Castle Eden Dene in November

Castle_Eden_Burn_Nov19

For the first time this year, I heard Castle Eden Burn before I saw it.  Walking down from the car park, the distant roar of water was apparent almost as soon as the canopy of largely leafless branches closed over me.  A few trees still held their leaves – spectacularly golden on beech and birch, in particular, and the Dene’s famous yews were still green, of course – but the forest was dressed for winter now, much as it was on my first visit this year, back in January (see “Castle Eden Dene in January”).  Then, I was surprised that there was no water in the Burn.  On this trip, however, I wore my chest waders.  Back in August, I had compared Castle Eden Burn to a wadi (see “The presence of absence in Castle Eden Dene”) so the heavy rain of the previous few weeks had led me to suspect that today would be different.

The water surging through the Dene was very turbid, so collecting stones to examine involved feeling around on the river bed with my hand until I located one that was not sufficiently bedded into the substratum to remove.   That’s not ideal, but needs must and I got the five cobbles I needed, each with a distinct biofilm, slimy to the touch.  This is the first time, after eleven months, that Castle Eden Burn’s substratum has looked and felt remotely like the substratum from most of the other rivers I know in this part of the world.

Under the microscope, I see lots of particulate matter but also plenty of algae.   Apart from a few filaments of the cyanobacterium Phormidium, these were mostly diatoms.   The green algae I described in “When the going gets tough …” back in May were not obvious.  The diatoms were mostly largely motile cells of Navicula, with a few sigmoid cells of Nitzschia clausii and some smaller cells whose identity I will need to confirm once I have cleaned the sample and prepared a permanent slide.  The Navicula species, in particular, are typical inhabitants of local rivers during winter and early spring, all tolerant to a wide range of conditions.   I suspect that the rainfall has washed a lot of fine particulate debris from the industrial estates in the upper catchment into the river, and these diatoms will have the resilience to cope with such types of pollution.  A large storm sewer overflow also empties into the burn about a kilometre upstream of where I was standing and this, I suspect, has been flowing over the past month or two.

I also saw a few cells of Achnanthidium minutissimum, which I generally associate with cleaner conditions.  I suspect, however, that numbers will be relatively low compared to its more pollution-tolerant brethren.   Again, I can give a more authoritative answer once I have cleaned the sample and performed a full analysis.

CEB_diatoms_Nov19

Diatoms from Castle Eden Burn, November 2019.  a., b.: Navicula trpunctata; c. – e.: Navicula lanceolata; f., g.: Rhoicosphenia abbreviata; h., i.: Nitzschia clausii; j., k.: Navicula gregaria; l. Achnanthidium minutissimum.   Scale bar: 10 micrometres (= 1/100thof a millimetre).   The photograph at the top of the post shows Castle Eden Burn just downstream from the point I sampled.

I originally set out to visit Castle Eden Burn six times during 2019 and this was the last of those. I’ve written about most of these visits already but not about my September visit.  There was, on that occasion, little new information to justify a separate post but I will include the sample I collected in my final overview of the algae of Castle Eden Burn, just as soon as I get this final sample cleaned and analysed.   Before then, I have one more post to write about the diatoms, based on some more detailed observations of a few of the species, and then it will be time to think about where to focus my observations during 2020.

Diatoms from the Troodos mountains

Troodos_snowscape_Apr19

Back in April, I wrote two posts about the algae from a stream draining a chromite mine in the Troodos mountains in Cyprus (see “Survival of the fittest (1)” and “Survival of the fittest (2)”).  I also planned to write a post about the diatoms growing in the stream but the slide I prepared has been sitting on my desk over the summer whilst I was distracted by other things.  However, I have just started looking at some samples from metal-enriched streams in the northern Pennines and, curious to see whether a Cypriot chromite mine had similar effects, I blew the dust off the slide and slipped it under my microscope.

The principal effect of toxic pollution is to reduce the number of species found and, in this respect, my sample from the outflow of the Hadjipavlou mine outflow was true to form, containing just eight species.  The most abundant of these was Meridion circulare, accounting for one in four of all the cells.  What is more, many of the cells were visibly distorted (see images a., c. and d., in particular, in the plate below).  This is quite a common phenomenon in metal-polluted streams (see “A twist in the tale”) though I have not seen it quite so obviously in Meridion circulare before. My own pet theory is that one of the enzymes involved in laying down the silica cell wall has a metal co-factor that is displaced by heavy metals.

Meridion_circulare_Hadjipavlou_Apr19

Meridion circulare from thepebbles from the stream draining Hadjipavlou chromite mine in the Troodos mountains, Cyprus, March 2019.  Scale bar: 10 micrometres ( = 1/100th of a millimetre).   The photograph at the top of the post shows snow on the Troodos mountains near the mine.

The only other diatom that was at all common in the sample was Hantzschia amphioxys, which also occurred alongside a smaller population of Hantzschia abundans.  I’ve not come across Hantzschia in metal-enriched streams before: it is a species that is most often associated with habitats that are not permanently submerged.  That may be the case at Hadjipavlou but the water that flows from mines comes from groundwater rather than rainfall so would not be subject to the strong seasonal variations that we associate with Mediterranean streams.  It is hard to draw a firm conclusion from a single visit.   Unlike Meridion circulare, however, neither population of Hantzschia showed any obvious distortion, perhaps due to the Hantzschia cells being more heavily silicified than those of Meridion circulare.

The extent to which cellular distortions are obvious does vary between species, as can be seen in “A twist in the tale …”  which compared three different representatives of the same genus in a metal-polluted stream.  I chose the word “obvious” with care as I do think that these phenomena are more easily seen in long thin cells than in shorter ones.  In the same Pennine streams where distorted Fragilaria are common, for example, I can also see distorted cells of smaller diatoms such as Achnanthidium minutissimum.  But you need a keen eye to spot these reliably.   Some other people have used fluorescent stains to look at other cellular irregularities, such as the position of the nucleus and damage to the nuclear membrane, but these require specialist approaches whereas distortions to cell outlines can be spotted from a standard analysis.

Hantzschia_spp_Hajipavlou_Apr19

Hantzschia abundans (k., l.) and Hantzschia amphioxys (m. – p.) in the from the stream draining Hadjipavlou chromite mine in the Troodos mountains, Cyprus, March 2019.  Scale bar: 10 micrometres ( = 1/100th of a millimetre). 

A few years ago I was involved in a study of diatoms from streams in Cyprus and I dug out some of these data in order to put the Hadjipavlou sample into context.  One immediate surprise was that many of the “reference” (i.e. pristine or near-pristine) samples in that survey also had relatively low diversity.   The 45 samples in this subset had, on average, nine species, and a mean Shannon diversity index of 1.7, compared to eight species and a Shannon diversity index of 1.42 for the Hadjipavlou sample.   I’ve never been a fan of diversity indices as measures of ecological quality (see “Baffled by the benthos (2) and links therein”) although I suspect that average diversity at Hadjipavlou measured over a period of time will always be low whereas average diversity at unimpacted sites is more likely to fluctuate. Equally, low diversity coupled with a second strand of evidence, such as distorted valves, is a useful sign to an ecologist that something untoward is happening.

diversity_indices

Number of taxa (left) and Shannon diversity (right) recorded in 45 samples from “reference” sites (i.e. minimal evidence of anthropogenic alteration) in Cyprus.  The arrows indicate the location of the Hadjipavlou stream within this dataset. 

The irony of writing about a heavily-polluted stream in the Troodos mountains is that the geological conditions which created the metal-rich veins hereabouts also create conditions for many plants endemic to Cyprus.   The serpentine and other ultramafic rocks create metal-rich soils within which few plants can survive (more about these here. I suspect that few of the plant enthusiasts drawn to Cyprus will ever cast more than a cursory glance at the green flocs adorning the abandoned mines of the Troodos mountains.

References

Licursi, M., & Gómez, N. (2013). Short-term toxicity of hexavalent-chromium to epipsammic diatoms of a microtidal estuary (Río de la Plata): Responses from the individual cell to the community structure. Aquatic Toxicology 134-135: 82-91.  https://doi.org/10.1016/j.aquatox.2013.03.007

Messy bedrooms …

Sand_Loch_May19

When I was tramping around the Shetland Islands earlier this year (see “Hyperepiphytes in the Shetland Islands“), looking at the algae that live in the freshwater lochs, I noticed some meandering hieroglyphs made from fine sediment on the tops of some of the stones in the littoral zone.   I see these occasionally at other places too, and know that they are the “galleries” of caseless caddis flies.  Caddis flies are close relatives of the butterflies and are best known because many of their larvae use “found materials” (in contemporary art jargon) to construct cases to protect themselves.  Some species use fine gravel, silt and sand, some use fragments of plants, some have cases that are very neat, some have a more haphazard approach to construction.  However, a few families of caddis flies eschew cases and, instead, build these galleries.

Many caddis fly larvae, whether cased or not, are grazers, scraping the algae off the rocks on the bed of the stream or lake.   There is evidence that the cases offer some protection against predators such as trout which, by increasing survival rate, means that it is worthwhile for the caddis larvae to divert some of their hard-earned energy into building these.   Presumably, their caseless cousins gain the same advantage to building their galleries but recent research has suggested that these galleries offer a further benefit.

Think of caddis larvae as adolescent caddis flies.  Now imagine that the caddis gallery is the equivalent of an adolescent’s bedroom.   Horribly messy, in other words.   Let’s leave that image of a teenager behind (as most human teenagers know their way to the bathroom) and consider what happens to all that waste material that emerges from the far end of a caddis larva’s digestive system.   This nutrient-rich “ manure” encourages algae, meaning that our caseless caddis flies are, in fact, gardeners and are able to tap into this extra energy resource within their galleries in order to grow.   That brings us back to the analogy with teenagers, as these also frequently graze in their bedrooms (the diatom Campylodiscus is even the same shape as a Pringle, whose empty containers litter the bedroom floor of my own progeny).   I guess it is a good thing that caddis larvae don’t wear socks as, with six legs and two prolegs, the mess inside the gallery would be indescribable.

Psychomiiddae_Sand_Loch_May19

Galleries of caseless caddis flies (possibly Psychomiidae) on the top surface of a cobble from Sand Loch, Shetland Islands with (right) a close-up of a single gallery. The photograph at the top of the post shows Sand Loch in May 2019.

A recent study in the Lake District has shown that this “gardening” means that the algae which grow in the fine sediment from which the galleries are constructed are different to those found elsewhere on the rock surface, with a greater proportion of diatoms, which are considered to be more palatable to invertebrates than other types of algae.  Some caddis flies are thought to go even further, and can selectively remove and discard the algae that are least palatable (some Cyanobacteira, for example).

It is possible that up to 40% of the larva’s energy needs are met from the gallery itself.   The tube is, in fact, not a static construction: the larva pokes its head out in order to graze the algae immediately in front of the gallery, and extends the gallery as the food supply within easy (and safe) reach is exhausted.   At the same time, it is consuming the alga-rich rear part of the gallery (reminiscent of Hansel and Gretel eating the gingerbread house?).   A gallery only has a life-span of 10 days in the laboratory; whether this is the same under field conditions is not clear but that gives us some idea of the transience of these structures.   This rapid turnover means that the caddis larva is always feeding on succulent early-succession species, rather than the tougher and less digestible algae that might appear in more mature biofilms.

I also see similar galleries on the bed of the River Ehen from time to time but have been told that these are formed by non-biting midge (chironomid) larvae, rather than by caddis.  I presume that the same processes are happening in these although I have not been able to find much written in the literature.

Organisms that can significantly alter the habitat in which they live, and affect the conditions experienced by other species in the habitat are termed “ecosystem engineers”.  Beavers are good examples, as their dams can have significant effects on organisms extending for hectares.  Yet, in their own small way, caseless caddis larvae are also ecosystem engineers.  As are adolescent boys.   Which makes me wonder, having only talked until now about the algae in their galleries, whether caseless caddis larvae also have patches of mould extending up their walls.

Chironomid_galleries_Ehen_March19

Galleries made by chironomid larvae on a boulder in the River Ehen, March 2019.

References

Hart, D. D. (1985). Grazing insects mediate algal interactions in a stream benthic community. Oikos 44: 40-46. https://doi.org/10.2307/3544041

Johansson, A. (1991). Caddis larvae cases (Trichoptera, Limnephilidae) as anti-predatory devices against brown trout and sculpin. Hydrobiologia 211: 185-194. https://doi.org/10.1007/BF00008534

Ings, N. L., Hildrew, A. G., & Grey, J. (2010). Gardening by the psychomyiid caddisfly Tinodes waeneri: Evidence from stable isotopes. Oecologia 163: 127-139. https://doi.org/10.1007/s00442-009-1558-8

Ings, N. L., Grey, J., King, L., McGowan, S., & Hildrew, A. G. (2017). Modification of littoral algal assemblages by gardening caddisfly larvae. Freshwater Biology 62: 507-518. https://doi.org/10.1111/fwb.12881

Otto, C., & Johansson, A. (1995). Why do some caddis larvae in running waters construct heavy, bulky cases? Animal Behaviour 49: 473-478. https://doi.org/10.1006/anbe.1995.0061

The devil lies in the detail …

Our latest ring test* slide took us on a vicarious journey to the beautiful River Don in Aberdeenshire.  Maybe because I have been doing this job for so long, but the quality of the landscape was clear to me as I peered through my microscope 500 kilometres away: the range of diatoms that I could see would not have thrived anywhere with more than the lightest touch from humankind.

One of the clues for me lay in some of the smallest diatoms on the slide.   It took some discussion amongst my fellow experts, but we eventually came up with a list of five different species of Achnanthidium (all illustrated below) which, together, constituted about a third of all the diatoms on the slide (admittedly, because they are small, they constitute rather less than a third of the total volume of diatoms, but that is another story ….).   The mere presence of several Achnanthidium species is, in my experience, usually a sign of high habitat quality (see “Baffled by the benthos (2)”) but unravelling the identities of the different species with a light microscope is challenging.

Achnanthidium-minutissimum-Medwin_WaterAchnanthidium minutissimum from Medwin Water, Scotland. Photographs from the Diatom Flora of Britain and Ireland by Ingrid Jüttner.  Scale bar: 10 micrometres (= 1/100thof a millimetre). 

Achnanthidium_pyrenaicum_Towie

Achnanthidium pyrenaicum from the River Don, Towie, Aberdeenshire.  Photographs by Lydia King.  Scale bar: 10 micrometres (= 1/100thof a millimetre). 

The genus Achnanthidium is a good example of the delicate co-existence between “identification” and “taxonomy” in the world of diatoms.   Individuals from this genus are usually small so anyone using a light microscope for routine analyses will be working right at the optical limits of their equipment whilst anyone with a serious interest in taxonomy will depend upon a scanning electron microscope for the insights needed for critical differentiation between species.

This divergence between the working methods of “identifiers” and “taxonomists” means that it is rarely possible to name every individual of Achnanthidium with complete confidence.  The ones that present clearly in valve view (i.e. face-up) can mostly be assigned to a species based on features we can see with a light microscope, but it is not always straightforward for those seen in girdle view (i.e. on their side) or which are partly obscured by other diatoms or extraneous matter on the slide.   In this example from the River Don, we also noticed that smaller individuals of A. gracillimum lost their characteristic rostrate/sub-capitate ends and were, as a result, not easy to differentiate from A. pyrenaicum.

Achnanthidium_gracillium_Towie_Water

Achnanthidium gracillimum from the River Don, Towie, Aberdeenshire.  Photographs by Lydia King.   Scale bar: 10 micrometres (= 1/100thof a millimetre). 

What continues to mystify me is why so many closely-related species can live in such close proximity. It is Achnanthidium that prompt this question here, but other genera display similar tendencies (see “When is a diatom like a London bus?”).  And this immediately generates another question: why are more people not asking this question of diatoms and, indeed, microscopic algae in general?

The answer to that question falls into two parts. The first is that understanding the precise ecological requirements of microscopic algae is not a trivial task, and assumes that you are able to get several closely-related species to live in culture (which, itself, assumes you know the precise ecological requirements of each … you see the problem?).   There is, as a result, a tendency to avoid experimental approaches and, instead, look for how species associate with likely environmental variables in datasets collected from sites exhibiting strong gradients of conditions.   However, this assumes that the forces that drive the differentiation between species work at the same scale at which we sample (see “Our patchwork heritage …” for more on this).

Underlying this, however, is a deeply-held belief, dating back at least forty years, that the niches of freshwater diatoms are determined primarily by the chemistry of the overlying water.   This is a dogma that has served us well when using diatoms for understanding the effects of environmental pollution but which is, ultimately, a limitation when trying to explain why we found five separate Achnanthidium species in a single sample, all exposed to the same stream water.

Achnanthidium_lineare_&_affine_Towie

Achnanthidium lineare (first three images from the left) and A. affine (two images on the right) from River Don, Towie, Aberdeenshire.  Photographs by Lydia King.  Scale bar: 10 micrometres (= 1/100thof a millimetre). 

I will go one step further: this dogma is so deeply held that referees rarely challenge the weak evidence that is produced to demonstrate different responses to environmental conditions between closely-related species.  There are certainly variations in environmental preferences between Achnanthidium species, but these are best expressed as trends rather than unambiguous differences and I have never seen such trends subject to rigorous statistical testing.

I blame better microscopes: greater magnification and resolution has revealed such a baffling amount of diversity that all the energy of bright diatomists is absorbed unravelling this rather than trying to explain what it all means (see “The meaning of … nothing”).  If we were bumbling along with the quality of equipment that Hustedt depended upon, then maybe we would be cheerfully lumping all these forms together and focussing on functional ecology instead.   Maybe.

* see “Reaching a half century” for more about the ring test scheme

A river runs through it …

Loire_at_Orleans_Sept19

I made an journey via Paris to Orleans last weekend to wave off my wife and daughter as they walk part of the Camino de Santiago.   The part of this austere Medieval pilgrimage route that they chose to follow happens to be one that meanders along the Loire Valley, passing chateaux and wineries en route.   Apparently, blisters are providing an element of self-mortification to keep the spirit of pious ascetism alive.

Orleans has a beautiful Medieval old town, dominated by a cathedral, along with plenty of reminders that it was here that Joan of Arc whupped the English.   More importantly, for a freshwater ecologist, a river runs through it.   The mighty Loire – the longest river in France – rises in the Masif Central and then flows north until it reaches Orleans, then it swings round to flow west to join the Atlantic Ocean at Nantes, a total length of just over 1000 km.  The river at Orleans is broad and shallow, divided into two channels by a wooded island; the left-hand chnnel is braided, with many small gravel islands, some with grass and herbaceous vegetation, and the water is generally shallow.  However, the bed of the river itself was a dark green in colour.  The Loire Valley is known as the “Garden of France” and this gives a clue to the scale of nutrient enrichment that we might expect in the river.  In the backwaters, this green backdrop was enlivened by patches of red due to the aquatic fern Azolla (see “Escape to Southwold”).

Azolla_Loire_Sept19

Patches of Azolla floating over algae-smothered substrata in a backwater of the River Loire at Orleans.  The picture at the top of the post shows a view looking downstream from the left side of the Pont George V. 

I brought a sample of the algae from some stones that were just within reach of the shore home on the Eurostar in a Perrier Water bottle for a closer inspection and was surprised by the diversity. In particular, I noticed several clumps of a narrow cyanobacterium that proved hard to photograph (possibly Homoeothrix janthina: see “Algae from the Alto Duoro”) along with many green algae.  I also saw Cladophoraglomerata, which is one of the classic indicators of nutrient rich conditions, along with Stigeoclonium, two forms of Spirogyra, two forms of Oedogonium and myriad green unicells and coenobia.  Stigeocloniumis another good indicator of nutrient enrichment,as  the filaments narrow to long colourless “hairs” when key nutrients are scarce – these cells are physiological adaptions to scavenge phosphorus and their absence at Orleans shows that this nutrient is not in short supply (see “A day out in Weardale”).

Stigeoclonium_Loire_Orleans_Sept19

Stigeoclonium cf. tenue (based on properties of erect filaments) from the River Loire at Orleans, September 2019.  Scale bar: 10 micrometres (1/100thof a millimetre). 

As well as green algae and cyanobacteria, there were also a lot of diatoms.  A few of these are illustrated below, and I’ll describe the diatoms in more detail in a future post.   As well as those I have photographed, I also saw long chains of a species of Fragilaria and another small araphid (possibly Staurosira) as well as Ulnaria ulna and some cells of Gomphonema and Navicula.   Note, in particular, the tube of Encyonema (possibly E. prostratum).   I’ve talked about tube-forming diatoms before (see “An excuse for a crab sandwich, really …”) but still can’t really explain what advantage this structure confers on a diatom.   What we can, perhaps, say, is that tube-dwelling is one of a several habits represented in the diatoms from Orleans – chains, erect, free-living motile, epiphytes  and more – and the mass of other algae create a rich diversity of microhabitats that the diatoms can exploit.

diatoms_Loire_Orleans

Some diatoms from the River Loire at Orleans: a. – d. Diatoma vulgare; e. Cocconeis pediculus; f. Encyonemasp.   Scale bar: 10 micrometres (= 1/100thof a millimetre).  The identity of the tube-dwelling form will have to wait until a cleaned sample is available. 

This abundance and diversity of green algae reminded me of some of the lush growths I had seen in UK rivers during the 2018 heatwave (see “Talking about the weather …”).  In a healthy river in the UK, I would expect to find less conspicuous growths than this, as invertebrate grazers would keep the algal biofilm shorn to a short stubble.   If, for any reason, the invertebrates cannot keep up with the algal growth, then a feedback loop is set up: the algae suck the valuable oxygen the invertebrates need from the water, the grazing reduces in intensity yet further, leading to a situation such as that I saw in the River Loire is the result.

Warm weather exacerbates the situation because water holds less oxygen at high temperatures.  In my posts about the River Wear last summer I commented that the plants in the river suggested that the river was more enriched with nutrients than was actually the case and I suspect that this was the result of these interactions.   The more southerly, more continental climate of the Loire Valley will experience these effects more often and it is possible that what I am looking at in Orleans may become the norm for UK rivers, as global warming intensifies.  Over the past decade I’ve worked on how to estimate the threshold concentrations of nutrients that a healthy river can endure.  However, nutrients rarely exert a direct effect on the plants and animals in a river but, instead, work through their effect on other factors such as oxygen. That will mean that global warming may wipe out any benefits of expensive nutrient reduction programs based on current estimates of the relationship between nutrients and river ecosystem health.  That’s a rather depressing prospect.

So I’ll end on a more cheerful note: the reason Heather and Rosie chose to start their Camino in Orleans was that they wanted to avoid a flight if possible.  At least that is how they sold it to me.   That they also chose to start their pilgrimage in a town close to the Sancerre vineyards may be pure coincidence.   Or maybe not ….

References

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

Wilco, C.E., Verbrak, P., Durance, I., Vaughn, I.P. & Ormerod, S.J. (2016).  Field and laboratory studies reveal interacting effects of stream oxygenation and warming on aquatic ectotherms.  Global Change Biology 22: 1769-1778.

Orleans_cathedral_Aug19

Orleans cathedral, August 2019.

Hilda Canter-Lund competition winners 2019

Zoe_Loffler_Synphony_of_Seaweed

This year’s Hilda Canter-Lund competition for the best algal-themed photograph has been won by Zoe Loffler for her image “Symphony of Seaweeds” taken on a at low tide near Apollo Bay, Victoria, Australia, while on a family camping trip.  She took the photo using a Google Nexus 5x Smartphone.  Zoe grew up diving in temperate waters near Melbourne, Australia. She completed her undergraduate degree and PhD at James Cook University in Townsville, Australia, studying the ecology of canopy-forming seaweeds (particularly Sargassum spp.) on coral reefs. She is now based in Sydney, and enjoys snorkelling and diving in temperate waters where there is such a wonderful diversity and abundance of seaweeds. The image meets Henri Cartier-Bresson’s maxim of the “decisive moment” (see “How to win the Hilda Canter-Lund prize”) and Zoe comments in her caption that the photo shows all who are unfamiliar with seaweeds that “they are not just brown and smelly!”.

Zoe_Loffler_portrait

Zoe Loffler: winner of the 2018 Hilda Canter-Lund prize for algal photography, for her image “Symphony of Seaweed”, shown at the top of the post.

Since 2016 we have also offered a second prize which is awarded to a photograph in a contrasting style to the overall winner.  This year, that prize goes to Damien Sirjacobs of the University of Liege in Belgium  for his image “Blue Haze”. This shows a bloom of benthic blue diatoms of the genus Haslea (H. ostrearia, H. provincialis) covering a community of macroalgae (Padina pavonica, Acetabularia acetabulum, Halopteris scoparia, Dictyota sp.) in the shallow water of Calvi Bay (Corsica, France). For scale, the circular caps on the end of the Acetabularia stalks are 5 – 10 mm in diameter.   The image was taken at a depth of four metres in May 2018 with a LUMIX TZ10 in an underwater housing, under natural light conditions, while scuba-diving along rocky shores of the Revellatta peninsula (Calvi Bay).

Damien_Sirjacobs_Blue_Haze

Damien Sirjacobs’ image: Blue haze”.  

There is a lot to interest readers of this blog in Damiens’s image; first of all, Acetabularia is another challenge to the generally-accepted view that multicellularity is the only option for large organisms.  Although the plant is quite large and, unlike Vaucheria is more elaborate than a simple tube of cytoplasm (see “The pros and cons of cell walls …”) .  The whole organism is, in fact, just one giant cell with a single nucleus.  

The diatom Haslea that grows over Acetabularia and the other macroalgae in Damien’s picture creates a blue haze due to a pigment called “marennine” which is found in vacuoles inside the cells (as you can see in the photograph below).   When marennine-containing species of Haslea are abundant around oyster beds (as is the case in parts of Brittany), then the pigment turns the gills of the oyster green and such oysters are highly sought after by gastronomes.   Whether or not these oysters really taste better is debatable but marennine certainly does have some antimicrobial properties.

Damien_Sirjacobs_portrait

Damien Sirjacobs: co-winner of the 2019 Hilda Canter-Lund prize for algal photography.

Haslea-Calvi-Julie-Seveno-2

Whilst Zoe’s image has direct visual appeal, and most people will recognise it almost straightaway as depicting seaweeds, Damien’s image has a more other-worldly quality.   Unless you are familiar with the habitats and organisms, then it is difficult to interpret what is portrayed (see “Abstracting from reality …”).   One of the challenges of photographing algae is that we are dealing with the real yet little understood aspects of biodiversity, creating a multi-layered problem: first, of capturing an impression of the organism(s) but, also,  of interpreting the image to a lay-audience.   In the case of Blue Haze we have that intriguing combination of beauty, mystery and economic relevance.   That is what makes phycology such a fascinating subject.

Reference

Gastineau, R., Prasetiya, F.S., Falaise, C., Cognie, B., Decottignies, P.,  Morançais, M., Méléder, V., Davidovich, N., Turcotte, F., Tremblay, R., Pasetto, P., Dittmer, J., Bardeau, J.-F., Pouvreau, J.-B. & Mouget, J.-B. (2018). Marennine-like pigments: blue diatom or green oyster cult?   pp. 529-551.  In: Blue Biotechnology: Production and Use of Marine Molecules (edited by Stéphane La Barre and Stephen S. Bates).  Wiley VCH Verlag GmbH & Co. KGaA

Out of my depth …

Castle_Eden_Dene_March19

I was about to start writing up an account of my latest visit to Castle Eden Dene, when I realised that I had forgotten to describe my previous visit, back in March.   I’ve already described a visit in January, when the stream was dry (see “Castle Eden Dene in January” and “Tales from a dry river bed”) and promised regular updates through the year.   It seems that, amidst all the travel that filled my life over the last three months, I overlooked the post that I should have written about the visit that I made in early March.

Whereas the river was dry in January, rain during February meant that, when I returned to the Dene on 11 March, some rather turbid water was flowing down the channel on its short journey to the North Sea.   There is, finally, something more like a stream habitat from which I can collect some diatoms.

Many of the diatoms that I found in March belonged to taxa that I had also seen in January; however, the proportions were quite different.   In some cases, species that were common in January were less common now (e.g. Humidophila contenta*) but there was a small Nitzschia species with a slightly sigmoid outline that was very sparse in the January sample but which was the most abundant species in the March sample.  I’ve called this “Nitzschia clausii” but the Castle Eden Dene population does not fit the description of this perfectly.   A lot can change in a couple of months, especially when dealing with fast-growing organism such as these, as my posts on the River Wear showed (see “A year in the life of the River Wear”).  Castle Eden Burn’s highly variable discharge just adds another layer of complication to this.

CED_diatoms_Mar19

Diatoms from Castle Eden Dene, March 2019:   a. – e.: Nitzschia cf clausii; f. Tabularia fasiculata; g. Tryblionella debilis; h. Luticola ventricosa; i. Luticola mutica; j. Ctenophora pulchella.  Scale bar: 10 micrometres (= 1/100thof a millimetre).   The picture at the top of the post shows Castle Eden Burn at the time that the sample was collected.   

Nitzschia clausii is described as being “frequent in brackish freshwater habitats of the coastal area and in river estuaries, as well as in inland waters with strongly increased electrolyte content”.   A couple of the other species from this sample – Ctenophora pulchella and Tabularia fasiculata (both illustrated in the diagram above) – have similar preferences.    My experience is that we do often find a smattering of individuals belonging to “brackish” species in very hard water, as we have in Castle Eden Burn.  Average conductivity (based on Environment Agency records) is 884 µS cm-1; however, values as high as 1561 µS cm-1.   The fluctuating discharge plays a role here, as any evaporation will serve to concentrate those salts that are naturally present in hard freshwater.   This should probably not be a big surprise: life in brackish waters involves adapting to fluctuating osmotic regimes so species that can cope with those conditions are also likely to be able to handle some of the consequences of desiccation.

Average values of other chemical parameters from 2011 to present, based on Environment Agency monitoring are: pH: 8.3; alkalinity: 189 mg L-1 CaCO3; reactive phosphorus: 0.082 mg L-1; nitrate-nitrogen: 1.79 mg L-1; ammonium-nitrogen: 0.044 mg L-1.   There is some farmland in the upper catchment, and the burn also drains an industrial estate on the edge of Peterlee but, overall, nutrient concentrations in this stream are not a major concern.   The Environment Agency classifies Castle Eden Burn as “moderate status” due to the condition of the invertebrates but does not offer any specific reason for this. I suspect that the naturally-challenging habitat of Castle Eden Burn may confound assessment results.

I’ve also been given some data on discharge by the Environment Agency which shows how patterns vary throughout the year.  The two sampling locations are a couple of kilometres above and below the location from which I collect my samples and both have more regular flow.  However, we can see a long period between April and September when discharge is usually very low.   The slightly higher values recorded in July are a little surprising, but are spread across a number of years.   It is also, paradoxically, most common for the burn to be dry in July too: clearly, a month of extremes.  As my own visits have shown, it is possible for the burn to be dry at almost any time of the year, depending on rainfall in the preceding period   The dots on the graph (representing ‘outliers’ – records that exceed 1.5 x interquartile range) show that it is also possible to record high discharges at almost any time during the year too.  I should also add that, as I am not a hydrologist, I am rather outside my comfort zone when trying to explain these patterns.  I would have said ‘out of my depth’ though that’s not the most appropriate phrase to use in this particular situation.

CED_discharge

Discharge in Castle Eden Burn, as measured by the Environment Agency between 2007 and present.   Measurements are from NZ 4136 2885 (‘upstream’) and NZ 45174039 (‘downstream’).  

* Note on Humidophila contenta:it is almost impossible to identify this species conclusively with the light microscope as some key diagnostic characters can only be seen with the scanning electron microscope.   However, all members of this complex of species share a preference for intermittently wet habitats so these identification issues are unlikely to lead to an erroneous ecological interpretation.  It is probably best to refer to this complex as “Humidophila contenta sensu lato” rather than “Humidophilasp.” order to distinguish them from those species within the genus that can be recognised with light microscopy.

Reference

Lange-Bertalot, H., Hofmann, G., Werum, M. & Cantonati, M. (2017).  Freshwater Benthic Diatoms of Central Europe: over 800 Common Species Used in Ecological Assessment. English edition with updated taxonomy and added species.  Edited by M. Cantonati, M.G. Kelly & H. Lange-Bertalot.  Koeltz Botanical books, Schmitten-Oberreifenberg.