Spotting spots …

A recurring theme of this blog has been that the process of “seeing” is one that requires a mind that is preconditioned to process the signals passed to it from the optic nerve.  For much of the natural history that we see on a country walk, that link is already in place: we notice trees, birds, butterflies and so on, even if we can’t always put a name on what we are looking at.   That means that natural history becomes a closed loop: noticing birds means we are more likely to notice differences amongst birds – whether in type or habit or the timing of their visits – which, in turn, stimulates greater mental awareness.   But not noticing something is equally self-perpetuating and many of my posts have focussed on the natural history that most of us routinely overlook.

A good reason for not noticing aspects of natural history is that the organisms in question do not even look “alive” in the first place.  Another is that they are so tiny that they are either microscopic or only just apparent with the naked eye.   As this post deals with a group of algae that fit both of these criteria, you can be forgiven for not knowing about these organisms before, but also for wondering what kind of person scratches away at submerged stones in the counterintuitive hope of finding life.

Earlier posts have talked about algae that form crusts on rocks, such as the red algae Hildenbrandia rivularis(see “More about red algae”) or the cyanobacteria of the genus Chamaesiphon (see “A bigger splash …”).  Chamaesiphon tends to form brown to almost black spots on rocks but there is one other type of alga that also forms dark brown patches of rock in streams that I have not previously written about.  This is the brown alga Heribaudiella fluviatilis.    Most of the brown algae are marine and largely beyond the scope of this blog (but see “Swimming in a sea of ignorance”) and there are just two genera recorded from UK freshwaters.   Both are rarely recorded, but probably more common than we think.  I’ll come back to this later in the post, but freshwater brown algae really do encapsulate the conundrum in the first paragraph: not noticing becomes self-perpetuating as we repeatedly fail to realise that there is something we ought to notice.

The photos at the top of the post – taken by Wolfgang Schülz in Germany – show characteristic colonies of Heribaudiella fluviatilis, with distinct margins, in contrast to Chamaesiphon crusts, whose margins are less clearly delimited.  The best way to confirm that you are looking at Heribaudiella rather than Chamaesiphon, however, is to scrape up the crust and examine it under a microsope.  Then the characteristic cells arranged in short filaments, each with several yellow-brown chloroplasts, should be obvious.   

A microscopic view of filaments of Heribaudiella fluviatilis.  Photo by Chris Carter.  The pictures at the top of the post show macroscopic views of Heribaudiella on stones from a river in Germany (courtesy of Wolfgang Schülz).

The late Nigel Holmes was good at spotting Heribaudiella colonies (which can be just a couple of millimetres across) on stones in rivers but the ability (or, perhaps, the inclination) to notice these has declined thereafter.   We know it is out there because the primers we use for molecular studies of diatoms also appear to pick up Heribaudiella as by-catch, and we were surprised at how often it appeared.   Curiously, these primers also differentiated between Heribaudiella and another brown algal genus, Bodenella.  I wrote about Bodenella in Depths of Imagination and also mentioned that there was some dispute about whether this was truly distinct from Heribaudiella.   Last summer, I revisited a stream from which this “Bodenella” was recorded but failed to get enough material to make a satisfactory examination of its microscopic features.   The jury is out on this, especially as the habitats where we’ve detected “Bodenella” are so different to the locations where it has been securely identified elsewhere in Europe.   Personally, I would be reluctant to add this as a new UK record without having backed up a molecular identification with some more traditional taxonomy.  

Having noticed that we’re not noticing Heribaudiella fluviatilis, one of the best arguments for stimulating interest in this organism is that it tells us useful information about the state of the stream in which it is growing.  Most of the chemical data describing its habitat that I’ve seen would suggest that it prefers “good status” water and is, thus, a sign that the ecosystem is healthy.   In the final analysis, though, records of Heribaudiellaprobably say more about the quality of the observer than of the habitat.   

References

Koletić, N., Alegro, A., Vuković, Rimac, A. & Šegota, V. (2018).  Spotting the spots: the freshwater brown algaHeribaudiella fluviatilis (Areschoug) Svedelius within stream communities of southeastern Europe.   Cryptogamie, Algologie 39: 449-463.

(this paper, interestingly, notes an association between Heribaudiella and Hildenbrandia that is also discussed in Depths of Imagination)

Wehr, J.D. (2011).  Phylum Phaeophyta.  pp. 354-357.   In: The Freshwater Algal Flora of the British Isles(edited by D.M. John, B.A. Whitton & A.J. Brook).  Cambridge University Press, Cambridge. 

Wehr, J.D. & Stein, J.R. (1985).  Studies on the biogeography and ecology of the freshwater phaeophycean alga Heribaudiella fluviatilis.  Journal of Phycology 21: 81-93.

The work that we described in Depths of Imagination was recently published:

Schülz, W., Kelly, M.G., King, L. & Cantonati (2021).  Did Zebra mussel fill the type habitat of a worldwide-rare freshwater brown macroalga?  Aquatic Conservation: Marine and Freshwater Ecosystems 31: 3657-3659.

Wrote this whilst listening to: Hildegaard of Bingen, and Nick Cave’s Murder Ballards.

Currently reading: Pen Vogler’s Scoff, about the history of British food.

Cultural highlight: Munich: Edge of War – film based on Robert Harris’ book set around the Munich crisis of 1938 and starring George Mackay and Jeremy Irons.  

Culinary highlight: mutton steak from our local organic butcher, with homemade chips and peppercorn sauce.

Unlikely bedfellows …

2022 started with enforced isolation due to a positive Covid test, so my first post of the year has to be desk- rather than field-based, picking up my tour of the major algal groups.   So far, I have covered Cyanobacteria (“Shuffling the pack …”), green algae (“The big pictures …”) and red algae (“Rhapsody in red …”).  This post will deal with the last of the major groups of algae, the Chromista, the kingdom which includes the diatoms, chrysophytes, brown algae and yellow-green algae.  Microscopic silica-encased diatoms and giant kelp may seem like unlikely relations, so read on …

Be prepared for a veritable porridge of terminology, reflecting the gradual evolution of ideas.   From the point of view of someone who wants to understand how different groups of algae are related, all are more-or-less synonyms.  However, each also has a precise usage, representing their place in a hierarchy that also includes non-algae.   The Chromista (also known as SAR (“Stramenopile, Alveolates and Rhizaria”) is a Kingdom (akin to “plants”, “animals” or “fungi”), within which the algae are spread across four Divisions.  Most of the algae that are found in freshwater benthic habitats (the main focus of this blog) are found in the Heterokontophyta (a subkingdom, also known as the Stramenopiles) and, more specifically, in the Ochrophyta (a superphylum).   However, some of the other groups are very abundant in freshwater and marine plankton.   All are thought to share a common ancestor, a red algal-type cell, although many have subsequently lost their chloroplasts.

A guide to the terminology of the Chromista.

TermDefinition
ChromistaAlgae with chlorophyll c but not b, evolved from a secondary endoysmbiosis  with a eukaryotic (red) alga, along with protists descended from these which have lost photosynthetic capability and plastids 
HeterokontaOrganisms possessing two flagellae of different lengths for at least part of the life cycle (hetero = different; kontos = punting pole)
StramenopilesOrganisms with a flagellum with short hair-like extensions.   The term is derived from Latin words stramen (= straw) and pilus (= hair).   However, I’ve also seen suggestions that  stramen refers to the yellow-brown colour of the chloroplasts.
OchrophytaRefers to the colour of the cells, from a Greek rather than a Latin root (okhra = yellow). 

One further surprise before we go on is that the non-algal representatives of this group include Cryptosporidium, responsible for gastrointestinal illness, Plasmodium, the parasite that causes malaria, and Phytophthora infestans, which causes potato blight. 

None of the groups in the subkingdom Hacrobia have been covered in this blog, mostly because they are not abundant in the habitats that I spend most of my time studying.   I do see cryptophytes (“Cryptista”) but never in large numbers, and when I do they are invariably moving too fast for me to photograph.  The Haptophyta include the Coccolithophores, which are very abundant in marine plankton, but not in freshwaters.  The only time one has appeared in this blog is in a feature on the Hilda Canter-Lund competition in 2017 (see below).  

Overview of the Chromista, following Ruggeiro et al. (2015).  Groups containing algae are indicated by arrows.  

Harosa, the other subkingdom, is divided into the Rhizaria, comprising non-photosynthetic  amoebae-like organisms and the Halvaria.  The Halvaria contains many important groups of algae.   The first of these is the Alveolata, which includes the dinoflagellates, again mostly found in plankton, both marine and freshwater.  Another major group is the Pseudofungi  (which includes the organism responsible for potato blight) and the final group is the Heterokontophyta, of whose algal interest lies in the superphylum Ochrophyta.

Overview of the Heterokontophyta, following Ruggeiro et al. (2015).  

There are four major groups of algae in the Ochrophyta of which the  diatoms, Bacillariophyceae, are widely-covered in this blog, needing no extra explanation here.  The Chrysophyceae have been encountered a few times but, as for some of the other Chromistan algae, much of their diversity is planktonic rather than benthic.

The best known representatives of the Phaeophyceae are the kelps and other brown seaweeds of marine littoral zones.  I have written about these, but not much.   Michiel Vos’ blog An Bollenssor is a good place to start exploring this group.   There are freshwater representatives, but their only appearance in this blog was to record their likely extinction from the Bodensee.   I was planning to write a post last summer but inadvertently deleted all my photos.   They are probably more common in rivers than most people realise but are easily overlooked and, as a result, rarely recorded.

Finally, the Xanthophyceae, the yellow-green algae, include important freshwater representatives such as Vaucheria and Tribonema. They differ from many of the other algae in the Chromista in that they are typically green in colour, lacking the extra pigments that give many other representatives yellow, orange and brownish hues.   The technical definition of the Chromista is that they contain chlorophyll c as well as chlorophyll a, and Xanthophyceae fulfil this criterion.  However, many of the other Chromista also contain extra pigments (carotenoids and xanthophylls) which are responsible for these other colours.   Xanthophyceae do have carotenoids and xanthophylls, but not the types or quantities that alter appearance.   For a long time, as a result, they were classified with the green algae.   That had a certain superficial logic until people started focussing on aspects of the organisms that were less obvious.  The outcome is a more natural classification but problems when faced with a small greenish cell that doesn’t fit any descriptions in the “green algae” section of an identification guide.   If you’ve managed to reach the end of this post, you’ll have realised by now that taxonomists and phylogeneticists don’t like to make life too easy for the rest of us. 

Further reading

Adl., S., Bass, D., Lane, C.E., Lukeš, J. et al. (2018).  Revisions to the classification, nomenclature, and diversity of eukaryotes.  Journal of Eukaryotic Microbiology 66: 4-119.   

https://doi.org/10.1111/jeu.12691

Derelle, R., López-García, P, Timpano, H. & Moreira, D. (2016).  A phylogenetic framework to study the diversity and evolution of the Stramenopoiles (= Heterokonts).  Molecular Biology and Evolution 33: 2890-2898.  https://doi.org/10.1093/molbev/msw168

Martin, W.F., Garg, S. & Verena, Z. (2015).  Endosymbiotic theories for eukaryotic origin.   Philosophical Transactions of the Royal Society of London Series B  3702014033020140330 http://doi.org/10.1098/rstb.2014.0330

Ruggiero, M.A., Gordon, D.P., Orrell, T.M., Bailly, N., Bourgoin, T., Brusca, R.T., Cavalier-Smith, T., Guiry, M.G., Kirk, P.M. (2015).  A higher level classification of all living organisms.  PLOSone DOI:10.1371/journal.pone.0119248

Appendix

Links to posts describing the major groups of Chromista found in freshwaters (and, in one case, marine habitats).  Only the most recent posts are included, but these should contain links to older posts (you can also use the WordPress search engine to find older posts).

GroupLink
CryptistaThe underwater world of Ennerdale Water …
HaptophytaHow to win the Hilda Canter-Lund Prize (4)
AlveolataSubmerged soap opera …
Invisible worlds at Malham Tarn
Heterokontophyta 
    BacillariophyceaeWho do you think you are?
    ChrysophyceaeThe little tarn of horrors …
Fade to grey …
A brief excursion to Norway
    PhaeophyceaeDepths of imagination …
Swimming in a sea of ignorance … 
Spotting spots …
    XanthophyceaeWhen a green alga is not necessarily a green alga …
The littoral ecology of Lough Down …