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PHD STUDENTS
Mehrnaz Asadnejad
PhD student at University of Gothenburg, 2026-2031.
(supervised by Lotta Kvarnemo)
Mehrnaz' PhD project focuses on evolution of female ornamentation. She will use modelling to explore it in general terms, together with Prof. Suzanne Alonzo, Univ of California Santa Cruz. Empirically, she will use the ornamented pipefish Nerophis ophidion as her model organism. To better understand the skin physiology of the ornament she will use light and electron microscopy, in collaboration with Docent Helen Sköld, Havets hus/University of Gothenburg. Finally, to investigate how costs and benefits of the ornament may change with female age and size, she will investigate that experimentally, together with Lotta Kvarnemo, Univ of Gothenburg and Prof. Gunilla Rosenqvist, Uppsala Univ. Prof. Staffan Andersson, Univ of Gothenburg, will be her internal co-supervisor.
Johanna Yliportimo
PhD student at Åbo Akademi University, Finland, 2019-2027.
(originally supervised by Kai Lindström; recently by Markus Öst, Katja Anttila and Lotta Kvarnemo)
Johanna Yliportimo is pursuing a PhD thesis on topics related to resource availability (such as food and nest sites) and female competition in sand gobies. Her work is focused on field-based experiments, set up in locations that differ naturally in food and/or nest site availability. She is combining these studies with measurements of carotenoid concentration in female gonads and eggs spawned in nests, as well as female expression of vitellogenin in liver. Overall her work is placed in the scientific "landscape" of operational sex ratio and sexual selection theory.
CURRENT AND FORMER MASTER & BACHELOR STUDENTS
Joana Cocoynacq
Intern and Master student from University of Lille, France, 2026-2027
(supervised by Lotta Kvarnemo & Helen Sköld)
Joana's project as an intern is on female ornamentation in Nerophis ophidion. She uses photos to investigate how the blue ornament of this pipefish developes before the breeding season starts, and how another ornament (skin folds) correlates with fecundity. For her Master's project, she collects data on the information value of the ornament by comparing females before (many mature eggs) and after mating (no mature eggs).
Beatriz Taboada
Master student, 60 hp, 2023-2024
(supervised by Lotta Kvarnemo & Gunilla Rosenqvist)
Pipefish females of Nerophis ophidion have a shiny blue ornament that males are lacking. Beatriz tackled some hard questions about the evolution of female ornamentation, related to differences between populations in terms of ornament costs, fecundity and condition. This excellent Master thesis has been submitted to a scientific journal.
Alice Anderstedt
Bachelor student, 2023
(supervised by Lotta Kvarnemo & Donald Blomqvist)
Alice investigated if olfaction correlates with on mating systems in birds, based on data from published studies. Mating system was measured as the proportion of broods with mixed paternity. She used three measures of olfactory ability: olfactory bulb diameter, olfactory bulb volume, and total number of olfactory receptor genes. Her study showed a negative correleation between mixed paternity and olfactory bulb diameter, indicating that mating patterns in birds may indeed be influenced by olfactory ability. Her study is published in Ecology and Evolution 2025: LINK .
Ayhushi Mahajan
Master student, India, 2022-2023
(supervised by Lotta Kvarnemo)
Sand goby females show a conspicuous dark pigmentation around the eyes (a "dark-eye display"). Manipulating adult sex ratio in experimental tanks, Ayushi has studied the potential effect of sex ratio on dark-eye displays. Her work helped us reach a better understanding of this enigmatic female ornamentation. Her study is published in Animal Behaviour 2025: LINK .
Lisa Carlsson & Maja Lagerqvist
Bachelor students, 30 hp, 2022
(Lisa supervised by Lotta Kvarnemo, Leon Green & Ola Svensson; Maja by Leon Green)
Round gobies are spreading into higher salinities along the Swedish westcoast, and as a consequence, there is increasing the overlap in range between round gobies and black grobies in this area. Lisa's and Maja's Bachelor theses focused on questions related to that overlap and potential competition between the species for preferred habitat, food and shelters. While Lisa worked in the lab, Maja's work was entirely field based. The overall result showed us that round gobies are (surprisingly) timid in comparison to the more aggressive black gobies. These interesting results will soon be written up and hopefully published in a good journal.
Anna Waffender
Erasmus student, Germany, 2022
(supervised by Lotta Kvarnemo & Ingrid Ahnesjö in Sweden, and Jonathan Henshaw in Germany)
Females of the broad-nosed pipefish show a B-shaped pattern along the upper body, an ornament that they can quickly turn on and off. Anna studied the effect of adult sex ratio on these female displays, by manipulating the sex ratio in experimental tanks and video recording behavioural interactions. Her work has helped us gain a better understanding of the function of this female ornament. Her study is published in Behavioral Ecology and Sociobiology 2024: LINK .
Petter Gustafsson
Bachelor student, 30 hp, 2021 (supervised by Lotta Kvarnemo, Ola Svensson & Eva-Lotta Blom)
Petter's Bachelor project focuses on noise in the sea and how it may affect reproductive behavior and mating success of sand gobies, breeding in shallow water. Male sand gobies make a sound during courtship, that appears to be important for their mating success. We therefore expected them to be It is sensitive to added noise, a question that Petter tested experimentally working in a bay close to Kristineberg Marine Research Center. Surprisingly, the noise created in his massive "noise park" had no effect on male nest building, female visits, number of spawnings, or number of invertebrates seen. Petter's work is about to be written up, and will hopefully published before long.
Martina Gríful
Erasmus student, Spain, 2021
(supervised by Leon Green & Lotta Kvarnemo)
The invasive fish round goby is common in the brackish waters of Gothenburg harbour. It is now expanding into coastal waters with higher salinity. To find out how far they are likely to spread, Leon has tested if round gobies caught at the expansion front are able to reproduce in 20 PSU (similar to where they were caught) and 30 PSU (fully marine water found along the Swedish west coast). Martina's Bachelor project focused on testing sperm swimming speed in these two salinities and sperm morphology after the fish had been acclimated for 2.5 months. She also compared egg development of clutches spawned in 20 and 30 PSU. This study is published published in Biological Invasions 2024: LINK. Another study on sperm traits in sneakers and breeding coloured sand gobies, that Martina was also part of, is published in Evolutionary Applications 2023: LINK
Felix Englund Örn
Master student 60 hp, 2016-2017 (supervised by Leon Green & Lotta Kvarnemo)
Andrew Holmes
Master student 60 hp 2016-2017 (supervised by Leon Green & Lotta Kvarnemo)
Round goby found in the Gothenburg harbour were presumably brought there as larvae by ballast water. The rivermouth of Göta Älv and offers a range of salinities for newly arrived fish. It is also likely that new larvae are brought repeatedly to this large harbour from multiple origins that differ in salinity.
Felix' project aimed to test if round goby individuals might sort themselves in this salinity gradient according to genetic origin. The project involved GBS (gene-by-sequencing) to test (a) if fish caught upstream in low salinity differ genetically from those caught down-stream in high salinity and (b) if their genetic profile can be matched to fish from other large harbours.
Andy's project overlapped with Felix' in that it focuses on round gobies caught at high versus low salinity sites in the Göta Älv river mouth. Andy's focus was to investigate if these fish differ in salinity tolerance, measured e.g. as metabolism when exposed to different salinities. He also looked at feeding preferences (stomach content) and size and age profiles of fish from the different sites.
Andy's and Felix' nice work is published in Evolutionary Applications 2023: LINK
Malin Nygård
Master student 30 hp
(supervised by Lotta Kvarnemo, Ingrid Ahnesjö & Ines Braga Goncalves)
Males of the broad nosed pipefish (Syngnathus typhle) provide care to the offspring in a brood pouch. Earlier work by Ines Braga Goncalves (see below) on the evolution of egg size and paternal care in pipefishes, has investigated oxygenation of the eggs inside the brood pouch. Malin's project, which builds on that knowledge and techniques, tests the effect of male size, egg size and egg density for the oxygen levels inside the brood pouch during embryo development. Malin's work is published in Behavioral Ecology 2019: LINK
It is also featured in a Research Outreach paper: https://researchoutreach.org/articles/pipefish-male-pregnancy-why-females-prefer-large-mates/
Aurora García-Berro
Erasmus student, Spain
(supervised by Lotta Kvarnemo)
The Swedish west coast differs from the Baltic sea in many ways that are expected to affect the investment into reproductive traits of fish living in the two habitats. Not only does the salinity differ, but also food abundance. Focusing on sand gobies, Aurora García-Berro investigated investment of females into ovaries, comparing fish from the Swedish west coast to fish from the Baltic sea. Her fine work is published in Royal Society Open Science, 2019: LINK
Susanne Andersson
Bachelor student, 15 hp
(supervised by Lotta Kvarnemo)
In previous work on the West Australian seahorse (Hippocampus subelongatus) (Kvarnemo et al 2007) we collected home range data for males and females of known mating and pair bond status. Susanne Andersson's Bachelor thesis focused on analyzing this data and testing e.g. if home range size, distances moved and overlap with other individuals differ between paired and unpaired male and female seahorses. Her work is published in Journal of Ethology, 2021: LINK
FORMER PHD STUDENTS
Leon Green | PhD student in Animal Ecology (2015-2020; Dept of Biological and Environmental Sciences, University of Gothenburg)
ResearchGate profile | University web page | Academia.edu
Eva-Lotta Blom | PhD student in Animal Ecology (2013-2017; Dept of Biological and Environmental Sciences, University of Gothenburg)
(co-supervised with Ola Svensson as main supervisor)
ResearchGate profile | Google Scholar profile | University web page
Gry Sagebakken | PhD student in Animal Ecology (2006-2012; Dept of Zoology, University of Gothenburg)
Inês Braga Gonçalves | PhD student in Animal Ecology (2006-2010; Dept of Zoology, University of Gothenburg)
Maria Prager | PhD student in Animal Ecology (2005-2010; Dept of Zoology, University of Gothenburg)
Under my supervision 2007-2010
Thesis: Phylogeny and Signal Diversity in Widowbirds and Bishops (Euplectes spp.)
Maria Norevik (formerly Lissåker) | PhD student in Animal Ecology (2002-2007; Dept of Zoology, Stockholm University)
Thesis: Paternal care, filial cannibalism and sexual conflict in the sand goby, Pomatoschistus minutus
Helena Strömberg | PhD student in Animal Ecology (1999-2004; Dept of Zoology, Stockholm University)
Under my supervision 2003-2004.
Thesis: Benthic cryptofauna and internal bioeroders on coral reefs
Ola Svensson | PhD student in Animal Ecology (2000-2004; Dept of Zoology, Stockholm University)
Thesis: Sexual selection in Pomatoschistus – nests, sperm competition, and paternal care
MEMBER OF SUPERVISORY COMMITTEES
Hanne Løvlie | PhD student in Ethology (2003-2008; Dept of Zoology, Stockholm University)
Thesis: Pre- and post-copulatory sexual selection in the fowl, Gallus gallus
Google Scholar Profile | ResearchGate | Academia.edu
Supervisors: Tom Pizzari & Sven Jakobsson
Helena Elofsson | PhD student in Functional Morphology, (2000-2005; Dept of Zoology, Stockholm University)
Thesis: Sperm motility in Gasterosteiform fishes: The role of salinity and ovarian fluid
Supervisor: Bertil Borg
Malin-Ah King | PhD student in Animal Ecology (1999-2003; Dept of Zoology, Stockholm University)
Thesis: Phylogenetic analyses of parental care evolution
Supervisor: Birgitta S. Tullberg
PREVIOUS MASTER AND BACHELOR STUDENTS
Beatriz Taboada (2024): Geographical variation of the coloration in the pipefish Nerophis ophidion (Master thesis, Univ of Gothenburg, 60 hp).
Alice Anderstedt (2023): Olfaction and paternity in birds - A correlation analysis of olfaction and multiple paternity (Bachelor thesis, Univ of Gothenburg, 15 hp). Supervised with Donald Blomqvist.
Petter Gustafsson (2023): Antropogena ljudföroreningars påverkan på liv under ytan. Bachelor thesis, Univ. of Gothenbrug, 30 hp). Supervised with Ola Svensson and Eva-Lotta Blom.
Ayushi Mahajan (2023): ‘Impact of varying adult sex ratio on temporary female ornamentation in Pomatoschistus minutus’ (Master thesis, Indian Institute of Science Education and Research, India). Main supervisor.
Lisa Carlsson (2022): ‘Black and round gobies: An experimental study of habitat choice and resource competition’ (Bachelor thesis, Univ of Gothenburg, 30 hp). Supervised with Ola Svensson and Leon Green.
Anna Waffender (2022): ‘Female ornamentation under biased adult sex ratios in the broad-nosed pipefish ’(Master thesis, University of Freiburg, Germany). Supervised with Jonathan Henshaw and Ingrid Ahnesjö.
Martina Griful (2021): ‘Reproductive dysfunction in seawater provides a barrier to the expansion of Neogobius melanostomus into the North Sea’. (Bachelor thesis, Erasmus student, Univ of Barcelona, Spain). Supervised with Leon Green.
Isabelle Dehkla (2018): ‘Effects of anthropogenic noise on Pomatoschistus microps’ (Master thesis, Univ of Gothenburg, 45 hp). Supervised with Eva-Lotta Blom.
Andrew Holmes (2017): ‘The effect of salinity on the round goby (Neogobius melanostomus)’ (Master thesis, Univ of Gothenburg, 60 hp). Supervised with Leon Green.
Felix Englund Örn (2017): ‘A look into salinity tolerance of the round goby: A tale of two salinities’. (Master thesis, Univ of Gothenburg, 60 hp). Supervised with Leon Green.
Susanne Andersson (2017): ‘Ensam sjöhäst söker? En undersökning av sjöhästars rörelsemönster’. (Bachelor thesis, Univ of Gothenburg, 15 hp).
Malin Nygård (2016): ‘Does egg size affect the oxygen levels in the male's brood pouch?’ (Master thesis, 30 hp). Supervised with Ingrid Ahnesjö and Inês Braga Goncalves.
Aurora García-Berro Navarro (2016). 'Reproductive allocation in competing females and parental males across environmental conditions in a marine fish'. Erasmus - Bachelor Student, University of Barcelona, Spain.
Sofie Schöld (2015): ‘Effects of salinity on reproduction in the sand goby (Pomatoschistus minutus)’. MSc Marine Science, University of Gothenburg, 45 hp
Jessica Åsbacka (2015): ‘The influence of salinity on behaviour and reproduction in the sand goby (Pomatoschistus minutes)’. MSc in Environmental Biology, Åbo Akademi University, 55 p
Sandra Johansson (2014): ‘Why do female sand gobies display black eyes during mating?’ BSc in Marine Science, University of Gothenburg, 15 hp
Emma Martinsson (2013): ‘Behaviour of brooding male pipefish in different oxygen conditions’. BSc in Animal Ecology, University of Gothenburg, 15 hp
Sara Sundström (2012): ‘Utveckling av pedagogiska skyltar på Borås djurpark’. Application work in Animal Ecology, University of Gothenburg, 15 hp
Johannes Björk (2011): 'Mechanisms behind sizeassortative nest choice in absence of intrasexual competition, in males of the sand goby Pomatoschistus minutes'. MSc in Animal Ecology, University of Gothenburg, 60 hp
Karin Olsson (2008). ‘An ecological niche? A study on nest material preference in sand gobies (Pomatoschistus minutus).’ Graduate work in Animal Ecology, University of Gothenburg, 15 hp
Josefin Sundin (2008): ‘Does male condition affect female choice in the sex-role reversed pipefish Syngnathus typhle?’. MSc in Animal Ecology, University of Gothenburg, 30 hp
Karin Olsson (2007): ‘Relative cost of courtship behaviour in nest-building sand gobies’. MSc in Animal Ecology, University of Gothenburg, 30 hp
Therese Stenlund (2006): ‘Effects on parental care in sand goby, Pomatoschistus minutus, caused by an egg predator and different oxygen levels’. MSc in Animal Ecology, University of Gothenburg, 30 hp
Anna Singer (2003): ‘Does nest site availability influence sneaking rate in the sand goby Pomatoschistus minutus?’. Graduate work in Animal Ecology, Stockholm University, 30 p
Andreas Nyman (2001-2002): ‘Intensitet i parningskonkurrens hos sandstubb i relation till äggtäckningsgrad i boet’. Graduate work in Animal Ecology, Stockholm University, 20 p
Maria Lissåker (2001): ‘Effects of a low oxygen environment on parental effort and filial cannibalism in the male sand goby, Pomatoschistus minutus’. Graduate work in Animal Ecology, Stockholm University, 20 p
Glenn Moore (1997-1999): ‘Mating patterns and sexual selection in the Western Australian Seahorse Hippocampus subelongatus’. MSc in Zoology, University of Western Australia
Ola Svensson (1995): ‘Parental behaviour depending on paternity assurance and food availability in the common goby Pomatoschistus microps’. Graduate work in Animal Ecology, Uppsala University, 20 p
Anna Karlsson (1993): ‘Why do female sand gobies (Pomatoschistus minutus) prefer males with eggs in their nests?’ Graduate work in Animal Ecology, Uppsala Univeristy, 20 p
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