The only way is up …

Ehen_Mill_Feb19

How does an alga move upstream?   I’m curious because, I am now seeing populations of Lemanea fluviatilisabout four kilometres further upstream in the River Ehen than when I first started my regular visits in 2013.   I can explain the presence of the organism partly through changes in the hydrology of the river: a small tributary, Ben Gill, that had been diverted into the lake in Victorian times was reconnected to the river in 2014 and this introduced periodic pulses of intense energy to the river that had immediate effects on the substrate composition.  Lemanea fluviatilisis a species that thrives in the fastest-flowing sections of streams so I am quite prepared to believe that even a small shift in the hydrology of this very regulated river might make the habitat more conducive.

But that does not explain how it got there in the first place.   If the alga was occurring a few kilometres further downstream we would not have any such problems: the upstream populations would provide innocula and, if the habitat conditions changed at the downstream location, then some of those propagules might be able to establish at the downstream locations.   But what about movement in the other direction?

There has been relatively little published on this topic in recent years.  I have a review by Jørgen Kristiansen from 1996 that considers the dispersal of algae but most of the references that he cites are quite a lot older than this and I have not seen much published subsequently.   He lists our options: dispersal by water, by organisms, by air currents and by human activity.   Let’s consider each in the context of Lemaneain the River Ehen.   Lemanea, like most red algae, has a complicated life cycle with the potential for dispersal in both the haploid and diploid phases, but that is probably more detail than we need right now.  We’ll just outline the options in broad terms:

Water:the linear flow of the river means that it is almost impossible for the downstream population to provide inocula for the new upstream locations.  It may be possible for populations from further upstream in the catchment to seed the new locations.  I have not seen Lemaneain any of the streams that flow into Ennerdale Water (from which the Ehen emerges) but my knowledge of the catchment is not exhaustive.   Likelihood: very low to low.

Lemanea_at_Mill_Feb19

Young shoots of Lemanea fluviatillis(bottom right) growing on a submerged boulder in the River Ehen at a location where I have not previously seen it.   These are growing alongside thick growths of diatoms (yellow-brown in colour) and patches of green filamentous algae.

Organisms:much of the older literature is concerned with the possibility of living algae or their propagules being transported in mud attached to bird’s feet or feathers and this cannot be ruled out.   There is also a recent study showing how mink may act as a vector for Didymosphenia geminata in Chile.  The Ehen also has aquatic mammals (such as otters) that could be acting as vectors for Lemanea, as well as migratory fish such as salmon and trout that could move propagules upstream.   There is also some evidence that some algae can survive passage through mammalian and invertebrate guts, and this, too, may provide a means for Lemaneato spread upstream.    Likelihood: low to medium.

Air currents / wind:quite a lot has been written about airborne dispersal of algae, with even Darwin making a contribution (see reference in Kristiansen).  The key hazard in airborne dispersal is desiccation so, in the case of Lemanea, the most likely lifecycle stages that could be dispersed in this way would be the diploid carpospores or haploid monospores. This, however, would assume that there were times during the year when the relevant life-cycle stages were exposed and, as Lemaneais a species that I usually find in the Ehen only fully-submerged, this is not very feasible.  Likelihood: low.

Human activity:there is evidence that Didymosphenia geminatacan be transported between sites attached to waders and new records often correspond with patterns of recreational use (references in Bergey & Spaulding – see below).   When we work in the Ehen we prefer to move downstream in order to minimise the risk of moving organisms on our kit, and we also clean our kit before we start.   However, a lot of people work in this part of the Ehen and it only takes one dirty wader to introduce a propagule.   Likelihood: low to medium.

We’ll almost certainly never know for sure why Lemanea fluviatilisis now thriving four kilometres further upstream than it was five years ago.  It is, however, worth bearing in mind that, given enough time, even a low probability may yield a positive result.   So none of the four hypotheses can be ruled out for sure.   Three of the possibilities are entirely natural, with one – movement by the stream itself – being constrained by the direction of flow.  Biological vectors look like a very plausible means of moving algal propagules around catchments but, for this to work, we need wildlife-friendly corridors around the river to support the animals and birds.  The upper Ehen has these, but many other rivers do not.

Actually, having a number of options all with a relatively low likelihood adds to the sense of mystery that every ecologist should have when they approach the natural world.  When cause and effect are too predictable, we tend to focus on engineering the right “solution”.  The truth, in our muddled and unpredictable world, is often that nudging several factors in the right direction will give us a more resilient outcome, even though we may have to wait longer for it to happen.

Reference

Bergey, E.A. & Spaulding, S.A. (2015). Didymosphenia: it’s more complicated.  BioScience65: 225.

Kristiansen, J. (1996).  Dispersal of freshwater algae – a review.  Hydrobiologia336: 151-157.

Leone, P.B., Cerda, J., Sala, S. & Reid, B. (2014).  Mink (Neovision vision) as a natural vector in the dispersal of the diatom Didymosphenia geminataDiatom Research29: 259-266.

Raven, J.A. (2009).  The roles of the Chantransia phase of Lemanea (Lemaneaceae, Batrachospermales, Rhodophyta) and of the ‘Mushroom’ phase of Himanthalia (Himanthaliaceae, Fucales, Phaeophyta).  Botanical Journal of Scotland46: 477-485.

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Life out of water …

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

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

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

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

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

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

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

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

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

Reference

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

Return to the Serra da Estrela

towards_manteigas

Back in October I wrote about the algae and other plants that I had found in a small stream draining the Serra da Estrela mountains in Portugal (see “Notes from the Serra da Estrela”).  I’ve now had a chance to look more closely at the diatoms that I found there, and can offer a few thoughts on the ecology of the stream.

I collected two samples from the stream: one by brushing the top surface of the granite stones with a toothbrush and the other from the darker patches that I described in the earlier post.   These were a mix of algae and mosses, with the former dominated by cyanobacterial filaments and diatoms.   I merged the two samples prior to digesting them, but the biofilm on the submerged rocks was very thin so it is the diatoms from the dark patches that dominate the slide that I prepared from this stream.   As my preliminary observations suggested, motile diatoms were very abundant in this sample, with Surirella roba, Navicula angustaand N. exilis all common, along with some Pinnularia and Nitzschia.   I do not often find motile diatoms to be quite so abundant in fast-flowing upland streams, but I suspect that this is because I look in the wrong places.   Our standard sampling method involves scrubbing the tops of submerged stones which, in this type of stream at least, are not situations where motile diatoms thrive.  By contrast, the tangle of cyanobacterial filaments and dead organic matter creates a very different environment, where an ability to adjust position in order to move away from densely-shaded areas and, perhaps, from situations where bacteria and fungi had used up all the available oxygen, was an advantage.

surirella_roba_unhais_sep18

Surirella robafrom the stream at Unhais de Serra, September 2018; a. – f.: valve views; g. – i.: girdle views. Scale bar: 10 micrometres (= 1/100thof a millimetre). The photo at the top of the post shows the view along the valley of the Rio Zêzere towards Mantiegas in the Serra da Estrela.

misc_diatoms_unhais_sep18

Miscellaneous diatoms from the stream at Unhais de Serra, September 2018: a. – d.: Cocconeis placentula, complete frustule, rapheless valve and two raphe valves; e. – g.: Navicula exilis; h. N. angusta; i. – k.: Pinnularia subcapitata, two valve views and a girdle view.  Scale bar: 10 micrometres (= 1/100thof a millimetre). 

A chain-forming species of Fragilariawas abundant in the original sample although, by the time I had prepared a slide, the chain had disintegrated into individuals or pairs of cells.  These all belonged to a member of the Fragilaria capucinacomplex, though I am not sure which one. There were also a few cells of the free-living (i.e. non-chain-forming) Fragilaria gracilis.    Eunotia minoror a close relative was also present, sometimes also forming short chains and, finally, I found a number of cells of Cocconeis placentula(possibly var. klinoraphis).

These are all diatoms that I would expect to find in a stream draining a hard rock such as granite in an area that is remote from any industrial or mining influences that might lead to artificial acidification.   There are mines in the area, but these are further south.  These do have a measurable effect on the biology of local streams, as the references at the end of this post attest.   However, this particular stream appears to be in rude health.

A curious side-effect of the years that I have spent looking at diatoms is that a sample such as this can evoke the environments from which it came: an assemblage of soft-water circumneutral diatoms conjures, in my mind, a particular landscape.   The label on the slide, of course, takes me straight back to our time in the Serra da Estrela but, in a more general sense, the diatoms capture an essence that transcends any one particular time or place.   Analysing diatom slides can become an escape from the humdrum and a chance to remember warmer days …

fragilaria_unhais_sep18

Fragilaria species from the stream at Unhais de Serra, September 2018: a. – g.: chain-forming member of Fragilaria capucina complex (a.-c.: valve views; d.-g.: girdle views); h.-j.Fragilaria gracilis.  Scale bar: 10 micrometres (= 1/100th of a millimetre).

eunotia_cf_minor_unhais_sep18

Eunotiacf. minorfrom the stream at Unhais de Serra, September 2018: j. – n.: valve views; o. valve view of a related species; p. girdle views. Scale bar: 10 micrometres (= 1/100thof a millimetre). 

References

Luis, A.T., Teixeira, P., Almeida, S.F.P., Matos, J.X. & Silva, E.F. (2004).  Environmental impact of mining activities in the Lousal area (Portugal): Chemical and diatom characterization of metal-contaminated stream sediments and surface water of Corona stream.  Science of the Total Environment409: 4312-4325.

Silva E.F., Almeida, S.F.P., Nunes, M.L. & Fredrik, A.T.L. (2009). Heavy metal pollution downstream the abandoned Coval da Mó mine (Portugal) and associated effects on epilithic diatom communities.  Science of the Total Environment407: 5620-5636.

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 …

 

 

Casting the net wide …

A  month or so ago I wrote a couple of posts about the green algae that were thriving in the River Wear this summer (see “Keeping the cogs turning …” and “More green algae from the River Wear”).  In one of those, I promised to write a post about a related genus, Hydrodictyon.   I did try to find some recent populations but ran out of time so have fallen back on some old pictures along with more of Chris Carter’s spectacular photography.

Hydrodictyon reticulatum is commonly called the “water net” and can form extensive, and sometimes nuisance, growths, either floating in a lake or a slow-flowing river, or as a mat at the edge (see photos below).   The cylindrical cells are arranged in pentagons or hexagons which can be visible with the naked eye (hence the net-like appearance).   These have a mode of asexual reproduction that results in tiny zoospores being formed inside each cell.  Each of these develops into a small daughter cell whilst still inside the mother cell, and the ends of these daughters then join together to form mini-nets.  You can see this happening in Chris’ image at the top of the post: there are some young cells in the foreground with a mature “mother” cell full of “daughter” cells forming their own nets inside.  Eventually, the wall of the mother-cell disintegrates and the daughter net is released.

A mat of Hydrodictyon reticulatum from the lower River Tweed; b. macro, and, c. microscopic views of coenobia of H. reticulatum from Thrapston Lake, Northampton (b. and c. by Chris Carter).

This is an extremely effective way of enabling Hydrodictyon reticulatum to spread quickly when conditions favour its growth and the image below shows just how extensive these mats can be.   It is a species that is more common in the warmer parts of the world but it does occur in the UK as far north as Scotland and Brian Whitton has predicted that it is a species that is likely to be favoured in some climate warning scenarios.   Some authors have suggested that Hydrodictyon favours nutrient-rich water, but some of the locations where I have found it (the River Tweed in Scotland, Sunbiggin Tarn in Cumbria) do not meet this criterion.  Rather, I suggest that well-lit, relatively undisturbed summer conditions are the key factor and that this is more likely to be the case in lowland areas that are, in many cases, also rich in nutrients.  It is more likely to be a correlation than a cause, in other words.   Whatever the cause, there is a huge dichotomy between the beauty of the organism under the microscope and the nuisance that it can cause.

A huge growth of Hydrodictyon reticulatum at Manor Farm Weir on the Jubilee River (a flood alleviation channel of the River Thames near Maidenhead).  Photo: Environment Agency.

This mode of asexual reproduction – in which the zoospores aggregate inside the parent – is also a feature of Pediastrum.  Even though the shapes and dimensions of the organisms are very different, they share some fundamental properties.   Molecular phylogenetic studies have also shown that there is a close affinity between Hydrodictyon, Pediastrum and the other genera I mentioned in “Keeping the cogs turning …”.   However, their habits and ecology are very different and that raises some interesting questions about a different matter entirely … the subject of my next post.

Reference

John, D.M., Pentecost, A. & Whitton B.A. (2001).  Terrestrial and freshwater eukaryotic algae.   pp. 148-149.  In: The Changing Wildlife of Great Britain and Ireland (edited by D.L. Hawksworth).   Taylor & Francis, London.

Krienitz, L. & Bock, C. (2012).  Present state of the systematics of planktonic coccoid green algae of inland waters.   Hydrobiologia 698: 295-326.

 

More green algae from the River Wear

Having discussed some of the recent name changes in green algae in the previous post, I thought that I would continue this theme using some of the other taxa that I found in the samples I collected from the River Wear a couple of weeks ago.   The plate below shows some specimens that, 20 years ago, I would not have hesitated to call Scenedesmus, characterised by coenobia of either four cells or a multiple of four cells arranged in a row.   Over 200 species, and 1200 varieties and forms have been recognised although there were also concerns that many of these so-called “species” were, in fact, variants induced by environmental conditions.  A further problem is that Scenedesmus and relatives do not have any means of sexual reproduction.  This means that any mutation that occurs and which does not have strong negative effects on the organism will be propagated rather than lost through genetic processes.  Working out what differences are really meaningful is always a challenge, especially when dealing with such tiny organisms.

Scenedesmus and Desmodemus species from the River Wear, Wolsingham, September 2018.  a. and b. Scenedesmus cf ellipticus; c. Desmodesmus communis.   Scale bar: 20 micrometres (= 1/50th of a millimetre).

The onset of the molecular era shed some new light onto these problems but, in the process, recognised differences within the genus itself that necessitated it being split into three, two of which are on the plate below.  Scenedesmus, in this modern sense, has cells with obtuse (rounded) apices and mucilage surrounding the cells whilst Desmodesmus has distinct spines at the apices of marginal cells and, sometimes, shorter ones elsewhere too.   In addition to these there is Acutodesmus, which is similar to Scenedesmus (i.e. without spines) but whose cells have more pointed (“acute”) ends and which does not have any surrounding mucilage.   A further genus, Pectinodesmus, has been split away from Acutodesmus on the basis of molecular studies, although there do not seem to be any features obvious under the light microscope which can differentiate these.

The genera Ankistrodesmus and Monoraphidium present a similar situation.  In the past, these long needle- or spindle-shaped cells would all have been considered to be Ankistrodesmus.   Some formed small bundles whilst others grew singly and this, along with a difference in their reproductive behaviour, was regarded as reason enough for splitting them into two separate genera.   Both were present in the Wear this summer, but only Monoraphidium presented itself to me in a manner that could be photographed.  Two species are shown in the plate below.   Recent molecular studies seem to not just support this division but also suggest that each of these could, potentially, be divided into two new genera, so we’ll have to watch out for yet more changes to come.

Monoraphidium species from the River Wear, Wolsingham, September 2018.  a. and b.: M. griffthii; c. M. arcuatum.  Scale bar: 20 micrometres (= 1/50th of a millimetre).

The final illustration that I managed to obtain is of another common coenobium-forming alga, Coelastrum microporum.   Though the three-dimensional form makes it a little harder to see, cell numbers, as for Pediastrum, Scenedesmus and Desmodesmus, are multiples of four.  I apologise if the picture is slightly out of focus, but it is a struggle to use high magnification optics on samples such as these.  The oil that sits between the lens and the coverslip conveys the slight pressure from fine focus adjustments directly to the sample, meaning that the cells move every time I try to get a crisper view.  That means it is impossible to use my usual “stacking” software.   The answer is to use an inverted microscope so that the lens was beneath the sample.  However, I do this type of work so rarely that the investment would not be worthwhile.

That’s enough for now.   I’m off on holiday for a couple of weeks, so the next post may be from Portugal and perhaps I will also find time to sample the River Duoro as well as the products of the vineyards in it’s catchment…

Coelastrum microporum from the River Wear,Wolsingam, Septmber 2018.  Scale bar: 20 micrometres (= 1/50th of a millimetre).

References

An, S.S., Friedl, T. & Hegewald, E. (2008).  Phylogenetic relationships of Scenedesmus and Scenedesmus-like coccoid green algae as inferred from ITS-2 rDNA sequence comparisons.   Plant Biology 1: 418-428.

Hegewald, E., Wolf, M., Keller, A., Friedl, T. & Krienitz, T. (2010).  ITS2 sequence-structure phylogeny in the Scenedesmaceae with special reference to Coelastrum (Chlorophyta, Chlorophyceae), including the new genera Comasiella and Pectinodesmus.   Phycologia 49: 325-355.

Krienitz, L. & Bock, C. (2012).  Present state of the systematics of planktonic coccoid green algae of inland waters.   Hydrobiologia 698: 295-326.

Krienitz, L., Bock, C., Nozaki, H. & Wolf, M. (2011).   SSU rRNA gene phylogeny of morphospecies affiliated to the bioassay alga “Selanastrum capricornutum” recovered the polyphyletic origin of crescent-shaped Chlorophyta.  Journal of Phycology 47: 880-893.

Trainor, F.R. & Egan, P.F. (1991).  Discovering the various ecomorphs of Scenedesmus: the end of a taxonomic era.   Archiv für Protistenkunde 139: 125-132.

Talking about the weather …

September is here.  When I visited this site two months ago we were in the midst of the heatwave and the samples I collected from the Wear at Wolsingham were different to any that I have seen at this location before, dominated by small green algae (see “Summertime blues …”).   As I drove to Wolsingham this time, I could see the first signs of autumn in the trees and the temperatures are more typical of this time of year.   We have had rain, but there has not been a significant spate since April and this means that there has been nothing to scour away these unusual growths and return the river to its more typical state.

That does not mean, however, that there have been no changes in the algae on the submerged stones.  Some of these differences are apparent as soon as I pick up a stone.  Last month, there was a thin crust on the surface of the stones; that is still here but now there are short algal filaments pushing through, and the whole crust seems to be, if anything, more consolidated than in July, and I can see sand grains amidst the filaments.   Biofilms in healthy rivers at this time of year are usually thin, due to intense grazing by invertebrates, so I’m curious to know what is going on here this year.

A cobble from the River Wear at Wolsingham, showing the thick biofilm interspersed with short green filaments.   Note, too, the many sand grains embedded in the biofilm.  The bare patch at the centre was created when I pulled my finger through it to show how consolidated it had become.  The cobble is about 20 centimetres across.

Many of the organisms that I can see when I peer at a drop of my sample through my microscope are the same as those I saw back in July but there are some conspicuous differences too.   There are, for example, more desmids, some of which are, by the standards of the other algae in the sample, enormous.   We normally associate desmids with soft water, acid habitats but there are enough in this sample to suggest they are more than ephemeral visitors.   And, once I had named them, I saw that the scant ecological notes that accompanied the descriptions referred to preferences for neutral and alkaline, as well as nutrient-rich conditions.  Even if I have not seen these species here before, others have seen them in similar habitats, and that offers me some reassurance.    In addition to the desmids, there were also more coenobia of Pediastrum boryanum and Coelastrum microporum compared to the July sample.

A view of the biofilm from the River Wear at Wolsingham on 1 September 2019. 

There were also more diatoms present than in my samples from July – up from about 13 percent of the total in July to just over 40 per cent in September.   The most abundant species was Achnanthidium minutissimum, but the zig-zag chains of Diatoma vulgare were conspicuous too.  The green filaments turned out to be a species of Oedogonium, not only a different species to the one I described in my previous post but also with a different epiphyte: Cocconeis pediculus this time, rather than Achnanthidium minutissimum.   I explained the problems associated with identifying Oedogonium in the previous post but, even though I cannot name the species, I have seen this form before (robust filaments, cells 1.5 to 2 times as long as broad) and associate it with relatively nutrient-rich conditions.  That would not normally be my interpretation of the Wear at Wolsingham but this year, as I have already said, confounds our expectations.   I did not record any Cladophora in this sample but am sure that, had I mooched around for longer in the pools at the side of the main channel, I would have found some filaments of this species too.

Desmids and other green algae from the River Wear at Wolsingham, 1 September 2019.  a. Closterium cf. acerosum; b. Closteriumcf. moniliferum; c. Cosmarium cf. botrysis; d. Closterium cf. ehrenbergii; e. Coelastrum microporum; f. Pediastrum boryanum.   Scale bar: 50 micrometres (= 1/20th of a millimetre).  

It is not just the differences between months this year that I’m curious about.  I did a similar survey back in 2009 and, looking back at those data, I see that my samples from August and September in that year had a very different composition.   There was, I remember, a large spate in late July or early August, and my August sample, collected a couple of weeks later had surprised me by having a thick biofilm dominated by the small motile diatom Nitzschia archibaldii.   My hypothesis then was that the spate had washed away many of the small invertebrates that grazed on the algae, meaning that there were few left to feed on those algae that survived the storm (or which had recolonised in the aftermath)..   As the algae divided and re-divided, so they started to compete for light, handing an advantage to those that could adjust their position within the biofilm.   This dominance by motile diatoms was, in my experience of the upper Wear, as uncommon as the assemblages I’m encountering this summer, though probably for different reasons.

Other algae from the River Wear at Wolsingham, September 2018.    The upper image shows Diatoma vulgare and the lower image is Oedogonium with epiphytic Cocconeis pediculus.   Scale bar: 20 micrometres (= 1/50th of a millimetre).

I suspect that it is the combination of high temperatures and low flows (more specifically, the absence of spates that might scour away the attached algae) that is responsible for the present state of the river.  This, along with my theory behind the explosion of Nitzschia archibaldii in August 2009, both highlight the importance of weather and climate in generating some of the variability that we see in algal communities in rivers (see “How green is my river?”).   The British have a reputation for talking about the weather.   I always scan the weather forecasts in the days leading up to a field trip, mostly to plan my attire and make sure that I will, actually, be able to wade into the river.  Perhaps I also need to spend more time thinking about what this weather will be doing to the algae I’m about to sample.