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MARINE INVERTEBRATES (Chapter 8)

Lead Authors: Alf B. Josefson and also Vadim Mokievsky

Contributing Authors:Melanie Bergmann, martin E. Blicher, Bodil Bluhm, Sabine Cochrane, Nina V. Denisenko, Christiane Hasemann, Lis L. Jørgensen, Michael Klages, Ingo Schewe, Mikael K. Sejr, cutting board Soltwedel, january Marcin We¸sławski and Maria Włodarska-Kowalczuk

SUMMARY

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Sea butterfly. Photo: Kevin LeeThis chapter brings together baseline details on the diversity of naval invertebrates in the Arctic Ocean and also discusses the importance of components that have shaped fads of biodiversity.

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The Arctic ocean is here characterized as the areas north of the Bering Strait ~ above the Pacific side and areas with constant seasonal sea ice cream cover ~ above the Atlantic side. The well-known marine invertebrate fauna the this area comprises c. 5,000 species, representing at the very least 24 phyla with representatives in all three naval realms: sea ice, pelagic and also benthic. Around 50% the the Arctic ocean overlays continent shelf locations at water depths ranging from 0-500 m. This Arctic Shelf constitutes 31% that the complete shelf area of the world. Much more than 90% that the well-known Arctic invertebrate types occur in the benthic realm. As for terrestrial environments, the most types rich taxon in all realms is Arthropoda, through most varieties among crustaceans, i.e. >1,500 varieties according to a current estimate. Various other species-rich taxonomic groups are Annelida, mostly bristle worms (Polychaeta), moss pets (Bryozoa) and Mollusca, including bivalves (Bivalvia) and also snails (Gastropoda). Amongst the meiobenthos (small-sized benthic metazoans,

“There are areas where the salmon is widening northto the high Arctic together the waters are getting warmerwhich is the instance in the Inuvialuit residence Settlement areaof the Northwest areas of Canada. Similar reportsare heard native the Kolyma river in the Russian Arcticwhere neighborhood Indigenous fishermen have captured sea medusaein their nets.Mustonen 2007.

The variety of known marine invertebrate varieties in the Arctic s is an extremely likely to increase in the future, because huge areas, particularly the deep-sea basins, space under-sampled. For example, a current estimate argues that several thousand benthic types have been missed to date. Contradictory to paradigms of one impoverished Arctic fauna due to a harsh environment, as seen in the terrestrial realm, the Arctic shelf fauna is not specifically poor, but considered to it is in of intermediary richness, similar in overall types richness come some other shelf faunas, such as the Norwegian shelf. The pattern of declining species richness with enhancing latitude, evident in the terrestrial realm, is controversial among marine invertebrates and conclusions count on the taxon and geographic range studied. A latitudinal decrease from the tropics to the Arctic was seen in shelf molluscs, while arthropods seem to show greater diversity in part Arctic locations compared through some non-Arctic areas.

Due to the turbulent geological background with repetitive glaciation occasions over the critical 3.5 million years, in addition to in ineffective isolation from nearby oceans, in situ evolution of species has been hampered, and also as a repercussion there are few Arctic endemics, at the very least on the continental shelves. However, bryozoans contain more endemics than many other groups, possibly partially related to bad dispersal in this group.The present-day invertebrate fauna in the Arctic is a mixture of types with various origins, whereby the majority have distribution reaching outside the Arctic, i.e. The boreal parts of the surrounding oceans. By and large the Arctic ocean is a sea that immigrants the have dispersed from adjacent oceans both in historical and in recent time.

Today’s biogeographic vehicle drivers of Arctic diversity are clearly seen in the distribution of beginnings in relationship to the two major gateways right into the Arctic, i.e. Indigenous the Atlantic Ocean and also the Pacific Ocean. On the continental shelves the proportions of present-day Pacific and also Atlantic species decrease with increasing distance indigenous the Bering Strait and also the NE Atlantic, respectively. Present inventories suggest that the Barents Sea has the highest types richness, gift ‘enriched’ through sub-Arctic and boreal species. Today’s Arctic deep-sea floor is most carefully related come the current North Atlantic fauna, i m sorry in a geological time perspective includes a solid Pacific influence.

Like other faunal aspects in the Arctic, naval invertebrates are affected by climate warming. The most noticeable effects will certainly be ~ above the fauna of the long-term ice (sympagic fauna) which will lose its habitat. However, detecting effects in the other realms is difficult, mainly due to the fact that there are only couple of time collection data available. It is expected that the fauna with solid boreal influence may display (perhaps temporarily) enhanced diversity, because of a combination of anticipated boosted food availability for the benthos and also immigration of types adapted come warmer waters. Indicators of borealization are already seen in marginal locations of the Actic Ocean. Long-term approximates of climate readjust effects top top diversity are daunting because of the facility interactions of transforms on multiple level of the Arctic system.

It is recommended the conservation actions space targeted towards whole systems fairly than individual species. Because system-focused preservation efforts typically focus on limited regions, we should know much more about diversity trends at a high spatial resolution, in certain the distribution of Arctic endemics in order to conserve as plenty of unique species as possible. Also we require to identify the ‘biodiversity hotspots’ – the locations which port high number of unique types due to habitat complexity and also other factors.There is a need for research study to gain a better understanding that the factors and processes that affect diversity. To accomplish this, local and taxonomic gaps should be closed and also time collection are essential to deal with temporal dynamics and changes in biodiversity. However, due to the fact that time is more than likely short before severe impacts of climate adjust will appear, us cannot wait because that a high frequency mapping the the whole Arctic. Instead we imply the establishment, or in some instances continuation, the time collection monitoring at selected website in types rich Arctic locations close to the major gateways, and in some locations distant from the gateways right into the Arctic. We likewise suggest defense of locations with the greatest proportion the Arctic endemic species, as well as the productive polynyas where pelagic-benthic coupling is strong and that are of high prominence for greater taxonomic life

INTRODUCTION

In this chapter, we think about the diversity the invertebrates native the whole benthic, pelagic and sea-ice realms of the Arctic Ocean, extensively defined as areas north of the Bering Strait ~ above the Pacific side and also areas with continuous seasonal sea ice cover top top the Atlantic next (Bluhm et al. 2011a). This corresponds generally to the delineation that the Arctic waters make in Fig. 6.4 in the fish chapter (Christiansen & Reist, chapter 6), however excluding the Bering and Norwegian Seas. Us recognize, however, the the literature cited below does not always follow this delineation.

The existing invertebrate diversity in the Arctic ocean area is the net result of countless factors acting both in historical and also recent time. Like in various other systems ~ above Earth, types diversity in the Arctic is affected by nichebased factors, such as adaptation to various environmental conditions and also by dispersal based factors, such as immigrant from species pools. The relative prestige of these two species of determinants is not always easy to disentangle and may vary v scale and also the level of connectivity to other ecosystems.

Niche-based components like adaptation to various environmental conditions are likely to account for a far-ranging part the biodiversity in the Arctic because it is much from homogeneous. In each of the three realms, invertebrate types inhabit a multitude of various habitats. The pelagic realm contains downwelling or upwelling areas, frontal zones and polynyas through a varying degree of coupling with the benthic kingdom below. The recent irreversible ice-cover in the central Arctic and seasonal ice in the rest of Arctic act as a specific habitat because that sea-ice associated life, and also within the ice realm habitats differ from highly productive ice edge areas to more oligotrophic area in brine networks in the ice, and also the ice-water interface on the underside the the ice.

The sea floor has considerable large scale topographic heterogeneity, for circumstances intertidal seaside areas, semi-enclosed fjords with fjord basins, estuaries of various sizes, an increased shelf zone with a number of canyons (Voronin, St. Anna) and also inner isolated depressions (like Novaya Zemlya Trench), and the deep sea v several containers separated by deep-sea ridges. At smaller sized scales, benthic areas contain different sediment habitats such as sand and mud as well as harder substrata like boulders and bedrocks. The Arctic ocean covers a large area, that which about 50% overlays shelf zones, which in turn constitute 31% of the complete shelf area of the people (Jakobsson et al. 2004). The is renowned that diversity normally increases with the extent of one area (MacArthur & Wilson 1967). If so, we would mean a high full diversity in specific of Arctic shelf fauna loved one to deep sea areas.

A conspicuous attribute of the sea locations of the Arctic is the solid gradient in salinity, both horizontally from river mouths out right into the open sea and also vertically, indigenous close to fresh near the surface to completely marine at depth. Hence, in addition to seasonal ice melt, salinity gradients are highly influenced by freshwater entry from mainly the Russian rivers, but additionally the MacKenzie and Yukon rivers in the western part of the Arctic Ocean. These big rivers together with smaller ones produce estuarine equipment of various spatial sizes which frequently harbor a peculiar collection of varieties adapted to cold water of short salinity. The area the most intensive fresh water influence is related to as a particular zoogeographical unit (Siberian brackish shallow province by Filatova 1957). A consequence of high freshwater inputs is likewise the permanent stratification of the central Arctic ocean with a surface salinity of less than 32‰ and a deep water salinity the 34‰ (Gradinger et al. 2010a), hence providing various habitats because that planktonic invertebrates, because pelagic organisms, like benthic ones, have various tolerances for low salinity.

Furthermore, different parts the the Arctic have various levels of productivity (Michel, thing 14), which likewise may influence diversity (Currie 1991). Productive locations often have much more species than unproductive areas, but the causal relationships are still uncertain (Currie et al. 2004) and also firm proof is likewise lacking for such effects on marine benthic diversity, although hump-shaped relationships have been reported in between chlorophyll a and Arctic benthos wealth (Witman et al. 2008). An instance of an oligotrophic area is the Beaufort Gyre, as contrasted with a fertile area in the Chukchi Sea shelf (Gradinger 2009) or Barents Sea shelf (Sakshaug 1997, Denisenko & Titov 2003).

The Arctic Ocean may be concerned as an open system where the toughness of the connections with adjacent oceans has adjusted over the critical 4 million years. Water currents facilitate dispersal indigenous sub-Arctic and also boreal components of adjacent oceans, v the frame Strait and also the Barents Sea from the Atlantic, and the Bering Strait native the Pacific s (e.g. We¸sławski et al. 2011). If the link with the Pacific has opened and also closed end time as result of varying sea levels, the deep Atlantic entrance has actually been extensively open. At present, over there is some 10 times an ext Atlantic water than Pacific water flowing into the Arctic ocean (Loeng et al. 2005).

In enhancement to habitat complexity and also the prestige of current dispersal from nearby oceans, the turbulent geological history has likewise been essential in shaping existing day diversity of Arctic invertebrates. In the comparatively young Arctic Ocean, the evolutionary beginning of naval invertebrates reflects a Pacific origin dating earlier to the opening of the Bering Strait 3.5 million years earlier (Adey et al. 2008). Throughout many of the Tertiary, the Arctic Ocean region supported a temperate biota, and completely Arctic conditions developed only during the latest component of this period. Sea ice cream cover developed c. 3-5 million years earlier (Briggs 2003). End the critical 3-5 million years, a series of glaciation periods with intermittent de-glaciations has produced an unstable setting with a collection of extinction and immigration occasions shaping current day diversity. These extinction occasions are believed to have precluded extensive local evolution or endemism top top the shelves (Dunton 1992). Furthermore, events throughout the critical 3.5 Myr have actually allowed great re-distributions of species in the boreal part of the northern hemisphere most likely still influence Arctic diversity today. The most pervasive readjust occurred during the so late ice-free Pliocene, after the opening of the Bering Strait, when comprehensive transgressions the invertebrates species across the Arctic arisen (Vermeij 1989, 1991, Mironov & Dilman 2010), largely from the species-rich Pacific center of diversity (Briggs 2003) come the northern Atlantic, an event called ‘The good Trans- Arctic Biotic Interchange’ (Briggs 1995). As contended by Briggs (2007), over there is small evidence native the naval realm that intrusions have reduced native diversity, but rather the they have included to the indigenous diversity, bring about an all at once increased diversity. A result of this significant transfer was as such likely one enrichment of the northern Atlantic pool of species with Pacific species. This swimming pool of species may be the resource of immigration right into the Arctic s in current time.

Against this elevator we mean that invertebrate diversity in the Arctic Ocean has been shaped to a high degree by dispersal based components like immigration and also a low degree of endemism. We mean the Arctic ocean to be conquered by wide-range boreal species. In this respect, that is exciting to compare the levels of endemism in the Arctic v those in the Antarctic, an additional cold an ar with similar glaciation history (Krylov et al. 2008), however which has been much an ext isolated from adjacent oceans by the solid Antarctic Circumpolar current (ACC). The ACC, formed in the Miocene, is the only current on earth extending from the sea surface to the sea floor, unimpeded by any type of landmasses (Hassold et al. 2009). We absolutely would guess a much higher degree that endemism in the Antarctic, which together we will see is in truth the case. Furthermore, provided that connectivity is solid between the Arctic Ocean and the boreal parts of the Pacific and also the Atlantic oceans, we would not suppose a markedly reduced richness in the Arctic, but fairly similar level of varieties richness as in the various other oceans, at least in proximity to the 2 gateways.

In enhancement to the natural structuring factors, diversity fads in the Arctic Ocean likely are influenced by sports in sampling methods and sampling frequency. For instance, some areas have been extensively investigated for more than a century (Barents Sea), while various other less accessible areas (deep Arctic basins) have actually been fairly poorly studied. This create a an obstacle when estimating complete numbers of types in the Arctic.

The main questions handle in this testimonial are:

Is the marine invertebrate diversity in the Arctic s impoverished contrasted with surrounding areas? space there large scale diversity fads within the AO area that deserve to be attributed to dispersal rather than niche adaptation? Is the unstable geological background and openness to adjacent oceans mirrored by a low degree of endemism? are there ‘hotspot’ areas that through virtue of their types diversity should be protected? deserve to we predict what the results of global warming on invertebrate varieties diversity?

CONCLUSIONS and RECOMMENDATIONS

Conclusions

The Arctic ocean area hosts c. 5,000 types of marine invertebrates, i beg your pardon is a similar level as is discovered in the various other polar environment, Antarctica, and is taken into consideration intermediate on a global scale. Arthropoda, greatly crustaeans, is the most speciose group and also does no exhibit the decreasing richness with boosting latitude as found in Mollusca.

Although the Arctic contains great morphological heterogeneity and a vast number of environmental gradients, offering the possibility for comprehensive niche adaptation, Arctic diversity seems greatly a an outcome of extinctions and dispersal occasions over the last c. 4 million years. Most types have origins from exterior the Arctic, and overall over there are couple of species endemic come the Arctic. The level of endemism varies greatly among different taxonomic groups, wherein bryozoans for instance seem to have actually a fairly high degree of endemism perhaps partly as result of their sessile habits and, maybe much more importantly, bad dispersal ability.

The glaciation background of the two polar seas seems reasonably similar, however unlike the Antarctic which has a long background of geographic isolation, the Arctic has actually been, and is, open up towards the two significant oceans, the Pacific and the Atlantic, back the toughness of the connections have varied end the last c. 4 million years. This is a likely explanation for the really low degree of endemism in the Arctic contrasted with the Antarctic. Today’s biogeographic vehicle drivers of Arctic diversity are clearly seen in the distribution of origins in relationship to the two major gateways into the Arctic, i.e. Indigenous the Atlantic and also Pacific Oceans, respectively. Top top the continent shelves, the proportions of present-day Pacific and also Atlantic species decrease with boosting distance native the Bering Strait and also the NE Atlantic, respectively. Present inventories show that the Barents Sea has the highest types richness, gift ‘enriched’ by boreal and also sub-Arctic species. Today’s Arctic deep-sea floor is most closely related to the existing North Atlantic fauna, i beg your pardon in a geology time perspective consists of a strong Pacific influence. The local species richness is highest possible in Arctic areas close come the two gateways, the Chukchi Sea for the Pacific and, also higher, the Barents Sea/ Kara Sea for the Atlantic. These observations together with the circulation patterns that zoogeographical affinities suggest the prestige of dispersal with the gateways right into the Arctic Ocean.

While areas within the Arctic through high types richness have been identified, such together the Barents Sea, it is unsure if there are real ‘hotspots’ the diversity, i.e. Areas with high diversity of unique or endemic varieties in the Arctic. This is because many of these species may be numerous in waters to the south and thus not unique. The polynyas, ice-free areas within the area that sea ice, might be hot spots in state of power flow (Michel, thing 14), wherein benthic and also pelagic invertebrates administer food for dense aggregations of birds and mammals.

There are already clear signs of an international warming effects on invertebrates, for circumstances northward growth of number of boreal species. As would be predicted, this borealization has actually so far arisen in the margins of the Arctic Ocean, generally at the two major gateways come the boreal parts of the Atlantic and Pacific. The quickly melting sea ice means loss of habitat for sympagic fauna.

In addition to temperature rise, an international change will certainly acidify the oceans, and also there is a great concern the this will certainly negatively impact calciferous invertebrates like several benthic and also pelagic molluscs. Experimental work shows that acidification hampers shell development in wing snails.

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recommendations

It is recommended the conservation actions are target towards totality systems quite than separation, personal, instance species. Specifics there space urgent requirements to record and understand Arctic biodiversity patterns and processes to have the ability to prioritize conservation efforts.

We need much more inventories

This contains the require to know where the highest diversity wake up in the Arctic, particularly for endemic species, in bespeak to conserve as numerous unique species as possible. Hence, there is a need for:

in-depth surveys the diversity in hitherto understudied areas like the east Siberian Sea and also the Canadian Arctic, together with deep-sea locations of the main Arctic Basin and also at the Arctic-Atlantic frontier. Research studies are likewise needed in the shallow subtidal to 12 meters, which still is one understudied area. Enhanced sampling and also taxonomic effort on poorly investigated groups, consisting of several among the meiofauna. Establishing and continuing several observation sites for permanent monitoring of marine ecosystems in different parts the the Arctic appropriate to attain a an ext holistic view of the changing Arctic. The existing biological stations together with marine protected areas could offer as a base for such long-term observations. A priority focus on continual time series monitoring at sites in the species-rich Arctic areas close to the major gateways, and also in some locations distant indigenous the gateways. Given the likelihood of small time before much more severe climate change effects will certainly be manifested, this involves both the establishment of some new sites and the extension of monitoring at existing website such as the White Sea biological Station, the Greenland Ecosystem monitoring in Godthåbsfjorden in W Greenland and Young Sund in NE Greenland, and also the HAUSGARTEN observatory west the Svalbard. The variety of observatories in both deep and shallow waters needs to be raised to encompass a large spectrum of experimentation areas and also communities. Repeated sampling should be conducted in the areas of previous studies, favor those the Golikov (1990, 1994a, 1994b, 1994c) in the Laptev and also West Siberian Seas. This studies carry out a enough background come evaluate any changes in recent neighborhood structure and composition.

We need research to understand maintenance of diversity so it is recommended:

to quantify immigration prices of boreal species into the Arctic and also investigate the feasible influence of worldwide warming on this rates. To investigate whether or not immigration that boreal varieties ‘enriches’ native diversity, and also whether immigrants have a an adverse influence top top the indigenous fauna. To additional implement molecule taxonomy to uncover the most likely presence that sibling types and come reveal historic migration patterns. The most optimistic approximates predict a diversity of ‘molecular work taxonomic units’ as much as three times the variety of described morphological species, also in such well studied teams as the Polychaeta (Carr et al. 2011). To investigate how increased primary production, which might be one repercussion of shrinking ice cream cover, affects varieties diversity both in the pelagic and the benthic systems. This could be performed in connection with polar fronts and also productive polynyas. Come investigate exactly how climate adjust influences alters in biogeographic distributions, specifically the borealization process, habitat loss for sympagic fauna and the circulation of calciferous fauna.

Based on existing knowledge us recommend defense of the adhering to areas:

Polynyas i m sorry are locations known to be vital for preserving seabird and also mammal populations. These locations should be closed because that fishing and petroleum extraction. The latter is necessary due to the fact that it is virtually difficult to clean up oil in waters with damaged ice. Huge estuaries, i m sorry harbor numerous of the distinctive Arctic species.