Kristjan Herkül

Biodiversity gradient in the Baltic Sea: a comprehensive inventory of macrozoobenthos data

Abstract In the Helsinki Commission Red List project 2009–2012, taxonomic and distributional data of benthic (macro) invertebrates were compiled by the present authors in a comprehensive checklist of the Baltic Sea fauna. Based on the most recent and comprehensive data, this paper presents the diversity patterns observed among benthic invertebrates in the Baltic Sea. As expected, the total number of species per sub-region generally declined along the salinity gradient from the Danish Straits to the northern Baltic Sea. This relationship is well known from the Baltic Sea and has resulted in a general assumption of an exponentially positive relationship between species richness and salinity for marine species, and a negative relationship for freshwater species. In 1934, Remane produced a diagram to describe the hypothetical distribution of benthic invertebrate diversity along a marine–freshwater salinity gradient. Our results clearly indicated the validity of this theory for the macrozoobenthic diversity pattern within the Baltic Sea. Categorisation of sub-regions according to species composition showed both separation and grouping of some sub-regions and a strong alignment of similarity patterns of zoobenthic species composition along the salinity gradient.

Large-scale variation in combined impacts of canopy loss and disturbance on community structure and ecosystem functioning

Ecosystems are under pressure from multiple human disturbances whose impact may vary depending on environmental context. We experimentally evaluated variation in the separate and combined effects of the loss of a key functional group (canopy algae) and physical disturbance on rocky shore ecosystems at nine locations across Europe. Multivariate community structure was initially affected (during the first three to six months) at six locations but after 18 months, effects were apparent at only three. Loss of canopy caused increases in cover of non-canopy algae in the three locations in southern Europe and decreases in some northern locations. Measures of ecosystem functioning (community respiration, gross primary productivity, net primary productivity) were affected by loss of canopy at five of the six locations for which data were available. Short-term effects on community respiration were widespread, but effects were rare after 18 months. Functional changes corresponded with changes in community structure and/or species richness at most locations and times sampled, but no single aspect of biodiversity was an effective predictor of longer-term functional changes. Most ecosystems studied were able to compensate in functional terms for impacts caused by indiscriminate physical disturbance. The only consistent effect of disturbance was to increase cover of non-canopy species. Loss of canopy algae temporarily reduced community resistance to disturbance at only two locations and at two locations actually increased resistance. Resistance to disturbance-induced changes in gross primary productivity was reduced by loss of canopy algae at four locations. Location-specific variation in the effects of the same stressors argues for flexible frameworks for the management of marine environments. These results also highlight the need to analyse how species loss and other stressors combine and interact in different environmental contexts.

Relating remotely sensed optical variability to marine benthic biodiversity.

Biodiversity is important in maintaining ecosystem viability, and the availability of adequate biodiversity data is a prerequisite for the sustainable management of natural resources. As such, there is a clear need to map biodiversity at high spatial resolutions across large areas. Airborne and spaceborne optical remote sensing is a potential tool to provide such biodiversity data. The spectral variation hypothesis (SVH) predicts a positive correlation between spectral variability (SV) of a remotely sensed image and biodiversity. The SVH has only been tested on a few terrestrial plant communities. Our study is the first attempt to apply the SVH in the marine environment using hyperspectral imagery recorded by Compact Airborne Spectrographic Imager (CASI). All coverage-based diversity measures of benthic macrophytes and invertebrates showed low but statistically significant positive correlations with SV whereas the relationship between biomass-based diversity measures and SV were weak or lacking. The observed relationships did not vary with spatial scale. SV had the highest independent effect among predictor variables in the statistical models of coverage-derived total benthic species richness and Shannon index. Thus, the relevance of SVH in marine benthic habitats was proved and this forms a prerequisite for the future use of SV in benthic biodiversity assessments.

Ecological niche differentiation between native and non-native shrimps in the northern Baltic Sea

Invasions of non-native species are modifying global biodiversity but the ecological mechanisms underlying invasion processes are still not well understood. A degree of niche separation of non-native and sympatric native species can possibly explain the success of novel species in their new environment. In this study, we quantified experimentally and in situ the environmental niche space of caridean shrimps (native Crangon crangon and Palaemon adspersus, non-native Palaemon elegans) inhabiting the northern Baltic Sea. Field studies showed that the non-native P. elegans had wider geographical range compared to native species although the level of habitat specialization was similar in both Palaemon species. There were clear differences in shrimp habitat occupancy with P. elegans inhabiting lower salinity areas and more eutrophicated habitats compared to the native species. Consequently, the non-native shrimp has occupied large areas of the northern Baltic Sea that were previously devoid of the native shrimps. Experiments demonstrated that the non-native shrimp had higher affinity to vegetated substrates compared to native species. The study suggests that the abilities of the non-native shrimp to thrive in more stressful habitats (lower salinity, higher eutrophication), that are sub-optimal for native shrimps, plausibly explain the invasion success of P. elegans.

Variability of Benthic Communities in Relation to Hydrodynamic Conditions in the North-Eastern Baltic Sea

ABSTRACT Herkül, K.; Torn, K.; Suursaar, Ü.; Alari, V., and Peterson, A., 2016. Variability of benthic communities in relation to hydrodynamic conditions in the north-eastern Baltic Sea. In: Vila-Concejo, A.; Bruce, E.; Kennedy, D.M., and McCarroll, R.J. (eds.), Proceedings of the 14th International Coastal Symposium (Sydney, Australia). Journal of Coastal Research, Special Issue, No. 75, pp. 867–871. Coconut Creek (Florida), ISSN 0749-0208. Wave forcing is an important hydrodynamic variable that influences spatial patterns and temporal dynamics of marine benthos. In this study, we used three principally different wave models – simplified wave model (SWM), spectral wave model SWAN (simulating waves nearshore), and locally calibrated point model (LCPM) – to assess the effects of wave forcing on the distribution and temporal dynamics of macrobenthos in the north-eastern Baltic Sea. Other important environmental variables like depth, salinity, turbidity etc., were also included into analyses of benthos distribution in order to quantify the relative importance of wave forcing. Following depth, SWM was the second most important environmental predictor of spatial distribution of benthos. The importance of SWAN was the lowest among environmental variables due to its low spatial resolution (1 nautical mile) compared to SWM (25 m). Considering the temporal dynamics of benthos, wave height significantly correlated with several benthic variables, but the direction and magnitude of effects were site-specific.

The environmental niche separation between charophytes and angiosperms in the northern Baltic Sea

Abstract Charophytes and angiosperms are the most important primary producers and habitat modifiers in shallow soft-bottom fresh and brackish water bodies. The sheltered bays of the low-salinity northern Baltic Sea provide unique habitats, where euryhaline macrophytes of both freshwater and marine origin can thrive. The regional cohabitation of several species, with generally similar biological traits, raises questions about the mechanisms underlying the coexistence. Generally, sympatric populations rely on niche separation to reduce competition pressure that, in turn, facilitates long-term coexistence. A large benthos biomass data-set, covering the whole extent of the Estonian sea area, the northern Baltic Sea, together with a set of 13 environmental variables (bathymetrical, hydrodynamical, chemical, biological) was used to test the potential niche separation between regionally coexisting charophytes and angiosperms. A constrained correspondence analysis indicated both the grouping of some species and a distinct niche separation of others. The niche centers of angiosperms Zostera marina, Zannichellia palustris, and Potamogeton perfoliatus diverged most strongly from all the other species, while those of Chara connivens and Myriophyllum spicatum nearly coincided. Depth, salinity, and duration of ice were the most influential environmental gradients that discerned the environmental niches of the species. Comparison of the breadths of the environmental niche spaces, occupied by the studied species, showed highly species-specific results. Within the studied species, Z. palustris and Z. marina had the lowest niche specialization, and C. connivens had the highest specialization. Altogether, among the angiosperms the variability of the environmental niche breadth was higher compared to the charophytes.

Mapping benthic biodiversity using georeferenced environmental data and predictive modeling

Biodiversity is critical for maintaining and stabilizing ecosystem processes. There is a need for high-resolution biodiversity maps that cover large sea areas in order to address ecological questions related to biodiversity-ecosystem functioning relationships and to provide data for marine environmental protection and management decisions. However, traditional sampling-point-wise field work is not suitable for covering extensive areas in high detail. Spatial predictive modeling using biodiversity data from sampling points and georeferenced environmental data layers covering the whole study area is a potential way to create biodiversity maps for large spatial extents. Random forest (RF), generalized additive models (GAM), and boosted regression trees (BRT) were used in this study to produce benthic (macroinvertebrates, macrophytes) biodiversity maps in the northern Baltic Sea. Environmental raster layers (wave exposure, salinity, temperature, etc.) were used as independent variables in the models to predict the spatial distribution of species richness. A validation dataset containing data that was not included in model calibration was used to compare the prediction accuracy of the models. Each model was also evaluated visually to check for possible modeling artifacts that are not revealed by mathematical validation. All three models proved to have high predictive ability. RF and BRT predictions had higher correlations with validation data and lower mean absolute error than those of GAM. Both mathematically and visually, the predictions by RF and BRT were very similar. Depth and seabed sediments were the most influential abiotic variables in predicting the spatial patterns of biodiversity.

Marine environmental vulnerability and cumulative risk profiles to support ecosystem-based adaptive maritime spatial planning

Original Article Marine environmental vulnerability and cumulative risk profiles to support ecosystem-based adaptive maritime spatial planning Robert Aps*, Kristjan Herkül, Jonne Kotta, Roland Cormier, Kirsi Kostamo, Leena Laamanen, Juho Lappalainen, Külli Lokko, Anneliis Peterson, and Riku Varjopuro Estonian Marine Institute, University of Tartu, Mäealuse 14, 12618 Tallinn, Estonia Helmholtz-Zentrum Geesthacht, Max-Planck-Stra e 1, 21502 Geesthacht, Germany Finnish Environment Institute, Mechelininkatu 34 A, 00260 Helsinki, Finland *Corresponding author: tel: þ3725062597; e-mail: robert.aps@ut.ee.

Environmental niche separation between native and non-native benthic invertebrate species: Case study of the northern Baltic Sea

Knowledge and understanding of geographic distributions of species is crucial for many aspects in ecology, conservation, policy making and management. In order to reach such an understanding, it is important to know abiotic variables that impact and drive distributions of native and non-native species. We used an existing long-term macrobenthos database for species presence-absence information and biomass estimates at different environmental gradients in the northern Baltic Sea. Region specific abiotic variables (e.g. salinity, depth) were derived from previously constructed bathymetric and hydrodynamic models. Multidimensional ordination techniques were then applied to investigate potential niche space separation between all native and non-native invertebrates in the northern Baltic Sea. Such an approach allowed to obtain data rich and robust estimates of the current native and non-native species distributions and outline important abiotic parameters influencing the observed pattern. The results showed clear niche space separation between native and non-native species. Non-native species were situated in an environmental space characterized by reduced salinity, high temperatures, high proportion of soft seabed and decreased depth and wave exposure whereas native species displayed an opposite pattern. Different placement of native and non-native species along the studied environmental niche space is likely to be explained by the differences in their evolutionary history, human mediated activities and geological youth of the Baltic Sea. The results of this study can provide early warnings and effectively outline coastal areas in the northern Baltic Sea that are prone to further range expansion of non-native species as climate change is expected to significantly reduce salinity and increase temperature in wide coastal areas, both supporting the disappearance of native and appearance of non-native species.

Effects of harbour dredging on soft bottom invertebrate communities: Does environmental variability affect the community responses?

The effect of dredging on the biomass structure of benthic invertebrate communities was examined at 9 sites in the northeastern Baltic Sea during 2002-2007. We analyzed whether and how bottom topography, depth and sediment type contributed to these relationships. In general, the effects of dredging on benthic invertebrates were weak. Dredging clearly increased the biomass of bivalves but the communities recovered in about a year. Bottom topography affected the response of invertebrates to dredging. Flat bottoms were more sensitive to dredging compared to sites situating on slopes. Spatial modelling was proved as a useful tool to predict spatial variability in the effects of dredging on benthic invertebrate communities.