Blinded by the light: victims of the night

In late October, the municipality of Buenavista del Norte on the Canary Island of Tenerife, celebrates the day of the Virgin of Los Remedios, including, among other features, a big light display. As a child, Airam Rodríguez noticed that many shearwaters would also drop in (literally) for the festivities, attracted by the bright lights, but unable, in many cases, to get back in the air. Many of these shearwaters died from a variety of causes, including the impact of flying into the ground, dehydration, predation and poaching. As an adult, Rodríguez collaborated with researchers around the world to evaluate the scope of light-induced shorebird fallout.


Fallout victim: grounded Short-tailed Shearwater. Credit: Airam Rodríguez

The researchers began their work by searching a science citation index – the Web of Science – for articles on light-induced seabird mortality. They used references from these articles to find additional articles. In addition, they used the internet and social media to find programs in which citizens are encouraged to report grounded birds, and contacted people associated with these programs to get qualitative and quantitative data.

Rodríguez and his colleagues discovered light induced seabird fatality on 47 islands, three continental locations and across all of the world’s oceans. Of 115 species of burrow-nesting petrels, 56 have been reported as grounded by light. Several other groups of birds, including puffins, auklet and eiders also suffer from light-induced fallout, and it is very likely that more species are unreported.


Numbers of reported grounded seabird fledglings across the globe.  Circle size = numbers of birds  reported. Numbers = number of species affected. Circle color = IUCN (endangerment) category for each species as follows: CR = critically endangered, EN = endangered, VU = vulnerable, NT = near threatened, LC = least concern.

Of deep concern is that 24 species are globally threatened. In addition, fallout has been reported at sea, induced by lights used for fisheries and by lights on oil platforms. All of the studies of light-induced fatalities on land documented the highest mortality in fledglings that are grounded during their first flights from their nests toward the ocean.


Numbers of species of threatened seabirds that were rescued across the globe.  Numbers were not available for species with ? symbol.

Researchers don’t know why birds are attracted to lights. Perhaps birds view lights as a source of food; for example some species eat bioluminescent prey. Alternatively, as cavity-nesting birds, the only light these chicks see is from their burrow entrance, particularly when their parents bring in food, so the fledglings might confuse light with a food source. Lastly, artificial lights might override any celestial light cues the birds normally use for navigation, confusing them and causing them to crash to the ground. Supporting this hypothesis, seabirds generally don’t crash into lights, which might be expected if they mistook a light for bioluminescent prey.

Cory's shearwater fledgling at their nest at Tenerife Canary Islands. Photo by Beneharo Rodríguez

Fledgling Cory’s Shearwater first sees the light of day after emerging from its burrow at Arona on southern Tenerife Island. Credit: Beneharo Rodríguez

So what can be done about this problem? Accurate data are hard to come by, as many estimates of fallout-induced mortality come from relatively untrained volunteers, who are less likely to report dead birds. As one example, on Kauai, surveys from a general public rescue program for Newell’s Shearwaters identified 7.7% mortality, whereas later systematic surveys by trained researchers indicated 43% mortality. In some rescue operations, birds are banded and released, which, in theory, allows researchers to estimate the survival rate of rescue birds, but, in practice, these data are usually insufficient for accurate estimates

Rodríguez and his colleagues recommend a multipronged approach to combat seabird fallout. Individuals grounded by artificial lights can be rescued so they don’t succumb to the common causes of death – dehydration, predation and vehicle collision. In many cases the general public takes birds to designated rescue stations, where they are cared for until judged to be ready to release. The first rescue program was set up on Kauai in 1978; since then, people working for 16 rescue programs have released over 40,000 birds.

Release of a grounded shearwater. Photo Nazaret Carrasco (1)

Beneharo Rodríguez releases a Cory’s Shearwater from a cliff at Buenavista del Norte on Tenerife Island. Credit: Nazaret Carrasco.

The birds would be best served if humans behaved in ways that minimized fallout. Researchers need to learn more about why birds are attracted to artificial lights so engineers can develop outside lights that don’t attract them. Existing lights can be turned off when not needed, and dimmed when they are essential. Special accommodation can be made for unusual cases; for example in Cilaos, Reunion, Indian Ocean, streetlights are turned off during the fledging period of Barau’s Petrel. Lights can also be shielded so they illuminate an area for humans, but minimize the light visible to birds. Degraded nesting and breeding habitat can be restored to help compensate for birds that are lost to fallout. Lastly, conservation efforts should benefit the local economies so that residents will be more likely to support conservation initiatives, such as reduced evening lighting, that they might otherwise oppose.

note: the paper that describes this research is from the journal Conservation Biology. The reference is Rodríguez, A., Holmes, N. D., Ryan, P. G., Wilson, K.-J., Faulquier, L., Murillo, Y., Raine, A. F., Penniman, J. F., Neves, V., Rodríguez, B., Negro, J. J., Chiaradia, A., Dann, P., Anderson, T., Metzger, B., Shirai, M., Deppe, L., Wheeler, J., Hodum, P., Gouveia, C., Carmo, V., Carreira, G. P., Delgado-Alburqueque, L., Guerra-Correa, C., Couzi, F.-X., Travers, M. and Corre, M. L. (2017), Seabird mortality induced by land-based artificial lights. Conservation Biology, 31: 986–1001. Thanks to the Society for Conservation Biology for allowing me to use figures from the paper. Copyright © 2017 by the Society for Conservation Biology. All rights reserved.

Seagrass scourge: when nutrient enrichment reaches the tipping point

Sean Connell has watched as south Australia has lost vast expanses of kelp forest and seagrasses over the past years. One of the primary culprits associated with loss of seagrass meadows is excessive nutrients, particularly nitrogen, which enters the ecosystem with runoff, and causes an increase in algal epiphytes (epiphytes are small plants that grow on other plants). Epiphytes can negatively affect seagrass by blocking sunlight needed for photosynthesis, and indirectly, by increasing the rate of cellular respiration within the ecosystem, thus using up oxygen needed by seagrass for metabolic processes.


Two dolphins swim above a bed of seagrass off the south Australian coast.

Connell and his colleagues noticed that seagrass loss was often sudden; a large seagrass meadow would appear to be in good shape, and then it would abruptly disappear. They suggested that there might be a threshold effect in nutrient levels that seagrasses can tolerate; that these systems function well until a certain threshold in nutrient levels is crossed, above which there is an abrupt loss of seagrasses. They tested this hypothesis by subjecting plots of the seagrass Amphibolis antarctica to seven different concentrations of dissolved inorganic nitrogen (DIN) over a 10 month period, and monitored the abundance of epiphytes and seagrass over that timespan.

The meadows were about two km offshore from Lady Bay, Fleurieu Penninsula, Australia, in about 5 meters of water. Different amounts of nitrogen fertilizer were wrapped in nylon bags (for slow continuous release of DIN) and staked to the ocean floor. Amphibolis antarctica grows by producing new leaves at the top of each leaf cluster, but at the same time it drops old leaves. Leaf turnover, the researchers’ measure of growth, is simply new leaf production minus old leaf drop. The researchers tied on a small nylon cable at known locations on selected plants, noted how many leaves were above and below each tie at the beginning of the experiment, and recounted leaf number 10 months later. Finally, the researchers measured epiphyte growth by microscopically viewing a sample of seagrass leaves, and counting the number seagrass leaf cells that were covered by epiphytes.

Seagrass growth was relatively unaffected by all tested DIN levels.


Leaf production per day in relation to concentration of DIN.

However, leaf drop showed a strong threshold effect; leaf drop rates increased sharply between 0.13 – 0.15 mg/L of DIN.


Leaf drop per day in relation to concentration of DIN.

Putting these two graphs together, you can see (below) that leaf turnover switched from positive to negative at 0.13 – 0.15 mg/L of DIN. Negative leaf turnover translates to a sudden loss of seagrass at that threshold. At least in this system, at this location, 0.13 – 0.15 mg/L of DIN is the tipping point, beyond which the seagrass system suddenly goes into decline.


Leaf turnover per day (left y-axis and red data), and Epiphyte cover (% – right y-axis and green data), in relation to concentration of dissolved inorganic nitrogen.

The graph also shows that the tipping point coincides with an epiphyte cover of approximately 60%. It is possible that increased epiphyte cover may reduce seagrass photosynthetic rates (particularly in lower leaves), so that leaf turnover suddenly shifts into the negative zone, but the study was not designed to identify the underlying mechanism.

Seagrass meadows perform important ecosystem services, such as absorbing excess nutrients from the sediment, and providing habitat and food for a diverse group of grazers and indirectly, for their consumers. Thus seagrass conservation is vital. The danger here is that moderate levels of nutrients do not appear to have much effect on seagrass populations, but there is an abrupt shift to seagrass loss once the nutrient threshold is crossed. This makes the system very difficult to manage, because the loss occurs without warning. Australian ecologists have repeatedly failed to restore lost seagrass meadows, as simply reducing nutrient levels does not reverse the process. Thus anticipating seagrass loss before it happens is the most viable management solution for this critical ecosystem.

note: the paper that describes this research is from the journal Conservation Biology. The reference is Connell, S. D., Fernandes, M., Burnell, O. W., Doubleday, Z. A., Griffin, K. J., Irving, A. D., Leung, J. Y.S., Owen, S., Russell, B. D. and Falkenberg, L. J. (2017), Testing for thresholds of ecosystem collapse in seagrass meadows. Conservation Biology, 31: 1196–1201. doi:10.1111/cobi.12951. Thanks to the Society for Conservation Biology for allowing me to use figures from the paper. Copyright © 2017 by the Society for Conservation Biology. All rights reserved.

Subsidized Shorelines: where pelagic meets benthic

Chris Harrod met his wife, Christina Dorador, while both were working at the Max Planck Institute for Limnology in Germany.   Subsequently Dorador began a postdoctoral position in her native Chile, and Harrod went for a visit over Christmas, 2007. He was impressed by the beautiful rocky inshore habitat that was dominated by kelp forests and many species of invertebrates and fish.

Macroalgal forest

Rocky shoreline near Tocopilla, Chile. Credit: Chris Harrod.

As a fisheries biologist, Harrod soon immersed himself in the inshore kelp forest-dominated ocean, and was stunned by the sheer volume of stuff floating around. The water was green rather than blue, and filled with decaying phytoplankton and zooplankton. Where did all this stuff come from? Harrod knew that off the Peruvian and north Chilean coasts, prevailing winds move surface waters away from the shoreline, inducing upwelling of deeper nutrient-rich waters to the surface. This nutrient flux is the basis of a huge anchoveta fishery, which feeds humans, fish, marine invertebrates and marine mammals. He wondered whether the mass of floating debris in the inshore habitat might originate from offshore waters brought in from upwelling, and if the debris actually fueled some of the larger fish and molluscs that dominate inshore kelp forests. The prevailing opinion was that the energy for these fish and molluscs originated from the inshore photosynthetic kelp, rather than from photosynthetic phytoplankton further offshore that get their nutrients from upwelling.


Two bilagay, Cheilodactylus variegatus, swim among green algae in a debris-laden inshore habitat off the coast of Chile. Credit: Chris Harrod.

While you can ask a fish or mollusc what they had for dinner, it is very difficult to get them to respond. Fortunately, ecologists can use stable isotope ratios – the ratio of a rare (and nonradioactive) isotope of an element to its standard common isotope – to help get the answers they need. Harrod collaborated with several researchers in this study, including his Master’s student, Felipe Docmac, who collected and analyzed much of the data and was the first author of the paper. Docmac and his colleagues used the ratio of heavy and light isotopes of carbon (C) and nitrogen (N) to calculate d13C (the ratio of 13C to 12C) and d15N (the ratio of 15N to 14N) to infer where inshore fish were getting their energy.

The basic question was whether the nutrients supporting the food web were primarily from pelagic or benthic sources. In this case, the pelagic source refers to phytoplankton from the offshore areas of upwelling, while the benthic source refers to kelp and green algae that grow on the bottom (benthos) of the inshore habitat. Docmac and his colleagues collected invertebrates and large fish from five different sites along the north Chilean Coast, and calculated 13C and 15N values from tissue samples. Invertebrates were divided into two groups: filter-feeders were represented by mussels, which fed on suspended materials (such as the prolific floating debris), while benthic grazers were gastropods (snails) that fed on benthic kelp and green algae.


Five collection sites along the northern coast of Chile.

I’m going to skip the precise details of how stable isotope analysis actually works; I’ll provide enough information so you can understand the findings. There are two important facts to keep in mind. First an animal’s stable isotope ratio is influenced by the stable isotope ratio of its food source. So an animal feeding on prey with high d13C and d15N will itself have higher stable isotope ratios than will an animal feeding on prey with lower d13C and d15N. Second, the stable isotope ratio increases as we go up the food chain in a predictable manner, because the lighter isotopes of carbon and nitrogen tend to be more readily excreted than are the heavier isotopes.

We are now ready to look at the data. First, notice that while there is some variation from location to location, the (Pelagic) mussels tend to have consistently lower d13C values than do the (Benthic) gastropods (X-axis of graph), but fairly equivalent d15N values (Y-axis of graph). The benthivorous fish have, as we would expect from animals higher up the food chain, much higher d15N values than either of the invertebrates. But here is the key. The benthivorous fish have a much lower d13C value than do the benthic invertebrates (gastropods). If gastropods (and presumably other grazers) were in the benthivorous fish food chain, then we would expect the fish to have a higher d13C value than do the benthic gastropods. The researchers thus conclude that these fish are deriving most of their energy from the pelagic debris that is washing in from ocean currents.


d13C (X-axis) and d15N (Y-axis) stable isotope values in benthivorous fish (black circles).  Bars emanating from each point indicate 95% confidence intervals (CI). Numbers inside symbols indicate the site of origin for each sample (see map above), Also shown are values for filter-feeding mussels (red up-pointing triangles) and grazing gastropods (blue down-pointing triangles). Gray lines indicated predicted values of diets that were based solely (100%) on pelagic or benthic sources.

The researchers were stunned by these findings. Going into the study, Harrod did not know what he would find, but would not have been surprised by a 15% contribution from pelagic sources, or maybe even 30%. But he was blown away that the data indicated estimates of greater than 90% pelagic contribution at most of the sites. Ecologists have long known that one ecosystem may subsidize a second ecosystem with resources. For example, salmon carcasses can provide nutrient subsidies to trees near riverways, or even deeper into the forest after being transported by bears. But the extent of the subsidy in this study is unprecedented. Docmac and his colleagues urge researchers to explore exactly how the pelagic materials get into the food web, and to see whether such subsidies are common near other upwelling zones worldwide so that coastal resources can be managed more effectively.

note: the paper that describes this research is from the journal Ecology. The reference is Docmac, Felipe, Miguel Araya, Ivan A. Hinojosa, Cristina Dorador, and Chris Harrod. 2017. Habitat coupling writ large: pelagic‐derived materials fuel benthivorous macroalgal reef fishes in an upwelling zone. Ecology doi:10.1002/ecy.1936. It was published online on Aug. 2, 2017, and should appear in print very soon. Thanks to the Ecological Society of America for allowing me to use figures from the paper. Copyright © 2017 by the Ecological Society of America. All rights reserved.

Meta-analysis measures multiple mycorrhizal benefits to plants

Plants and fungi sometimes live together in peace and harmony. Arbuscular mycorrhizal associations are associations between plant roots and fungi, in which the fungal hyphae (usually branched tubular structures) grow between root cells, penetrating some cells with a network of branches or arbuscules.  Oftentimes these are mutualistic associations with both the plants and the fungi benefiting from living together. Though plants with arbuscular mycorrhizal fungi (AMF) tend to grow better than plants without AMF, it not always clear what causes them to do so.


Kura clover, Trifolium ambiguum, grown with AMF (left) and without AMF (right). Credit: Liz Koziol.

Ecologists have traditionally viewed arbuscular mycorrhizal associations as a straightforward nutrient-carbon exchange. Fungal hyphae, with their vast surface area, pick up nutrients (such as nitrogen and phosphorus compounds) from the soil, which they deliver to the root cells in exchange for plant-produced carbon molecules.

But recently researchers have identified numerous other potential ways that the fungi help the plants, including the following: (1) promoting water uptake and transport, (2) helping to spread allelochemicals – toxic chemicals that some plants release to rid themselves of nearby competitors, (3) inducing chemical defenses against herbivores, (4) enhancing disease resistance, and (5) promoting soil aggregation or clumping, which stabilizes the soil near the roots, reduces erosion and promotes stable water flow.

Ecology Fig1

Camille Delavaux and her colleagues wondered whether these other plant benefits might actually be more important than we originally thought. Delavaux was planning to write a review paper for a 1 credit independent study, but she found so many papers on this topic that she decided to collaborate with fellow students Lauren Smith-Ramesh and Sara Kuebbing on a full-scale meta-analysis.

A meta-analysis is a systematic analysis of data collected by many other researchers. Delavaux and her colleagues used the Web of Science database to find 4410 studies on how AMF supplied plants with nutrients and 1239 studies on how AMF provided other plant benefits. That’s a lot of studies! But for the meta-analysis, the authors only used a small fraction of these studies because they set certain restrictions. For example, to be used in the meta-analysis the authors required each study to show some measure of variation for the data (such as standard deviation or standard error). In addition, the authors required each study to compare plants grown under two conditions: with AMF and without AMF.  In many studies the researchers collected soil, which they sterilized in a hot oven, and then set up a test group, which they inoculated with AMF spores or a plug of soil or root fragments that contained AMF. In addition, these studies also had a control group of plants that received only sterilized soil with no AMF added.


A collection of eight different species of AMF spores. Credit: Liz Koziol.

Delavaux and her colleagues compared how plants performed with and without an AMF. Because each study was different, one might only have been looking at the effects of AMF on nitrogen uptake performance, while a second study might consider how AMF influenced soil aggregation. Effect size (Hedges d+) compares mean performance of the AMF plant to mean performance of non-AMF plants for a particular variable (such as nitrogen uptake or soil aggregation). A positive effect size means that the AMF plant did better. Of course we need to know how much better is biologically meaningful, so for each variable the researchers calculated the 95% confidence intervals of the mean effect size. If the 95% confidence intervals were positive, then Delavaux and her colleagues could be 95% confident that there was a biologically important effect of AMF on plants for that particular measure of performance.

As expected, the researchers found a positive effect of AMF on plant nitrogen uptake. The mean effect size was 0.674 with a 95% confidence interval of 0.451- 0.912. We can interpret this to mean that we are 95% confident that the true mean effect size on nitrogen uptake is between 0.451 and 0.912. But the greatest effect of AMF on plants was on soil aggregation (mean effect size = 1.645, 95% confidence interval = 1.032 – 2.248). AMF also had significant positive effects on phosphorus uptake, water flow and disease resistance.


Mean effect size (Hedges’ d+) of AMF on different factors considered in the meta-analysis.  The horizontal error bars are the 95% confidence intervals. n = number of observations.  If the error bars do not cross zero, inoculation with AMF had a significant positive effect relative to plants without AMF.

This meta-analysis shows that AMF help plants in many different ways. Researchers knew about the AMF impact on nitrogen and phosphorus uptake, but may be surprised to learn of equally strong effects on water flow, disease resistance and soil aggregation. Consequently, AMF may be very useful for forest management, agriculture, conservation and habitat restoration. As examples, conservation biologists and forest managers may need to consider adding AMF to soils that have suffered severe burns from fires, which may kill the existing soil fungi. Or agriculturalists intent on growing a particular crop may want to inoculate the soil with a specific group of AMF spores that enhance soil aggregation and water uptake, so their crop may thrive in a habitat that might otherwise not be suitable.

More than 3/4 of land plants form associations with AMF. Consequently, any attempts to restore habitats or to maintain high levels of species diversity in existing ecosystems require understanding what types of AMF inhabit the soils, and how these AMF influence ecosystem functioning.

note: the paper that describes this research is from the journal Ecology. The reference is Delavaux, C. S., Smith-Ramesh, L. M. and Kuebbing, S. E. (2017), Beyond nutrients: a meta-analysis of the diverse effects of arbuscular mycorrhizal fungi on plants and soils. Ecology, 98: 2111–2119. doi:10.1002/ecy.1892. Thanks to the Ecological Society of America for allowing me to use figures from the paper. Copyright © 2017 by the Ecological Society of America. All rights reserved.

Limpet larvae and their fantastic voyage

As he began his PhD program, Takuya Yahagi was puzzled by some laboratory findings. Juvenile red blood limpets, Shinkailepas myojinensis, seemed to survive and grow extraordinarily well at temperatures between 15-25° C. Adult limpets live in deep sea vent communities, where temperatures generally range between 6-11° C.

limpet photo

Adult Shinkailepas myojinensis.  These are approximately 6 mm in length. Credit: Takuya Yahagi.

Yahagi and his colleagues wondered why limpets are making babies that survive and grow at much higher temperatures than they are likely to experience after hatching.

Screen Shot 2017-07-04 at 10.48.26 AM

Deep sea hydrothermal vent community at 795 meters depth at Myojinsho Caldera in the northwest Pacific. White patches on the rocks are vast communities of chemosynthetic bacteria which are being grazed by purple/pinkish limpets. You can also see the white feathery feeding legs of a barnacle population in the upper portion of the photo. Credit: JAMSTEC

Yahagi reasoned that perhaps, in the natural world, the limpet juveniles live in different (warmer) environments than do their parents. If they migrated closer to the sea surface, their world would be somewhat warmer. But limpet babies are microscopic, so capturing them near the sea surface (and knowing that you had captured them!) is very challenging. Working with three other researchers, Yahagi decided to collect indirect evidence to test the hypothesis that baby limpets migrate to the surface where they feed and grow before returning to the ocean depths.


Larval S. myojinensis limpet 156 days after hatching. sh=shell, f =foot, e=eye, vl=velar lobe.

Initially, the researchers needed to determine what temperatures these growing limpets preferred. With the help of a remotely operated submarine, they collected adult limpets laden with egg capsules, and placed newly hatched larvae into separate containers under different conditions. Some larvae were fed and raised at one of six different temperatures: 5, 10, 15, 20, 25 and 30° C. Other larvae were starved at 5, 15 or 25° C to see how long they survived at different temperatures. If the larvae were migrating upwards to warmer waters, it was important to see how long they could survive until they arrived at the richer food sources near the surface.

Starved larvae survived up to 150 days at the lowest temperature, and for more than three weeks at 25° C, which provided ample time for upward migration (even at very mellow baby limpet swimming speeds). Fed larvae grew much more quickly at warmer temperatures, with best growth at 25° C, and no growth at 5-10° C, which is the approximate temperature at hydrothermal vents.. Larvae initially grew quickly at 30° C, but long term exposure to that temperature killed them.


Growth (shell length) of fed larvae at different temperatures.

These temperature profiles corresponded to temperatures at the sea surface down to about 100 meters, which ranged between 19-28° C. This correspondence supported the hypothesis that juveniles migrated upwards in the water column after hatching. But could Yahagi and his colleagues find any direct evidence for this vertical migration? To answer this question, they video-recorded new hatchlings in a clear plastic bath, and measured how fast these limpets swam, and what direction they preferred. They discovered that new hatchlings constantly swam upward in their test bath, and swimming speed was considerably faster at warmer temperatures.

The sea surface is a wonderful place to find food, because sunlight is abundant, so there are abundant phytoplankton to satisfy even the most voracious juvenile limpets. But sea surfaces also have very strong currents which can whisk juvenile limpets hundreds or thousands of kilometers away. The upshot is that vertical migration and wide dispersal of juveniles by ocean currents can introduce new genes into far-away limpet populations.

Screen Shot 2017-07-04 at 10.48.05 AM

A hot vent animal community at 700 meters depth at Minami-Ensei Knoll in the northwest Pacific. Prevalent groups include lobsters (white), two species of shrimp, mussels and two different limpet species. Credit: JAMSTEC.

Gene flow – the movement of genes from one population to another – has some important genetic impacts. Without gene flow, two populations that are separated from each other can become genetically distinct. But the mixing of genes from long-distance dispersal can prevent this from occurring. The researchers compared 1218 base pairs of the COI gene from 77 adult limpets that were collected from four different sites which were separated, in some cases, by more than 1000 kilometers. In support of the gene flow hypothesis they found no evidence of any genetic differentiation among the four populations.

Yahagi Fig1

Hydrothermal vent fields in the northwest Pacific Ocean.  Black squares are limpet collection sites for this study.  Notice the vast distances separating these populations.

Gene flow requires long distance dispersal, and the adult limpets travel very little along the sea floor. This finding of no genetic differentiation among the geographically separated populations supports the hypothesis that the juveniles migrate upwards, feed on abundant phytoplankton, and are carried to new distant environments. There, they mature and settle into new ocean vent communities where they can feed on the superabundant chemosynthetic bacteria associated with the ocean vents. But we still don’t know how limpets find a new ocean vent community – do they migrate, checking out possible vent habitats, while they are still juveniles and still capable of swimming? Do they have sense organs that pick up environmental cues such as hydrogen sulfide content, water temperature, turbulence or noise from vent emissions, to help them complete their fantastic ocean voyage?

note: the paper that describes this research is from the journal Ecology. The reference is Yahagi, Takuya, Hiromi Kayama Watanabe, Shigeaki Kojima, and Yasunori Kano. 2017. Do larvae from deep‐sea hydrothermal vents disperse in surface waters? Ecology 98: 1524-1534Thanks to the Ecological Society of America for allowing me to use figures from the paper. Copyright © 2017 by the Ecological Society of America. All rights reserved.

Light levels limit lake phytoplankton response to fertilization

One might naively think that because we humans are land-dwelling creatures, our impact on aquatic ecosystems might be relatively minor. Unfortunately, this assumption is incorrect, as human activities are changing aquatic environments in profound ways that influence how aquatic species survive and interact. Global warming is increasing lake and river temperatures, uncontrolled development is causing some streams to run dry and others to flood, and agricultural practices are adding nutrients to many lakes and streams. Because these human impacts occur simultaneously, it is difficult to evaluate how each factor contributes to the observed changes in species relations.

In northern Sweden, lakes vary naturally in the amount of dissolved organic carbon (DOC) they contain. DOC comes from runoff of decaying plant matter, so lakes surrounded by substantial vegetation, or that experience a great deal of water input (runoff) from the surrounding area, would have higher DOC than other lakes. DOC is potentially very important to lakes, because DOC tends to discolor a lake, which reduces light penetration and slows down photosynthesis. On the positive side, carbon may bond to other molecules such as phosphorus and nitrogen, which are important nutrients that may be in short supply in these relatively infertile lakes.   Anne Deininger and her colleagues focused their studies on two factors: DOC and nitrogen. Most lakes have too much nitrogen, a result of excessive use of nitrogen fertilizers that run off into lakes, so these relatively low-nitrogen lakes provided the researchers with a unique opportunity to see how these two factors, DOC and nitrogen, interacted in a natural ecosystem.

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Low DOC control lake. Credit: M. Klaus

The researchers selected six lakes that varied naturally in DOC levels: two low (~7 mg DOC/liter), two medium (~11 mg DOC/liter), and two high (~20 mg DOC/liter). In 2011 they measured everything possible about each lake: abundance of all of the life forms, DOC, temperature, light levels, nutrients and photosynthetic rates. In 2012 and 2013, they supplemented one of each pair of lakes with nitrogen compounds every one to two weeks. The added nitrogen was equivalent to the higher nitrogen inputs that are experienced by lakes in southern Sweden. And, as you might expect, the researchers continued measuring all factors of interest in both the experimental (fertilized) and control (unfertilized) lakes throughout the year – at least until the lakes froze over.


Anne Deininger (in orange) and Sonja Prideaux collect samples from a lake. Credit: M. Deininger.

Deininger and her colleagues were most interested in differences in the abundance of phytoplankton – small free-floating photosynthetic organisms, because these are the primary producers – the organisms that produce the chemical energy (via photosynthesis) that enters food webs. There are many different types or groups of these phytoplankton; some are flagellated, with hair-like processes that allow them to navigate in the water column. Some are exclusively autotrophs, producing their own energy from photosynthesis, some are primarily hetrotrophic, eating other organisms or the remains of dead organisms, while others are mixotrophs, using both strategies to produce energy. Cyanobacteria are photosynthetic bacteria, while picophytoplankton are phytoplankton of unusually small size.

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Flagellated phytoplankton (Cryptomonas). Illustration by Anne Deininger.

Many important findings are summarized in the graph below. “B” represents the year before fertilization (2011), while “A1” is 2012 (after fertilization – 1st year) and “A2” is 2013 (after fertilization – 2nd year). Remember only the N-lakes were fertilized; the control lakes were simply monitored all three years. One finding is that in 2011, the high DOC lakes had the lowest phytoplankton abundance.  A second is that the low and medium DOC lakes had both flagellated and non-flagellated phytoplankton, while the high DOC lakes were dominated by flagellated phytoplankton.

Moving to the years after fertilization (A1 and A2), you can see that nitrogen fertilization increased phytoplankton abundance, but more so for the low-DOC lake. However, fertilization had little impact on the types of phytoplankton found in each lake; rather it simply increased the abundance of already existing groups.


Mean biomass of major phytoplankton groups in relation to DOC.  Recall that B refers to 2011 (the year before fertilization), while A1 and A2 refer to the two years after fertilization (2012, 2013).

The data can be organized so we can get a better view of what is happening quantitatively. Fertilization increases phytoplankton biomass, but much more for lakes with low DOC levels. In addition DOC appears to decrease phytoplankton abundance.


Deininger and her colleagues conclude that in these northern lakes, phytoplankton production is nutrient-limited at low DOC levels, but becomes limited by light availability in more murky waters. So adding nitrogen increases phytoplankton abundance to a greater extent in low DOC lakes. High DOC lakes have more flagellated autotrophs, as these species can swim to the top of the water column where there is more light for photosynthesis. As needed, flagellated phytoplankton can move lower in the water column where nutrients are more abundant.

The researchers emphasize that the nitrogen experiments were only conducted for two years. They don’t know if, for example, the types of species would change if fertilization continued for more than two years. They also don’t know if after 2013, the communities reverted to their pre-fertilization state, or if biomasses remained higher when nitrogen fertilization stopped. These types of questions are important to pursue because we humans are making drastic changes to most of our aquatic systems in a very uncontrolled manner. We need to understand the effects of these changes to the aquatic environment, and also how we can reverse the effects should they prove to be highly detrimental.

note: the paper that describes this research is from the journal Ecology. The reference is Deininger, A., Faithfull, C. L., & Bergström, A. K. (2017). Phytoplankton response to whole lake inorganic N fertilization along a gradient in dissolved organic carbon. Ecology98(4), 982-994. Thanks to the Ecological Society of America for allowing me to use figures from the paper. Copyright © 2017 by the Ecological Society of America. All rights reserved.

Fires foster biological diversity on the African savanna

As an ecology student back in days of yore, I was introduced to the classic mutualism between ants and swollen-thorn acacia trees. In this mutually beneficial relationship, ants protect acacia trees by biting and projecting very smelly substances at hungry herbivores, and by pruning encroaching branches of plant competitors. In return for these services, acacia trees provide the ants with homes in the form of swollen thorns, and in some cases also provide food for their defenders.


Swollen thorns of Acacia drepanlobium occupied by C. nigriceps. Credit: Ryan L. Sensenig.

I always assumed there were limits to what these ants could do. I knew that elephants were a constant problem for trees trying to get established on the African savanna. I figured, wrongly, that ants could not do much to counter a determined thick-skinned elephant. But as Ryan Sensenig describes, ants will swarm any intruding elephant, rushing into the elephant’s very sensitive trunk and mouth, biting it and, in some cases, exuding a chemical compound that is very offensive to an elephant’s keen sense of smell. So don’t mess with these ants if you can help it!


The Laikipia Plateau has one of the few growing elephant populations in East Africa. Credit: Ryan L. Sensenig.

Fires play an important role in savanna ecosystems, killing many trees before they can get established, and creating a mosaic of burned and unburned areas which vary in grass quality and quantity, and in the abundance of acacia trees (and other species as well). Recently burned grasslands tend to be lower in grass abundance and higher in grass nutrient levels. In a previous study of controlled burns, Sensenig and his colleagues showed that larger animals, such as elephants, tended to graze in unburned areas, which had more grass – albeit of lower quality. Returning seven years after the burn, he was surprised to find that elephants, which eat both trees and grass, had shifted to the burned sites in preference to unburned sites. He thus wondered whether fire was having an impact on the ant-acacia mutualisms that defend acacias from elephants and other large herbivores.


Sunset strikes an Acacia xanthophloea on Mpala Research Centre in Laikipia, Kenya. Credit: Ryan L. Sensenig.

Ants do not share trees. In Mpala Research Centre in the Laikipia Plateau of Kenya, there are four mutually-exclusive species of ants that live in Acacia drepanolobium trees: Crematogaster sjostedti, C. mimosae, C. nigriceps, and Tetraponera penzigi.

Sensenig and his colleagues wanted to know whether the controlled burns had a long-lasting effect on ant species distribution on acacia trees. The researchers surveyed 12 plots that had been burned seven years previously and an equal number of unburned plots to see how burns affected which ant species were present.


Goshen College research students estimate ant densities on Acacia drepanolobium trees in the Kenya Longterm Exclosure Experiment. Credit: Ryan L. Sensenig.

They found that C. nigriceps was more common in acacias from burned areas while the other three species were more common in trees from unburned areas.


Why were there more C. nigriceps ants in previously burned areas? One explanation is that perhaps C. nigriceps is better at avoiding getting burned by fire, or else is better at recolonizing after a fire. To look for species difference in response to fire, the researchers simulated fires by burning elephant dung and dried grass in 3-gallon metal buckets, creating a small sustained smoke source. They stationed observers every 50 meters along a 500 meter transect for the first experiment, and a 1.8 km transect for the second experiment. They then measured ant-evacuation rate by counting the number of ants moving down the trunk. There were some very pronounced differences, with C. nigriceps having the highest evacuation rate, C. mimosae also showing a strong smoke response, and the other two species showing little evidence of any response.


Evacuation rate for each species in response to smoke.

C. mimosae generally prevails when it battles a colony of C. nigriceps. These results indicate that the subordinate C. nigriceps is able to maintain its presence in the community, in part, by taking advantage of its superior performance when it encounters a fire. The researchers also found some evidence that C. nigriceps is better at recolonizing after a fire than is C. mimosae. So despite being the subordinate species, C. nigriceps is abundant in this ecosystem.

Returning to those elephants, the researchers describe one final experiment in which some plots had a series of fences that excluded herbivores, while other plots were open to herbivores, including elephants.


In this experiment, as well, there were burned and unburned plots. In general, there were more ants present when herbivores were present, as the trees invested more in swollen thorns and in ant food (in the form of nectar) to attract protective ants. In addition, ants were more abundant in unburned plots than in plots that had been previously burned, with the exception of C. nigriceps when herbivores were excluded.

Ecologists have long known that fire maintains savanna ecosystems by preventing the grasslands from being overgrown by trees. This study shows that fires shift ant community structure in favor of the subordinate ant species (C. nigriceps), resulting in a higher diversity of ant species overall. The researchers suggest that if fires become more common in savannas, elephants may become more attracted to acacias that harbor a reduced (or nonexistent) cast of defenders, which could lead to a further reduction in the abundance of acacia trees and their mutualistic ants.

note: the paper that describes this research is from the journal Ecology. The reference is Sensenig, R. L., Kimuyu, D. K., Ruiz Guajardo, J. C., Veblen, K. E., Riginos, C., & Young, T. P. (2017). Fire disturbance disrupts an acacia ant–plant mutualism in favor of a subordinate ant species. Ecology, 98(5), 1455-1464.Thanks to the Ecological Society of America for allowing me to use figures from the paper. Copyright © 2017 by the Ecological Society of America. All rights reserved.