17 May 2024. Members of our Evolutionary Fish Ecology Lab had a blast attending this years 47th Larval Fish Conference in Huron, OH. Hannes, Emma, Max and alumnus Chris Murray (now at WHOI), went on a road trip from Connecticut to Lake Eerie to present and learn about all things larval fish. On Tuesday morning, the conference crowd enjoyed excursion or recreation options, all the while catching up with good old colleagues and making new connections and friends. A particular achievement: each of us presented research on a different fish species; while Hannes showed the first data emerging from his sabbatical research on Chilean silversides, Emma talked about baby California grunion development. Max presented a poster and two talks, the first about CO2 effects on the onset of schooling in Atlantic silversides and a second one one Black Sea Bass overwintering dynamics. Last, Chris Murray gave a fascinating first look into gene expression data from our most recent sand lance CO2 experiment. All around, a strong showing of our lab!
Contributions from our lab to the 47th Larval Fish Conference in Huron, OH
Baumann, H., Gallardo, A., Gallardo, C., and Urbina, M. 2024. First evidence for countergradient growth variation in the Chilean silverside Odontesthes regia. Oral presentation
Siegried, E. and Johnson, D. 2024. Eyes bigger than your stomach: developmental inaccuracy in larval California grunion. Oral presentation.
Zavell, M.D., Mouland, M., Barnum, D., Matassa, C., Schultz, E.T., and Baumann, H. 2024. Overwintering dynamics of northern stock Black Sea Bass, Centropristis striata, juveniles. Oral presentation.
Zavell, M.D., O'del, J., Mouland, M., Webb, J.F., and Baumann, H. 2024. Ontogeny of larval schooling and effects of ocean acidification in Atlantic Silversides (Menidia menidia). Oral presentation.
Murray, C.S., Jones, L., Siegfried, E., Zavell, M.D., Baumann, Z., Wiley, D., Therkildsen, N., Aluru, N., and Baumann, H. 2024. Examining the effect of ocean acidification on hatching enzyme gene expression in Northern sand lance (Ammodytes dubius). Oral presentation.
24 June 2023. We are overjoyed to be able to announce today that NSF's Division of Integrative Organismal Systems has awarded our proposed research to better understand sand lance CO2 sensitivity!
With a sense of pride and humility we will take on this intriguing case, follow it down some rabbit holes, while keeping in mind the big picture. This fall, our collaborative team will begin its renewed work, now on both congeneric sandlance species (Ammodytes dubius, A. americanus).
We already have two talented PhD students recruited to the task, Lucas Jones and Emma Siegfried. With curiosity and anticipation, we look forward to the next years of eco-evolutionary research on some of the most important forage fish species on the Northwest Atlantic Shelf.
NSF-ORCC (Organismal Response to Climate Change): Collaborative Research: Mechanisms underpinning the unusual, high CO2 sensitivity of sand lances, key forage fishes on the Northwest Atlantic Shelf (#2307813, 2023-2026, $1,050,000)
*** Why are sand lance so sensitive to future high CO2 conditions in the ocean? ***
Public Award Abstract
Ocean warming and acidification are direct, predictable consequences of man-made climate change with likely vast but still insufficiently understood consequences for marine life.
So far, most tested fish species appear only mildly sensitive to ocean acidification, but sand lances are an exception. Sand lances are small, eel-like, schooling fishes of enormous importance as food for marine fish, seabirds and mammals in temperate to polar ecosystems, and recent research conclusively demonstrated that many sand lance embryos have trouble developing and hatching under predicted future ocean conditions.
This project uses modern experimental and molecular tools to understand exactly WHY sand lance embryos are so unusually sensitive and which genes and enzymes are responsible for this. Genes will also reveal whether some specific genotypes are less sensitive to warming and acidification, which can then be used to predict whether the species could evolve to be more tolerant over time.
Another important objective is to test a closely related sand lance species to find out, whether the high climate sensitivity might be of general concern in this important group of forage fishes. This project combines innovative ecological, evolutionary, and genomic research to help society anticipate looming marine ecosystem changes in the 21st century, while equipping the next generation of scientists with the needed tools and expertise to succeed in the challenges ahead.
The project also creates opportunities for high school students from underprivileged Connecticut schools to accompany the team on sand lance sampling trips to Stellwagen Bank National Marine Sanctuary.
American sand lance (Ammodytes americanus) swimming in surface waters of Wells Harbor, ME in November 2021
Technical Award Abstract
Two recent studies on Northern sand lance (Ammodytes dubius), a key forage fish on offshore sand banks across the Northwest Atlantic shelf (NWA), have robustly demonstrated that predicted future CO2 conditions induce some of the most severe reductions in embryo survival and hatching success seen yet among tested fish species. This project has four objectives for revealing the mechanisms underpinning this unusual, high CO2-sensitivity as well as the ubiquity and genetic basis of this phenomenon.
[1] For the first time, we will rear A. dubius offspring produced from wild spawners to late larval stages at factorial CO2 × temperature conditions to test whether sand lance larvae are as CO2-sensitive as embryos.
[2] For the first time, we will use transcriptomic tools (RNAseq, RT-qPCR) to elucidate mechanisms causing ‘CO2-impaired hatching’, focusing specifically on hatching enzymes, to better understand a newly discovered mortality mechanism due to high CO2 in fishes.
[3] Modern genomic approaches (low-coverage whole genome sequencing; allele frequency shifts, relatedness analyses) will reveal whether high CO2-sensitivity has a genetic basis in sand lance and could therefore evolve.
[4] And for the first time, we will extend CO2 × temperature experiments to a congener, the American sand lance (A. americanus), which provides an important scientific contrast between nearshore vs. offshore species CO2-sensitivities and will yield critical insights whether high CO2-sensitivity is a wider concern within the sand lance family.
5 December 2022. We are proud to announce that the ICES Journal of Marine Science just published our latest sand lance study! The work spearheaded by Lucas Jones and subject of his Masters thesis research has brought together a large, international group of collaborators to better understand the genetic relationships between disparate sand lance populations across their large geographical range. This is an Open Access publication that will hopefully be of use to researchers studying sand lance everywhere.
Genetic Barriers, a Warming Ocean, and the Uncertain Future for an Important Forage Fish
In the vast oceans, one would assume their inhabitants can travel far and wide and, as a result, populations of a species would mix freely. But this doesn’t appear to be the case for a vital forage fish called the sand lance.
Sand lance are small schooling fish impressively rich in lipids, which makes them a fantastic and significant food source for at least 70 different species ranging from whales and sharks to seabirds, says UConn Associate Professor of Marine Sciences Hannes Baumann.
The Northern sand lance can be found from the waters off New Jersey all the way north to Greenland. Researchers, including Baumann and Ph.D. student Lucas Jones, were interested to see if sand lance constitute a massive, homogenous population, or whether there are genetically distinct groups. Their findings are published in the ICES Journal of Marine Science.
Baumann explains these are important questions to answer when considering conservation and sustainable management of the species, especially since the regions where sand lance live are warming faster than many areas of the planet due to climate change.
Sampling fish from such a broad range is no small task, but two years ago, Baumann and Jones began reaching out to other researchers to see if they had tissue samples to spare. Baumann credits the work to the international group of colleagues who contributed samples including co-authors from Canada and Greenland, and who helped sequence and analyze the data including co-authors from Cornell University.
In all, Baumann, Jones, and the team were able to sequence and analyze nearly 300 samples from a variety of locations across the sand lance’s range using a technique called low-coverage whole genome sequencing. They also sequenced the first reference genome for sand lance.
In a nutshell, Baumann says they found an area on the Scotian Shelf, off the coast of Nova Scotia, where a genetic break occurs. The researchers distinguished two distinct groups, one north and one south of the divide, with parts of the genome differing quite dramatically – namely on chromosomes 21 and 24. Without obvious physical barriers like a mountain range separating the groups, Baumann says it’s logical to ask how these differences are possible.
“That is the scientific conundrum,” says Baumann, and the answer, it appears, lies in the currents.
“When fish from the north reproduce and drift south, they are genetically less adapted to warmer southern waters, even if it’s five or six degrees warmer in the winter, they are just not surviving,” Baumann says. “These populations may be linked by the ocean currents, but the realized connectivity is basically zero.”
Separation of 3 sand lance species based nuclear and mitochondrial DNA (Jones et al. ICES JMS 2022)
This finding is a first for the sand lance, but it has been shown in other species such as lobsters, cod, and scallops, and this research adds further evidence to an apparent temperature divide at the Scotian Shelf, and helps demonstrate that temperature is an important factor in survival.
“Example after example shows that the ocean is not as homogeneous a place as expected, and there are all kinds of things that prevent that constant mixing,”Baumann says. “We found another striking example of that.”
When researchers find adaptation in an environment where mixing is continuous, like in the ocean, Baumann says, the question is how it is possible that groups stay different, even though they are constantly encountering other genotypes. That is where powerful genomic methods, like the ones used in this paper, come in handy.
“Parts of the genome in many species have what we call a ‘genetic inversion,’ which means that the genes on the chromosome from one parent have a certain order and the genes on the same chromosome that come from the other parent that code for the same thing, and they’re the same area, but they’re flipped,” Baumann says.
These inversions mean recombination cannot occur; therefore, the genes are passed down through the generations and play an important role in adaptation.
“We discovered on chromosomes 21 and 24 there are whole regions that are completely different and that is like the trademark signature of what we call an inversion because there’s no recombination going on.”
Baumann says that knowing there are genetic and ecological barriers on the Scotian Shelf is important, because with climate change, this barrier may move north and while that may be good news for southern fish, it’s bad news for the fish currently there.
The researchers were also a little relieved in finding two clusters, because had there been many smaller clusters, it could make management and conservation more challenging, especially considering scenarios like the construction of offshore wind parks. Areas potentially well situated for wind turbines can also be habitats for sand lance, and construction disrupts habitats. If there were many, smaller population clusters, a single construction project could pose the risk of completely wiping out a cluster, whereas with more widely dispersed populations, though the local population may be temporarily disturbed, it will not be long before they are able to re-establish after construction is completed.
Baumann plans to focus further research on studying the genetic basis of the thermal divide.
“We want to make sure that this fish is productive and resilient, despite climate change, so we should make sure these areas where they are occurring are protected,” Bauman says. “These decisions should include experts to ensure if there’s an area that is very critical to sand lance, that any disturbance is temporary.”
It isn’t an unsolvable conflict, but it is something that we need to do, says Baumann, who also notes that it is possible that sand lance north of the thermal divide are already suffering more from warming because the region is warming faster.
“It could be that these two clusters have different vulnerabilities to climate change,” he says. “We don’t know that yet but that’s something that should be pursued.”
When carbon is emitted into the atmosphere, about a quarter of it is absorbed by the earth’s oceans. As the oceans serve as a massive ‘sink’ for carbon, there are changes to the water’s pH – a measure of how acidic or basic water is. As oceans absorb carbon, their water becomes more acidic, a process called ocean acidification (OA). For years, researchers have worked to understand what effect this could have on marine life.
While most research so far shows that fish are fairly resilient to OA, new research from UConn, the University of Washington, the National Oceanic and Atmospheric Administration (NOAA), and Southern Connecticut State University, shows that an important forage fish for the Northwest Atlantic called sand lance is very sensitive to OA, and that this could have considerable ecosystem impacts by 2100. The team’s findings have just been published in Marine Ecology Progress Series 687.
Sand lance spawn in the winter months in offshore environments that tend to have stable, low levels of CO2, explains UConn Department of Marine Sciences researcher and lead author Hannes Baumann.
“Marine organisms are not living in a uniform ocean,” Baumann says. “In near shore environments, large CO2 fluctuations between day and night and between seasons are the norm, and the fish and other organisms are adapted to this variability. When we stumbled upon sand lances we suspected they are different. We thought that a fish that lives in a more open-ocean offshore environment might be more sensitive than the near-shore fish because there’s just much less variability.”
The project was a collaboration with physical oceanographers, including Assistant Professor of Marine Sciences Samantha Siedlecki and Michael Alexander from NOAA’s Physical Sciences Laboratory in Boulder, Colorado, who modeled CO2 levels in 2050 and 2100 for a specific part of the Gulf of Maine where sand lance spawn. Then Baumann and his team reared sand lance embryos in the lab under experimentally higher CO2 levels matching the projected levels.
There are instances of direct fish mortality as result of elevated CO2, but they are rare, says Baumann. However, sand lance embryos proved to be exceptionally sensitive, and fewer embryos hatched under future oceanic CO2 conditions. The researchers repeated the experiments three more times to avoid jumping to conclusions but each time they observed the same result.
“We found that embryo survival-to-hatch decreased sharply with increasing CO2 levels in the water, concluding that this is one of the most CO2-sensitive fish species studied thus far,” Baumann says.
Sand lances are surely one of the most important forage fish here on the Northwest Atlantic shelf… The humpback whales, sharks, tuna, cod, shearwaters, terns — you name it — they are all relying on sand lance.
With this interdisciplinary approach combining model forecasts and serial experimentation the researchers arrived at a picture that is much more specific.
“We consequently applied principles of serial experimentation, which is a most timely and important topic in ocean acidification research right now,” Baumann says. “Because our findings are backed up by repeated independent evidence, they are more robust than many published ocean acidification studies to date.”
In addition to preventing many sand lance embryos from developing normally, the researchers document a second negative, and novel, response to elevated CO2. Higher CO2 levels appear to make it harder for embryos to hatch.
Baumann explains the lowered pH likely renders enzymes needed for successful hatching less effective, leaving the embryos unable to break through their eggshell (chorion) to hatch.
The results show that by 2100, due to acidification, sand lance hatching success could be reduced to 71% of today’s levels. Since sand lance are such a critical component of the food web of the Northwest Atlantic, this marked decrease in sand lance would have profound impacts throughout the ecosystem.
“Sand lances are surely one of the most important forage fish here on the Northwest Atlantic shelf,” Baumann says. “Their range spans from the Mid Atlantic Bight all the way to Greenland. Where we studied them, on Stellwagen Bank, they are called the backbone of the ecosystem. The humpback whales, sharks, tuna, cod, shearwaters, terns — you name it — they are all relying on sand lance, and if sand lance productivity goes down, we will see ripple effects to all these higher trophic animals. Even though we humans don’t fish for sand lance, we need to take care of the species because it has such a huge effect on everything else.”
Baumann says this study supports the hypothesis that offshore, high latitude marine organisms like the sand lance may be among the most vulnerable to OA. As a result, these organisms and food webs will likely be impacted first and soon, and we must act now.
Previous research has focused on opportunistically chosen species when testing their sensitivity for ocean acidification, says Baumann, but this should change.
“We need strategic thinking about what species we are testing next, because we cannot test every marine fish species, that’s an impossible task. We should concentrate on fish species that are likely the most vulnerable, and therefore the ones that are probably being affected first and this research makes a compelling argument that those are the fish species at higher latitudes and in more offshore than nearshore environments.”
4 March 2022. Hannes was the invited speaker at today's Friday seminar of the Department of Marine Sciences. His talk gave an overview of the research highlights of our multi-disciplinary and multi-institutional efforts to better understand basic ecological facts, population connectivity & structure, and the unusually high CO2-sensitivity of sand lance embryos. The remotely given presentation was attended by 62 people, some of which listened in from as far away as Norway!
The talk was recorded and can be accessed via the public link below.
The unusual ecology and climate sensitivity of sand lance, a key forage fish on the Northwest-Atlantic Shelf
No matter how you look at these small, slender-bodied fishes that at times live buried in sediment or emerge as dense pelagic schools, northern sand lance (Ammodytes dubius) easily awe even the most hard-to-impress scientist or naturalist. Their unusual behavior, patchy occurrence, and reproductive timing are paralleled by their extraordinary importance as forage fish that sustain well-known hotspots of iconic predators (cod, tuna, sharks, seabirds, whales) all across the Northwest Atlantic shelf. And yet, despite their recognized role as the ‘backbone’ of many shelf ecosystems, we still don’t understand many basic aspects of sand lance ecology, population structure and their vulnerability to manmade climate change. Over the past years, our lab has been working alongside other US and Canadian research groups on multiple sand lance projects that have produced stunning new insights into these enigmatic fish. This seminar will outline some of the highlights. We discovered that the seasonal growth of these fish relies heavily on the lipid-rich copepod Calanus finmarchicus and showed that after a dormancy period in summer they spawn on Stellwagen Bank for just a brief period at the end of fall. To resolve questions of connectivity between sand lance areas, we performed large-scale Lagrangian drift simulations that suggested areas of high, low and negligible retention of sand lance offspring and showed overlaps with planned offshore wind lease areas. A large collaborative effort succeeded in obtaining specimens from across the entire distributional range (Greenland to Mid-Atlantic Bight), and subsequent whole genome sequencing newly revealed a stark genomic differentiation between northern and southern population clusters. Last, we performed multiple years of rearing experiments on embryos that consistently showed an unusual sensitivity of sand lance to future, high CO2 oceans. When coupled with regional, end-of-century pCO2 projections we estimate that rising CO2 levels alone could reduce sand lance hatching success to 71% in 2100 relative to today. Warming, acidification, and habitat exploitation therefore emerge as key factors lining up against the future productivity of this forage fish, which is so critically important across Northwest-Atlantic shelf ecosystems.
Monday, November 22nd 2021. Big and heartfelt congratulations to Lucas Jones, who presented his Master thesis to his peers at the institute and colleagues national and international. Well done, Lucas!
A link to his recorded presentation will be posted here soon.
The UConn Department of Marine Sciences
Presents a Master’s Thesis Presentation by
Lucas Jones
B.A., University of Connecticut, 2018
4:00 p.m., Monday, November 22, 2021
Marine Sciences Building, Seminar Room 103
Using Low-Coverage, Whole Genome Sequencing to Study Northern Sand Lance (Ammodytes dubius) Population Connectivity in the Northwest Atlantic
Northern sand lance (Ammodytes dubius) are key forage fish in Northwest Atlantic (NWA) shelf ecosystems, where they exclusively occur on coarse-grain, offshore sand banks. This patchy occurrence may result in genetically more fragmented, less connected populations, but traditional morphological or genomic approaches have so far been unsuccessful in fully resolving the species’ population structure and connectivity. My study pursued an alternative genomic approach, using low-coverage, whole genome sequencing (LcWGS) to address these important questions. I extracted DNA from 273 A.dubius specimens collected by collaborators from sevenregions across the species geographical range, from Greenland to New Jersey, USA. From LcWGS data, I identified 11,558,126 single nucleotide polymorphisms (SNPs) that allowed quantifying genetic differentiation between populations (FST), thereby revealing the genetic structuring of populations throughout the NWA. Despite the potentially homogenizing influence of the general north to south ocean circulation, I found a clear genetic break around Nova Scotia that delineated a northern from a southern A. dubius supergroup. Only within the southern supergroup, genetic distances increased with the geographic distance between sample sites. At the focal site of Stellwagen Bank (southern Gulf of Maine), A. dubius samples collected over several years (2014 – 2019) revealed small but significant temporal genetic differences that imply varying occupation of this offshore habitat by genetically different sand lance contingents. Inclusion of samples from the inshore congener A. americanus confirmed the clear genetic separation between both species and further determined that all sand lance caught on Stellwagen Bank are exclusively A. dubius. Overall, my work suggests the existence of two spatially distinct A. dubius populations with little ‘realized’ connectivity, which is critical knowledge to aid protection and management of offshore marine resources.
20 March 2020. We are happy to announce that the prestigious journal Fish & Fisheries just published a comprehensive review about the role of sand lance in the Northwest Atlantic Shelf ecosystem. The article, which came out of a workshop on this topic three years ago, reviews the the current state of knowledge about these enigmatic and important forage fish and urges continued efforts to better understand their role in the ecosystem and sensitivity to climate stressors.
This work represents the first comprehensive assessment of this important forage fish in the Northwest Atlantic, though similar efforts have been carried out in the Pacific Northwest and Europe. In the Atlantic, sand lance are observed to be a significant food source for the federally endangered Roseate tern, Atlantic sturgeon and cod, Harbor and Grey seals and Minke and Humpback whales. “This paper is a call to our peers and colleagues that there is a big gap in knowledge, and to bring more attention to these species as unmanaged forage fish,” says Staudinger.
18 December 2019. Hannes and Lucas just returned from a spontaneous road trip to visit our good friends and collaborators at the University of Quebec in Rimouski (UQAR), Canada. We drove for over 10 hours (one-way) through snowstorms and icy voids to meet with Prof. Dominique Robert, who had collected sand lance samples from the Gulf of St. Lawrence and from Nova Scotia to be included in our new genomic study on the population connectivity of this species. Hannes gave a talk about our sand lance work and we saw a new institute in a new place, while frantically trying to stay warm amidst the brutal cold. Seeing the St. Lawrence in its icy, majestic beauty was a truly amazing experience.
Afterwards, we drove back through Maine and then repeated the fin-clipping of samples in Scituate at the Stellwagen Bank National Marine Sanctuary office, so we now have almost all samples in hand to start the DNA extraction and sequencing.
We are excited for the next steps!
Sand lance samples to be included in the genomic study
Rimouski on the south shore of the mighty and icy St. Lawrence River on 12/13/19
Lucas and Hannes listened to Corinne Burns talking about her PhD research at UQAR on 16 Dec 2019
Hannes gave a talk about sand lance research at UConn
The icy beauty of the St. Lawrence River
Sun glistening on the ice on the banks of the St. Lawrence River on 16 Dec 2019
Snow storm on the I91 in Vermont on 15 Dec 2019
Driving back through Maine on 17 Dec 2019 ... 6h of snowstorm
Sand lance sample thawing to be fin-clipped
Hannes and Lucas fin-clipping specimens in the Stellwagen Bank NMS office in Scituate on 18 Dec 2019
21 November 2019. We are excited to announce the Chris Murray‘s paper on the unusual, high sensitivity of early life Northern sand lance to acidification and warming has just been published in the journal of Conservation Physiology! This is the first publication of our extensive work on this enigmatic species.
Sand lance species play a key ecological role in most temperate to polar shelf ecosystems of the northern hemisphere, but they have remained unstudied with respect to their sensitivity to predicted future CO2 levels in the ocean. For the past three years (2016 – 2018), we have sampled and spawned with northern sand lance (Ammodytes dubius) from Stellwagen Bank National Marine Sanctuary and subsequently reared their embryos under factorial CO2 x temperature conditions to hatch and early larval stages. Our results were striking, in all years, high CO2 conditions severely reduced embryo survival up to 20-fold over controls, with strong synergistic reductions under combined high CO2 and temperature conditions. High CO2 also delayed hatching, reduced remaining endogenous energy reserves at hatch, and in combination with higher temperatures, reduced embryonic growth.
Indeed, given the observed effect sizes, northern sand lance might be the most CO2 sensitive fish species tested to date.
15 November 2018. After a stretch of foul weather kept us from going out to Stellwagen Bank last week, this time all the stars aligned for Emma and Mackenzie. Due to their success in catching spawning ripe Northern sandlance, we are now embarking on our third year of CO2 x temperature experiments on this species!
Here is how Mackenzie Blanusa experienced her first trip to these enigmatic waters:
“This particular sandlance cruise was a day filled with firsts and is definitely a trip to remember. I accompanied Emma, Hannes’ postdoc, up to Scituate the night before the cruise and was given a rundown of what needed to be accomplished. I was a bit overwhelmed at first, because I’ve never dealt with sandlances before and did not know a lot about these fish. Nevertheless, I was eager to learn something new and was ready to help out wherever needed.
The goal of the sandlance cruise was to collect running ripe males and females to do a fertilization via strip spawning. Emma and I were a bit doubtful at first because we got less than 10 sandlance on the first two trolls. However, things got much better by the afternoon, and our most successful trawl caught 147 sand lance. I helped out with the fertilization and deploying the trawl, two things I have never done before. The most exciting part of the day was getting to see humpback whales. Usually they are in the distance but today they were right next to the boat. Everyone on board said that this never happens and it was very unusual so I felt very lucky to have seen whales at such a close proximity.”
Overall, the trip was a huge success and it was very refreshing to see everything go as planned. The only downside to the day was driving back home through a snowstorm. I later found out that there was a 73% fertilization success and we got 27,000 embryos for Emma’s experiment. I am very grateful to have gotten the opportunity to help out on this sampling cruise and am looking forward to doing this again in the future!