Introduction
Large subtidal brown algal species form extensive underwater forests along the Irish coastline. Dominant species include the orders Fucales (Himanthalia elongata (Linnaeus) S.F. Gray 1821 andCystoseira spp.), Tilopteridales (Sacchoriza polyschides(Lightfoot) Batters), and Laminariales (Laminaria digitata(Hudson) J.V. Lamouroux, Alaria esculenta (Linnaeus) Greville,Saccharina latissima (Linnaeus) C.E. Lane, C. Mayes, Druehl & G.W. Saunders, and L. hyperborea (Gunnerus) Foslie). Recently, non-native species have arrived in these subtidal ecosystems via poleward distribution shifts (e.g., Laminariales - L. ochroleucaBachelot de la Pylaie; Schoenrock et al., 2019) or from aquaculture and/or shipping practices (Fucales - Sargassum muticum (Yendo) Fensholt 1955; Laminariales - Undaria pinnatifida (Harvey) Suringar; Kraan, 2017). Along semi-exposed rocky coastlines, L. hyperborea forms dense forests, whereas in the calmer, shallow regions of tidal loughs and fjords, S. latissima can form small forests on hard substratum. The paucity of subtidal research makes it difficult to determine the current and historical scope of these habitats along Ireland’s shorelines, but the synergistic threats of ocean warming and increased commercial harvesting present a critical need to better understand the historical and present day distribution of these important species in order to protect current and future marine forests.
As ecosystem engineers (Jones et al., 1994), kelp provides structure for shallow marine habitats as a resource and a habitat for many organisms (> 300 macrofauna species in L. hyperborea forests in Ireland throughout the year; K. Schoenrock et al., personal communication). In other parts of Europe, kelp forests are foraging habitats for marine birds and fish (Norderhaug et al., 2003; 2005), provide substrata for diverse marine assemblages (Bengtsson et al., 2012; H. Christie et al., 2007; H. P. Christie et al., 2003; Norderhaug et al., 2002; K. Schoenrock et al., personal communication), and are the basis of multiple food chains from coastal habitats to the deep sea (Filbee-Dexter et al., 2018), even providing a carbon resource for phytoplankton (Fredriksen, 2003). These habitats can also dampen wave energy, protecting coastlines from erosion (Lovas & Tørum, 2001), and modify parameters of the marine environment, including carbonate chemistry and light attenuation (Dean, 1985; Hofmann et al., 2011; Krause-Jensen et al., 2015). In Ireland, observations indicate kelp forests are seasonal homes to commercial species like the edible crab (Cancer pagurus Linnaeus), European lobster (Homarus gammarus Linnaeus) and multiple species of juvenile fish that inhabit the kelp canopy (Schoenrock et al., personal communication). There are natural seasonal patterns in these marine communities, but certain species inhabit kelp forests throughout the year including echinoderms, such as the common star (< 3 cm in diameter, Asterias rubens Linnaeus), spiny sea star (Marthasterias glacialisJullien), the common urchin (Echinus esculentus Linnaeus), a full summary of species is provided in Table 1, (K. Schoenrock et al., personal communication). The constant presence of these taxa in kelp forests over two years of monitoring suggests that they are indicators of healthy ecosystems in the west of Ireland. Regions to the north, south, and east have similar communities (K. Schoenrock, personal communication), but these regions have not been as thoroughly surveyed as the west of Ireland.
Kelp forest decline has been observed world-wide driven by warming oceans and heatwave events, anthropogenic inputs (harvesting and eutrophication), and herbivore pressure (Krumhansl et al., 2016; Reed et al., 2016; T. Wernberg et al., 2019). In Europe, the distribution of kelp species has changed over time with climate forcing (from the last glacial maximum), and species are predicted to continue retracting their southern range and move northwards with ocean warming (Assis et al., 2016; Assis, Araújo, et al., 2018; Assis, Serrão, et al., 2018a). Currently, warmer-water subtidal forests dominated by L. ochroleuca have retracted from their southern range edges in Morocco to current limits mid-Portugal (Assis, Serrão, et al., 2018b), while the cold-water kelp L. hyperborea has retracted its southern range edge from the Portuguese coastline to the Spanish coastline of the Bay of Biscay (Assis, Araújo, et al., 2018). Marine forests from the Mediterranean coast to southern Portugal are poorly understood, though it seems many species ranges are retracting at their lower latitude range limits. On the other hand, at higher latitudes abundances of some species, such as S. latissima and S. polyschides , are increasing at their northern range edges, while the spread of invasive species, such as U. pinnatifida, is increasing (Araujo et al., 2016). In addition, herbivore pressure is devastating L. hyperborea at its northern ranges (northern Norway), though urchin populations can fluctuate from year to year, allowing regrowth of some populations (Hagen, 1995). Harvesting is also a threat to kelp forests across Europe, particularly France and Norway where the commercial exploitation of L. hyperborea and L. digitata has been occurring for decades (Valero et al., 2001). The mechanical removal ofL. hyperborea results in the removal of whole individuals. Recovery can take greater than five years (Lorentsen et al., 2010), and populations in the heaviest areas of harvest may be important reservoirs of genetic diversity (e.g., Brittany, France: Robuchon et al., 2014).
Kelp have been of great interest in the past, in flux with economic, socio-political, and technical advances beginning during the 1700s. Therefore, there are many historical accounts of seaweed from sea captains, fisheries, and naturalists along the Irish coastline from the late 1700s to the present day. These qualitative historical records provide perspective on the value of kelp along Ireland’s coastline, but given the pressures outlined above, we need a better understanding and documentation of kelp ecosystems. In this review, we focus on the presence of the subtidal species L. hyperborea from 1700s Ireland to present day, collating disparate historical records for the first time. This review significantly contributes to our understanding of kelp forest function in Ireland at a time when interest in kelp harvesting is increasing despite the fact we do not understand basic ecological and evolutionary processes at work in these systems. We show that regular and systematic monitoring is urgently needed in order to conserve and inform policy makers to foster resilience, defined as the ability to recover from a disturbance and maintain ecosystem function, in this vital coastal resource.