Once upon a time, measuring the quantity of algae growing on the beds of streams and rivers was a painstaking, slow process that invariably revealed large amounts of spatial and temporal variation that, very often, obscured the ecological signals you were looking for. That has changed in the last decade thanks to the availability of field fluorimeters such as the BenthoTorch. This makes it much quicker and easier to measure chlorophyll concentrations, the usual proxy for algal quantity. Thanks to devices such as this it is now much easier to discover that your ecological signal is masked by spatial and temporal variation.
We’ve generated a lot of data about the fluxes of algae in the River Ehen using a BenthoTorch over the past five years and are in a position where we can start to make some generalisations about how the quantity of algae vary over the course of a year. In broad terms, the results I showed in “The River Ehen in January” back in 2014 have not varied greatly over subsequent years, with peak biomass in mid-winter and low biomass in the summer (due, we presume, to intense grazing by invertebrates). Curiously, we see a much less distinctive seasonal pattern in the nearby Croasdale Beck, but that is a story for another day….
The BenthoTorch uses an algorithm to partition the fluorescence signal between three major algal groups and, though this is not without issues (see below), I thought it might be interesting to see how these groups varied with biomass trends, and consider how this links to ecological theory. The first group I’m considering are the green algae which, in this river, are mainly filamentous forms. The general pattern, seen in the graph below, is for a gradual increase in the proportion of green algae, which fits with the current understanding of thicker biofilms having greater structural complexity with filamentous algae out-competing attached single celled algae to create a “canopy” of algae that are more effective at capturing light and other resources. The relationship is, however, strongly wedge-shaped so, whilst many of the thickest biofilms have a lot of green algae, there are also thick biofilms where green algae are scarce or even non-existent. Croasdale Beck shows a similar, but less pronounced, trend.
Relationship between the proportion of green algae and the total quantity of benthic algae (expressed as chlorophyll concentration) in the River Ehen (a.) and Croasdale Beck (b.). The blue lines show quantile regression fits at p = 0.8, 0.5 and 0.2. The image at the top of the post shows Ben Surridge using a BenthoTorch to measure algal biomass beside Croasdale Beck in Cumbria.
The second graph shows that this pattern of a gradual increase in proportion is also the case for diatoms and, once again, there is a broad wedge of points with an upward trend. But, once again, there are also samples where biomass is high but diatoms are present in very low numbers or are even absent. What is going on?
The problem is clear I think, if one looks at the final image in “The only way is up …” where the very patchy nature of algal communities in the River Ehen (and, indeed, many other rivers). There are plenty of algae on this boulder, but not organised in a homogeneous manner: some zones on the boulder are almost pure diatom whilst others are almost pure green algae (and there are also zones that are almost pure Lemanea– I’ll come to that in a future post). We try to sample the stones as randomly as possible so you can see the potential for getting very different numbers depending on where, on a stone, we point the BenthoTorch’s sensor.
Relationship between the proportion of diatoms and the total quantity of benthic algae (expressed as chlorophyll concentration) in the River Ehen (c.) and Croasdale Beck (d.). The blue lines show quantile regression fits at p = 0.8, 0.5 and 0.2.
With experience, you can make an educated guess about the types of algae present in a biofilm. I’ve tried to capture this with my watercolours, using washes of raw sienna for the diatoms and a grass-green for the green algae, which roughly matches the colour of their respective growths in the photo in my earlier post. The two groups of algae a are relatively distinct on that particular boulder. The top row roughly matches the upper “edge” of the graph showing variation in diatoms, whilst the bottom row emulates the upper “edge” of the graph showing variation in green algae. These are the two extreme situations; however, we also often see darker brown growths in the field, which can be recreated by mixing the raw sienna and grass-green together. When I peer through a microscope I often see green algae smothered in diatoms: genera such as Oedogoniumare particularly prone as they have less mucilage than some of the others we find in the Ehen. Their filaments often host clusters of Fragilariacells as well as Achnanthidium minutissimum, whilst stalked Gomphonemaand chains of Tabellaria flocculosaoften grow through the tangle of green filaments. The dark brown colour is deepened yet further by the colour of the underlying rock, so my effort on white watercolour paper is a little misleading.
A colour chart showing how different proportions of green algae and diatoms influence the colour of biofilms.
The final graph shows how, as the average biomass increases in the River Ehen, so the variability in biomass also increases. The River Ehen is one of the cleanest rivers I know but I suspect that this pattern in benthic algal quantity could be reproduced in just about any river in the country. What I would not expect to see in any but the purest and most natural ecosystems is quite so much variation in the types of algae present. Once there is a little enrichment, so I would expect the algae to become more of a monoculture of a dominant filamentous alga plus associated epiphytes. Like much that happens in the microscopic world of rivers, it is easier to describe than it is to measure.
That, however, is only part of the story but I’ll come back to explain the patterns in the other main groups of algae in the Ehen and Croasdale Beck in my next post.
The relationship between mean chlorophyll density and the standard deviation (based on measurements from five separate stones) for samples from the River Ehen and Croasdale Beck.