hypothesis.  Until then …

 

 

Life in the deep zone …

The view above – looking along Ennerdale Water from the western end – is one that I’ve used before in this blog.  The difference today is that there is about twenty metres of foreshore exposed.  Normally, water covers all the area in the foreground.   Not today: Ennerdale Water is one more victim of our present drought conditions.  During the winter, we often see water splashing over the weir at the outfall; today, the weir head is a metre above the lake level and flow in the River Ehen is maintained only by pumps installed by United Utilities.

When W.H. Pearsall visited Ennerdale Water in the 1920s, he considered it to be one of the most primitive of the Lake District’s lakes (see “The power of rock …”).   However, this supposedly wild lake had been tamed by a weir since the middle of the 19th century in order to provide drinking water for Whitehaven and Workington and surrounding areas.   That, in turn, has consequences for the river downstream, especially at times such as this when, unless augmented by pumps, there would be no water in the River Ehen below the outfall.   At some point in the next decade, a new water infrastructure project will pipe water to west Cumbria from Thirlmere, after which the weir can be removed and fluctuations in water level in both lake and river will be more natural.

The weir at the outfall of Ennerdale Water, with the fish pass at the far end. 

Meanwhile, however, I was able to explore areas of the lake littoral zone that would normally be hidden from me.  My notebook, for example, records my observations that this part of the lake shore has a stony bottom yet, as can be seen from the picture above, these form a belt about 20 metres wide, after which there is firm sand.   Normally, this would be close to the limit of safe wading but, today, I could walk out with just a pair of thin neoprene shoes.   Looking down, I could see a number of tufts of the alga Nitella flexilis growing in this sand.   I’ve written about this species before (see “Finding the missing link in plant evolution …”) and have seen it in the lake before, but not in this particular location. Standing with the lake water lapping against my shins I could bend down and take some photographs of these with my underwater camera that give this usually chilly location a semi-tropical feel.

It is a useful reminder to those of us who dabble in lake littoral zones and think that we understand their ecology that a lot happens beyond the depth in which we can safely wade.   Marco Cantonati and colleagues, for example, have found big changes in the composition of the algal flora of Alpine lakes when they used Scuba diving to explore the depths of their littoral zones.  No doubt, we would see similar changes if we were to try the same in the Lake District.

References

Cantonati, M., Scola, S., Angeli, N., Guella, G. & Frassanito, R. (2009).  Environmental controls of epilithic diatom depth-distribution in an oligotrophic lake characterized by marked water-level fluctuations.   European Journal of Phycology 44: 15-29.

Cantonati, M. & Lowe, R.L. (2014).  Lake benthic algae: toward an understanding of their ecology.  Freshwater Science 33: 475-486.

 

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.

 

Desmid diversity …

Back in September, I wrote about a joint British Phycological Society and Quekett Microscopical Club field weekend looking at desmids in the Lake District (see “Desmid Masterclass”, “Lessons from School Knott Tarn” and “Different tarn, different desmids …”).  Dave John sent some of the samples that we collected to David Williamson, the UK’s leading expert on desmids but, at 92, too frail to join us, and he has now sent back some fine drawings illustrating the range of desmids that he encountered.

Two of the tarns (Long Moss Tarn, Kelly Hall Tarns) are already recognised as Internationally Important Plant Areas (IPAs) for desmids because of their desmid diversity and containing internationally very rare desmids (based largely on David Williamson’s records) so their diversity is not a complete surprise to us.  Nonetheless, David found a total of 129 desmid taxa in the three tarns, whilst another desmid specialist, Marien van Westen, identified almost 160 desmids in another set of samples from the same tarns.

The drawings are arranged in three plates, one for each tarn.   Desmids identified by David Williamson from the three tarns are illustrated.  The desmids have been numbered and the captions prepared by David John who is analysing the findings and comparing them with surveys dating back to the 1970s.   David Williamson has drawn the taxa at different scales to roughly balance the arrangement on the collage, and adjusted the sizes so important details are visible.   No details of the chloroplasts are given since all samples had been preserved in formalin.  A few of the desmids, particularly those that are very long, have not been included in the plates.

Desmids from Long Moss Tarn (SD 292 936), September 2017.   Long Moss Tarn is shown in the photograph at the top of this post.

Desmids from Kelly Hall Tarn (SD 289 933), September 2017.

Desmids from School Knott Tarn (SD 427 973), September 2017.

Key

1-Actinotaenium diplosporum; 2-Actinotaenium turgidum;  3-Bambusina borreri;  4-Closterium acerosum var. borgei; 5-Closterium angustatum;  6-Closterium archerianum var. pseudocynthia;  7-Closterium archerianum; 8-Closterium attenuatum;  9-Closterium baillyanum var. alpinum; 10-Closterium baillyanum; 11-Closterium closterioides; 12-Closterium costatum; 13-Closterium dianae var. arcuatum; 14-Closterium dianae var. minus;  15-Closterium didymotocum; 16-Closterium incurvum; 17-Closterium intermedium; 18-Closterium kuetzingii;  19-Closterium lunula; 20-Closterium navicula;  21- Closterium setaceum; 22-Closterium striolatum; 23-Cosmarium amoenum; 24-Cosmarium anceps; 25-Cosmarium binum; 26-Cosmarium brebissonii; 27-Cosmarium contractum;  28-Cosmarium davidsonii; 29-Cosmarium debaryi;  30-Cosmarium depressum; 31-Cosmarium formosulum; 32-Cosmarium hostensiense; 33-Cosmarium incrassatum var. schmidlei; 34-Cosmarium margaritatum; 35-Cosmarium margaritiferum; 36-Cosmarium monomazum var. polymazum;  37-Cosmarium obtusatum;  38-Cosmarium ornatum; 39-Cosmarium ovale;  40-Cosmarium pachydermum; 41-Cosmarium pachydermum var. aethiopicum; 42-Cosmarium perforatum var. skujae; 43-Cosmarium portianum; 44-Cosmarium punctulatum;  45-Cosmarium quadratum; 46-Cosmarium quadrum; 47-Cosmarium subochthodes var. majus; 48-Cosmarium subtumidum var. groenbladii;  49-Cosmarium subundulatum; 50-Cosmarium tetragonum var. ornatum ; 51-Cosmarium tetraophthalmum; 52-Cosmarium variolatum;  53-Cosmocladium tuberculatum; 54-Desmidium aptogonum; 55-Desmidium swartzii; 56-Docidium baculum; 57-Euastrum ampullaceum; 58-Euastrum ansatum;  59-Euastrum bidentatum var. speciosum; 60-Euastrum gemmatum; 61-Euastrum luetkemulleri; 62-Euastrum oblongum; 63-Euastrum pectinatum; 64-Euastrum pulchellum; 65-Euastrum verrucosum; 66-Gonatozygon aculeatum; 67-Gonatozygon brebissonii; 68-Groenbladia undulata; 69-Haplotaenium minutum;  70-Hyalotheca dissiliens;  71- Micrasterias americana var. boldtii; 72-Micrasterias compereana; 73-Micrasterias crux-melitensis; 74-Micrasterias denticulata; 75-Micrasterias furcata; 76-Micrasterias pinnatifida;  77-Micrasterias radiosa; 78-Micrasterias rotata; 79-Micrasterias thomasiana; 80-Micrasterias truncata; 81-Netrium digitus; 82-Netrium digitus var. latum; 83-Netrium interruptum;  84-Penium exiguum; 85-Penium margaritaceum; 86-Pleurotaenium coronatum var. robustum;  87-Pleurotaenium ehrenbergii; 88-Pleurotaenium truncatum; 89-Sphaerozosma filiforme; 90-Staurastrum arachne;  91-Staurastrum arctiscon; 92-Staurastrum bieneanum; 93-Staurastrum boreale var. robustum; 94-Staurastrum cristatum; 95-Staurastrum dilatatum; 96-Staurastrum inconspicuum; 97-Staurastrum kouwetsii; 98-Staurastrum lapponicum; 99-Staurastrum maamense; 100-Staurastrum polytrichum; 101-Staurastrum productum; 102-Staurastrum quadrangulare; 103-Staurastrum striolatum; 104-Staurastrum teliferum; 105-Staurastrum tetracerum; 106-Staurodesmus convergens; 107-Staurodesmus convergens var. wollei; 108-Staurodesmus cuspidatus var. curvatus; 109-Staurodesmus megacanthus; 110- Xanthidium antilopaeum; 111-Xanthidium antilopaeum var. laeve; 112-Xanthidium antilopaeum var. polymazum; 113-Xanthidium cristatum.

More about Platessa oblongella and Odontidium mesodon

As my last post used the conventions of figurative art to describe algal ecology, I thought I would stick to graphs – science’s very own school of abstract art – for this one.   I spent some time in “Small details in the big picture” discussing the ecology of Platessa oblongella (including P. saxonica) but without saying very much about the types of streams where these species were found.  So I am going to take a step away from the Ennerdale catchment in this post and, instead, collate environmental data a large number of sites to get a broader understanding of their habitat preferences.  As these species are often associated with Odontidium mesodon (see “A tale of two diatoms …”), I will summarise the preferences of this species at the same time (but see Annex 1 for a graph of this species’ preferences for still versus standing water).

The first set of graphs show the response of these species to pH and alkalinity and establish both as species typical of circumneutral soft water.  Platessa oblongella can be abundant in more acid conditions (i.e. to the left of the green vertical lines) but most of the records where it is abundant have pH values between 6.5 and 7.5.   Note that P. oblongella can also be found in humic waters, where lower pH thresholds apply (see Annex 2).

Distribution of Odontidium mesodon and Platessa oblongella (including P. saxonica) to pH and alkalinity in UK streams.   Vertical lines for pH indicate threshold values that should support high (blue), good (green), moderate (orange) and poor (red) ecological status classes.  See Annex 2 for more explanation.

The second set of graphs shows how these species respond to inorganic nutrients.   Both are most abundant when inorganic nutrients are present in low concentrations, though the trend is stronger for phosphorus than it is for nitrate-nitrogen.   The graphs for Platessa oblongella, however, both have a few outliers.   I have seen P. oblongella in a few situations where I did not expect it – I remember finding it in the Halebourne, a stream draining heathland around Aldershot and Bagshot in Surrey, where the water was well buffered (mean alkalinity: 61.3 mg L-1 CaCO3) and nutrient concentration were high (mean total oxidised nitrogen: 4.01 mg L-1; dissolved phosphorus: 0.25 mg L-1) and Carlos Wetzel and colleagues note some other anomalous records from the literature in their paper (cited in my earlier post), including a few from high conductivity and even brackish environments.   So we should treat these plots as indicative of the ecological preferences rather than definitive.

Distribution of Odontidium mesodon and Platessa oblongella (including P. saxonica) to nitrate-N and dissolved phosphorus in UK streams.   Vertical lines indicate threshold values that should support high (blue), good (green), moderate (orange) and poor (red) ecological status classes.  See Annex 2 for more explanation.

The final pair of plots show how the relative abundance of these two species changes over the course of the year.  These plots show the months when each taxon is abundant, by the standards of that taxon.  Because Platessa oblongella tends to be very numerous in samples, the threshold for this taxon (the 90th percentile of all records) is higher than that for O. mesodon.   This reveals a very clear pattern of O. mesodon thriving in Spring whilst P. oblongella is abundant throughout the year, but with a slight preference for summer and autumn.  We need to reconcile these patterns with the observations in A tale of two diatoms that show that P. oblongella is associated with thinner biofilms than O. mesodon and try to work out whether season is driving the patterns or whether the seasonal patterns are the manifestation of other forces.   My suspicion is that P. oblongella is a classic pioneer species but also has a low-growing prostrate habit which means that it should be resistant to heavy grazing, which may confer an advantage in the summer and autumn when grazers are most active.  However, I may be getting ahead of myself, as we are in the process of analysing data on grazer-algae interactions in the River Ehen and Croasdale Beck that may throw more light on this.  There are clearly more layers to this story yet to be revealed …

Distribution of Odontidium mesodon (i.) and Platessa oblongella (j., including P. saxonica). The solid lines represent relative sampling effort (i.e. the proportion of samples in the dataset collected in a particular month) and the vertical bars represent samples where the relative abundance of taxon in question exceeded the 90th percentile for that taxon (20% for P. oblongella/P. saxonica and 5% for O. mesodon).

Reference

The dataset used for these analyses is that used in:

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.

Annex 1: Odontidium mesodon’s preference for still or standing water

As I included a graph showing the preference of Platessa oblongella / P. saxonica for still or standing water in “A tale of two diatoms …”, I have included a similar graph for Odontidium mesodon here.   I have not included any data from the streams that flow into Ennerdale Water’s north-west corner in this graph as this would give a distorted picture.  To date, I have only seen a single valve of O. mesodon during analyses of 14 samples from these streams but I have not yet sampled these in spring which, as the graph above shows, is the time when O. mesodon is most abundant.   Like Platessa oblongella, O. mesodon is predominately a species of running, rather than standing waters.

Differences in percentage of Odontidium mesodon in epilithic samples from Ennerdale Water and associated streams.  Data collected between 2012 and 2018.

Annex 2: notes on species-environment plots

These are based on interrogation of a database of 6500 river samples collected as part of DARES project.  Vertical lines show UK environmental standards for conditions necessary to support good ecological status: blue = high status; green = good status, orange = moderate status and red = poor status.  Note that there are no environmental standards for alkalinity and the vertical lines show a rough split of the gradient into low alkalinity (“soft water”: < 10 mg L-1 CaCO3), low/moderate alkalinity (³ 10, < 75 mg L-1 CaCO3), moderate/high alkalinity (³ 75, < 150 mg L-1 CaCO3) and high alkalinity (“hard water”: ³ 150 mg L-1 CaCO3).

pH thresholds are for clear water (see UK TAG’s Acidification Environmental Standards.  The corresponding thresholds for humic waters are lower (high/good: 5.1; good/moderate: 4.55; moderate/poor: 4.22; poor/bad: 4.03).

Phosphorus thresholds are based on UK TAG’s A Revised Approach to Setting WFD Phosphorus Standards.   Current UK phosphorus standards 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 the position of boundaries based on the average alkalinity and altitude measurements in the DARES database.

Note, too, that phosphorus analyses use the Environment Agency’s standard measure, which is unfiltered molybdate reactive phosphorus.  This approximates to “soluble reactive phosphorus” or “phosphorus as orthophosphate” in most circumstances but the reagents will react with phosphorus attached to particles that would have been removed by membrane filtration.

Nitrate-nitrogen: There are, currently, no UK standards for nitrates in rivers.  Values plotted here are derived in the same way as those for phosphorus (see “This is not a nitrate standard”)

 

Small details in the big picture …

I’ve written about Platessa oblongella, a small diatom common in low alkalinity environments, before (see “A tale of two diatoms …” and links therein) but my travels around west Cumbria are gradually revealing more and more about the ecology of this organism, so bear with me as I explain my latest findings.

My first graph shows how the distribution of this diatom varies in different types of water body in the Ennerdale catchment.   I have analysed 223 samples from this small area over the past few years and, within this dataset, there is a very clear distinction between situations where Platessa oblongella is abundant and situations where it is very rare.   I have very few records from Ennerdale Water itself (present in just two out of 27 samples, and never comprising more than 2.7% of all diatoms in the sample) nor from the River Ehen, which flows out of the lake (present in just 16 out of 164 samples, and always £ 1% of all diatoms).  By contrast, in Croasdale Beck and in streams that flow into the north-west corner of the lake, it is present in 28 out of 32 samples, with a maximum relative abundance of 69%.   In ten samples it forms more than 10% of all diatoms present.   Several of my samples from the small streams were collected from just a few metres above the point where they joined the lake, which makes the distinction between these streams and the lake that much more intriguing.

My theory – based on data I showed in A  tale of two diatoms  is that Platessa oblongella is a species of disturbed habitats and that the littoral zone of a lake, whilst subject to some turbulence, is less disturbed than the rough world of an unregulated stream.  The contrast between the River Ehen immediately below the dam at the outfall of the lake and the various small tributary streams also supports this idea.

Differences in percentage of Platessa oblongella (including P. saxonica) in epilithic samples from Ennerdale Water and associated streams.  Data collected between 2012 and 2018 (along with one sample from River Ehen collected in 1997).   The photograph at the top of the post shows Ennerdale Water, photographed in January 2018.

Some of the populations I looked at seemed to consist of two distinct forms, one broader than the other.   This variability is quite common in Platessa oblongella and Carlos Wetzel and colleagues recently published a paper which suggests that these are, in fact, two distinct species.   When I first started looking at diatoms, John Carter, my mentor, used the name Achnanthes saxonica, but Krammer and Lange-Bertalot, in the revised Süsswassserflora, regarded this as a synonym of Achnanthes oblongella, a species first found in Thailand.   Wetzel’s study shows, as well as the difference in valve width, differences in the fine details of the striae between the two species.   They also decided that both species belonged in the genus Platessa, rather than Achnanthes.

Platessa oblongella (top) and P. saxonica (bottom) from Croasdale Beck, October 2017.  Scale bar: 10 micrometres (= 1/100th of a millimetre).

Valve width is, however, a very useful criterion, as the histograms below show.   The left hand graph shows a distinctly bimodal distribution of widths in specimens from Croasdale Beck, whilst the right hand graph shows a much tighter, and clearly unimodal, range.   This comes from another tributary stream flowing into the Ehen about 500 metres below the lake itself.  Quite why two species can co-exist in one stream but only one is present in another is not clear.

The modes of these populations are very close to the median widths for P. saxonica (narrow, ± 4/5 – 5 mm) and P. oblongella (broader, ± 6.5 mm) respectively but, as the left hand histogram shows, there is some overlap.    You might have trouble, for example, deciding whether a valve that was 5.5 mm wide was a “fat” P. saxonica or a “thin” P. oblongella.   My standard advice in situations such as this is that we should identify populations not individuals although, in the case of Croasdale Beck, this will still leave a grey area between the “fat” and “thin” valves where a judgement call is necessary.   In this case I think I could have done it because the P. saxonica valves in this stream tended to have a greater length:breadth ratio than those of P. oblongella, though I have not actually quantified this.

Width of valves in populations of “Achnanthes oblongella” from a) Croasdale Beck, and b) an unnamed tributary stream of the River Ehen, October 2017. 

There is more to say about the ecology of these species, but I have probably written enough for now.  I will leave you, for now, to bask in the rare sensation that occurs when diatom taxonomists make a situation clearer rather than more opaque, and return to this subject in a future post.

References

Carter, J.R. (1970).   Observations of some British forms of Achnanthes saxonica Krasske.  Microscopy: Journal of the Quekett Microscopical Club 31: 313-316.

Wetzel, C.E., Lange-Bertalot, H. & Ector, L. (2017).  Type analysis of Achnanthes oblongella Østup and resurrection of Achnanthes saxonica Krasske (Bacillariophyta).  Nova Hedwigia, Beiheft 146: 209-227.