Assessing ecological functioning in marine
benthic communities
A thesis submitted to the University of Newcastle upon Tyne for the degree of Doctor of Philosophy
by
Julie Bremner
Dove Marine Laboratory
School of Marine Science and Technology
University of Newcastle upon Tyne
January 2005
Abstract
With the advent of an ecosystem approach to marine management, the importance of developing methods to investigate ecological functioning is receiving increasing attention. This thesis develops a novel approach for describing ecological functioning in marine benthic systems. Biological traits analysis (BTA) uses a suite of life history, morphology and behaviour characteristics of species to describe aspects of their functioning. Comparison of BTA with two other approaches proposed for describing functioning in marine ecosystems established that BTA identified a range of biological attributes important for differentiating benthic communities and was better able to describe spatial patterns in assemblage composition than the other measures. Appraisal of the analytical tools proposed for use in BTA revealed they provided similar views of assemblage functioning, with the nonparametric tool being appropriate for providing a general picture of functioning, while the more complex parametric tools had greater power to detect anthropogenic impacts. Evaluation of the type and number of traits included in BTA showed it was sensitive to the number of traits selected for analysis, with optimal results being gained by maximising trait number. Examination of the relationship between functioning and environmental variability revealed that trait composition was related to changes in a number of environmental factors, although this relationship was complex and the nature of associations between traits and specific environmental factors varied depending on the location of assemblages. Further analyses focussed on the impacts of anthropogenic activities on benthic assemblage functioning. These revealed that assemblage functioning was impacted by fishing disturbance in both subtidal and intertidal assemblages. A number of traits were impacted by fishing, including some associated with vulnerability to physical stress and others related to resistance to disturbance, while other aspects of functioning remained unaffected. The thesis has increased our understanding of biological traits analysis as a tool for describing functioning in marine benthic systems. It has also contributed to some interesting ecological and management issues, such as the relationship between species and functioning and the importance of, information required for, and strategies available for conservation of ecological functioning in marine ecosystems.
Contents
Abstract.............................................................................................................................i
Acknowledgements........................................................................................................ii
Contents.............................................................................................................................iii
List of Tables and Figures
TABLES ......................................................................................................................vii
FIGURES.....................................................................................................................ix
Chapter 1: Biomonitoring and human impacts on marine ecosystems
INTRODUCTION ......................................................................................................................2
DISCRETE APPROACHES TO BIOMONITORING..........................................................................3
INTEGRATIVE APPROACHES
Ecological processes .................................................................................................5
Ecological functioning..............................................................................................7
Chapter 2: Ecological functioning in marine benthic ecosystems
ABSTRACT ..........................................................................................................14
INTRODUCTION ...............................................................................................14
METHODS
Data collection ....................................................................................................17
Data analysis
Relative taxon composition analysis.................................................................18
Trophic group analysis ......................................................................................19
Biological traits analysis .....................................................................................19
Comparison of approaches.................................................................................22
RESULTS
Relative taxon composition analysis....................................................................23
Trophic group analysis.........................................................................................27
Biological traits analysis.......................................................................................30
iv
Comparison of approaches................................................................................35
DISCUSSION ......................................................................................................35
Chapter 3: Methods for conducting biological traits analysis in marine
benthic ecosystems
ABSTRACT .........................................................................................................44
INTRODUCTION ...............................................................................................45
METHODS
Dataset 1....................................................................................................................48
Dataset 2....................................................................................................................48
Comparison of analytical tools .................................................................................49
Power of the analytical tools.....................................................................................52
Sensitivity of BTA ....................................................................................................52
RESULTS
Comparison of analytical tools .................................................................................54
Power of the analytical tools.....................................................................................55
Sensitivity of BTA ....................................................................................................58
DISCUSSION .........................................................................................................60
Chapter 4: Does ecological functioning in marine benthic assemblages
change with environmental conditions?
ABSTRACT ........................................................................................70
INTRODUCTION..............................................................................71
METHODS
Invertebrate data.................................................................................74
Environmental data ............................................................................76
Data analysis
Environment.......................................................................................76
Taxa and the environment.................................................................77
Functioning and the environment ..................................................78
v
RESULTS
Environmental variables ................................................................79
Taxa and the environment..............................................................82
Functioning and the environment ...............................................82
DISCUSSION ................................................................................91
Chapter 5: Biological traits of the north sea benthos – does fishing affect
benthic ecosystem functioning?
ABSTRACT ...................................................................................97
INTRODUCTION..........................................................................98
METHODS
Study area...................................................................................100
Fishing pressure ..........................................................................100
Biological traits analysis..............................................................105
RESULTS .....................................................................................107
DISCUSSION ................................................................................114
Chapter 6: The effects of human disturbance on ecological functioning in
different intertidal habitats
ABSTRACT .....................................................................................................121
INTRODUCTION .........................................................................................122
METHODS
Experimental sites...........................................................................................125
Experimental design........................................................................................127
Sediment parameters.......................................................................................127
Faunal sampling...............................................................................................128
Description of ecological functioning.............................................................128
Statistical analysis............................................................................................130
vi
RESULTS
Seal Sands
Sediments ......................................................................................................131
Taxon analysis ..............................................................................................131
Ecological functioning ..................................................................................139
Ballochmartin Bay
Sediments ......................................................................................................140
Taxon analysis ..............................................................................................147
Ecological functioning ..................................................................................149
DISCUSSION ...........................................................................................................................154
Chapter 7. General discussion...................................................................................165
References.........................................................................................................................173
Appendices .......................................................................................................................198
Chapter 7
General discussion
This thesis has clearly demonstrated the utility of biological traits analysis for characterising aspects of ecological functioning in marine benthic communities. It has shown that the approach performs well in comparison to more traditional measures of ecological functioning (Chapter 2), displays the attributes of a good biomonitoring tool (it can be easily applied (Chapter 3), it is applicable to different geographical areas and habitat types (Chapters 2 and 4) and it is sensitive to and can identify the effects of anthropogenic activities (Chapters 5 and 6)) and provides useful information on the functioning of benthic assemblages. In addition, the findings presented in this thesis have raised some issues that are of both intellectual interest and practical importance for marine ecosystem ecology and management.
The ultimate purpose of ecosystem management is to maintain the ‘goods and services’ provided by marine ecosystems, with the former generally translating to the abundance of target species and the latter to maintaining the rates of ecological processes. We know that some anthropogenic activities can impact on receiving systems by altering the species richness, diversity or composition of local assemblages. The phenomenon is well documented in marine benthic systems, particularly in respect to anthropogenicallydriven physical disturbances of the seabed such as fishing (Hutchings, 1990; Jennings & Kaiser, 1998; Johnson, 2002).
The need to conserve ecological processes, coupled with the occurrence of anthropogenically-driven changes in species abundances, prompts the question ‘do changes in biodiversity lead to changes in ecological processes?’, or, in other words, ‘how does biodiversity relate to ecosystem functioning (actually ecosystem processes, see Raffaelli et al., 2003)?’. Biodiversity and ecosystem ‘functioning’ is a well-established subject area in terrestrial circles, where much attention has been directed towards answering the question (see Tilman, 1999; Loreau et al., 2001), but it is a very new field of study in the realm of marine ecology.
Initial marine research has documented complex relationships between species biodiversity and ecological processes, with changes in biodiversity causing process-changes in some situations but not others (Bolam et al., 2002; Raffaelli et al., 2003). This result adds weight to the assertion that the specific biological attributes exhibited by species are of more importance to the maintenance of ecological processes than simply species number itself (Bengtsson, 1998; Diaz & Cabido, 2001; Giller et al., 2004), because these attributes determine the extent to which species contribute to ecological processes (in simple terms, all species are not equal in relation to ecological processes). A logical extension of this argument is that the biological attributes expressed by an assemblage (i.e. the sum total of attributes expressed by all component species) will determine how that assemblage contributes to ecological processes.
This, in turn, raises two further questions; (1) ‘do changes in the attributes expressed by an assemblage affect ecological processes?’ and on a more basic level, given that the same attribute can be expressed by a number of taxa, (2) ‘do changes in the species complement always lead to changes in the attributes expressed by an assemblage?’. The answer to the second question will clearly impact on the first as, if changes in the species complement do not lead to changes in the attributes expressed by assemblages, logic would suggest that the answer to the first question is of little practical importance for marine ecosystem management.
This thesis cannot directly address the first question as ecological processes have not been measured here, but the findings can contribute towards the second because BTA provides a measure of the attributes expressed by assemblages (biological trait composition). Chapters 2 and 4 provided evidence that, at least in some cases, changes in the species complement did not lead to changes in the biological trait composition of benthic assemblages in the southern North Sea, eastern Channel and Irish Sea regions. If this is a general phenomenon, it suggests that ecological processes will not always be affected by changes in species diversity, because these do not translate into changes in the attributes expressed by the assemblage.
However, extreme caution must be used in generalising these observations. Although some stability of trait composition in the face of changes in the species complement was documented in the thesis, it did not occur universally. In some cases, differences in species composition did translate into differences in trait composition. For example, in Chapter 2, megafauna assemblages off the north-eastern French coast (south of Boulogne, see Figures 2.2 and 2.7) were as variable when classified by their biological traits as they were when classified by species composition, and changes in the species complement of intertidal assemblages in a north-eastern English estuary resulted in changes in the attributes expressed by the assemblages (Chapter 6).
In theory, the attributes expressed by an assemblage remain stable when diversity changes because a given attribute will be expressed by several different species, so reductions or removals of one of these species will be compensated for by increases in one or more of the others (see Walker, 1992; Frost et al., 1995; Naeem, 1998). Logic, however, dictates that a juncture will be reached where species change must impact on the attributes expressed by an assemblage. At some stage, if reductions in biodiversity are severe enough, all of the species expressing a given trait will be removed and compensation becomes impossible. This point is obvious. What is not clear is the stage at which more subtle changes in the species complement, such as changes in the relative proportions of component species, will lead to changes in the attributes expressed by assemblages.
This is not a trivial issue. Compensation is the mechanism by which stability in the biological attributes of assemblages is maintained. If we cannot determine the extent of compensation, we cannot determine when changes in the species complement will lead to changes in the attributes expressed by assemblages (question 2). This will, consequently, make it difficult to determine how anthropogenically-driven changes in the species complement will affect ecological processes. This thesis has provided evidence that the attributes expressed by benthic assemblages may sometimes remain stable in the face of changes in the species complement, but the extent of the phenomenon, and the mechanisms underlying it, although extremely important for ecosystem ecology and management, are far from clear and require appropriate quantification.
The changes in functioning documented in response to human impacts in Chapters 5 and 6 signal that ecosystem management needs to be applied at the level of ecological functioning as well as at the species level. Ecological functioning must be conserved in benthic assemblages if ecological processes are to be maintained. However, it would be extremely difficult to conserve ecological functioning in it’s entirety without ceasing anthropogenic activities in the sea. As this is unlikely, priority may be given to identifying which aspects of functioning to preserve, or what actions to take to conserve as much functioning as possible. Clearly, ecosystem-based conservation should protect those aspects of functioning that have the strongest links to ecological processes.
In benthic assemblages, faunal attributes such as tube-dwelling, burrow-dwelling and burrowing have potentially important implications for the regulation of ecological processes (see Chapter 6). Protecting tube-dwelling, burrow-constructing and burrowing fauna should then, theoretically, make a large contribution towards maintaining the rates of ecological processes.
At the present time, the links between specific faunal activities and processes are, largely, based on theoretical predictions and many unknowns remain. For instance, although recent work has provided experimental evidence of the links between faunal activities and specific ecological processes (Biles et al., 2002; Howe et al., 2004; Lohrer et al., 2004; Webb & Eyre, 2004a, b), the extent of the contributions these taxa make to processes is still unclear. Also, we do not know if all taxa sharing these traits have the same impact on processes, whether the degree to which taxa express a trait affects their contributions, or whether other traits expressed by the taxa also affect their involvement in the regulation of ecological processes. If we are to conserve functioning and, therefore, ecological processes we must have more information on how specific traits determine the maintenance of these ecological processes.
However, at a practical level, ecosystem management must operate in the absence of perfect knowledge and it is better to employ a cautionary strategy based on logic and theory than to apply none at all. Chapters 4 and 6 showed that, when the attributes expressed by assemblages were examined, the traits tube- and burrow-dwelling most often exhibited negative responses to fishing impacts. If the traits are fundamental to the regulation of ecological processes, these anthropogenically-mediated changes in functioning could have far-reaching consequences for marine ecosystems. Uncertainty over the importance of their role in process-regulation should not prevent these traits from being protected because, as shown in this thesis, this potentially crucial aspect of functioning is vulnerable to physical seabed disturbance.
Another option for preserving ecological functioning in the light of imperfect knowledge is to adopt one of the strategies used in conservation based on taxonomic identities. Here, communities are classified on the basis of their species composition, with community types that are rare within a given area being selected for protection. This is a conservative approach that protects communities which may otherwise be lost, even though the implications of their loss at the ecosystem level may be, as yet, unknown. An analogous strategy would be relatively simple to implement at the functional level, being conducted in much the same manner as at the taxonomic level, only using the biological attributes present in assemblages instead of species lists.
This approach has the advantage that it can be applied alongside species-level classification, which is essential as conservation must operate at both levels. These two approaches may well identify different communities for protection, as those communities that are designated as rare using species-level classifications may not be rare in terms of their functioning, or vice versa. Chapter 2 provides a good example of this situation. When biological trait composition was considered, a group of five assemblages was classified as rare throughout the southern North Sea and eastern Channel region (group 2, Figure 2.7). However, on the basis of species composition, these five assemblages were classified as common (Figure 2.2). This confirms that conservation focussed at the species level is not enough to protect ecological functioning in marine ecosystems. While protection of ecological functioning will add another level of complexity to ecosystem management strategies, it is necessary and can be fitted into the framework already in place for species and habitat conservation.
One requirement for successful ecosystem management that is currently lacking is consensus on terminology. Ecosystem investigation of this form incorporates different disciplines such as biology, chemistry and physical sciences and it is a relatively new area for many researchers, even in the comparatively advanced field of terrestrial ecosystem ecology. Ecologists from different research disciplines may use different terminology to refer to what may be, essentially, the same thing (see Emmerson & Huxham, 2002).
This thesis has followed Naeem et al. (1999) in defining ecological functioning as the maintenance and regulation of ecological processes. Other authors have defined ecosystem functioning as the ‘processes occurring in a system’ (Biles et al., 2002), i.e. ‘biogeochemical activities such as production, community respiration, decomposition, nutrient cycling or nutrient retention’ (Naeem & Wright, 2003). Yet more authors define some of these ‘activities’, such as decomposition rates and nutrient cycling, as ‘ecosystem properties’ (Loreau et al., 2001).
These different terminologies are justifiable and understandable in what is an emerging field of interdisciplinary science. They do, however, create a great deal of confusion, and confusion in terminology can make it difficult to locate relevant information and share new findings and ideas in the interdisciplinary arena. While these differences in terminology do not, and should not be allowed to, prevent the development of new research in marine ecosystems, consensus in terminology at some point in the near future would greatly aid understanding and the advancement of the field.
The focus of this thesis has been on developing biological traits analysis as a tool for describing ecological functioning of marine benthic assemblages, and investigating changes in functioning in response to both natural and anthropogenic factors. However, the work has also raised and contributed to our knowledge of some interesting ecological and management issues, such as the relationship between species and the attributes expressed by assemblages, the information required for conservation of functioning and, ultimately, ecological processes, and the need for, and strategies for implementation of, conservation of ecological functioning. The thesis has therefore both contributed to our understanding of biological traits analysis in ecosystem biomonitoring and opened up a number of interesting avenues for consideration and future research, with regards to the relationship between biodiversity, functioning and ecological processes.
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