Survival of the fittest (2) …

As well as the bright green flocs of Tribonema, the stream draining the Hadjipavlou chromite mine also had bright orange-red growths on some of the pebbles on its bed.  These seemed to be composed primarily of the Cyanobacterium Chamaesiphonthough I am still not sure what species.   Using the limited literature I have, from the UK and Germany, I would opt for either Chamaesiphon polymorphusor C. polonicus.   This particular alga was very easy to remove from stones, compared to other epilithic Chamaesiphon species (see “A bigger splash …”).  This is a feature of C. polymorphus, though the colour is more typical of C. polonicus*.  On the other hand, that bright colour could be the response to high solar radiation, so maybe my north European guides are not that reliable.  It could be something else altogether.

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Chamaesiphon growths on pebbles in the stream draining Hadjipavlou chromite mine in the Troodos mountains, Cyprus, March 2019.

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Colonies of Chamaesiphon from Hadjipavlou chromite mine under the microscope.   Scale bar: 10 micrometres (= 1/100thof a millimetre). 

In addition to the Chamaesiphon, there were a few diatoms, mostly Achnanthidium minutissimumand Meridion circulare.   These are typical species of metal-rich streams, as is the general lack of diversity that was evident.   There were also a few filaments of the cyanobacterium Phormidium, along with quite a few Paramecium and Vorticella.  As these are both heterotrophs that feed on organic matter, their abundance is probably at least partly a reflection of the long time that the sample spent in my suitcase between collection and analysis.  The latter is a fascinating organism to watch: it is a goblet-shaped cell with a fringe of cilia around the lip (or “peristome”).  These beat in unison to create water currents that draw small particles towards the cell.   These particles mostly at least an order of magnitude smaller than the algae)  are then collected in food vacuoles where they are digested.   A few of these vacuoles can be seen in the image of Vorticella below.

Vorticella is attached to its substrate by a stalk which contains contractile filaments, giving it spring-like qualities.  Watching a Vorticella is a beguiling experience, with the undulating rows of cilia drawing food into the vestibule (as the opening is known).  At intervals, the whole cell lurched across the field of view as the “spring” in the stalk suddenly contracted, shortening the stalk.  After this, the stalk would gradually extend again, the cilia not having missed a beat meanwhile.   This process may simply be a device that enables the Vorticellato exploit its locality to the full, as well as creating some additional turbulence to keep a steady flow of particles towards the peristome.  To be honest, I haven’t seen a more convincing explanation but, even if we don’t know why it does what it does, Vorticella is a fascinating organism to watch, whether or not I understand what is going on.

I’ll be coming back to talk more about the diatoms in a future post, and writing these posts has also reminded me that I’ve never written about the interesting mine sites almost on my own doorstep.  I cut my ecological teeth looking at these habitats back in the 1980s and they are striking examples of natural selection in action.   So, plenty of potential for more left-field natural history …

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Other organisms present in the Hadjipavlou chromite mine. a. – d.: Meridion circulare; e. Phormidiumsp.; f. Vorticellasp.   Scale bar: 10 micrometres (= 1/100thof a millimetre). 

* Note: after I had written this post Brian Whitton confirmed that it was, most likely, Chamaesiphon polonicus.

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Survival of the fittest (1) …

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When I signed up to a trip to Cyprus in late March I was anticipating feeling some warm Mediterranean sun on my skin after the ravages of the British winter.  I did not expect snow and sleet.   However, as one of our destinations was the Troodos mountains, the location of Cyprus’ only ski resort, maybe it was a case of unrealistic expectations.   Fortunately, we realised our mistake just in time to pack some warm clothes, and the unseasonable weather did, at least, mean that the spring flowers at lower altitudes were, even by Cypriot standards, particularly impressive.

I was in Cyprus primarily as a camp follower on a reconnaissance trip for a geology and botany excursion next year.   Cyprus is, to put it in layman’s terms, the outcome of a collision between the African and European continental plates.   The Troodos mountains are a geologist’s paradise, having a wide range of features arising from this and from associated volcanic activity.   As the molten rocks cooled, minerals precipitate out and the resulting geological strata reflect differences in the melting points of these minerals.   Some of these minerals, such as chromite, are commercially valuable and have been mined for centuries.   Indeed, the name Cyprus itself is derived from cuprous, the Greek word for copper, which was mined here since 4000 BC.

The Hadjipavlou mine is set amidst pine forests close to the highest point of the Troodos.  It was an active chromite mine from 1950 to 1954 and from 1979 to 1982 but was abandoned when cheaper sources of chromite became available in South Africa.   Over a million tonnes of ore were extracted in the period when the chromite mines in the area were active, but now there are few obvious signs apart from this adit driven into the hillside.   A small stream bearing water that has percolated through the rocks and collected in the mine’s galleries emerges from the mine entrance and tumbles down the hillside to join the stream below.   This, on closer inspection, has some quite interesting microbial growths.

First of all, having been told that this is a chromite mine, you might expect the water to carry toxic concentrations of heavy metals.   So you might also be surprised to see abundant growths of bright green algae thriving in the stream immediately downstream of the mine entrance.   This is, in fact, a common phenomenon in mine waters and happens, we think, because the fast-growing algae evolve metal tolerance whilst the animals that feed on them are slower to adapt.   This is, literally, survival of the fittest and, with nothing to eat them, the algae grow prolifically.

These filaments belong to the genus Tribonemawhich, despite being bright green in colour, actually belongs to the yellow-green algae, the Xanthophyta, rather than to the green algae.  This group is actually more closely related to the diatoms than to the green algae, though it can be hard to understand why simply by peering through a microscope.  One useful test is to add a little iodine  solutionto the slide: this binds to the starch inside green algae cells, staining them a dark brown colour.   The Xanthophyta, by contrast, do not have starch as their storage product so the cells are not stained by iodine.   The only other member of this group that I have discussed in this blog is Vaucheria, a very different alga (see “Who do you think you are?”).

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Tribonema cf affinein the channel draining the Hadjipavlou chromite mine in the Troodos mountains, Cyprus, March 2019.   a. close-up of the alga in situ; b.  microscopic view of filaments; c. fragments of disintegrated filaments showing the H-shaped cell endings.  Scale bar: 10 micrometres (= 100thof a millimetre).   

Tribonemahas simple, unbranched filaments with two or more plate-like chloroplasts arranged around the cell periphery.   One other feature is the arrangement of the cell wall, which tends to consist of two overlapping halves.  When filaments disintegrate (as they often do) the fragments have an H-shape, with each end forming half the cell wall of a different cell.   The cells are, in fact, cylindrical but this is not apparent with the flattened perspective of a high magnification objective.   This feature is not universal in the Xanthophyta, nor is it unique to this group (a few green filamentous algae show the same characteristic) but it is a useful hint that you may be looking at Tribonema.

Whilst lush growths of algae is a common feature of streams draining mines, the species that form these growths can vary.   In the northern Pennines, I am used to seeing green algae in these habitats, but there are at least three different genera that I find.  Typically there is just one filamentous alga in this location, and they tend to be  constant over time: they are reliable sources for material for undergraduate practical classes as a result.  There is more to this story but I will have to come back to it at some point in the future.  .

There is also more to the algal flora of the Hadjipavlou chromite mine but, again, that will have to wait for another post.  I should also confess that, although I visited the mine briefly last year, these samples were collected by Heather, whilst I was sitting snugly below the snow line.

Hug a Brexiteer …

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I was hoping to start this blog, written on the original date for Brexit, noting that, in contrast to most other UK citizens, I had begun the day outside the EU but had, during the course of the morning, re-joined the Union.  The delay in the date for Brexit messes up that neat little opener but the experience of walking across the Green Line in Nicosia, from the Turkish Republic of Northern Cyprus to the Republic of Cyprus is a sobering reminder of the way that festering resentments within communities can spiral out of control.

There are few certainties in UK politics at the moment but, based on voting patterns in the referendum, it is very likely that over 40 per cent of the population is going to be dissatisfied with the outcome of the Brexit negotiations.   I have made my own views clear in this blog and I know that some of my readers disagree with my views.   This post is not about the rights and wrongs of Brexit but about the aftermath, and how the country as a whole treats that large proportion who will almost certainly be disappointed by the outcome.

The situation in Cyprus is complex but there are parallels to Brexit in that, after 1945, the key political question concerned a union (with Greece in this case) that would have left a significant minority of the population feeling disenfranchised.  On the other hand, there is one key difference from the UK in 2019 in that the disenfranchised minority were ethnically distinct.  In 1974 the failure to find a mutually-acceptable settlement led eventually to invasion by Turkish forces and the partition of the island which persists to this day.   We in the UK should be thankful that there is no such clear “them” and “us” distinction as our politicians pick their way through the morass of possibilities.

But the absence of a physiognomic, linguistic or religious differentiator in the Brexit debate does not mean that differences – and resentments – will not persist long after a final settlement is agreed.  That means the country, once it has resolved the present Brexit stalemate, will need to think seriously about a reconciliation process to heal the divisions.   Time, alone, will not necessarily be enough; indeed, time may even sharpen the divisions, especially if the economy is not buoyant in the post-Brexit years.  I live in a liberal bubble where almost everyone I encounter is pro-EU and opposed to Brexit; however, if the UK does end up leaving the EU, there is no point in brooding over what might have been.  We will need to pick ourselves up, dust ourselves down and move on.   And hug a Brexiteer.  Judging by the press reports, many of them are going to be just as disappointed as the Remainers.   At least we will have that in common

 

Return to Cyprus …

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

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

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

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

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

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

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

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

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

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

Reference

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

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

Cypriot delights …

I could not return from my visit to Cyprus without an algal sample and a fine opportunity presented itself early last week when we visited the Avgás Gorge, on the Akámas peninsula at the west coast of Cyprus, just north of Pathos.  This is a spectacular limestone ravine whose steep sides offered welcome relief from the Mediterranean sun.  At points, as in the photograph above, the ravine narrowed to just a few metres wide, reminiscent of the siq which guards the entrance to Petra except that instead of exquisite carvings we stumbled across a Russian team conducting a glamour shoot.

A small stream made its way down the gorge.  The presence of woody debris at intervals suggested a considerable head of water during the winter months but, at this time of year the water has reduced in power, tumbling across a series of boulders into stagnant pools, interspersed with short runs shaded either by the high cliffs or the vegetation that flourished away from the harsh glare of the sun.

In the sections where water was still running there were clumps of Chara tangled up with filamentous algae, with plenty of bubbles of oxygen as evidence that both were busily photosynthesising away.  The filamentous algae was a coarse unbranched filament that was clearly a relative of Cladophora but which did not match any of the genera or species that I had encountered before (see “Fieldwork at Flatford” for similar situation).   There were pebbles and cobbles between these clumps, their surfaces criss-crossed by the galleries of caseless caddis larvae – probably Psychomyiidae, according to Richard Chadd.

Chara growing in the stream at Avgás Gorge in western Cyprus, April 2018.

Cladophora or a relative growing in the stream at Avgás Gorge in western Cyprus, April 2018.  The scale bar is 50 micrometres (= 1/20th of a millimetre). 

There were a number of diatoms present too, the most abundant of which was a chain-forming Ulnaria.  Unfortunately, despite having just co-authored a paper on Ulnaria from Cyprus, I cannot name the species, as I saw no cells in valve view.  I will have to return to this subject once I have prepared a permanent slide from the sample that I brought back.   The chloroplasts in the illustration below are not in a very healthy state because the sample lived in a fridge for almost a week before I was able to get it under a microscope.  Had I looked at this sample 20 years ago, I would have assumed that I was looking at a species of Fragilaria, as most keys then stated categorically that Synedra (the former generic name) were solitary rather than chain-forming.  However, we now know that there are several Ulnaria species that form chains although most that I see in my regular haunts in the UK do not.   Our paper states that the species we describe form short chains although, as we worked from cleaned samples collected by other people, I now wonder if that was an artefact of the preparation process and whether these, too, formed longer chains in their living state.

A chain of Ulnaria from the stream at Avgás Gorge in western Cyprus, April 2018.  Scale bar: 20 micrometres (= 1/50th of a millimetre).

Though it is rare for me to stray into describing the invertebrate life of streams, the Psychomyiidae are actually an important part of the story here, as the larvae graze algae from the surface of the stones.  They create a silk tube and this, in turn, becomes covered with fine sediment to create the galleries that are visible in the picture.   We know that invertebrates can change the composition of the attached algae by grazing but I sometimes wonder if the caseless caddis larvae also change the composition by creating myriad patches of very fine sediment across the rock surface.   If you look closely you can also see a couple of Simuliidae (blackfly) larvae.  These attach themselves to the rock by a circle of hooks at their last abdominal segment (I think that is “bum” in entomological language) and then use fan-like structures around their mouths to filter tiny particles from the water.  However, I have also seen Simuliidae larvae bent double on rock surfaces in order to hoover up particulate matter and algae that live there.

That, I thought, was just about enough natural history for one day.   I put my toothbrush and bottle back into my rucksack and made my way back down until, turning the corner, I stumbled across the Russian glamour models.   I’ve often written about the similarities between freshwater ecosystems in different parts of Europe but you don’t often see a bikini – or less – in April in my cold, damp corner.

Galleries of Psychomyiidae larvae on the top of a limestone cobble from the stream at Avgás Gorge in western Cyprus, April 2018.

Reference

Cantonati, M., Lange-Bertalot, H., Kelly, M.G. & Angeli, N. (2018).  Taxonomic and ecological characterization of two Ulnaria species (Bacillariophyta) from streams in Cyprus.   Phytotaxa 346: 78-92.

Wallace, I. (2003).   The Beginner’s Guide to Caddis (order Trichoptera).  Bulletin of the Amateur Entomologists’ Society 62: 15-26.

Letter from Cyprus

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Travel can be one of the most rewarding forms of introspection ….
Laurence Durrell, Bitter Lemons, 1957

When I stepped off flight EZY1973 from Manchester to Paphos on Saturday night I passed a personal milestone. Arriving in Cyprus means that I have now visited all 28 Member States of the European Union. Starting with (West) Germany in 1972 on an exchange visit before the UK was even a member of the European Economic Community, followed shortly after by a family holiday to southern Austria (where my father had been stationed just after the war) with a day trip to Slovenia (then part of Yugoslavia), the number started to increase in the late 1990s when I became involved in the work of CEN, the European Standards Agency and, from the mid-2000s onwards, with the intercalibration exercise associated with the Water Framework Directive. A few years ago I made a list and realised just how many I had visited, after which, I have to admit, my choice of conference and holiday destinations was driven by this rather childish whim. Latvia, Malta and Bulgaria, all subjects of posts on this blog, were ticked off, leaving just Cyprus. This year, a family holiday to celebrate my mother’s 80th birthday provided the opportunity and, after some shameless lobbying, we had booked a villa near Paphos via AirBnB and were on our way.

How Europe has changed in the 47 years since my first overseas trip. Twelve countries were behind the Iron Curtain, three of the remainder were right-wing dictatorships. Two have merged (East and West Germany) whilst seven have become disentangled from previous relationships (the Baltic States from the USSR, Slovenia and Croatia from the former Yugoslavia and the two former constituents of Czechoslovakia from each other). Cyprus, from where I am writing, was in political chaos in the early 1970s. A former British colony whose territory was argued over by Greece and Turkey, it was soon to be split into two, separated by a buffer zone. I used to browse my Collins World Atlas assuming national borders to be fixed and immutable; the older and wiser me wonders where (and when) the next changes will come from.

The intercalibration exercise, in particular, was an opportunity for an exchange of ideas and I counted co-authors from 23 of the 28 EU states on my publication list. Looking back, these papers show remarkable consistency in some aspects of ecology across Europe whilst, in other respects, I am much more cautious about assuming that knowledge gained in my damp corner of north-west Europe can be applied to warmer and more continental regions. This publication list includes, incidentally, two papers about Cyprus, despite never having either visited before or having a native Cypriot on my list of co-authors. In the first paper, we worked with an Austrian employed by the Ministry of the Environment but, for the second, the samples were collected and analysed by Italians and Germans whilst I helped out with data analysis. Scientific colonialism is not, perhaps, dead?

My favourite? I don’t think I should single one of the 28 out. The food and culture of the warm lands of the Mediterranean basin draw me but I think that the parched summer landscapes would lose their appeal if I was there for too long. I find the grey, damp climate of my own corner of Europe wearisome but the greenness of the Spring and Summer, and the Autumn colours almost compensate. My ideal, in other words, seems like it should be a semi-nomadic existence but that, too, would pale with time. The truth is that, for me, elsewhere, being wanted, is always more wondered at ….

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