**2. Ecosystem services provided**

**Category of macrobenthic community**

Macro-algae dominated (MA)

Unconsolidated Sediment dominated (US)

Hard substrate dominated (HS)

Reef species dominated (RS)

**Examples of subunits identified by** 

Calcareous algal bed Canopy-forming algal bed

Coralline/crustose algal bed

Tunneling megafauna Burrowing anemones

Other non-molluscan invertebrate bed

Mineral/wood boring

Diverse colonizers Attached tubebuilding fauna Mobile crustaceans and gastropods on hard or mixed substrates Sessile/attached molluscs and/ or non-molluscan invertebrate communites

Branching/columnar/ foliose/plate/table coral reef

Encrusting coral reef Massive coral reef Shallow molluscan dominated Mesophotic reef

ecosystems services provided [2, 4, 5].

fauna

**Ecosystem services provided Direct/**

Pharmaceutical compounds derived from marine

Capturing and filtering sediments and organic wastes in transit from inland regions to the ocean

Pharmaceutical compounds derived from marine

Capturing and filtering sediments and organic wastes in transit from inland regions to the ocean

Primary production of benthic algae, high levels of secondary production and great diversity in benthic animals, provide forage for crabs, finfish

Pharmaceutical compounds derived from marine

Capturing and filtering sediments and organic wastes in transit from inland regions to the ocean Hard substrate for attached animals, provides finfish, crustacean and shorebird forage Filters suspended material from the water for

Sediment Stabilization erosion control via wave

High levels of secondary production and great diversity in benthic animals, forage for crabs,

Pharmaceutical compounds derived from marine

Soil formation, photosynthesis and nutrient cycling

Scuba diving and other nature-based tourism

**Table 3.** Marine coastal macro-biotic assemblages that comprise the benthic component for CMECS standards and

**Provisioning services**

algae and invertebrates **Regulating services**

**Provisioning services**

algae and invertebrates **Regulating services**

Sediment stabilization

**Provisioning services**

algae and invertebrates **Regulating services**

improved water quality.

finfish and shorebirds

**Provisioning services** Provides building materials Fisheries and associated industries

algae and invertebrates **Supporting services**

**Cultural services**

reduction.

and shorebirds

**indirect**

Indirect [17]

Indirect [18]

Indirect [19–21]

Direct and indirect [22–30]

**Supporting literature**

**CMECS**

66 Ecosystem Services and Global Ecology

Kelp forest

Bivalve bed

Ecosystem services provided by coastal macroinvertebrates assemblages include both direct and indirect benefits (**Table 3**). Marine ecosystem services provided by these groups of macrofauna and flora that directly provide benefit encompass the services that provide food, medicine, recreation, support of fisheries, and storm protection. Other ecosystem services are less tangible, and so more difficult to documents, such as the habitat's role in absorbing carbon from the atmosphere—a positive effect on our global climate. In addition to the economic supports coastal areas provide, human attitudes, beliefs, behaviors, customs, and traditions are often associated with the surrounding nature and environmental quality. These cultural ecosystem services are often neglected but are a significant feature of the services that could be lost if the biodiversity of these assemblages becomes threatened.

Ecosystem services provided by marine coastal zones are classified by four categories (as they are for most identified ecosystem services). The four categories identified are Provisional, Supporting, Regulating, and Cultural Services. Provisioning services include food, water, and products such as building materials from mangrove and coral reef, and pharmaceutical compounds derived from marine algae and invertebrates. Supporting services include soil formation, primary productivity, and nutrient cycling; coastal habitats such as seagrass beds and mangroves are important nursery areas for the young stages of fishes and invertebrates that support coastal communities and commercial and recreational fisheries. Regulating service**s** include regulation of climate; natural hazards such as floods, disease, wastes, and water quality, coastal wetlands play an important role in water quality regulation by capturing and filtering sediments and organic wastes in transit from inland regions to the ocean. On a global scale, fixation of atmospheric carbon by oceanic algae and its eventual deposition in deep water represents an important part of the global carbon cycle and thus influences climate trends. Cultural services include recreational, esthetic, and spiritual benefits derived from nature. Coastal tourism is the fastest-growing sector of the global tourism industry [9], and is a major part of the economies of many small island-developing nations. Moreover, the cultures and traditions of many coastal peoples are intimately tied to the marine ecosystems on which they depend.

Coastal marine ecosystem services are also provided directly, through human use or experience of the service or indirectly, via impacts of supporting and regulating services on other services and environments. Cultural ecosystems services of a variety provided by macroinvertebrate communities near the coasts include those tied to the culture and traditions of coastal peoples in many developing nations by supporting local small scale fisheries, recreational and esthetic services across the globe as a source of natural interest and exploration for people of all ages, scientific and sociological endeavors, and ecotourism opportunities like scuba diving and sport fishing.

**Vascular plant dominated habitat (VP)**

CMECS biotic group: sea grass

CMECS biotic subclass: emergent tidal marsh and biotic group: brackish marsh

CMECS biotic group: tidal mangrove forest and tidal mangrove shrubland biotic group. Mangrove forests

these ecosystems.

bed

**Important species of the** 

The approximately 72 species of sea grasses are commonly divided into four main groups: Zosteraceae, Hydrocharitaceae, Posidoniaceae and Cymodoceaceae. The major sea grasses include Cymocedea sp., Halodule sp., Thalassia sp., Halophilla sp., Vallisnera sp., Ruppia sp., Phyllospadix sp., and **Assemblage biotic structure**

Ecosystem Services Provided by Benthic Macroinvertebrate Assemblages in Marine Coastal Zones

Seagrass beds are complex structural habitats that provide refuge and foraging opportunities for abundant and diverse faunal communities. Slow moving mollusks, larger crustaceans, sponges and echinoderms are all commonly found associated with these areas

Fish and shrimp come into salt marshes looking for food or for a place to lay their eggs. Larger decapods and oysters are also key species that depend on the tidal

marshes

The list of common species supported by mangels is line and includes: barnacles, oysters, mussels, sponges, worms, snails and small fish live around the roots. Mangroves water contain crabs, jellyfish and are a nursery to many juvenile fish

This VP category of biota include these groups—biotic group: seagrass bed—tidal aquatic vegetation beds dominated by any number of seagrass or eelgrass species; biotic subclass: emergent tidal marsh—communities dominated by emergent, halophytic, herbaceous vegetation; and biotic group: tidal mangrove forest—tidally influenced, dense, tropical or subtropical forest with a shore zone dominated by true mangroves (and associates) that generally are 6 m or taller [4].

**Table 4.** Vascular plant dominated habitat (VP) CMECS definition and important species and dominancy relations in

**Key biodiversity aspects of assemblage**

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69

There are six seagrass bioregions according to Short et al. [2, 32] which is the current standard used by the international seagrass research community. These six bioregions are Temperate North Atlantic (I), Tropical Atlantic (II), Mediterranean (III), Temperate North Pacific (IV), Tropical Indo-Pacific (V), and Temperate Southern Ocean (VI), and are based on assemblages of taxonomic groups of seagrasses in temperate and tropical areas and the physical

separation of the world's oceans

Marine and freshwater species occur in the intertidal zone of coastal estuaries. These areas and are usually intermixed with intertidal mudflats that are rich with invertebrates and seaweeds. These transitional zones are key nursery areas for many commercial species

Tidally influenced, dense, tropical or subtropical forest with a shore zone dominated by true mangroves (and associates) that generally are 6 m or taller. Dwarf shrub and short mangroves are placed in the tidal mangrove shrubland biotic group. Mangrove forests occur along the sheltered coasts of tropical latitudes of the Earth, and are commonly found on the intertidal mud flats along the shores of estuaries, usually in the region between the salt marshes and seagrass beds and may extend inland along river courses where tidal amplitude is high. Also, mangrove cays may occur within the lagoon complex of

barrier reefs

**assemblage**

Zostera sp.

Salt bushes and **grasses** are the dominant plants, with Sparina sp., Juncus sp. and Salicornia sp. common in the plant Communities. The plants are dominated by emergent, halophytic, herbaceous vegetation (with occasional woody forbs or shrubs) along low-wave-energy, intertidal areas of estuaries and rivers. Also brackish marshes dominated by species with a wide range of salinity tolerance

Mangroves are not a taxonomic group but identified by their salt tolerance. Several tree and shrub species are the structural basis for these tropical vegetation that supports many diverse invertebrate species as

juveniles

Macroinvertebrate assemblages form the basis for the majority of the coastal marine services as illustrated by the biotopes that are defined by the species that characterize the biotic components.

Changes in the local coastal marine environments following perturbations are myriad and occur in both the short term and long term spatial and temporal realms [10–16]. Changes to these environments, either by a natural or human induced physical change can impact the resident macro-fauna assemblages and the ecosystem services they provide in a numerous of ways. The majority of the threats identified to these communities is heightened after an environmentally perturbing event that is of a large scale, and as documented are altered long-term for certain near shore biotopes (see **Table 3**).

Delineating the impacts of large scale events on coastal marine benthic invertebrate assemblages are identified in literature from natural hazards such as hurricane and earthquake events [17–19].

To examine the global effects that result in terms of the macro-benthic assemblages, one needs to characterize each major habitat type and synthesize current findings with related environmental disturbance known impacts. Based on the CMECS, major macroinvertebrate assemblages can be categorized as follows to examine the ecosystem services provided and possible impacts after a major change (see **Table 1**).

#### **2.1. Vascular plant dominated habitat (VP)**

Three subsets make up the VP biota (see **Table 4**). Sea grass beds, tidal marshes, and mangels globally provide significant ecosystem services but also experience the greatest threats from human activity. Seagrass beds, are a lesser known area for many, given their submergence and often hidden location for most observers. Rooted flowering aquatic grasses dominate this assemblage of biota. These sea grasses are significant refugia for the macroinvertebrate assemblages that depend on their bioprocessed and are dominated by turtle grass species in the tropical zones (*Thalassia* spp., *Halodule* spp., *Syringodium* spp., etc.), and *Posidonia* spp., *Ruppia* spp., and *Zostera* spp. in the more temperate waters [20]. These habitats stabilize and protect the shorelines, but additionally support a diverse array of macroinvertebrates. These various community members in tern support the higher order consumers and thus, support fisheries in both adult and juvenile stages. A significant feature of these (VP) areas is that they provide a complex structural habitat that serves as a nursery area for many commercially important species that might not depend on these areas beyond the nursery stages. An often overlooked global ecosystem service provided by these VP assemblages are the carbon stored in sediments from these coastal ecosystems and is known as "blue carbon" because it is stored in the marine environs.


Coastal marine ecosystem services are also provided directly, through human use or experience of the service or indirectly, via impacts of supporting and regulating services on other services and environments. Cultural ecosystems services of a variety provided by macroinvertebrate communities near the coasts include those tied to the culture and traditions of coastal peoples in many developing nations by supporting local small scale fisheries, recreational and esthetic services across the globe as a source of natural interest and exploration for people of all ages, scientific and sociological endeavors, and ecotourism opportunities like scuba diving and sport fishing.

Macroinvertebrate assemblages form the basis for the majority of the coastal marine services as illustrated by the biotopes that are defined by the species that characterize the biotic

Changes in the local coastal marine environments following perturbations are myriad and occur in both the short term and long term spatial and temporal realms [10–16]. Changes to these environments, either by a natural or human induced physical change can impact the resident macro-fauna assemblages and the ecosystem services they provide in a numerous of ways. The majority of the threats identified to these communities is heightened after an environmentally perturbing event that is of a large scale, and as documented are altered long-term

Delineating the impacts of large scale events on coastal marine benthic invertebrate assemblages are identified in literature from natural hazards such as hurricane and earthquake

To examine the global effects that result in terms of the macro-benthic assemblages, one needs to characterize each major habitat type and synthesize current findings with related environmental disturbance known impacts. Based on the CMECS, major macroinvertebrate assemblages can be categorized as follows to examine the ecosystem services provided and possible

Three subsets make up the VP biota (see **Table 4**). Sea grass beds, tidal marshes, and mangels globally provide significant ecosystem services but also experience the greatest threats from human activity. Seagrass beds, are a lesser known area for many, given their submergence and often hidden location for most observers. Rooted flowering aquatic grasses dominate this assemblage of biota. These sea grasses are significant refugia for the macroinvertebrate assemblages that depend on their bioprocessed and are dominated by turtle grass species in the tropical zones (*Thalassia* spp., *Halodule* spp., *Syringodium* spp., etc.), and *Posidonia* spp., *Ruppia* spp., and *Zostera* spp. in the more temperate waters [20]. These habitats stabilize and protect the shorelines, but additionally support a diverse array of macroinvertebrates. These various community members in tern support the higher order consumers and thus, support fisheries in both adult and juvenile stages. A significant feature of these (VP) areas is that they provide a complex structural habitat that serves as a nursery area for many commercially important species that might not depend on these areas beyond the nursery stages. An often overlooked global ecosystem service provided by these VP assemblages are the carbon stored in sediments from these coastal ecosystems and is known as "blue carbon" because it is stored

components.

68 Ecosystem Services and Global Ecology

events [17–19].

in the marine environs.

for certain near shore biotopes (see **Table 3**).

impacts after a major change (see **Table 1**).

**2.1. Vascular plant dominated habitat (VP)**

This VP category of biota include these groups—biotic group: seagrass bed—tidal aquatic vegetation beds dominated by any number of seagrass or eelgrass species; biotic subclass: emergent tidal marsh—communities dominated by emergent, halophytic, herbaceous vegetation; and biotic group: tidal mangrove forest—tidally influenced, dense, tropical or subtropical forest with a shore zone dominated by true mangroves (and associates) that generally are 6 m or taller [4].

**Table 4.** Vascular plant dominated habitat (VP) CMECS definition and important species and dominancy relations in these ecosystems.

Perturbations to sea grass beads, and impacts of large-scale weather events (such as tsunamis for example) have indicated that seagrass beds are resilient to perturbations. The findings regarding the macroinvertebrate diversity of major taxonomic groups is less positive as it is most likely that the biota that are part of these VP areas are tied to density of vegetation [33, 34].

Another category of VP are the salt marshes of the temperate and tropic areas. These prominent vegetated coastal habitats and their proximal coastal areas are well known to of high value as a nursery grounds. Their value as land run off filters is significant. Lesser recognized for the importance of these areas is the high diversity of macroinvertebrate species. As a nursery grounds these areas are significant to both commercial and sport fishing activities. Perturbing events that shift the sediments and inundate the area with fresher water draining of streams or highly saline off shore water, can load toxic land run off, scour vegetative areas, and/or deposit debris that compromises the health of these habitats and thus the macroinvertebrate assemblages [22]. The transitional nature of these areas between land and ocean make them particularly subject to physical changes such as those often seen by development.

Mangels, also known as mangrove habitats, are a group of coastal tropical halophytes that provide structural complexity and protect the shoreline by stabilizing sediments. Because the halophytes that form the basis of these assemblages are from various taxonomic groups, different environmental factors (beyond salinity) can impact their viability. Development of these areas often occurs given the tropical climate and attractiveness for tourism, the development of these shorelines destroys these areas. Tsunami impacts have been examined for some habitats, and it appears mangroves may never fully recover from events that result in the extirpation of these halophytes [23, 35]. Loss of the mangroves mean loss of the ecosystem services they provide in addition to losing the associated macroinvertebrate fauna. As with the salt marshes of the temperate zones, mangels are a significant nursery for many fish, both sport and commercially important fisheries can be impacted by their loss.

#### **2.2. Macroalgae dominated habitat (MA)**

**Table 5** offers an overview of the macroalgae dominated habitat. Kelp forests are temperate near-shore habitats that support diverse macro-invertebrate assemblages. There are other MA assemblages but the kelp forest are the most dominant example from a global perspective and also provide significant ecosystems services. Both the primary productivity and the structural complexity of their fronds are key factors in support of the whole ecosystem. Kelp, in particular the brown kelps, are well adapted to be resilient against strong currents, they are tolerant to storm surges. Interestingly, they appear to be prone to concentration radioactive material, after the tsunami of the Indian Ocean in 2010 radioactive were found in the kelp off the California coast in the weeks after the tsunami event in Japan. The materials did not remain in the kelp for a long period of time. This suggests they are able to be expelled into the biotope, but as a result presumably to be taken up by other organisms [24, 25].

macrobenthos that provides key services. These small and relatively overlooked groups of invertebrates turn the sediments and process organics. These fine soils and the high degree of organics and detritus associated can be harmed by strong surges and deposited elsewhere smothering other areas with hypoxic sludge [26]. These US areas are frequently dredged to replenish shorelines and considered to be unattractive. Overlooking the services they provide would be an error. Sand habitats are teeming with diversity despite the common assumption that they do not, the macro-invertebrates present in these tidal zones show resilience to storm events and recover quickly after a Tsunami event [27]. The recovery of the macro-invertebrates in these assemblages can be quick if recruitment areas adjacent are not impacted. The planktonic nature of the larvae of most invertebrates living in these areas allows for quick recruitment and recovery after a large environmental change like the shifting of sediments from a beach restoration

**Table 6.** Biotic class: soft sediment fauna—areas that are characterized by fine unconsolidated substrates (sand, mud) and that are dominated in percent cover or in estimated biomass by infauna, sessile epifauna, mobile epifauna, mobile fauna that create semi-permanent burrows as homes, or by structures or evidence associated with these fauna [4].

**Assemblage biotic structure Key biodiversity aspects of** 

**Assemblage biotic structure Key biodiversity aspects of** 

**assemblage**

substrates

Subtidal soft bottom habitats are diverse based on distinct organism assemblages that are influenced by differences in substrate type (sand vs. mud), organic content and bottom depth. Most of these fauna possess specialized organs for burrowing, digging, embedding, tube-building, anchoring, or locomotory activities in soft

Kelp forests provide both primary productivity and a structural base for many species. The holdfasts as well as the surface mats of kelp fronds support thousands of invertebrate individuals, including polychaetes, amphipods, decapods, and ophiuroids. Larger vertebrates

Ecosystem Services Provided by Benthic Macroinvertebrate Assemblages in Marine Coastal Zones

frequent these areas

Macroalgal communities can exist at all depths within the photic zone, on diverse substrates, and across a range of

**Table 5.** MA biotic subclass: benthic macroalgae—aquatic beds dominated by macroalgae attached to the substrate, such

Species tunnel freely within the sediment or embed themselves wholly or partially in the sediment (e.g., tilefish burrows, lobster burrows). Other organisms such as crustaceans, echinoderms and mollusks may be locally abundant

**assemblage**

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71

Many macroalgal types and communities have low temporal persistence and can bloom and die-back within short periods. This aspect of macroalgae impact the nature of the ecosystem services at an given time

or large-scale weather event.

**Macroalgae dominated habitat (MA)**

CMECS biotic subclass: benthic macroalgae

**Unconsolidated sediment dominated habitat** 

CMECS soft sediment fauna

**(US)**

**Important species of the** 

Aquatic beds dominated by macroalgae attached to the substrate, such as kelp (*Fucus* sp., *Macrocystis* sp.), intertidal fucoids, and other calcareous algae

**Important species of the** 

Often dominated in percent cover or in estimated biomass by infauna, sessile epifauna, mobile epifauna, mobile fauna that create semipermanent burrows as homes, or by structures or evidence associated with

**assemblage**

these fauna

**assemblage**

energy and water chemistry regimes [4].

as kelp, intertidal fucoids, and calcareous algae.

#### **2.3. Unconsolidated sediment dominated habitat (US)**

Perhaps one of the most overlooked macro-faunal assemblages are mud flats and other fine sediment habitats (**Table 6**). Although not at all evident to most, these areas support infaunal


Perturbations to sea grass beads, and impacts of large-scale weather events (such as tsunamis for example) have indicated that seagrass beds are resilient to perturbations. The findings regarding the macroinvertebrate diversity of major taxonomic groups is less positive as it is most likely that the biota that are part of these VP areas are tied to density of vegetation [33, 34].

Another category of VP are the salt marshes of the temperate and tropic areas. These prominent vegetated coastal habitats and their proximal coastal areas are well known to of high value as a nursery grounds. Their value as land run off filters is significant. Lesser recognized for the importance of these areas is the high diversity of macroinvertebrate species. As a nursery grounds these areas are significant to both commercial and sport fishing activities. Perturbing events that shift the sediments and inundate the area with fresher water draining of streams or highly saline off shore water, can load toxic land run off, scour vegetative areas, and/or deposit debris that compromises the health of these habitats and thus the macroinvertebrate assemblages [22]. The transitional nature of these areas between land and ocean make them particularly subject to physical changes such as those often seen by development.

Mangels, also known as mangrove habitats, are a group of coastal tropical halophytes that provide structural complexity and protect the shoreline by stabilizing sediments. Because the halophytes that form the basis of these assemblages are from various taxonomic groups, different environmental factors (beyond salinity) can impact their viability. Development of these areas often occurs given the tropical climate and attractiveness for tourism, the development of these shorelines destroys these areas. Tsunami impacts have been examined for some habitats, and it appears mangroves may never fully recover from events that result in the extirpation of these halophytes [23, 35]. Loss of the mangroves mean loss of the ecosystem services they provide in addition to losing the associated macroinvertebrate fauna. As with the salt marshes of the temperate zones, mangels are a significant nursery for many fish, both

**Table 5** offers an overview of the macroalgae dominated habitat. Kelp forests are temperate near-shore habitats that support diverse macro-invertebrate assemblages. There are other MA assemblages but the kelp forest are the most dominant example from a global perspective and also provide significant ecosystems services. Both the primary productivity and the structural complexity of their fronds are key factors in support of the whole ecosystem. Kelp, in particular the brown kelps, are well adapted to be resilient against strong currents, they are tolerant to storm surges. Interestingly, they appear to be prone to concentration radioactive material, after the tsunami of the Indian Ocean in 2010 radioactive were found in the kelp off the California coast in the weeks after the tsunami event in Japan. The materials did not remain in the kelp for a long period of time. This suggests they are able to be expelled into the biotope,

Perhaps one of the most overlooked macro-faunal assemblages are mud flats and other fine sediment habitats (**Table 6**). Although not at all evident to most, these areas support infaunal

sport and commercially important fisheries can be impacted by their loss.

but as a result presumably to be taken up by other organisms [24, 25].

**2.3. Unconsolidated sediment dominated habitat (US)**

**2.2. Macroalgae dominated habitat (MA)**

70 Ecosystem Services and Global Ecology

Macroalgal communities can exist at all depths within the photic zone, on diverse substrates, and across a range of energy and water chemistry regimes [4].

**Table 5.** MA biotic subclass: benthic macroalgae—aquatic beds dominated by macroalgae attached to the substrate, such as kelp, intertidal fucoids, and calcareous algae.


**Table 6.** Biotic class: soft sediment fauna—areas that are characterized by fine unconsolidated substrates (sand, mud) and that are dominated in percent cover or in estimated biomass by infauna, sessile epifauna, mobile epifauna, mobile fauna that create semi-permanent burrows as homes, or by structures or evidence associated with these fauna [4].

macrobenthos that provides key services. These small and relatively overlooked groups of invertebrates turn the sediments and process organics. These fine soils and the high degree of organics and detritus associated can be harmed by strong surges and deposited elsewhere smothering other areas with hypoxic sludge [26]. These US areas are frequently dredged to replenish shorelines and considered to be unattractive. Overlooking the services they provide would be an error.

Sand habitats are teeming with diversity despite the common assumption that they do not, the macro-invertebrates present in these tidal zones show resilience to storm events and recover quickly after a Tsunami event [27]. The recovery of the macro-invertebrates in these assemblages can be quick if recruitment areas adjacent are not impacted. The planktonic nature of the larvae of most invertebrates living in these areas allows for quick recruitment and recovery after a large environmental change like the shifting of sediments from a beach restoration or large-scale weather event.

#### **2.4. Hard substrate dominated biotopes (HB)**

This category includes artificial reefs (human places). Macroinvertebrates that colonize hard substrates are generally in competition for space to attach to in the larval stages (**Table 7**). After a large weather event with strong currents or storm surges, boulders and cobble are scattered, and rocky shores could be scoured by these water movements or also by thermal pollution. New human created habitat can also occur in the form of unintentional deposition of sediments of large size and intentional artificial reef type habitat (many recreational charter captains create and maintain their own reefs by submerging solid structures as a base such as old chicken coops or shopping carts to create a reef that they can locate to support their businesses). Little is known about the specific effects on these types of macroinvertebrate assemblages that populate the HB areas. The high larval settling needs and competition for hard places for larvae to settle, these coastal assemblages may be the first to recover after a storm event [28, 29].

> coral and other carbonate producers and physical processes such as waves and currents that modulate ecological processes and redistribute carbonate material within reef systems." [4]. These areas are well known as the most diverse and likely also provide the most significant oceanic ecosystems services as a result. These areas additionally provide the most varied in terms of type of services as they provide more esthetic and ecotourism support to a greater degree than any other macroinvertebrate assemblages. Yet these significant areas are also some of the most delicate and threatened habitats. Coral bleaching can occur as the result of numerous stressors and large-scale weather events can devastate large regions from both abiotic and biotic stressors [27, 30, 36, 37]. Of all the marine coastal biotopes, literature suggests it is the coral dependent fauna that can be devastated from large-scale changes such as those that occur after a tsunami event, but more investigation is needed to determine if recovery is possible.

> In order to be classified as reef biota, colonizing organisms must be judged to be sufficiently abundant to construct

**Table 8.** Biotic class: reef biota areas dominated by reef-building fauna, including living corals, mollusks, polychaetes

Macroinvertebrates from all taxonomic groups comprise

the assemblages

identifiable biogenic substrates. When not present in densities sufficient to construct reef substrate [4].

**Assemblage biotic structure Key biodiversity aspects of** 

Ecosystem Services Provided by Benthic Macroinvertebrate Assemblages in Marine Coastal Zones

**assemblage**

fauna as well

Nearly 25% of all known marine species are associated with coral reefs the rich biodiversity covers most taxonomic groups and has many complex interactions with adjacent

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73

**Coral reef dominated habitat (CS)**

CMECS biotic subclass: shallow/mesophotic coral reef biota

or glass sponges.

**Important species of the assemblage**

Stony (scleractinian) corals and crustose coralline red algae

Although much more study is needed to determine specific impacts in local areas, globally speaking, macroinvertebrate assemblages do recover after severe natural environmental perturbing events in general, but do so differentially. Anthropogenic perturbations that destroy the physical support of the biotic assemblages are less likely to recover, generally due to development of the shoreline and drainage of these areas. More work is needed to verify the longer term impacts that natural events have from habitat perturbation to ecosystem service losses, anthropogenic impacts have yet to be documented to a great degree from a global perspective. Human impacts are more often permanent, so prevention of further threats are the main reason more knowledge and awareness of the ecosystem services is crucial [32, 38–49].

In general, vascular plant dominated biotopes (VP) seem resilient (except for mangroves) after an environmentally perturbing events with recovery well underway in one annual cycle. Macroalgae/protista dominated biotopes (MA) may be impacted even at great distance from source of perturbations or related contamination little is known about the effects on the fauna they support. Both unconsolidated substrate dominated (US) and hard substrate dominated biotopes (HB) are noted to have recovery times close to that identified for sea grass areas. Reef species dominated areas (CS) are subject to many environmental stressors, the physical and chemical changes that result from an environmental perturbing event impact the corals species negatively but the fauna that rely on the physical structural components may shift in diversity but do persist. Defining recovery in terms of the macro invertebrate assemblage would seem

#### **2.5. Coral reef dominated habitat (CS)**

The highest biodiversity in the list of coastal macro-invertebrate assemblages is not surprising to be the coral reefs and related invertebrate reef macroinvertebrate assemblages (**Table 8**). As identified by CMECS "The Shallow/Mesophotic Coral Reef Biota are largely based on the growth form of the dominant corals that (a) reflect differences in environmental conditions and (b) provide varied habitat circumstances (such as increased cover) for associated fish and invertebrate species. The same coral species can present different growth forms under different environmental circumstances. For example, *Acropora* sp. can have both branching and table growth forms, depending on the environment. To reflect the differences in the physical and biological environments, the same species may be used to define communities in more than one coral group the interaction between ecological processes responsible for the growth of


**Table 7.** Biotic subclass: attached fauna—areas characterized by rock substrates, gravel substrates, other hard substrates, or mixed substrates that are dominated by fauna which maintain contact with the substrate surface, including firmly attached, crawling, resting, interstitial, or clinging fauna [4].


**2.4. Hard substrate dominated biotopes (HB)**

72 Ecosystem Services and Global Ecology

**2.5. Coral reef dominated habitat (CS)**

**Important species of the assemblage**

Dominated by fauna which maintain contact with the substrate surface, including firmly attached, crawling, resting, interstitial, or clinging fauna. Fauna may be found on, between, or under rocks or other hard substrates or substrate

mixes

attached, crawling, resting, interstitial, or clinging fauna [4].

**Hard substrate dominated (HB)**

CMECS biotic subclass: attached fauna/ anthropogenic origin hard substrates

This category includes artificial reefs (human places). Macroinvertebrates that colonize hard substrates are generally in competition for space to attach to in the larval stages (**Table 7**). After a large weather event with strong currents or storm surges, boulders and cobble are scattered, and rocky shores could be scoured by these water movements or also by thermal pollution. New human created habitat can also occur in the form of unintentional deposition of sediments of large size and intentional artificial reef type habitat (many recreational charter captains create and maintain their own reefs by submerging solid structures as a base such as old chicken coops or shopping carts to create a reef that they can locate to support their businesses). Little is known about the specific effects on these types of macroinvertebrate assemblages that populate the HB areas. The high larval settling needs and competition for hard places for larvae to

settle, these coastal assemblages may be the first to recover after a storm event [28, 29].

The highest biodiversity in the list of coastal macro-invertebrate assemblages is not surprising to be the coral reefs and related invertebrate reef macroinvertebrate assemblages (**Table 8**). As identified by CMECS "The Shallow/Mesophotic Coral Reef Biota are largely based on the growth form of the dominant corals that (a) reflect differences in environmental conditions and (b) provide varied habitat circumstances (such as increased cover) for associated fish and invertebrate species. The same coral species can present different growth forms under different environmental circumstances. For example, *Acropora* sp. can have both branching and table growth forms, depending on the environment. To reflect the differences in the physical and biological environments, the same species may be used to define communities in more than one coral group the interaction between ecological processes responsible for the growth of

> Depending on water depth, light penetration, wave energy, and other physical and biological processes, algae and macroalgae can provide extensive or sporadic cover and food for other species in the nearshore subsystem. Many attached fauna are suspension feeders and feed from the water column. Other attached fauna are benthic feeders, including herbivores, predators, detritivores, deposit feeders, and omnivores

**Table 7.** Biotic subclass: attached fauna—areas characterized by rock substrates, gravel substrates, other hard substrates, or mixed substrates that are dominated by fauna which maintain contact with the substrate surface, including firmly

**Assemblage biotic structure Key biodiversity aspects of** 

**assemblage**

pipelines

Rocky subtidal habitat includes all hard substrate areas of the ocean bottom. Anthropogenic reefs include any areas where hard, persistent material has been placed either purposely or accidentally by humans. Examples include rock jetties at the entrance to many bays, shipwrecks, anchoring systems for renewable energy projects, and unburied portions of underwater cables or

In order to be classified as reef biota, colonizing organisms must be judged to be sufficiently abundant to construct identifiable biogenic substrates. When not present in densities sufficient to construct reef substrate [4].

**Table 8.** Biotic class: reef biota areas dominated by reef-building fauna, including living corals, mollusks, polychaetes or glass sponges.

coral and other carbonate producers and physical processes such as waves and currents that modulate ecological processes and redistribute carbonate material within reef systems." [4]. These areas are well known as the most diverse and likely also provide the most significant oceanic ecosystems services as a result. These areas additionally provide the most varied in terms of type of services as they provide more esthetic and ecotourism support to a greater degree than any other macroinvertebrate assemblages. Yet these significant areas are also some of the most delicate and threatened habitats. Coral bleaching can occur as the result of numerous stressors and large-scale weather events can devastate large regions from both abiotic and biotic stressors [27, 30, 36, 37]. Of all the marine coastal biotopes, literature suggests it is the coral dependent fauna that can be devastated from large-scale changes such as those that occur after a tsunami event, but more investigation is needed to determine if recovery is possible.

Although much more study is needed to determine specific impacts in local areas, globally speaking, macroinvertebrate assemblages do recover after severe natural environmental perturbing events in general, but do so differentially. Anthropogenic perturbations that destroy the physical support of the biotic assemblages are less likely to recover, generally due to development of the shoreline and drainage of these areas. More work is needed to verify the longer term impacts that natural events have from habitat perturbation to ecosystem service losses, anthropogenic impacts have yet to be documented to a great degree from a global perspective. Human impacts are more often permanent, so prevention of further threats are the main reason more knowledge and awareness of the ecosystem services is crucial [32, 38–49].

In general, vascular plant dominated biotopes (VP) seem resilient (except for mangroves) after an environmentally perturbing events with recovery well underway in one annual cycle. Macroalgae/protista dominated biotopes (MA) may be impacted even at great distance from source of perturbations or related contamination little is known about the effects on the fauna they support. Both unconsolidated substrate dominated (US) and hard substrate dominated biotopes (HB) are noted to have recovery times close to that identified for sea grass areas. Reef species dominated areas (CS) are subject to many environmental stressors, the physical and chemical changes that result from an environmental perturbing event impact the corals species negatively but the fauna that rely on the physical structural components may shift in diversity but do persist. Defining recovery in terms of the macro invertebrate assemblage would seem to suggest that recovery occurs relatively quickly, with mangroves being the exception as it is suggested that they may never fully recover once the integrity of the habitat is destroyed.

There are many environmentally perturbing threats both natural and human that can limit the ecosystem services provided by marine coastal zone assemblages (**Table 9**). There are new research areas that focus on different regions and habitats, and more large scale methods are beginning to allow a picture of ecosystem services and the complex ways these macroinvertebrate assemblages provide them [21, 42, 44, 46–51].

**3. Conclusions**

**Identified threats to coastal marine macro-invertebrate communities**

Declining fishery resources

Limiting the human environmental changes to the coastlines from the decision-making perspective are one significant way that the ecosystem services of the marine coastal zone macroinvertebrate and associated macro-flora can be sustained. The macroinvertebrate biodiversity of these areas is resilient overall but the basis for the ecological assemblages, either the physical aspects or the biotic bases, must be able to provide the structure needs for refuge or attachment to support them. Additional challenges in considering the ecosystem services provided by macroinvertebrate assemblages in marine coastal zones resides in the policy makers, the planning decisions, coastal development, and most importantly of building consensus around ecosystem services in a locality. Research is needed that explores the application of a consensus approach across different land and seascape units. Assessment of the coastal zone biota still requires much research and practical work; finding ways to incorporate ecosystem services and its myriad values into the work of planners and policy makers in the marine and

**Table 9.** Threats and potential for heightened effects to macro invertebrate near shore communities after an environ-

**Mechanisms of impact Potentially heightened by an** 

Ecosystem Services Provided by Benthic Macroinvertebrate Assemblages in Marine Coastal Zones

**environmentally perturbing event**

http://dx.doi.org/10.5772/intechopen.73150

75

Yes—significant for breached coastal nuclear and industrial facilities

Yes—death assemblages and large numbers of eggs or fry of certain fish

species

Further scientific and societal endeavors are needed to identify ecosystem services in a locality and to then identify effects to ecosystem services provided by the macroinvertebrate assemblages specifically. Globally a picture of services and negative impacts on the services provided are identified in general. Specific impacts for categories of macroinvertebrate assemblages are lesser known, even as the body of research grows (**Table 3**). To maintain the ecosystem services provided by marine coastal zones macroinvertebrate assemblages (Provisioning, Supporting, Regulating, and those relating to Cultural Services) will `require an understanding and collaborative approach among researchers, planners, and those that ultimately rely on these services. Ultimately, more research is needed to identify which actions can be taken to lessen the loss and speed the recovery of these communities after large-scale events originating from both natural and human impacts to restore these important human related ecosystem services. The most significant gains could be made in determining further what recovery after an event is possible can be made in the different biotic assemblages, and what methods

coastal environment is as important as it is challenging (**Table 9**) [8].

Oil pollution Human activities, including the flushing of ocean

mentally (human impacts or natural) perturbing event.

vessel bilges, leakage from undersea oil wells, and runoff or discharges from land-based facilities

Marine environmental change and the fishery industry effects on environmental disruption

to safeguard against human impact can be possible.



**Table 9.** Threats and potential for heightened effects to macro invertebrate near shore communities after an environmentally (human impacts or natural) perturbing event.
