Papers by Veijo Jormalainen
Botanica Marina, 1989
... null Enriquez Dominguez, Susana / Heimann, Kirsten / Pohnert, Georg / Poulin, Michel / Sheare... more ... null Enriquez Dominguez, Susana / Heimann, Kirsten / Pohnert, Georg / Poulin, Michel / Shearer, Carol A. / Amsler, Charles D. / Beardall, John / Berges, John A. / Campbell, Jinx / Dawes, Clinton J. / Gomez, Ivan / Hoppenrath, Mona / Wynne, Michael J. / Yoon, Hwan Su. ...
Ambio, 2007
In diverse littoral communities, biotic interactions play an important role in community regulati... more In diverse littoral communities, biotic interactions play an important role in community regulation. This article reviews how eutrophication modifies biotic interactions in littoral macroalgal communities. Eutrophication causes blooms of opportunistic algae, increases epibiotism, and affects regulation by grazers. Opportunistic algae and epibionts harm colonization and growth of perennial algae. Grazing regulates the density and species composition of macroalgal communities, especially at the early stage of algal colonization. Eutrophication supports higher grazer densities by increasing the availability and quality of algae to grazers. This may, on the one hand, enhance the capability of grazers to regulate and counteract the increase of harmful, bloom-forming macroalgae; on the other hand, it may increase grazing pressure on perennial species, with a poor tolerance of grazing. In highly eutrophic conditions, bloom-forming algae may also escape grazing control and accumulate. Increasing epibiotism and grazing threaten in particular the persistence of habitat-forming perennials such as the bladderwrack. An interesting property of biotic interactions is that they do not remain fixed but are able to evolve, as the traits of the interacting species adapt to each other and to abiotic conditions. The potential of plants and grazers to adapt is crucial to their chances to survive in changing environment.
Animal Behaviour, 1989
... Est. Coast. Shelf ScL, 20, 105-116. Jansson, AM. Matthiesen, AS. 1971. ... Biol. Ecol., 2, 24... more ... Est. Coast. Shelf ScL, 20, 105-116. Jansson, AM. Matthiesen, AS. 1971. ... Biol. Ecol., 2, 24 36. Kangas, P., Autio, H., H/illfors, G., Luther, H., Niemi, A. Salemaa, H. 1982. A general model of the decline of Fucus vesiculosus at Tvfirminne, south coast 0f Finland in 1977-81. Acta Bot. ...
Ecology letters, 2012
Despite the importance of consumers in structuring communities, and the widespread assumption tha... more Despite the importance of consumers in structuring communities, and the widespread assumption that consumption is strongest at low latitudes, empirical tests for global scale patterns in the magnitude of consumer impacts are limited. In marine systems, the long tradition of experimentally excluding herbivores in their natural environments allows consumer impacts to be quantified on global scales using consistent methodology. We present a quantitative synthesis of 613 marine herbivore exclusion experiments to test the influence of consumer traits, producer traits and the environment on the strength of herbivore impacts on benthic producers. Across the globe, marine herbivores profoundly reduced producer abundance (by 68% on average), with strongest effects in rocky intertidal habitats and the weakest effects on habitats dominated by vascular plants. Unexpectedly, we found little or no influence of latitude or mean annual water temperature. Instead, herbivore impacts differed most consistently among producer taxonomic and morphological groups. Our results show that grazing impacts on plant abundance are better predicted by producer traits than by large-scale variation in habitat or mean temperature, and that there is a previously unrecognised degree of phylogenetic conservatism in producer susceptibility to consumption.
Marine environmental research
The brown alga Fucus vesiculosus is a foundation species in the Baltic Sea littoral, hosting a ri... more The brown alga Fucus vesiculosus is a foundation species in the Baltic Sea littoral, hosting a rich faunal community. We compared the species composition and diversity of invertebrate macrofauna living on F. vesiculosus between sites differing in their eutrophication status and exposure to waves at three different times during a season. We determined the size, nitrogen and phlorotannin content of the alga. The invertebrate community differed substantially between sites near fish farms and those in more pristine environment. Snails and bivalves were more abundant on the Fucus stands near fish farms than on control stands, where crustaceans were more abundant. The abundance of molluscs decreased with the increasing shore exposure, while gammaridean amphipods dominated on the exposed shores. Abundance of several taxa increased during the proceeding growing season. The density of the most important herbivore of F. vesiculosus, Idotea balthica, varied 100-fold during the season being the lowest in June and the highest in August when the generation born in the summer started to feed on Fucus. Thus, the diversity and composition of Fucus-associated invertebrate fauna varies both with environmental conditions of the stand and seasonally. Although the negative effects of eutrophication on distribution and abundance of Fucus stands are well documented, a moderate increase of nutrients was found to increase the species richness of Fucus-associated fauna in early summer.
Oecologia, 2010
In the marine littoral, strong grazing pressure selects for macroalgal defenses such as the const... more In the marine littoral, strong grazing pressure selects for macroalgal defenses such as the constitutive and inductive production of defense metabolites. Induced defenses are expected under spatiotemporally varying grazing pressure and should be triggered by a reliable cue from herbivory, thereby reducing grazing pressure via decreased herbivore preference and/or performance. Although induced resistance has frequently been demonstrated in brown macroalgae, it is yet to be investigated whether induced macroalgal resistance shows genetic variation, a prerequisite for evolutionary responses to selection. In addition, consequences of induced resistance on herbivore performance have rarely been tested while the role of brown algal phlorotannins as inducible defense metabolites remains ambiguous. Using preference bioassays, we tested various cues, e.g., natural grazing, waterborne cues or simulated grazing to induce resistance in the brown alga Fucus vesiculosus. Further, we investigated whether there are induced responses in phlorotannin content, genetic variation in induced resistance or incurred performance costs to the mesoherbivore isopod, Idotea baltica. We found that both direct grazing and waterborne grazing cues decreased the palatability of F. vesiculosus, while increasing the total phlorotannin content. Since the sole presence of the herbivore also increased the total soluble phlorotannins, yet failed to stimulate deterrence, we concluded that phlorotannins alone do not explain increased resistance. Induced resistance varied between algal genotypes and thus showed potential for evolutionary responses to variation in grazing pressure. Induced resistance also incurred performance costs for female I. baltica via reduced egg production. Our results show that the induced resistance of F. vesiculosus decreases grazing pressure by deterring herbivores as well as impairing their performance. Resistance may be induced in advance by waterborne cues and spread effectively throughout the F. vesiculosus belt. Through lowering herbivore performance, induced resistance may also reduce future grazing pressure by decreasing the population growth rate of I. baltica.
Behavioural processes, 2008
Habitat choice of herbivores is expected to be a resolution of a trade-off between food and shelt... more Habitat choice of herbivores is expected to be a resolution of a trade-off between food and shelter. The resolution of this trade-off may, however, be dynamic within a species because distinct phenotypes may value these factors differently and the value may vary temporally. We studied this hypothesis in the marine herbivore Idotea balthica (Isopoda), by simultaneously manipulating both food and shelter, and investigated whether the resolution of the trade-off differed between sexes, colour morphs and day and night (i.e. high and low predation risk). Isopods chose between exposing and concealing backgrounds in which the quantity or quality of food varied. When choosing between the backgrounds in the absence of food, females preferred the concealment more than males did. However, in a trade-off situation the isopods traded shelter for food, and females more so than males. Thus, males' lower preference for the shelter was not counterbalanced by a stronger preference for food. The microhabitat use also differed between night and day showing adaptation to diurnally fluctuating predation risk. We suggest that microhabitat utilization of females is more strongly tied to variation in risk and resources than that of males, for whom other factors, such as seeking mates, may be more important.
Journal of evolutionary biology, 2009
We studied ecological divergence of host use ability in a generalist marine herbivore living in t... more We studied ecological divergence of host use ability in a generalist marine herbivore living in two distinct host plant assemblages. We collected Idotea balthica isopods from three populations dominated by the brown alga Fucus vesiculosus and three dominated by the seagrass Zostera marina. In two reciprocal common garden feeding experiments for adult and laboratory-born juvenile isopods, we found that isopods from both assemblages performed better with their sympatric dominant host species than did isopods allopatric to this host. This indicates parallel divergence of populations according to the sympatric host plant assemblage. Furthermore, initial body size and body size-dependent mortality differed between populations from the two assemblages. In nature, this may result in lower fitness of immigrants compared with that of residents and consequently reinforce divergence of the populations. Finally, we discuss how phenotypic plasticity and maternal and random effects may associate with the results.
Oecologia, 2008
Macroalgae have to cope with multiple natural enemies, such as herbivores and epibionts. As these... more Macroalgae have to cope with multiple natural enemies, such as herbivores and epibionts. As these are harmful for the host, the host is expected to show resistance to them. Evolution of resistance is complicated by the interactions among the enemies and the genetic correlations among resistances to different enemies. Here, we explored genetic variation in resistance to epibiosis and herbivory in the brown alga Fucus vesiculosus, both under conditions where the enemies coexisted and where they were isolated. F. vesiculosus showed substantial genetic variation in the resistance to both epibiosis and grazing. Grazing pressure on the alga was generally lower in the presence than in the absence of epibiota. Furthermore, epibiosis modified the susceptibility of different algal genotypes to grazing. Resistances to epibiosis and grazing were independent when measured separately for both enemies but positively correlated when both these enemies coexisted. Thus, when the enemies coexisted, the fate of genotypes with respect to these enemies was intertwined. Genotypic correlation between phlorotannins, brown-algal phenolic secondary metabolites, and the amount of epibiota was negative, indicating that these compounds contribute to resistance to epibiosis. In addition, phlorotannins correlated also with the resistance to grazing, but this correlation disappeared when grazing occurred in the absence of epibiota. This indicates that the patterns of selection for the type of the resistance as well as for the resistance traits vary with the occurrence patterns of the enemies.
Advances in marine biology, 2011
PloS one, 2013
Nutrient enrichment in coastal marine systems can have profound impacts on trophic networks. In t... more Nutrient enrichment in coastal marine systems can have profound impacts on trophic networks. In the Baltic Sea, the population of Great Cormorant (Phalacrocorax carbo sinensis) has increased nearly exponentially since the mid-1990s, and colonies of these seabirds can be important sources of nitrogen enrichment for nearby benthic communities due to guano runoff. In this study we used stable isotope analyses and diet mixing models to determine the extent of nitrogen enrichment from cormorant colonies, as well as to examine any possible changes in herbivore diet preferences due to enrichment. We found significantly higher levels of d 15 N in samples from colony islands than control islands for producers (the dominant macroalga Fucus vesiculosus, filamentous algae, and periphyton) and herbivores, as well as a positive correlation between enrichment and nest density in colony sites. We also found that enrichment increased over the breeding season of the cormorants, with higher enrichment in late summer than early summer. While the amount of total nitrogen did not differ between colony and control sites, the amount of guano-based nitrogen in algae was .50% in most sites, indicating high nitrogen enrichment from colonies. Herbivores (the isopod Idotea balthica and the gastropod Theodoxus fluviatilis) preferred feeding upon the dominant macroalga Fucus vesiculosus rather than on filamentous algae or periphyton in both control and colony, and there was a significant increase in periphyton consumption near colony sites. Overall, guano from cormorant colonies seems to have effects on both producers and herbivores, as well as the potential to modify algae-herbivore interactions.
Oikos, 2003
. Induction of phlorotannin production in a brown alga: defense or resource dynamics? -Oikos 103:... more . Induction of phlorotannin production in a brown alga: defense or resource dynamics? -Oikos 103: 640-650. Increase of phenolic secondary metabolites, phlorotannins, in brown algae due to gastropod grazing has been interpreted as an anti-herbivore adaptation. Here we tested whether such a response could be due to changes in truly available resources for the alga, not by the grazing activity of snails as such. We allowed two species of snails, Theodoxus flu6iatilis and Physa fontinalis to graze on Fucus 6esiculosus. These species feed on epibiota and particulate matter on the thallus but do not eat the thallus of F. 6esiculosus. We further simulated snail grazing by nutrient enhancement, removal of epibiota and by a combination of the two. Manipulations of nutrient and light availability revealed the crucial role of epibiota in mediating resource availability for F. 6esiculosus. Nutrient enhancement alone increased epibiota and decreased phlorotannins. Cleaning the thallus resulted in increased growth, and together with nutrient enhancement also in a trade-off with phlorotannins. Presence of T. flu6iatilis on the thallus induced phlorotannin production, a response differing from the simulations of snail grazing. However, we suggest that the increase in phlorotannins may not be an induced defense but rather a consequence of a specific way of resource manipulation by this snail species. T. flu6iatilis removes hyaline hairs that facilitate nutrient uptake. P. fontinalis did not remove hyaline hairs and the response of the alga to its grazing was similar to the treatment where we mechanically removed epibiota suggesting that cleaning of the thallus is the major mechanism how this snail species affects F. 6esiculosus. Genetic variation in phlorotannin concentrations highly exceeded the induced responses of simulated or real snail grazing. This casts doubt for the efficiency of induced phlorotannin production to act as a defense, but is not contradictory with the interpretation of phlorotannins responding to variation in resource availability.
Oecologia, 1997
Intraspecific microhabitat segregation is expected to arise when there are age- or sex-specific d... more Intraspecific microhabitat segregation is expected to arise when there are age- or sex-specific differences in predation risk. The degree to which conspecific predation (cannibalism) can generate this risk, however, is poorly understood. In this paper, we examine microhabitat use, cannibalism, and individual responses to the presence of conspecifics in Thermosphaeroma thermophilum, an endangered isopod crustacean species that is endemic to a single, thermal spring in Socorro, N.M. USA. In samples from the natural habitat, juveniles (mancas) were found mainly on vegetation, whereas adults were found predominantly on bottom sediments. Females were found on vegetation more often than males. In laboratory containers without refuges, males cannibalized females, males and females cannibalized mancas, and mancas cannibalized each other, even in the presence of alternative food. When placed in containers provided with refuges, mancas actively avoided adults. We suggest, therefore, that cannibalism in T. thermophilum generates age-, size-, and sex-specific predation risks which are responsible for microhabitat segregation between mancas and adults, and between males and females. Since interspecific predation in the spring is negligible, cannibalism appears to play a significant role in population regulation and behavioral evolution in this species. We recommend, given the current “endangered” status of this species, that microhabitat heterogeneity be maintained in its native spring because it provides refuges from cannibalism and may support a larger and more viable natural population.
Journal of Evolutionary Biology, 2004
Selection in a brown alga 809 J . E V O L . B I O L . 1 7 ( 2 0 0 4 ) 8 0 7 -8 2 0 ª 2 0 0 4 B L ... more Selection in a brown alga 809 J . E V O L . B I O L . 1 7 ( 2 0 0 4 ) 8 0 7 -8 2 0 ª 2 0 0 4 B L A C K W E L L P U B L I S H I N G L T D Selection in a brown alga 819 J . E V O L . B I O L . 1 7 ( 2 0 0 4 ) 8 0 7 -8 2 0 ª 2 0 0 4 B L
Phytochemical Analysis, 2007
Separating individual compounds by HPLC represents an effective method for the detection and quan... more Separating individual compounds by HPLC represents an effective method for the detection and quantification of phenolic compounds and has been widely utilised. However, phlorotannins are commonly quantified using colorimetric methods, as the total amount of the whole compound group. In the present paper the separation of a set of individual soluble phlorotannins from the phenolic crude extract of Fucus vesiculosus was achieved by HPLC with UV photodiode array detection. Different gradient programs for reversed- and normal-phase HPLC methods were developed and tested. Normal-phase (NP) conditions with a silica stationary phase and a mobile phase with a linear gradient of increasing polarity were found to separate 16 individual components of the phenolic extract. The suitability of the NP-HPLC method for mass spectrometric application was preliminarily tested. Sample preparation was found to be a critical step in the analysis owing to the rapid oxidation of phlorotannins; ascorbic acid was used as an antioxidant. Copyright © 2007 John Wiley & Sons, Ltd.
International Journal of Plant Sciences, 2002
We investigated the level of within-alga integration, the consequent ability to compensate for si... more We investigated the level of within-alga integration, the consequent ability to compensate for simulated herbivory, and the effect of nutrient availability on the ability to compensate in Fucus vesiculosus (L.). Although F. vesiculosus lacks vascular connections, resource translocation occurs on a local scale. Vegetative apical parts and reproductive structures were dependent on the resources provided by the mature thallus. Thus, damage from herbivory has the potential to decrease the growth and reproductive success of F. vesiculosus. In contrast to vascular plants, F. vesiculosus did not show any plasticity in response to different types of damage. Removal of an apical part terminated the growth of the focal branch but had no effect on the growth of the neighboring branch. Removal of the older part of the thallus had no effect on the branching of the apical meristems. Thus, the within-alga growth pattern remained similar whether the old thallus or an apical part was damaged. Increased resource availability did not enhance the ability of algae to compensate for lost thallus. The strong dependence of the apical part on the older thallus and the low plasticity in the type of response to damage indicate that in F. vesiculosus the tolerance of herbivory may be low.
Biological Journal of The Linnean Society, 1995
Variable selection, including spatio-temporal variation, frequency-dependent selection and differ... more Variable selection, including spatio-temporal variation, frequency-dependent selection and differential selection due to habitat choice, may maintain polymorphism in heterogeneous environments. We studied predation as a selective agent on colour polymorphism of the aquatic isopod I baltica. Variable predation on this species can arise from at least three sources. First, apostatic selection was studied by testing the formation of preferences on colour morphs in the perch, a common predator of I baltica. Such acquired preferences should induce apostatic selection. While our results indicate some acquired preferences, there was significant heterogeneity in the behaviour of predator individuals. Second, temporal variation in selection can arise due to habitat shift from the green algae juvenile habitat to the bladderwrack adult habitat, and the consequent change in the crypsis of the morphs. Different crypsis between sexes probably promoted high predation mortality among females in the juvenile habitat. The high rate of male mortality during the breeding period, on the other hand, was presumably due to their high mate-searching activity. Third, the sex-dependent habitat choice of I baltica leads to sexual differences in the susceptibility of morphs to predation. Predators preferred the white-spotted morph over the uniform one in males but not in females, supporting the ‘dimorphic niche’ hypothesis as an explanation of sexual differences in morph frequencies. Finally, no evidence was found that the colouration patterns were under sexual selection. We therefore conclude diat variable predation is the most promising explanation for the maintenance of polymorphism in I. baltica.
Uploads
Papers by Veijo Jormalainen