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Tomas Brodin

Tomas Brodin
Tomas Brodin is an aquatic ecologist, studying ecological effects of pharmaceuticals in aquatic systems. Tomas has a background in evolutionary- and behavioral ecology, and is now using this knowledge to increase the ecological relevance of chemical risk-assessment in general, and pharmaceutical risk-assessment in particular. His research bridges the gap between the lab and the real world by combining lab-experiments, for mechanistic understanding, with large-scale field studies that answers the million dollar question “What happens in the lake or stream?”


My research is currently focused on four main tracks.  

I. Ecological effects of pharmaceuticals in the environment (2010 – present)

Humans consume more pharmaceuticals than ever and consumption is set to rise. As a consequence, increasing amounts of pharmaceuticals are released into waterways worldwide with virtually no knowledge of how they might affect aquatic ecosystems. Some conspicuous effects of these emerging contaminants are already evident including the feminization of fish by contraceptive residue. However, recent work suggests that important effects of pharmaceuticals in aquatic environments are much more widespread than currently believed, and that these effects may result in major changes in species interactions, population survival and ecosystem functioning. In several earlier laboratory studies, I have shown that concentrations of pharmaceuticals presently found in waterways alter important behavioural traits in both aquatic macroinvertebrates and wild fish, and that this in turn affects both feeding efficiency and predation risk. These results suggest that pharmaceutical contamination of aquatic environments may change species interactions, in particular predator-prey interactions, with severe ecosystem-effects as potential consequence. Recently my research focus has turned towards realistic large-scale studies in lakes and rivers using a combination of acoustic telemetry and physiological biologgers to test if findings from the lab also hold in natural settings. The overall finding of the studies suggests that effects of pharmaceutical contamination of natural systems might be much more widespread than we predict based on conventional ecotoxicological tests. Our results highlight the importance of validating lab-results with field-studies, since the increased complexity of real ecosystems can produce unexpected effects of contamination.

When I first started this multidisciplinary collaboration to study ecological effects of pharmaceuticals in the environment I did not expect that the reactions to the results would be so intense. The first paper was published in the journal Science (Brodin et al. 2013) and showed that environmentally relevant concentrations of an anxiolytic drug, that are found in surface waters all over the world, alters ecologically important behavioral traits. The paper was selected to represent Science, and was presented as the “main attraction”, at the annual AAAS-meeting in Boston where I was invited to hold a press conference.  After this I continued to study the ecological effects of pharmaceuticals in aquatic environments and have since published an additional 16 medium to high-impact papers (e.g. Nature Communications). One important reason for the very high interest from both press and public was my focus on ecological relevance. For example, I decided early on to only use common species of wild fish from northern populations to study potential ecological effects. As it turns out the effects of pharmaceuticals are asymmetrical, even between fish species, and interestingly aquatic invertebrates (food for fish) can bio-concentrate up to 10000 times higher concentrations of pharmaceuticals in their tissue than is present in the surrounding water. While many fish species are affected (e.g. perch - Perca fluviatilis, roach – Rutilus rutilus and Atlantic salmon – Salmo salar) by for example anti-anxiety medication some species (e.g. crucian carp - Carassius carassius, pike – Esox lucius) seem unaffected and the mechanistic explanation for this is currently under investigation.

II. Causes for, and consequences of, the spread of invasive species (2011 – present)

Within this project I study relationships between behaviorally dependent dispersal, performance and ecological impact. In particular, I focus on if dispersers (at the invasion front) have higher performance and impact in new patches than non-dispersers (in older parts of the invasion succession), and how that is affected by density, competition and predation. So far I have mainly used the round goby (Neogobius melanostomus) as a study species since it is an extremely successful invasive that currently is spreading north in the Baltic Sea. Another important reason to increase our knowledge of the round goby is that they readily invade freshwater rivers and streams and is considered a future major threat for wild trout and salmon populations along the Swedish east coast.

This part of my research profile has already generated many interesting results and received international attention. Among other things, we have shown that individuals from newly founded populations of this invasive fish species display a different behavioural repertoire (i.e. more active, more risk-taking and less social) compared to individuals in older populations. This behavioural difference translates into individuals at the invasion front being better at consuming novel food, which could have grave implications for what ecological consequences to expect at the front of biological invasions. Indeed, we just recently published a paper showing that dispersing invaders invoke a more pronounced ecological effect on the invertebrate community compared to non-dispersing invaders.

I am currently involved in two collaborative projects, one national and one international (Sweden, Denmark, Switzerland), that study different aspects of the ecological consequences that round goby can have on northern aquatic ecosystems. 

III. Mechanisms driving dispersal and/or migration of fish (2013 – present)

As a result of large-scale environmental alterations due to climate change, many species are experiencing range-margins shifting northward (or southward on the southern hemisphere). As new suitable habitat is made available to the north, some habitats in the south become less suitable. Hence it is imperative for long-term persistence that organisms, especially at the range-margins, are able to disperse and colonize new patches. While many previous studies have focused on species characteristics that might explain between-species differences in dispersal/migration tendency, I chose to look at how individual variation in dispersal associated traits within species might influence dispersal propensity. Furthermore, I am particularly interested in how anthropogenic influences (e.g. chemical pollution, climate change, and habitat alterations like dams) might affect the distribution of traits within a population and how this in turn affects dispersal and/or migration.

Within this project I use a combination of lab assays and field studies to disentangle the multiple factors influencing dispersal and/or migration decisions at any given time. For example, we recently published a paper (Hellström et al. 2016) where we combined migration studies in the lab with studies of migration in a natural stream and showed for the first time that fear is an important factor determining migration speed in Atlantic salmon smolt. Another recently published interesting discovery is that the age since first establishment of a population is important for what traits induce dispersal (Thorlacius & Brodin 2017).

IV. Biodiversity in air


Selected publications

1.     Vossen, L., Cerveny, D., Osterkrans, M., Thörnqvist, P-O., Jutfelt, F., Fick, J., Brodin, T., Winberg, S. 2020. Chronic Exposure to Oxazepam Pollution Produces Tolerance to Anxiolytic Effects in Zebrafish (Danio rerio). Environmental Science & Technology 54: 1760-1769.

2.     Cerveny, D., Brodin, T., Cisar, P., McCallum, E., Fick, J. 2020. Bioconcentration and behavioral effects of four benzodiazepines and their environmentally relevant mixture in wild fish. Science of the Total Environment 702, 134780.

3.     Vossen, L., Cerveny, D., Sen Sarma, O., Thörnqvist, P.O., Jutfelt, F., Fick, J., Brodin, T and Winberg, S. 2020. Low concentrations of the benzodiazepine drug oxazepam induce anxiolytic effects in wild-caught but not in laboratory zebrafish. Science of the Total Environment 703, 134701.

4.     Therry, L., Raffard, A., Finn, F., Koch, K., Brodin, T., Simon Blanchet, S., Cote, J. 2020. Does range expansion modify trait covariation? A study of a northward expanding dragonfly. Oecologia 192: 565–575.

5.     Behrens, J., von Friesen, L., Brodin, T., Ericsson, P., Hirsch, Persson, A., Sundelin, A., van Deurs, M., Nilsson, A. 2020. Personality- and size-related metabolic performance in invasive round goby (Neogobius melanostomus). Physiology & Behavior 215, 112777. 

6.     Michelangeli, M., Cote, J., Chapple, D., Sih, A., Brodin, T., Fogarty, S., Bertram, M., Eades, J., Wong, B. 2020. Sex-dependent personality in two invasive species of mosquitofish. Biological Invasions 22: 1353–1364

7.     Ågerstrand, M., Balshine, S., Brodin, T., Brooks, B., Maack, G., McCallum, E., Pyle, G., Saaristo, M., Ford, A. 2020. Use of behavioural endpoints in regulation of chemicals. Environmental Science: Processes & Impacts (accepted)

8.     Karlsson, E., Johansson, A-M., Ahlinder, J., Lundkvist, M., Singh, NJ., Brodin, T., Forsman, M., Stenberg, P. 2020. Airborne microbial biodiversity and seasonality in Northern and Southern Sweden. PeerJ 8, e8424.

9.     Leander, J., Klaminder, J., Jonsson, M., Brodin, T., Leonardsson, K. and Hellström, G. 2020. The old and the new: evaluating performance of acoustic telemetry systems in tracking migrating Atlantic salmon (Salmo salar) smolt and European eel (Anguilla anguilla) around hydropower facilities. Canadian Journal of Fisheries and Aquatic Sciences. (Early online)

10.  Späth, J., Nording, M., Lindberg, R., Brodin, T., Jansson, S., Yang, J., Wan, D., Hammock, B., Fick, J. 2020. Novel metabolomic method to assess the effect-based removal efficiency of advanced wastewater treatment techniques. Environmental Chemistry 17: 1-5.

11.  Luttbeg, B., Hammond, J., Brodin, T. and Sih A. 2020. Predator hunting modes and predator-prey space games. Ethology 126: 476-485.

12.  McCallum, E., Cerveny, D., Fick J., Brodin, T. 2019. Slow-release implants for manipulating contaminant exposures in aquatic wildlife – a new tool for field ecotoxicology. Environmental Science and Technology 53: 8282−8290.

13.  Sundin, J., Jutfelt, F., Thorlacius, M., Fick, J., Brodin, T. 2019. Behavioural alterations induced by the anxiolytic pollutant oxazepam are reversible after depuration in a freshwater fish. Science of the Total Environment 665: 390-399.

14.  McCallum, E., Lindberg, R., Andersson, A., Brodin, T. 2019. Stability and uptake of methylphenidate and ritalinic acid in nine-spine stickleback (Pungitius pungitius) and water louse (Asellus aquaticus). Environmental Science and Pollution Research 26: 9371–9378.

15.  McCallum, E., Sundelin, A., Fick, J., Alanara, A., Klaminder, J., Hellström, G., Brodin, T. 2019. Investigating tissue bioconcentration and the behavioural effects of two pharmaceutical pollutants on sea trout (Salmo trutta) in the laboratory and field, Aquatic Toxicology 207: 170-178.

16.  Brodin, T., Fogarty, S., Sih, A., Cote. J. 2019. Personality-dependent survival of the invasive mosquitofish: being social can be deadly. Aquatic Invasions 14: 465–477.

17.  Klaminder, J., Fahlman, J., Leander, J., Jonsson, M., Brodin, T., Fick, J., Hellström, G. 2019. Less anxious salmon smolt become easy prey during downstream migration. Science of the Total Environment 687: 488–493.

18.  Hellström, G., Brodin, T., Jonsson, M., Olsen, H., Leander, J., Fahlman, J., Fick, J., Klaminder, J. 2019. Environmentally relevant concentrations of the common anxiolytic pharmaceutical oxazepam do not have acute effect on spawning behavior in mature male Atlantic salmon (Salmo salar) parr. Journal of Applied Ichthyology (Early online)

19.  Jonsson, M., Andersson, M., Fick, J., Brodin, T., Klaminder, J., Piovano, S. 2019. High-speed imaging reveals how antihistamine exposure affects escape behaviors in ambush predator’s prey. Science of the Total Environment 648: 1257–1262.

20.  Saaristo, M., Lagesson, A., Bertram, M.G., Fick, J., Klaminder, J., Johnstone, C.P., Wong, B.B.M, Brodin, T. 2019. Behavioural effects of anxiolytic exposure, temperature and predation-risk on the European perch (Perca fluviatilis). Science of the Total Environment 655: 1311–1320.

21.  Hellström, G., Brodin, T., Palm, D., Rivinoja, P., 2019. Effects of boulder addition on European grayling (Thymallus thymallus) in a channelized river in Sweden. Journal of Freshwater Ecology 34: 559-573.

22.  Lagesson, A., Saaristo, M., Brodin, T., Fick, J., Klaminder, J., Martin, J., Wong B. 2019.  Fish on steroids: temperature dependent effects of 17β-trenbolone on predator escape, boldness, and exploratory behaviors. Environmental Pollution 245: 243-252.                          

23.  Richmond, E.K., Rosi, E., Walters D.M., Fick, J., Brodin, T., Sundelin, A., Grace, M.R. 2018 Medicated stream and riparian food webs: 69 pharmaceuticals contaminate stream insects and their consumers. Nature Communications 9:4491.

24.  Thorlacius, M., Brodin, T. 2018. Investigating large‐scale invasion patterns using‐small scale invasion successions—phenotypic differentiation of the invasive round goby (Neogobius melanostomus) at invasion fronts. Limnology and Oceanography 63: 702-713.

25.  Rodriguez, A., Zhang, H., Klaminder, J., Brodin, T., Andersson, P., Andersson, M. 2018. ToxTrac: a fast and robust software for tracking organisms. Methods in Ecology and Evolution 9: 460-464.

26.  Fahlman, J. Fick, J. Karlsson, M. Jonsson, M. Brodin, T. Klaminder J. 2018. Using laboratory incubations to predict the fate of pharmaceuticals in aquatic ecosystems, Environmental Chemistry 15: 463-471.

27.  van den Brink, P.J.V., Boxall, A., Maltby, L., Brooks, B.W., Rudd, M.A., Backhaus, T., Spurgeon, D., Verougstraete, V., Ajao, C., Ankley. G.T., Apitz, S.E., Arnold, K., Brodin, T., Cañedo-Argüelles, M., Chapman, J., Corrales, J., Coutellec, M-A., Fernandes, T.F., Fick, J., Ford, A.T., Giménez Papiol, G., Groh, K.J. Hutchinson, T.H., Kruger, H., Kukkonen, J.V.K., Loutset, S., Marshall, S., Muir, D., Ortiz-Santaliestra, M.E., Paul, K.B., Rico, A., Rodea-Palomares, I., Römbke, J., Rydberg, T., Segner, H., Smit, M., van Gestel, C.A.M., Vighi, M., Werner, I., Zimmer, E.I., van Wensem, J. 2018. Towards Sustainable Environmental Quality: Priority Research Questions for Europe. Environment Toxicology and Chemistry 37: 2281-2295.

28.  Pohl, J. Björlenius, B., Carlsson, G., Fick, J., Norrgren, L., Örn, S., Brodin, T., Larsson, D.G.J. 2018. Effects on reproduction and behavior in zebrafish (Danio rerio) exposed to an ozonated sewage effluent. Aquatic Toxicology 200: 93-101.

29.  Wang, H., Sikora, P., Rutgersson, C., Lind, M., Brodin, T., Björlenius B., Larsson, J., Norder, H. 2018. Differential removal of human pathogenic viruses from sewage by conventional and ozone treatments. International Journal of Hygiene and Environmental Health 221: 479-488.

30.  Lagesson, A., Brodin, T., Fahlman, J., Fick, J., Jonsson, M., Persson, J., Byström, P. Klaminder, J. 2018. No evidence of increased growth or mortality in fish exposed to oxazepam in semi-natural ecosystems. Science of the Total Environment 615: 608-614.

31.  Saaristo M. Brodin T. et al. 2018. Direct and indirect effects of chemical contaminants on the behaviour, ecology and evolution of wildlife. Proceedings of the Royal Society B: Biological Sciences 285: 20181297.                                                                     

32.  Rodriguez, A., Zhang, H., Klaminder, J., Brodin, T., Andersson, M. 2017. ToxId: an efficient algorithm to solve occlusions when tracking multiple animals. Scientific Reports 7: 14774.

33.  Brodin, T., Nordling, J., Lagesson, A., Klaminder, J., Hellström, G., Christensen, B., Fick, J. 2017. Environmental relevant levels of a benzodiazepine (oxazepam) alters important behavioural traits in a common planktivorous fish, (Rutilus rutilus), Journal of Toxicology and Environmental Health, Part A, DOI:10.1080/15287394.2017.1352214

34.  Cote, J., Brodin, T., Fogarty, S., Sih, A. 2017. Non‐random dispersal mediates invader impacts on the invertebrate community. Journal of Animal Ecology 86: 1298-1307.

35.  McCallum, E.S., Krutzelmann, E., Brodin, T., Fick, J., Sundelin, A., Balshine, S. 2017. Exposure to wastewater effluent affects fish behaviour and tissue-specific uptake of pharmaceuticals. Science of the Total Environment 605–606: 578–588.

36.  Fick J, Brodin T, M. Heynen, J. Klaminder, M. Jonsson, K. Grabicova, T. Randák, R. Grabic, J. Slobodnik, A. Sweetman, M. Earnshaw, R. Loos. 2017. EU-wide monitoring survey of anxiolytics in surface water. Chemosphere 176: 324-332.

37.  Samson, E., Hirsch, P., Palmer, S., Behrens, J., Brodin, T., Travis, J. 2017. Early Engagement of Stakeholders with Individual-Based Modeling Can Inform Research for Improving Invasive Species Management: The Round Goby as a Case Study. Frontiers in Ecology and Evolution 5: 149.

38.  Johansson, F., Brodin, T., Stracevicius, D., Fürstenberg-Hägg, E. 2017. Shift in migration phenology of a wintering population of Dippers Cinclus cinclus in northern Sweden. Ornis Svecica 27: 57–63.

39.  Rodriguez, A., Zhang, H., Wiklund, K., Brodin, T., Klaminder, J., Andersson, P., Andersson, M. 2017. Refining particle positions using circular symmetry. PLoS ONE 12: e0175015.                                                                               

40.  Hirsch, P., Thorlacius, M., Brodin, T., Burkhardt-Holm, P. 2016. An approach to incorporate individual personality in modelling fish dispersal across in-stream barriers. Ecology and Evolution 7: 720–732.

41.  Klaminder, J., Hellström, G., Fahlman, J., Jonsson, M., Fick, J., Lagesson, A., Bergman, E., Brodin, T. 2016. Drug-Induced Behavioral Changes: Using Laboratory Observations to Predict Field Observations. Frontiers Environmental Science 4:81. doi: 10.3389/fenvs.2016.00081

42.  Hellström, G., Klaminder, J., Finn, F., Persson, L., Alanärä, A., Jonsson, M., Fick, J., Brodin, T. 2016. GABAergic anxiolytic drug in water increases migration behaviour in salmon. Nature Commuication. 7, 13460 doi: 10.1038/ncomms13460.

43.  Heynen, M., Backström T., Fick J., Jonsson M., Klaminder J. & Brodin T. 2016. Home alone—The effects of isolation on uptake of a pharmaceutical contaminant in a social fish. Aquatic Toxicology 180: 71-77.

44.  Lagesson, A., Fahlman, J., Brodin, T., Fick, J., Jonsson, M., Persson, J., Byström, P. Klaminder, J. Bioaccumulation of five pharmaceuticals at multiple trophic levels in an aquatic food web - Insights from a field experiment. Science of the Total Environment 568: 208-215.

45.  Heynen, M., Fick J., Jonsson M., Klaminder J. & Brodin T. 2016. Effect of bioconcentration and trophic transfer on realized exposure to oxazepam in two predators, the dragonfly larvae (Aeshna grandis) and the Eurasian perch (Perca fluviatilis). Environmental Toxicology and Chemistry (DOI: 10.1002/etc.3368).

46.  Hellström, G., Klaminder J., Jonsson, M., Fick J. & Brodin T. 2016. Upscaling behavioural studies to the field using acoustic telemetry. Aquatic Toxicology 170: 384-389.

47.  Heynen, M., Brodin T., Jonsson M., Klaminder J. & Fick J. 2016. Tissue-specific uptake of the benzodiazepine oxazepam in adult Eurasian perch (Perca fluviatilis). Environmental Chemistry (accepted).

48.  Drotz, M.K., Brodin, T. & Nilsson, A.N. 2015. Changing names with changed address: Integrated taxonomy and species delimitation in the Holarctic Colymbetes paykulli group (Coleoptera: Dytiscidae). PLoS ONE 10(11): e0143577. doi:10.1371/journal.pone.0143577.

49.  Klaminder, J., Brodin, T., Sundelin, A., Anderson, N.J., Fahlman, J., Jonsson, M. & Fick, J. 2015. Long-Term Persistence of an Anxiolytic Drug (Oxazepam) in a Large Freshwater Lake. Environmental Science and Technology 49: 10406-10412.

50.  Eggermont, H., Balian, E., Azevedo, J., Beumer, V., Brodin, T. et al. 2015. Nature-based solutions: framing an emerging term that may deeply influence future environmental management and research in Europe. GAIA - Ecological Perspectives for Science and Society 24(4): 243-248.

51.  Jonsson, M., Ershammar, E., Fick, J., Brodin, T. & Klaminder J. 2015. Effects of an antihistamine on carbon and nutrient recycling in streams. Science of the Total Environment 538: 240-245.

52.  Thorlacius, M., Hellström, G. & Brodin, T. 2015. Behavioral dependent dispersal in the invasive Round goby (Neogobius melanostomus) depends on population age. Current Zoology 61: 529-542.                              

53.  Brodin, T., Heynen, M., Fick, J., Klaminder, J., Piovano, S. & Jonsson, M. 2014. Inconspicuous effects of pharmaceuticals in aquatic systems – ecological impacts through behavioural modifications at dilute concentrations. Philosophical Transactions of the Royal Society B 369: 20130580.

54.  Klaminder, J., Jonsson, M., Fick, J., Sundelin A. & Brodin T. 2014. The conceptual imperfection of aquatic risk assessment tests: highlighting the need for tests designed to detect therapeutic effects of pharmaceutical contaminants. Environmental Research Letters 9: 084003. 

55.  Jonsson, M., Fick, J., Klaminder, J. & Brodin, T. 2014. Antihistamines and aquatic insects: bioaccumulation and impacts on behavior in damselfly larvae (Zygoptera). Science of the Total Environment 472: 108-111.

56.  Brodin, T. & Drotz, M.K. 2014. Individual variation in dispersal associated behavioural traits of the invasive Chinese mittencrab (Eriocheir sinensis, H. Milne Edwards, 1854) in Lake Vänern. Current Zoology 60: 410–416.

57.  Brodin, T., Fick, J., Jonsson, M. & Klaminder, J. 2013. Dilute Concentrations of a Psychiatric Drug Alter Behavior of Fish from Natural Populations. SCIENCE 339: 814-815.

58.  Cote, J., Fogarty S., Tymen B., Sih A. & Brodin, T. 2013. Personality-dependent dispersal cancelled under predation risk. Proceedings of the Royal Society B – 280: 20132349.

59.  Brodin, T., Lind, M.I., Wiberg, M.K. & Johansson, F. 2013.  Personality trait differences between mainland and island populations in the common frog (Rana temporaria). Behavioural Ecology and Sociobiology 67: 135-143.

60.  Drotz, M.K., Brodin, T., Saura, A. & Giles, B.E. 2012. Ecotype differentiation in the face of gene flow within the diving beetle Agabus bipustulatus (Linnaeus, 1767) in northern Scandinavia. PLoS ONE 7(2): e31381. doi:10.1371/journal.pone.0031381.

61.  Hammond, J., Luttbeg, B., Brodin, T. & Sih, A. 2012. Spatial scale influences the outcome of the predator-prey space race between tadpoles and predatory dragonflies. Functional Ecology 26: 522-531. doi: 10.1111/j.1365-2435.2011.01949.x. 

62.  Drotz, M.K., Brodin, T. & Berggren, M. 2012. Distribution patterns of the Swedish Chinese Mitten Crab (Eriocheir sinensis, H. Milne Edwards, 1853) in Lake Vänern. Aquatic Invasions 7: 243-249.                      

63.  Cote, J., Fogarty S. Brodin, T., Smith, K.L. & Sih, A. 2011. Personality-dependent dispersal in the invasive mosquitofish: group composition matters. Proceedings of the Royal Society B, 278: 1670-1678.

64.  Cote, J., Fogarty, S., Smith, K., Brodin, T. & Sih, A. 2010. Personality traits and dispersal tendency in the invasive mosquitofish (Gambusia affinis). Proceedings of the Royal Society B 277:1571-1579. – Highlighted in NATURE 463: 273 doi:10.1038/463273a.

65.  Conrad, L., Smith, K.L., Brodin, T., Saltz, J. & Sih, A. 2011. Behavioural syndromes in fish – a review with implications for ecology and fisheries management. Journal of Fish Biology 78: 395-435.

66.  Brodin, T. & Drotz, M.K. 2011. Larval behavioral syndrome does not affect emergence behavior in a damselfly (Lestes congener). Journal of Ethology 29: 107-113.                  

67.  Cote, J., Clobert, J., Brodin, T., Fogarty, S. & Sih, A. 2010. Personality-dependent dispersal: characterization, ontogeny and consequences for spatially structured populations. Philosophical Transactions of the Royal Society B, 365: 4065-4076. 

68.  Johansson, F., Sniegula, S. & Brodin, T. 2010. Emergence patterns and latitudinal adaptations in development time of dragonflies (Odonata) in north Sweden and Poland. Odonatologica 39: 97-106. 

69.  McCauley, S., Brodin, T. & Hammond, J. 2010. Foraging rates of larval dragonfly colonists are positively related to habitat isolation: results from a landscape-level experiment. American Naturalist 175: E66–E73. doi: 10.1086/650444s. 

70. Drotz, M.K., Brodin, T. & Nilsson, A.N. 2010. Multiple Origins of Elytral Reticulation Modifications in the West Palearctic Agabus bipustulatus Complex (Coleoptera, Dytiscidae). PLoS ONE 5: e9034. doi:10.1371/journal.pone.0009034.