Offspring sizes vary within populations but the reasons are unclear. Game‐theoretic models predict that selection will maintain offspring‐size variation when large offspring are superior competitors (i.e., competition is asymmetric), but small offspring are superior colonizers. Empirical tests are equivocal, however, and typically rely on interspecific comparisons, whereas explicit intraspecific tests are rare.
In a field study, we test whether offspring size affects competitive asymmetries using the sessile marine invertebrate, Bugula neritina. Surprisingly, we show that offspring size determines whether interactions are competitive or facilitative — large neighbours strongly facilitated small offspring, but also strongly competed with large offspring. These findings contradict the assumptions of classic theory — that is, large offspring were not superior competitors. Instead, smaller offspring actually benefit from interactions with large offspring— suggesting that asymmetric facilitation, rather than asymmetric competition, operates in our system.
We argue that facilitation of small offspring may be more widespread than currently appreciated, and may maintain variation in offspring size via negative frequency‐dependent selection.
Offspring size theory has classically viewed offspring interactions through the lens of competition alone, yet our results and those of others suggest that theory should accommodate positive interactions in explorations of offspring‐size variation.
Cameron H, Marshall DJ (2019) Can competitive asymmetries maintain offspring size variation? A manipulative field test. Evolution PDFDOI
It’s party time in the Centre for Geometric Biology: We’ve reached 500 generations of experimental evolution in our large and small algal cells!
Three and a half years ago Martino Malerba set up the first culture of the single-celled marine phytoplankton Dunaliella tertiolecta and begun to artificially select for large and small cells. The selection process of separating the largest and the smallest cells has continued twice a week ever since. Martino is now the proud ‘father’ of algal cells where the big cells are more than 10 times the size of the smaller cells.
The evolved algae lines have enabled Martino and his colleagues to look at the consequences of being a particular size without having to compare different species, which vary in many other ways than just size. This research directly supports the fundamental question the Centre for Geometric Biology seeks to understand “why do organisms grow to the size they do?”
The team has looked at the effects of size at both the level of the cell and the population. They have found that altering the size of a cell profoundly alters many fundamental traits of algal physiology and ecology. This affects how cells of different sizes will cope with fluctuating resources and, in turn, how those populations of cells use energy and grow.
Exciting research underway looks at community-level effects. What happens when grazers are only offered either large or small algal cells: does changing algal size have repercussions up the food chain?
Understanding how individuals grow and regulate their energy is critical in helping scientists predict how large-scale impacts, such as climate change, affect organisms.
Phytoplankton have critical roles in the ocean; they form the base of most food webs and fix large amounts of carbon. So, while the research stemming from these artificially evolved algal cells is theoretically interesting, it also has a very direct and immediate application to how we understand and manage climate change.
About half of the coastline of Europe, the United States and Australasia is modified by artificial structures. In newly published research, we identified a new effect of marine urbanisation that has so far gone unrecognised.
When we build marinas, ports, jetties and coastal defences, we introduce hard structures that weren’t there before and which reduce the amount of sunlight hitting the water. This means energy producers such as seaweed and algae, which use light energy to transform carbon dioxide into sugars, are replaced by energy consumers such as filter-feeding invertebrates. These latter species are often not native to the area, and can profoundly alter marine habitats by displacing local species, reducing biodiversity, and decreasing the overall productivity of ecosystems.
Incorporating simple designs in our marine infrastructure to allow more light penetration, improve water flow, and maintain water quality, will go a long way towards curbing these negative consequences.
We are used to thinking about the effects of urbanisation in our cities – but it is time to pay more attention to urban sprawl in the sea. We need to better understand the effects on the food web in a local context.
Most animals that establish themselves on these shaded hard structures are “sessile” invertebrates, which can’t move around. They come in a variety of forms, from encrusting species such as barnacles, to tree-shaped or vase-like forms such as bryozoans or sponges. But what they all have in common is that they can filter out algae from the water.
In Australian waters, we commonly see animals from a range of different groups including sea squirts, sponges, bryozoans, mussels and worms. They can grow in dense communities and often reproduce and grow quickly in new environments.
How much energy do they use?
In our new research, published in the journal Frontiers in Ecology and the Environment, we analysed the total energy usage of invertebrate communities on artificial structures in two Australian bays: Moreton Bay, Queensland, and Port Phillip Bay, Victoria. We did so by combining data from field surveys, laboratory studies, and satellite data.
We also compiled data from other studies and assessed how much algae is required to support the energy demands of the filter-feeding species in commercial ports worldwide.
In Port Phillip Bay, 0.003% of the total area is taken up by artificial structures. While this doesn’t sound like much, it is equivalent to almost 50 soccer fields of human-built structures.
We found that the invertebrate community living on a single square metre of artificial structure consumes the algal biomass produced by 16 square metres of ocean. Hence, the total invertebrate community living on these structures in the bay consumes the algal biomass produced by 800 football pitches of ocean!
Similarly, Moreton Bay has 0.005% of its total area occupied by artificial structures, but each square metre of artificial structure requires around 5 square metres of algal production – a total of 115 football pitches. Our models account for various biological and physical variables such as temperature, light, and species composition, all of which contribute to generate differences among regions.
Overall, the invertebrates growing on artificial structures in these two Australian bays weigh as much as 3,200 three-tonne African elephants. This biomass would not exist were it not for marine urbanisation.
How does Australia compare to the rest of the world?
We found stark differences among ports in different parts of the world. For example, one square metre of artificial structure in cold, highly productive regions (such as St Petersburg, Russia) can require as little as 0.9 square metres of sea surface area to provide enough algal food to sustain the invertebrate populations. Cold regions can require less area because they are often richer in nutrients and better mixed than warmer waters.
In contrast, a square metre of structure in the nutrient-poor tropical waters of Hawaii can deplete all the algae produced in the surrounding 120 square metres.
Does it matter?
Should we be worried about all of this? To some extent, it depends on context.
These dense filter-feeding communities are removing algae that normally enters food webs and supports coastal fisheries. As human populations in coastal areas continue to increase, so will demand on these fisheries, which are already under pressure from climate change. These effects will be greatest in warmer, nutrient-poor waters.
But there is a flip side. Ports and urban coastlines are often polluted with increased nutrient inputs, such as sewage effluents or agricultural fertilisers. The dense populations of filter-feeders on the structures near these areas may help prevent this nutrient runoff from triggering problematic algal blooms, which can cause fish kills and impact human health. But we still need to know what types of algae these filter-feeding communities are predominantly consuming.
Our analysis provides an important first step in understanding how these communities might affect coastal production and food webs.
In places like Southeast Asia, marine managers should consider how artificial structures might affect essential coastal fisheries. Meanwhile, in places like Port Phillip Bay, we need to know whether and how these communities might affect the chances of harmful algal blooms.
One hectare of ocean in which fishing is not allowed (a marine protected area) produces at least five times the amount of fish as an equivalent unprotected hectare, according to new research published today.
This outsized effect means marine protected areas, or MPAs, are more valuable than we previously thought for conservation and increasing fishing catches in nearby areas.
Previous research has found the number of offspring from a fish increases exponentially as they grow larger, a disparity that had not been taken into account in earlier modelling of fish populations. By revising this basic assumption, the true value of MPAs is clearer.
Marine Protected Areas
Marine protected areas are ocean areas where human activity is restricted and at their best are “no take” zones, where removing animals and plants is banned. Fish populations within these areas can grow with limited human interference and potentially “spill-over” to replenish fished populations outside.
Obviously MPAs are designed to protect ecological communities, but scientists have long hoped they can play another role: contributing to the replenishment and maintenance of species that are targeted by fisheries.
Yet fishers remain sceptical that any spillover will offset the loss of fishing grounds, and the role of MPAs in fisheries remains contentious. A key issue is the number of offspring that fish inside MPAs produce. If their fecundity is similar to that of fish outside the MPA, then obviously there will be no benefit and only costs to fishers.
Big fish have far more babies
Traditional models assume that fish reproductive output is proportional to mass, that is, doubling the mass of a fish doubles its reproductive output. Thus, the size of fish within a population is assumed to be less important than the total biomass when calculating population growth.
But a paper recently published in Science demonstrated this assumption is incorrect for 95% of fish species: larger fish actually have disproportionately higher reproductive outputs. That means doubling a fish’s mass more than doubles its reproductive output.
When we feed this newly revised assumption into models of fish reproduction, predictions about the value of MPAs change dramatically.
Fish are, on average, 25% longer inside protected areas than outside. This doesn’t sound like much, but it translates into a big difference in reproductive output – an MPA fish produces almost 3 times more offspring on average. This, coupled with higher fish populations because of the no-take rule means MPAs produce between 5 and 200 times (depending on the species) more offspring per unit area than unprotected areas.
Put another way, one hectare of MPA is worth at least 5 hectares of unprotected area in terms of the number of offspring produced.
We have to remember though, just because MPAs produce disproportionately more offspring it doesn’t necessarily mean they enhance fisheries yields.
For protected areas to increase catch sizes, offspring need to move to fished areas. To calculate fisheries yields, we need to model – among other things – larval dispersal between protected and unprotected areas. This information is only available for a few species.
We explored the consequences of disproportionate reproduction for fisheries yields with and without MPAs for one iconic fish, the coral trout on the Great Barrier Reef. This is one of the few species for which we had data for most of the key parameters, including decent estimates of larval dispersal and how connected different populations are.
We found MPAs do in fact enhance yields to fisheries when disproportionate reproduction is included in relatively realistic models of fish populations. For the coral trout, we saw a roughly 12% increase in tonnes of caught fish.
There are two lessons here. First, a fivefold increase in the production of eggs inside MPAs results in only modest increases in yield. This is because limited dispersal and higher death rates in the protected areas dampen the benefits.
However the exciting second lesson is these results suggest MPAs are not in conflict with the interests of fishers, as is often argued.
While MPAs restrict access to an entire population of fish, fishers still benefit from from their disproportionate affect on fish numbers. MPAs are a rare win-win strategy.
It’s unclear whether our results will hold for all species. What’s more, these effects rely on strict no-take rules being well-enforced, otherwise the essential differences in the sizes of fish will never be established.
We think that the value of MPAs as a fisheries management tool has been systematically underestimated. Including disproportionate reproduction in our assessments of MPAs should correct this view and partly resolve the debate about their value. Well-designed networks of MPAs could increase much-needed yields from wild-caught fish.
Marine urbanization is a term that describes the increasing proliferation of structures such as piers, jetties, marinas, sea walls and other coastal defences in the marine environment. Martino Malerba, Craig White and Dustin Marshall from the Centre for Geometric Biology are proposing that this type of urban spread into the ocean could rob local systems of some of their productivity, changing local food web structure as well as function.
Many artificial structures, including floating platforms, barges, piers, pontoons seawalls and port quays, decrease the access of direct sunlight into the water. This means that instead of ecological communities dominated by ‘energy-producing’ seaweeds that require light to photosynthesise, there tends to be a shift toward dense assemblages of ‘energy-consuming’ filter-feeding invertebrates. The research team are particularly interested in finding out how this additional filter-feeding biomass affects energy flow and the productivity of coastal food webs.
Martino and his colleagues have combined data from field surveys, laboratory studies and satellite data to analyse total energy usage of invertebrate communities on artificial structures in Port Phillip and Moreton Bays. The team then used estimates from other studies to estimate the amount of primary production required to support the metabolic demands of the entire filter-feeding biomass living on artificial structures of all main commercial ports worldwide: the ‘trophic footprint’.
In order to do this, Martino and colleagues first used satellite photos to estimate how much area available for colonisation is created by artificial structures in Port Phillip and Moreton Bays. They then used field surveys to estimate the total invertebrate biomass occurring on all artificial structures in the two bays. In Port Phillip Bay, the estimated biomass on artificial structures is the equivalent of 3,151 female African bush elephants.
The next step was to transport communities back to the laboratory to measure mean daily energy consumption per unit area and then scale this up to artificial structures within the whole bay. Based on their estimates they found that biomass on artificial structures can consume between 0.005% and 0.05% of the total yearly energy production in Port Phillip Bay and Morton Bay respectively. This means that each square metre of artificial structure will consume 6 to 20 m² of marine primary productivity in these bays.
When they went through essentially the same process using data available from the scientific literature, the team found, on average, each metre of port consumes 26 m²of ocean primary productivity. But there are stark differences between ports in different parts of the world. For example, one square metre of artificial structure in cold, highly productive regions can require as little as 0.9 m² of ocean (e.g. St. Petersburg, Russia), whereas a square metre of artificial structure in the nutrient-poor tropical waters of Hawaii can deplete all of the productivity in the surrounding ~120 m².
Martino, Craig and Dustin point out that a large percentage of ocean shoreline is now modified by engineering, with associated impacts on primary production. Burgeoning coastal human populations are expected to increase demands on fisheries in the future and food web productivity is already forecasted to decrease, due to the effects of climate change. Understanding the impacts of marine urbanisation on food webs is becoming increasingly important so that design of artificial structures minimises impacts on food webs and remediation efforts can be undertaken if necessary.
Our recent research has found that we are systematically underestimating the true value of marine protected areas (MPAs) to fisheries.
An important function of MPAs is to protect both representative and unique ecological communities, but scientists have long hoped they can play another role: contributing to the replenishment and maintenance of exploited fish stocks.
Wild fisheries are under intense pressure and landings of fish catches have flattened out despite an ever-increasing fishing effort. The most effective kind of MPAs are areas we set aside as ‘no take’ zones, where removal of animals and plants is banned. Fish populations within these areas can grow with limited human interference and potentially ‘spill-over’ to replenish fished stocks outside of MPAs. Despite the potential benefit, anglers remain sceptical that any spill-over will offset the loss of fishing grounds and the role of MPAs in fisheries remains contentious.
When we calculate how much a protected area contributes to a fishery, we work out the average length of both fished and unfished populations. Fish inside MPAs are bigger, on average, than those outside and so will produce more offspring than their smaller relatives outside MPAs. Generally, fisheries scientists relate the size of the fish to the reproductive output, whereby one unit increase in size equates to one unit increase in egg production. They estimate how many juveniles will ‘spill-over’ and enter the fishing grounds.
It turns out there are a number of problems with the way the spill-over effect is currently calculated. Length is generally the measure of fish size recorded in a survey, but focusing on length risks underestimating the differences in reproduction inside and outside of MPAs. Length and mass do not change at the same rate, so a 28% increase in fish length results in a 109% increase in mass. It can seem counter-intuitive that increasing fish length by around one quarter more than doubles the mass because the human brain tends to struggle when thinking about non-linear relationships.
This is compounded when we consider another non-linear relationship; fish mass and reproductive output. Our research team from the Centre for Geometric Biology and collaborator Ross Robertson from the Smithsonian Tropical Research Institute, recently found that bigger fish produce disproportionately more eggs than smaller fish in all fish species they looked at. This research made us realise that we need to be focused on protecting the biggest fish in a fishery.
And that is not all. In 1906, Danish mathematician Johan Jensen described the ‘fallacy of the average’, now known as Jensen’s inequality. Jensen pointed out that when relationships are non-linear we can’t assume that the average performance is equal to the performance under average conditions.
In our example, Jensen’s inequality means we further under-estimate reproductive output from inside the MPA. This is because fish size relates to reproductive output in a non-linear way so the reproductive output at average size is not the same as the average reproductive output. The inequality is greater inside the MPA where fish sizes are bigger and so this makes a further contribution to our under-estimate of reproductive output.
When we take Jensen’s inequality into account, and add it to the underestimates relating to the non-linear relationships already discussed, we find that there is a 175% increase in reproductive output for fish inside MPAs compared to those outside.
While this translates to a much smaller ‘spill-over’ effect, (more like a 12% increase in tonnes of caught fish per year for the coral trout fishery when MPAs are included in the management of the fishery), it is still a substantial increase in yield.
MPAs represent an essential tool for protecting larger fish, and the research team hope that a more accurate accounting of the value of MPAs will increase support for their use by a wide variety of stakeholders, including anglers themselves.
Authors: Martino E Malerba, Craig R White, and Dustin J Marshall
Published in: Frontiers in Ecology and the Environment
Artificial structures are proliferating along coastlines worldwide, creating new habitat for heterotrophic filter feeders. The energy demand of this heterotrophic biomass is likely to be substantial, but is largely unquantified.
Combining in situ surveys, laboratory assays, and information obtained from geographic information systems, we estimated the energy demands of sessile invertebrates found on marine artificial structures worldwide.
At least 950,000 metric tons of heterotrophic biomass are associated with commercial ports around the world, emitting over 600 metric tons of carbon dioxide into the atmosphere and consuming 5 million megajoules of energy per day.
We propose the concept of a trophic “footprint” of marine urbanization, in which every square meter of artificial structure can negate the primary production of up to 130 square meters of surrounding coastal waters; collectively, these structures not only act as energy sinks and carbon sources, but also potentially reduce the productivity of coastal food webs.
Malerba ME, White CR, Marshall DJ (2019) The outsized trophic footprint of marine urbanization. Frontiers in Ecology and the EnvironmentPDFDOI