Nutrient enrichment and water depth modify consumer control in rocky shore macroalgal communities

The objective of my thesis is to assess mechanisms of ecological community control in macroalgal communities in the Baltic Sea. In the top-down model, predatory fish feed on invertebrate mesograzers, releasing algae partly from grazing pressure. Such a reciprocal relationship is called trophic cascade. In the bottom-up model, nutrients increase biomass in the food chain. The nutrients are first assimilated by algae and, via food chain, increase also abundance of grazers and predators. Previous studies on oceanic shores have described these two regulative mechanisms in the grazer - alga link, but how they interact in the trophic cascades from fish to algae is still inadequately known. Because the top-down and bottom-up mechanisms are predicted to depend on environmental disturbances, such as wave stress and light, I have studied these models at two distinct water depths. There are five factorial field experiments behind the thesis, which were all conducted in the Finnish Archipelago Sea. In all the experiments, I studied macroalgal colonization - either density, filament length or biomass - on submerged colonization substrates. By excluding predatory fish and mesograzers from the algal communities, the studies compared the strength of the top-down control to natural algal communities. A part of the experimental units were, in addition, exposed to enriched nitrogen and phosphorus concentrations, which enabled testing of bottom-up control. These two models of community control were further investigated in shallow (<1 m) and deep (ca. 3 m) water. Moreover, the control mechanisms were also expected to depend on grazer species. Therefore different grazer species were enclosed into experimental units and their impacts on macroalgal communities were followed specifically. The community control in the Baltic rocky shores was found to follow theoretical predictions, which have not been confirmed by field studies before. Predatory fish limited grazing impact, which was seen as denser algal communities and longer algal filaments. Nutrient enrichment increased density and filament length of annual algae and, thus, changed the species composition of the algal community. The perennial alga Fucus vesiculosusA and the red alga Ceramium tenuicorne suffered from the increased nutrient availabilities. The enriched nutrient conditions led to denser grazer fauna, thereby causing strong top-down control over both the annual and perennial macroalgae. The strength of the top-down control seemed to depend on the density and diversity of grazers and predators as well as on the species composition of macroalgal assemblages. The nutrient enrichment led to, however, weaker limiting impact of predatory fish on grazer fauna, because fish stocks did not respond as quickly to enhanced resources in the environment as the invertebrate fauna. According to environmental stress model, environmental disturbances weaken the top-down control. For example, on a wave-exposed shore, wave stress causes more stress to animals close to the surface than deeper on the shore. Mesograzers were efficient consumers at both the depths, while predation by fish was weaker in shallow water. Thus, the results supported the environmental stress model, which predicts that environmental disturbance affects stronger the higher a species is in the food chain. This thesis assessed the mechanisms of community control in three-level food chains and did not take into account higher predators. Such predators in the Baltic Sea are, for example, cormorant, seals, white-tailed sea eagle, cod and salmon. All these predatory species were recently or are currently under intensive fishing, hunting and persecution, and their stocks have only recently increased in the region. Therefore, it is possible that future densities of top predators may yet alter the strengths of the controlling mechanisms in the Baltic littoral zone.

Open Access - current developments

Presentation at the Nordic Perspectives on Open Access and Open Science seminar, Helsinki, October 15, 2013

Do Open Access journals qualify?

Presentation at the Nordic Perspectives on Open Access and Open Science seminar, Helsinki, October 15, 2013

Anti-predator adaptations in aquatic environments

Predation is an important selective force that has led to the evolution of a variety of fascinating anti-predator adaptations, such as many types of protective coloration and prey behaviours. Because the evolution of life has begun in the aquatic environment and many anti-predator adaptations are found already in relative primitive taxa, it is likely that many of these adaptations evolved initially in the aquatic environment. Yet, there has been surprisingly little research on the mechanisms and function of antipredator adaptations in aquatic systems. To understand the function of anti-predator adaptations and natural selection imposed on prey appearance and behaviour, I have investigated how protective coloration can be used, either as such or together with behavioural adaptations, to manipulate predator behaviour and decrease predation risk. To this end I conducted a series of behaviour ecological laboratory experiments in which I manipulated the visual appearance of artificial backgrounds and prey items. In paper I of this thesis, I investigated background choice as an anti-predator strategy, by observing the habitat choice of the least killifish (Heterandria formosa) between pairs of artificial backgrounds, both in the presence and absence of predation threat. It has been suggested that prey could decrease their risk of being detected by predators either by preferring backgrounds into which they blend or by preferring visually complex backgrounds. The least killifish preferred a background that matched their patterning to a background that mismatched it, showing that they are able to respond to cues of visual similarity between their colour pattern and the surrounding environment. Interestingly however, in female least killifish visual complexity of the background was a more important cue for habitat safety and may override or act together with background matching when searching for a safe habitat. It is possible that in females, preference for visually complex backgrounds is associated with lower opportunity costs than preference for matching backgrounds would be. Generally, the least killifish showed stronger preference while under predation threat, indicating that their background choice behaviour is an antipredator adaptation. Many aquatic prey species have eyespots, which are colour patterns that consist of roughly concentric rings and have received their name because they for humans often resemble the vertebrate eye. I investigated the anti-predator function of eyespots against predation by fish in papers II, III and IV. Some eyespots have been suggested to benefit prey by diverting the strikes of predators away from vital parts of the prey body or towards a direction that facilitates prey escape. Although proposed over a century ago, the divertive effect of eyespots has proven to be difficult to show experimentally. In papers II and III, I tested for divertive effect of eyespots towards attacking fish by presenting artificial prey with eyespots to laboratory reared three-spined sticklebacks (Gasterosteus aculeatus). I found that eyespots strongly influenced the behaviour of attacking sticklebacks and effectively drew their strikes towards the eyespots. To further investigate this divertive effect and whether the specific shape of eyespots is important for it, I tested in paper III the response of fish also to other markings than eyespots. I found that eyespots were generally more effective in diverting the first strikes of attacking fish compared to other prey markings. My findings suggest that the common occurrence of eyespots in aquatic prey species can at least partly be explained by the divertive effect of the eyespot shape, possibly together with the relative simple developmental mechanisms underlying circular colour patterns. An eyebar is a stripe that runs through the eye, and this pattern has been suggested to obscure the real eyes of the prey by visually blending parts of the eyes and head of the prey and by creating false edges. In paper III, I show that an eyebar effectively disrupts an eyelike shape. This suggests that eyebars provide an effective way to conceal the eyes and consequently obstruct detection and recognition of prey. This experiment also demonstrates that through concealment of the eyes, eyebars could be used to enhance the divertive effect of eyespots, which can explain the common occurrence of eyebars in many species of fish that have eyespots. Larger eyespots have been shown to intimidate some terrestrial predators, such as passerine birds, either because they resemble the eyes of the predator’s own enemy or because highly salient features may have an intimidating effect. In papers II and IV, I investigated whether the occurrence of eyespots in some aquatic prey could be explained by their intimidating effect predatory fish. In paper IV, I also investigated the reason for the intimidating effect of eyelike prey marks. In paper II, I found no clear intimidating effect of eyespots, whereas in paper IV, using a different approach, I found that sticklebacks hesitated to attack towards eyelike but not towards non-eyelike marks. Importantly, paper IV therefore presents the first rigorous evidence for the idea that eye mimicry, and not merely conspicuousness, underlies the intimidating effect. It also showed that the hesitation shown by fish towards eyelike marks is partly an innate response that is reinforced by encounters with predators. Collectively, this thesis shows that prey colour pattern and the visual appearance of the habitat influence the behaviour of fish. The results demonstrate that protective coloration provides numerous distinctive ways for aquatic prey to escape predation. Thus, visual perception and behaviour of fish are important factors shaping the appearance and behaviours of aquatic prey.

Vahinkoeläinsodasta psykologiseen omistajuuteen. Petokonfliktien historiallinen tausta ja nykypäivän hallinta

Suurpetojen kannanhoitoon, ja etenkin suden kannanhoitoon liittyvä monitahoinen konfl ikti esiintyy jännitteinä paikallisyhteisöjen ja keskushallinnon, maaseudun ja kaupunkien sekä maallikoiden ja tutkijoiden välillä. Suurpetopolitiikan legitimiteettikriisi näkyy kansallisen kannanhoidon tavoitteiden ja toimenpiteiden kiistämisenä passiivisin ja aktiivisin toimin. Tällä ajankohtaisella ongelmalla on mittavat historialliset juuret. Luonnon ja ihmisen välinen suhde on ollut jatkuvaa tasapainottelua konfl iktien ja niiden hallinnan kanssa, ja suurpedot ovat väistämättä olleet konfl iktin keskiössä lajityypillisten piirteidensä vuoksi. Tämän tutkimuksen tavoitteena on osoittaa ympäristöhistoriallisen tutkimusotteen kautta suomalaisen vahinkoeläinhistorian keskeiset toimet ja toimijat, sekä ymmärtää vahinkoeläinten kannanhoidon tavoite- ja toimenpideasettelun vaikutukset eläinpopulaatioihin historiallisessa ja ekologisessa kontekstissa. Tutkimuksen toinen tavoite on tuottaa synteesi nykypäivän suurpetojen kannanhoidon tavoite- ja toimenpiteiden tarkoituksenmukaisuudesta ja vaikuttavuudesta historiallisessa kontekstissa ja esittää uudenlainen lähestymismalli sosiaalisesti kestävämmälle suurpetopolitiikalle. Vahinkoeläinhistoriaa käsitellään deduktiivisella ja rekonstruktiivisella tutkimusotteella käyttäen aineistona lakitekstejä, maksettuja tapporahoja, sanomalehtikirjoituksia, sidosryhmien kirjoituksia sekä eläintieteellisiä julkaisuja. Etenkin laeissa ja asetuksissa eri eläimistä asetetut tapporahat ja toisaalta saalistilastojen perusteella maksetut palkkiot on nostettu tässä tutkimuksessa merkittävään rooliin asenteiden tulkinnan välineenä. Tarkastelu ulotetaan nisäkkäiden lisäksi myös lintuihin, jotka ovat edellä mainitun aineiston valossa olleet merkittävä vainottu eläinryhmä. Nykypäivän petokonfl iktin tarkastelussa keskiöön nostetaan suurpetoasenteet, niiden muodostuminen ja vaikutus suurpetopolitiikan tavoitteiden ja toimenpiteiden kiistämisessä. Suurpetopolitiikan legitimiteettikriisiä tarkastellaan paikallisten uhmakkuutena hallintoa kohtaan ja tulkinta keinoista puuttua sosiopoliittiseen rikokseen rakennetaan tätä ajatusta vasten. Suurpetojen salakaatajat toimivat yhteisönsä puolesta ja saavat tukensa toimilleen yhteisöltään. Sosiopoliittinen rikos ei siten ole ainoastaan salakaatajien vaan sen sijaan yhteisön yhteinen. Salakaatajat ja paikallisyhteisö pyrkivät häivyttämään yhteiskunnassa viranomaisten ja sidosryhmien taholta toimintaan liittämää stigmaa ja siten oikeuttamaan tekonsa vallitsevissa olosuhteissa. Arvoristiriitoihin perustuvat konfl iktit ovat haasteellisia hallita ja lähtökohtana on oltava ymmärrys siitä, että etenkin suteen liittyvät jännitteet ja ristiriidat ihmisten välisissä suhteissa ovat väistämättömiä. Tunnistamalla historiallisen aineiston merkittävyys nykypäivän ongelmakeskeisen kannanhoidon ratkaisun tietopohjana on mahdollista rakentaa sosiaalisesti kestävämpää suurpetojen kannanhoitoa. Tässä väitöskirjatyössä painotetaan suurpetoihin liittyen ajatusta, jonka mukaan paikallisilta suurpetoalueilla eläviltä ihmisiltä tai sidosryhmien edustajilta ei edellytetä arvomaailman muuttumista, vaan hyväksytään heidän käsityksensä suurpetojen roolista vahingontekijöinä. Sen sijaan vahvistetaan suurpetojen välineellistä roolia paikallisyhteisöjen resurssina ja oleellisten sidosryhmien roolia yhteisön toimijoina psykologisen omistajuuden rakentamisen kautta. Suurpetojen kestävä kannanhoito vaatii siten riistahallinnon toimintakulttuurin muutosta ja uudenlaista suhdetta yhteiskunnan eri toimijoihin.

Spatial and temporal variation in fish populations and assemblages in coastal waters of the northern Baltic Proper

The distribution and traits of fish are of interest both ecologically and socio-economically. In this thesis, phenotypic and structural variation in fish populations and assemblages was studied on multiple spatial and temporal scales in shallow coastal areas in the archipelago of the northern Baltic Proper. In Lumparn basin in Åland Islands, the fish assemblage displayed significant seasonal variation in depth zone distribution. The results indicate that investigating both spatial and temporal variation in small scale is crucial for understanding patterns in fish distribution and community structure in large scale. The local population of Eurasian perch Perca fluviatilis L displayed habitat-specific morphological and dietary variation. Perch in the pelagic zone were on average deeper in their body shape than the littoral ones and fed on fish and benthic invertebrates. The results differ from previous studies conducted in freshwater habitats, where the pelagic perch typically are streamlined in body shape and zooplanktivorous. Stable isotopes of carbon and nitrogen differed between perch with different stomach contents, suggesting differentiation of individual diet preferences. In the study areas Lumparn and Ivarskärsfjärden in Åland Islands and Galtfjärden in Swedish east coast, the development in fish assemblages during the 2000’s indicated a general shift towards higher abundances of small-bodied lower-order consumers, especially cyprinids. For European pikeperch Sander lucioperca L., recent declines in adult fish abundances and high mortalities (Z = 1.06–1.16) were observed, which suggests unsustainably high fishing pressure on pikeperch. Based on the results it can be hypothesized that fishing has reduced the abundances of large predatory fish, which together with bottom-up forcing by eutrophication has allowed the lower-order consumer species to increase in abundances. This thesis contributes to the scientific understanding of aquatic ecosystems with new descriptions on morphological and dietary adaptations in perch in brackish water, and on the seasonal variation in small-scale spatial fish distribution. The results also demonstrate anthropogenic effects on coastal fish communities and underline the urgency of further reducing nutrient inputs and regulating fisheries in the Baltic Sea region.