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  • 1.
    Corral-López, Alberto
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Eckerström-Liedholm, Simon
    Stockholm University, Faculty of Science, Department of Zoology.
    Van der Bijl, Wouter
    Stockholm University, Faculty of Science, Department of Zoology.
    Kotrschal, Alexander
    Stockholm University, Faculty of Science, Department of Zoology.
    Kolm, Niclas
    Stockholm University, Faculty of Science, Department of Zoology.
    No association between brain size and male sexual behavior in the guppy2015In: Current Zoology, ISSN 1674-5507, Vol. 61, no 2, p. 265-273Article in journal (Refereed)
    Abstract [en]

    Animal behavior is remarkably variable at all taxonomic levels. Over the last decades, research on animal behavior has focused on understanding ultimate processes. Yet, it has progressively become more evident that to fully understand behavioral variation, ultimate explanations need to be complemented with proximate ones. In particular, the mechanisms generating variation in sexual behavior remain an open question. Variation in aspects of brain morphology has been suggested as a plausible mechanism underlying this variation. However, our knowledge of this potential association is based almost exclusively on comparative analyses. Experimental studies are needed to establish causality and bridge the gap between micro-and macroevolutionary mechanisms concerning the link between brain and sexual behavior. We used male guppies that had been artificially selected for large or small relative brain size to study this association. We paired males with females and scored the full known set of male and female sexual behaviors described in guppies. We found several previously demonstrated associations between male traits, male behavior and female behavior. Females responded more strongly towards males that courted more and males with more orange coloration. Also, larger males and males with less conspicuous coloration attempted more coerced copulations. However, courting, frequency of coerced copulation attempts, total intensity of sexual behavior, and female response did not differ between large-and small-brained males. Our data suggest that relative brain size is an unlikely mechanism underlying variation in sexual behavior of the male guppy. We discuss these findings in the context of the conditions under which relative brain size might affect male sexual behavior

  • 2.
    Eckerström-Liedholm, Simon
    Stockholm University, Faculty of Science, Department of Zoology.
    Behavioural, physiological and morphological correlates of life-history in killifishes − a macroevolutionary approach2019Doctoral thesis, comprehensive summary (Other academic)
    Abstract [en]

    Life-histories commonly evolve along a continuum from short-lived and fecund, to long-lived and less fecund. Because life-history traits are mostly components of reproduction and survival, understanding the causes and consequences of life-history variation is at the core of evolutionary biology. This thesis aims to identify what other key traits (e.g. behavioural, physiological and morphological traits) covary with life-history, and why. Numerous hypotheses describe how life-history might be associated with other traits, with life-history trade-offs often considered to be a primary driver of any such relationships. For example, since resources are limited, increased investment in one trait must lead to decreased investment in one or several other traits, all else equal. Hypotheses on the relationship between life-history and other traits have been tested in many studies, but empirical studies in controlled experimental settings are rare. In this thesis I explore how behaviour, physiology and morphology relate to variation along the life-history continuum from fast to slow, in a system with substantial variation in life-history traits - the killifishes.

    I began by exploring the patterns of egg to body size allometry in killifishes (Paper I), where species with faster life-histories showed indications of constraints on the independent evolution of egg size and body size. Furthermore, I found evidence of differences in variance and in the rates of evolution of egg size and body size across species, potentially caused by the colonisation of ephemeral habitats, which could have selected for adaptations that lead to differences in size.

    I then performed a comparative common garden study (Paper II) of the pace-of-life syndrome hypothesis, which predicts that species with fast life-histories should take larger risks in order to maintain their increased reproductive rate. I obtained data on risk taking behaviours, including movement, tendency to enter an open environment, and aggressiveness, in addition to metabolic rate, for 20 species of killifish, with multiple replicates per species. The results indicated trait dependent associations with life-history, where aggression seemed to correlate positively with speed of life-history, in congruence with our prediction.

    Next, my colleagues and I assessed the association between life-history and sexual selection (Paper III), in order to determine if investment in secondary sexual traits might be traded off against survival in killifish. Fin size was found to be negatively associated with escape performance in a simulated predator attack, suggesting survival costs for individuals with large fins. Importantly, fin size was also positively associated with the speed of life-history, supporting the hypothesis that costs to survival probability is lower in fast-living species.

    Lastly, I tested the hypothesized negative covariation between relative brain size and speed of life-history, by collecting and analysing brain size measurements for 21 species of killifish (Paper IV). Surprisingly, a positive relationship between speed of life-history and relative brain size was found for adults, although juveniles did not differ in relative brain size. This implies at least one of two things: either there is no need to trade off brain size with life-history since resource acquisition is higher, or brain size and life-history are traded-off with other traits.

    In conclusion, I show that previously found trade-offs between life-history and investment in other costly traits are only sometimes present, when tested in a system with substantial divergences in the speed of life-history. I also provide evidence for a trait dependent association between life-history and among species differences in risk-taking and metabolic rate.

  • 3.
    Eckerström-Liedholm, Simon
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Stockholm University, Faculty of Science, Department of Zoology.
    Gonzalez-Voyer, Alejandro
    Stockholm University, Faculty of Science, Department of Zoology. Universidad Nacional Autónoma de México, Mexico.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Time-limited environments affect the evolution of egg-body size allometry2017In: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 71, no 7, p. 1900-1910Article in journal (Refereed)
    Abstract [en]

    Initial offspring size is a fundamental component of absolute growth rate, where large offspring will reach a given adult body size faster than smaller offspring. Yet, our knowledge regarding the coevolution between offspring and adult size is limited. In time-constrained environments, organisms need to reproduce at a high rate and reach a reproductive size quickly. To rapidly attain a large adult body size, we hypothesize that, in seasonal habitats, large species are bound to having a large initial size, and consequently, the evolution of egg size will be tightly matched to that of body size, compared to less time-limited systems. We tested this hypothesis in killifishes, and found a significantly steeper allometric relationship between egg and body sizes in annual, compared to nonannual species. We also found higher rates of evolution of egg and body size in annual compared to nonannual species. Our results suggest that time-constrained environments impose strong selection on rapidly reaching a species-specific body size, and reproduce at a high rate, which in turn imposes constraints on the evolution of egg sizes. In combination, these distinct selection pressures result in different relationships between egg and body size among species in time-constrained versus permanent habitats.

  • 4.
    Eckerström-Liedholm, Simon
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Kotrschal, Alexander
    Näslund, Joacim
    Stockholm University, Faculty of Science, Department of Zoology.
    Rowiński, Piotr
    Stockholm University, Faculty of Science, Department of Zoology.
    Gonzalez-Voyer, Alejandro
    Stockholm University, Faculty of Science, Department of Zoology.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Fast life-histories are associated with larger brain size in killifishesManuscript (preprint) (Other academic)
    Abstract [en]

    Comparative studies suggest a negative relationship between pace of life-history, and relative energetic investment into brain size. However, since brain size typically evolves as a correlated response to selection on body size, any lag in brain size evolution will result in a shift in relative brain size (e.g. small body – large relative brain size).Coevolution between body size and life-history hence has the potential to drive secondary associations between relative brain size and life-history, when body size is correlated with life history. However, as far as we know, the relationship between relative brain size and life-history strategy has not been examined in systems that simultaneously present marked contrasts in life-history but no concordant shifts in body size. Using a common garden approach, we test the association between relative brain size and life-history in 21 species of killifish; a study system that fulfils the aforementioned requirements. Contrary to the prediction that brain size evolves through energetic trade-offs with life-history, we found that adults, but not juveniles, of fast-living species had larger relative brain sizes. Rather than an energetic link to life-history, our results suggest that fast- and slow-living species differ in terms of how cognitively demanding environments they inhabit are, or alternatively in the ontogenetic timing of somatic vs. neural growth.

  • 5.
    Eckerström-Liedholm, Simon
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Stockholm University, Faculty of Science, Department of Zoology.
    Morozov, Sergey
    van der Bijl, Wouter
    Rowiński, Piotr
    Stockholm University, Faculty of Science, Department of Zoology.
    Gonzalez-Voyer, Alejandro
    Stockholm University, Faculty of Science, Department of Zoology.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Macroevolutionary evidence suggests trait-dependent coevolution between behaviour and life-historyManuscript (preprint) (Other academic)
    Abstract [en]

    Species with fast life-histories prioritize current over future reproduction, which ought to require greater energetic resources, but also results in a shorter time-period to realize their reproductive potential, compared to slow life-histories, which prioritize future reproduction. Hence, behaviours that increase access to both resources and mating opportunities, at a cost of increased mortality risk, are thought to coevolve with the pace of life-history. However, whether this prediction holds across species, is yet to be tested under standardized conditions. Here, we test how potentially risky behaviours, which facilitate access to resources and mating opportunities (i.e. activity, boldness and aggression), along with metabolic rate, correlates with the pace of life-history across 20 species of killifish, which present a remarkable divergence in the pace of their life-histories. We found a positive correlation between the pace of life-history and aggression, but not with any other behavioural traits or metabolic rate. Aggression is often expressed in the context of mating, while the other behaviours we measured might be more relevant for access to energetic resources. Our results therefore suggest that the trade-off between current and future reproduction plays a more prominent role in shaping mating behaviour, while behaviours related to acquisition of energetic resources may be more affected by ecological factors.

  • 6.
    Eckerström-Liedholm, Simon
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Stockholm University, Faculty of Science, Department of Zoology.
    Morozov, Sergey
    van der Bijl, Wouter
    Stockholm University, Faculty of Science, Department of Zoology. University of British Columbia, Canada.
    Rowiński, Piotr K.
    Stockholm University, Faculty of Science, Department of Zoology.
    Gonzalez-Voyer, Alejandro
    Stockholm University, Faculty of Science, Department of Zoology. Universidad Nacional Autónoma de México, Mexico.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Macroevolutionary evidence suggests trait-dependent coevolution between behavior and life-history2019In: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 73, no 11, p. 2312-2323Article in journal (Refereed)
    Abstract [en]

    Species with fast life-histories typically prioritize current over future reproductive events, compared to species with slow life-histories. These species therefore require greater energetic input into reproduction, and also likely have less time to realize their reproductive potential. Hence, behaviors that increase access to both resources and mating opportunities, at a cost of increased mortality risk, could coevolve with the pace of life-history. However, whether this prediction holds across species, remains untested under standardized conditions. Here, we test how risky behaviors, which facilitate access to resources and mating opportunities (i.e., activity, boldness, and aggression), along with metabolic rate, coevolve with the pace of life-history across 20 species of killifish that present remarkable divergences in the pace of life-history. We found a positive association between the pace of life-history and aggression, but interestingly not with other behavioral traits or metabolic rate. Aggression is linked to interference competition, and in killifishes is often employed to secure mates, while activity and boldness are more relevant for exploiting energetic resources. Our results suggest that the trade-off between current and future reproduction plays a more prominent role in shaping mating behavior, while behaviors related to energy acquisition may be influenced by ecological factors.

  • 7.
    Rowiński, Piotr
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Laurila, Anssi
    Gotthard, Karl
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Stockholm University, Faculty of Science, Department of Zoology.
    Lind, Martin
    Richter-Boix, Alex
    Eckerström-Liedholm, Simon
    Stockholm University, Faculty of Science, Department of Zoology.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Thermally induced parental effects influence life history traits and covary with an environmental cline in common frog populationsManuscript (preprint) (Other academic)
    Abstract [en]

    Across latitudinal clines, the juvenile developmental rates of ectotherms often covary with the length of the growing season, due to life-history trade-offs imposed by time-constrained environments. However, as the start of the growing season often varies substantially across years, adaptive parental effects on juvenile developmental rates may mediate the costs of a delayed spring. By employing a meta-analysis, we tested whether larval developmental rates across a latitudinal cline of common frogs (Rana temporaria) are affected by fluctuating onsets of breeding, across years. We predicted that larval developmental rate will be inversely related to the onset of breeding, and that northern populations will be more prone to shorten their developmental rate in response to late breeding, as the costs of delayed metamorphosis should be highest in areas with a shorter growing season. We found that the larval period of both northern and southern populations responded to parental environmental conditions to similar degree in absolute terms, but in different directions. In northern populations, a late season start correlated with decreased development time, suggesting that the evolution of parental effects aid population persistence in time-constrained environments. In southern populations, late season start correlated with increased development time, which could potentially be explained as a predator avoidance strategy. Our findings suggest that local ecological variables can induce adaptive parental effects, but responses are complex, and likely trade-off with other ecological factors.

  • 8.
    Rowiński, Piotr
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Näslund, Joacim
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Stockholm University, Faculty of Science, Department of Zoology.
    Eckerström-Liedholm, Simon
    Stockholm University, Faculty of Science, Department of Zoology.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    The evolution of a placenta is not linked to increased brain size in poeciliid fishesManuscript (preprint) (Other academic)
    Abstract [en]

    Maternal investment traits are considered to have a direct influence on the size of energetically costly organs, including the brain. In placental organisms, offspring are supplied with nutrients during pre-natal development, which in turn may modulate brain size evolution. While this hypothesis has received some support in mammals (i.e. in the marsupial/placental transition), how the evolution of the placenta affects brain size in other taxa is largely unknown.Here, we use eight poeciliid fish species to test if species with placental transferred nutrients, invest more resources into offspring brain development than species with no placental structures. We predicted that the evolution of the placenta would be associated with larger relative brain size in fry, and possibly also shallower ontogenetic brain size allometry, if cognitive demands are similar in adults across placental and non-placental species. We tested these hypotheses by taking non-invasive brain size measurements during the first four weeks of life, and relating these to corresponding somatic growth. Contrary to our expectations, we did not find any differences in brain size between the two maternal strategies. Furthermore, we did not find any differences in how relative brain size changed over ontogenetic development between placental and non-placental species. Elsewhere, maternal investment traits have been commonly linked to brain size, however the species investigated here only exhibit pre-natal provisioning, which may reduce the potential for maternal investment into brain size. Our results suggest that coevolution between placental structures and juvenile brain size is not a general pattern.

  • 9.
    Rowiński, Piotr
    et al.
    Stockholm University, Faculty of Science, Department of Zoology.
    Sowersby, Will
    Stockholm University, Faculty of Science, Department of Zoology.
    Näslund, Joacim
    Stockholm University, Faculty of Science, Department of Zoology.
    Eckerström-Liedholm, Simon
    Stockholm University, Faculty of Science, Department of Zoology.
    Gotthard, Karl
    Stockholm University, Faculty of Science, Department of Zoology.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Variation in developmental rates is not linked to environmental unpredictability in annual killifishesManuscript (preprint) (Other academic)
    Abstract [en]

    Comparative evidence suggests that adaptive plasticity may evolve as a response to predictable environmental variation. However, receiving less attention is unpredictable environmental variation, which is considered to affect evolutionary trajectories by increasing phenotypic variation (bet-hedging). If increased variance in development time evolved as an adaptation to unpredictable environmental conditions, we would expect species inhabiting locations with unpredictable conditions to have a higher variance in development time. Here, we examine the occurrence of bet-hedging in egg developmental rate in seven species of annual killifish, originating from a gradient of precipitation rate variation, under three different incubation temperatures (21°C, 23°C, and 25°C). These fish species persist as dormant eggs buried in the soil, in seasonal environments with regular habitat desiccation. At the onset of the rainy season, the eggs must be sufficiently developed in order to hatch, and complete their life-cycle. We found substantial differences among species in both mean and variation of egg development rates, as well as species-specific plastic temperature responses. However, there was no clear relationship between variation in egg development time and variation in precipitation (environmental predictability). Hence, if species specific variances are adaptive, they do not diverge in accordance with simple linear relationship to variation in precipitation.

  • 10. Sowersby, Will
    et al.
    Eckerström-Liedholm, Simon
    Stockholm University, Faculty of Science, Department of Zoology.
    Rowiński, Piotr
    Stockholm University, Faculty of Science, Department of Zoology.
    Balogh, Julia
    Eiler, Stefan
    Upstone, Joseph
    Gonzalez-Voyer, Alejandro
    Stockholm University, Faculty of Science, Department of Zoology.
    Rogell, Björn
    Stockholm University, Faculty of Science, Department of Zoology.
    Costly sexual ornaments coevolve with fast life-histories in killifishesManuscript (preprint) (Other academic)
    Abstract [en]

    Sexually selected ornaments constitute an important investment into reproduction, increasing current mating success, at a potential cost to survival. Theory suggests that exaggerated sexual ornaments may coevolve with the life-history trade-off between current and future reproduction, however this hypothesis has remained unexplored. Here, we determine how the size of secondary sexual traits coevolves with the pace of life-histories (slow or fast), using a clade of killifishes, where independent adaptations to ephemeral environments have resulted in substantial divergences in life-history strategy. In addition, we assess costs to swimming performance driven by enlarged, ornamental fins. We predict that killifishes with fast life-histories, which inhabit time-limited environments and prioritize current reproduction, will have a greater tendency to evolve enlarged fins, compared to killifishes with slow life-histories. Indeed, we found that species with fast life-histories had more pronounced sexual size dimorphism, with males from these species having exaggerated dorsal and anal fins, compared to species with slow life-histories. Furthermore, males from species with fast life-histories and larger ornaments exhibited lower swimming performances compared to both conspecific females, and individuals from species with slow life-histories. Our results indicate that the trade-off between current and future reproduction, can be an evolutionary driver of costly sexual ornaments.

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