[Oceanography and Marine Biology - an Annual Review] R. N

OCEANOGRAPHY and
MARINE BIOLOGY

AN ANNUAL REVIEW

Volume 44

OCEANOGRAPHY and
MARINE BIOLOGY

AN ANNUAL REVIEW

Volume 44

Editors

R.N. Gibson

Scottish Association for Marine Science The Dunstaffnage Marine Laboratory
Oban, Argyll, Scotland [email protected]

R.J.A. Atkinson

University Marine Biology Station Millport
University of London
Isle of Cumbrae, Scotland r .[email protected]

J.D.M. Gordon

Scottish Association for Marine Science The Dunstaffnage Marine Laboratory
Oban, Argyll, Scotland [email protected]

Founded by Harold Barnes

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Contents

Preface

vii

ix

Correction to Volume 43

Review of three-dimensional ecological modelling related to the North Sea shelf system. Part II: model validation and data needs

Günther Radach & Andreas Moll

1
61

Role, routes and effects of manganese in crustaceans

Susanne P. Baden & Susanne P. Eriksson

Macrofaunal burrowing: the medium is the message

85

Kelly M. Dorgan, Peter A. Jumars, Bruce D. Johnson & Bernard P. Boudreau

Mediterranean coralligenous assemblages: a synthesis of present knowledge

123

Enric Ballesteros

Defensive glandular structures in opisthobranch molluscs — from histology to ecology

Heike Wägele, Manuel Ballesteros & Conxita Avila

197 277

T a xonomy, ecology and behaviour of the cirrate octopods

Martin A. Collins & Roger Villanueva

The ecology of rafting in the marine environment. III. Biogeographical and evolutionary consequences

Martin Thiel & Pilar A. Haye

323 431

Potential effects of climate change on marine mammals

J.A. Learmonth, C.D. MacLeod, M.B. Santos, G.J. Pierce, H.Q.P. Crick & R.A. Robinson

Author Index

465

497 515
Systematic Index Subject Index

Preface

The forty-fourth volume of this series contains eight reviews written by an international array of authors that, as usual, range widely in subject and taxonomic and geographic coverage. The editors welcome suggestions from potential authors for topics they consider could form the basis of future appropriate contributions. Because an annual publication schedule necessarily places constraints on the timetable for submission, evaluation and acceptance of manuscripts, potential contributors are advised to contact the editors at an early stage of preparation. Contact details are listed on the title page of this volume.
The editors gratefully acknowledge the willingness and speed with which authors complied with the editors’ suggestions, requests and questions, and the efﬁciency of Taylor & Francis in ensuring the timely appearance of this volume.

Correction to Volume 43

Carney, R.S. 2005. Zonation of deep biota on continental margins. Oceanography and Marine Biology: An

Annual Review 43, 211–278.

In reviewing various ideas put forward to explain deep benthic depth zonation (Carney 2005), the TROX, Trophic-Oxygen, model for foraminiferan distributions received special mention. This conceptual model incorporates in an easily understood manner several lines of thought concerning the importance of oxygen and labile carbon influx in controlling surficial and interstitial microhabitats. The origin of the TROX model, however, is misattributed to Loubere and his associates (page 228). The TROX model was proposed in the context of microhabitats with Adriatic transect data used as an example by Jorissen et al. (1995). An important feature of the TROX model is the explicit link between oxygen and carbon flux, such that the ecological importance of food availability changes with oxygen concentration. Given adequate oxygen, food is the primary factor. When microbial consumption of higher food levels reduces microhabitat oxygen, then low oxygen can become the primary factor. There have been other attempts to create general foraminiferan distribution models that incorporate both oxygen and carbon flux somewhat similar to TROX. An algebraically formal model of gradient distribution employing the concept of ‘r’ and ‘K’ selected species was developed by Sjoerdsma & van der Zwaan (1992) and tested with mixed success on archived distribution data from the Gulf of Mexico. Unfortunately, the geochemical relationship between oxygen and carbon flux was omitted. Bottom oxygen was estimated from water column profiles, and flux estimated only on the basis of depth. The conceptual model proposed by Loubere et al. (1993) was an assemblage model incorporating both production and taphonomy. Like the TROX model, it is an important contribution to the understanding of trophic control of geographic distribution. That mode links the geochemistry of oxygen and carbon flux. Flux was estimated from sedimentary oxygen consumption; samples were analysed from the western Gulf of Mexico.

REFERENCES

Carney, R.S. 2005. Zonation of deep biota on continental margins. Oceanography and Marine Biology: An

Annual Review 43, 211–278.

Jorissen, F.J., de Stigter, H.C. & Widmark, J.G.V. 1995. A conceptual model explaining benthic foraminiferal

microhabitats. Marine Micropaleontology 26, 3–15.

Loubere, P., Gary, A. & Lagoe, M. 1993. Generation of the benthic foraminiferal assemblage — theory and

preliminary data. Marine Micropaleontology 20, 165–181.

Sjoerdsma, P.G. & van der Zwaan, G.J. 1992. Simulating the effect of changing organic ﬂux and oxygen content on the distribution of benthic Foraminifera. Marine Micropaleontology 19, 103–150.

OCEANOGRAPHY and
MARINE BIOLOGY

AN ANNUAL REVIEW

Volume 44

Oceanography and Marine Biology: An Annual Review, 2006, 44, 1-60

REVIEW OF THREE-DIMENSIONAL ECOLOGICAL MODELLING RELATED TO THE NORTH SEA SHELF SYSTEM.
PART II: MODEL VALIDATION AND DATA NEEDS

GÜNTHER RADACH & ANDREAS MOLL

Institut für Meereskunde (IfM), Universität Hamburg (ZMK-ZM A W ),
Bundesst r . 5 3, D-20146 Hamburg, Germany

E-mail: [email protected], [email protected]
Abstract The aim of this review is to provide an overview of the status of validation of eleven biogeochemical and ecological models of the greater North Sea (COHERENS, CSM-NZB, DCM- NZB, DYMONNS, ECOHAM, ELISE, ERSEM, FYFY, GHER, NORWECOM, POLCOMS- ERSEM) showing the realism achieved as well as the problems hindering a better degree of validity of the models. Several of the models were able to reproduce observations of the state variables correctly within an order of magnitude, but all models are not capable of reproducing every simulated state variable in the range of observations. None of the models can be called a valid model. Comparison of results from different models with datasets are evaluated according to the different spatial and temporal scales, for which data products were available, namely for regional distributions, annual cycles, long-term developments and events. The higher the trophic level, the greater was the discrepancy with the data. Problems still exist in determining the necessary complexity of the ecosystem model. More complexity in the model does not necessarily improve the simulations. Special attention should be devoted to the regeneration mechanisms in the sediments. Species’ groups have been simulated so far with rather limited success. The ecological model simulations did not reproduce fully the observed variability. Possible sources of lacking coincidence with observations originating from the spatial and temporal resolution of the internal dynamics, the trophic resolution, or the resolution of the forcing functions are discussed. Most of the models still need to be evaluated more intensively for their predictive potential to be judged. They have not yet been tested to a degree which is possible today using the various existing datasets from the northwest European shelf seas (presented in the Appendix). Common datasets for the necessary annual cycles of forcing functions are needed.

Introduction

The overall aim of this review is to give an overview on the state-of-the-art biogeochemical and ecological three-dimensional modelling related to the marine ecosystem of the greater North Sea. The goal is to provide guidelines for the further development of marine ecosystem models which will be used in the future for making predictions about how the marine ecosystem of the North Sea functions and how concentrations and ﬂuxes of biologically important elements like carbon, nitrogen, phosphorus, silicon and oxygen vary in space and time, throughout the shelf over a timescale of years, in response to physical forcing. Three-dimensional physical oceanographic modelling, which forms the basis of the ecological models, was recently reviewed by Jones (2002).

1

GÜNTHER RADACH & ANDREAS MOLL

With respect to future environmental problems occurring in the North Sea, a decadal frame has to be envisaged for modelling. Models are the only tools which can be used as a means to investigate developments of annual cycles and interannual variability in shelf sea systems to be expected in future decades, by simulating scenarios with different initial settings and boundary conditions.
This review consists of two parts. Part I (Moll & Radach 2003) described the existing ecological models dealing with the greater North Sea; it characterised the complexity and the achievements of eleven ecosystem models for the whole North Sea or parts of it. Seven of these models are threedimensional models (NORWECOM, GHER, ECOHAM, ERSEM, ELISE, COHERENS, POLCOMS-ERSEM) designed for describing the ecosystem dynamics of the North Sea, based on the physical oceanographic circulation, in more or less detail. Several of the models were used for specific applications, some of which have already been described in the literature. The information about these models and their applications from all available papers were aggregated under the heading of the names of the eleven ecological models in Moll & Radach (2003; their Table 1).
This Part II provides an overview of the status of validating these models of the greater North
Sea area. The term ‘status of validation’ denotes the current state of realism which a model has reached, as shown by the process of validating the model results by appropriate methods. The status of validation is considered to be a very important property of these models, because it is of great importance for simulating future scenarios that the simulations can realistically reproduce a series of observed annual cycles and especially their observed variability. The aim of this part is not only to show the realism achieved, but also to find out the problems hindering a better degree of validity of the models.
The basis for both parts is the published literature on biogeochemical and ecological threedimensional modelling related to the marine ecosystem of the greater North Sea. Although this review is concentrated on the North Sea, the problems arising in modelling other shelf seas and also the oceans in temperate climatic regions are very similar, as the corresponding literature shows. The quality of the simulations is reported on the basis of judgements of the facts given in the publication, in so far as they help to elucidate the achievements and failures of the models. Being restricted to the knowledge contained in the publications, very often questions about causes of failures must remain open.
The order of the paper is as follows: after a short recapitulation of the published attempts at validating the models, the applied validation methods and the quantitative measures of the goodness of fit used for evaluating the comparison of simulations and data are briefly reported. Then the validational status of the selected models is discussed in detail with respect to defined space and timescales: The simulation results presented in the available literature are analysed with respect to regional horizontal distributions, to annual cycles and long-term developments, and also the temporal event scale. The discussion brings out common features of success and failure of the models and suggests measures to overcome the problems. A few important datasets available for testing the models are located and briefly described in the Appendix.

Validational efforts for ecological models for the North Sea

Before presenting the validational state of the marine ecological North Sea models listed in Table 1, those publications which contain information on validational efforts are summarised. For most of the models their results were compared more or less thoroughly to observations. In a few cases comparisons to common datasets were performed. Nearly all models presented time-series of simulated state variables vs. observed data in graphical form and they presented annual cycles of various state variables in comparison to either observed time-series at special locations or aggregated and averaged box data. While older simulations concentrated on the mean situation, newer papers also compared their simulations with data from actual years.

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VALIDATION OF THREE-DIMENSIONAL ECOLOGICAL MODELLING

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