EEOS 630 Biol. Ocean. Processes Chapter 5 Class 8-9: 9/25/08, 9/30 Revised: 9/16/08 ©2008 E. D. Gallagher GLOBAL PATTERNS OF BENTHIC COMMUNITY STRUCTURE ESPECIALLY DEEP-SEA DIVERSITY TABLE OF CONTENTS Page: List of Tables .................................................................................... 2 List of Figures ................................................................................... 3 Assignment...................................................................................... 3 Required ................................................................................ 3 Etter, R.J. and L. S. Mullineaux. 2001. ................................................. 3 Gallagher, E. D. & K. E. Keay. 1998. .................................................. 3 Jumars, P. A. and E. D. Gallagher. 1982 ................................................ 3 Supplemental............................................................................. 4 Etter, R. J. and J. F. Grassle. 1992 ..................................................... 4 Huston, M. 1979 .................................................................. 4 Mills, E. L. 1969 .................................................................. 4 Rex, M. A., C. R. McClain, N. A. Johson, R. J. Etter, J. A. Allen, P. Bouchet and A. Warén. 2005. ......................................................................... 4 Sanders, H. L. 1968 ................................................................ 4 Snelgrove, P. V. R. and C. R. Smith. 2002. ............................................. 4 Trueblood, D. D., E. D. Gallagher, and D. M. Gould. 1994 . 4 Witman, J. D., Ron J. Etter, and F. Smith. 2004. ......................................... 4 Comments on the papers ........................................................................... 4 What is a community?...................................................................... 5 Continua vs. discrete entities ......................................................... 5 Community structure ............................................................... 7 Community function ................................................................ 8 Parallel-level bottom communities ........................................................... 12 Diversity ............................................................................... 14 Measuring richness and evenness .................................................... 14 Sanders rarefaction and Hurlbert’s E(Sn ) .............................................. 18 Classification and ordination ............................................................... 21 Factors controlling benthic community structure ................................................ 22 Biogeography .................................................................... 22 Salinity and temperature ........................................................... 23 The Intermediate Disturbance Hypothesis ............................................. 25 Why are there so many species in the deep sea?................................................. 29 Basic patterns in deep-sea communities ............................................... 29 List of ideas on deep-sea diversity .................................................... 31 Terms and Concepts.............................................................................. 37 Faunal similarity and dissimilarity indices ..................................................... 41 EEOS 630 Biol. Ocean. Processes Benthos, P. 2 of 96 Outlines of papers ............................................................................... 43 Required and Recommended ............................................................... 43 Mills, E. L. 1969 ................................................................. 43 Web Resources.................................................................................. 45 References ..................................................................................... 46 General Benthic Ecology .................................................................. 46 General community ecology ................................................................ 47 Sampling the benthos ..................................................................... 48 Coring & profiling ................................................................ 48 Processing samples ............................................................... 49 Classification analysis..................................................................... 49 Books and methods papers ......................................................... 49 Benthic clustering ................................................................ 53 Clustering physico-chemical variables ................................................ 54 Phylogenetic clustering ............................................................ 54 Miscellaneous classification analysis ................................................. 55 Ordination .............................................................................. 55 Classics ................................................................................ 56 The effects of salinity and grain size on benthic community structure . 56 Other references on benthic community structure ............................................... 58 Historical succession papers ................................................................ 60 Shallow soft-bottom succession ............................................................. 61 Long-term (> 5 years) benthic sample surveys .................................................. 66 Hard-substrate succession.................................................................. 67 Species diversity ......................................................................... 70 Components of diversity: á, â, ã [and ù] .............................................. 70 Deep-sea diversity and succession ................................................... 71 Latitudinal diversity gradients ....................................................... 75 Explanations for patterns of diversity (e.g., intermediate disturbance hypothesis) .............. 76 Diversity Indices ................................................................. 80 How many species? ............................................................... 84 Sampling Distributions per se ............................................................... 86 Similarity indices ........................................................................ 89 Miscellaneous ........................................................................... 91 Index ......................................................................................... 93 List of Tables Table 1. 24 Ecosystem attributes that change during succession (Odum 1969) ................................. 9 Table 2. Soft-bottom benthic community attributes that change during the course of succession (Rhoads et al. 1978). 10 Table 3 Three hypothetical communities, I, II, and III, showing the ambiguous nature of diversity. 20 Table 4 Some characteristics of four Pacific deep-sea communities (Jumars & Gallagher 1982)................. 30 Table 5. A list of key dates and ideas in the study of deep-sea diversity. Many of these ideas are reviewed in Jumars & Gallagher (1982), Rex (1983), and Grassle (1989).............................................. 31 Table 6. Web resources on community structure........................................................ 45 EEOS 630 Biol. Ocean. Processes Benthos, P. 3 of 96 List of Figures Figure 1. General model of distribution of benthic infaunal successional stages along a gradient of increased environmental disturbance from left to right (after Pearson & Rosenberg 1978) and the associated Benthic Habitat Quality (BHQ) index... Sediment profile images assigned to a successional stage are mounted above the general model, where oxidized sediment is rust-brown and reduced sediment is grey or black. In the bottom of the figure the generalized species (S), abundance (A), biomass (B) diagram are illustrated. Fig. 2 from Rosenberg (2001) ........................................................................ 11 Figure 2. A metric scaling of Sander’s Station R data, using PCA-H ordination of CNESS (m=7) as described in Trueblood et al. (1994). The sampling period (1!17) and replicate number are indicated. The results of a cluster analysis of the same CNESS matrix is superimposed on the ordination. The point where a sample with equal abundances of all species, Hm ax, is also shown. ............................................. 13 Figure 3. A metric scaling of Sander’s Station R data, using PCA-H ordination of CNESS (m=7) as described in Trueblood et al. (1994). The results of a cluster analysis of the same CNESS matrix is superimposed on the ordination. The point where a sample with equal abundances of all species, H max, is also shown. 14 Figure 4 The salinity clock for the US East Coast. The cosine of the angle among species vectors (“arrows”) indicates the probability that species will co-occur. Salinity is the major factor controlling the distribution of species in the neritic zone. Abbreviations: ACTECANA (Acteocina canaliculata), AMPEABVA (Ampelisca abdita & A. vadorum), CYATPOLI (Cyathura polita), HETEFILI (Heteromastus filiformis), LEPTPLUM (Leptocheirus plumulosa), LIMNHOFF (Limnodrilus hoffmeisteri), MAREVIRI (Marenzelleria viridis), MEDIAMBI (Mediomastus ambiseta), MULILATE (Mulinia lateralis),