Page 106 - Introduction to Paleobiology and The Fossil Record
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PALEOECOLOGY AND PALEOCLIMATES  93


             contains more dissolved carbon dioxide          although other factors such as the substrate
             (CO 2) providing a means to corrode carbon-     were also infl uential.
             ate. At given depths in the world’s oceans,       Paleontologists were slow to adopt these
             carbonate material begins to dissolve at        insights from modern marine biology. There
             so-called compensations depths. Below the       were a few pioneer studies on Carboniferous
             carbonate compensation depth (CCD) the          assemblages in the 1930s, but it was the classic
             dissolution of calcium carbonate exceeds        work by Alfred Ziegler in the 1960s that
             supply and at about 4–5 km calcite is not       really brought these methods to the attention
             preserved. The depth is shallower for arago-    of paleontologists. He identifi ed  fi ve  depth-
             nite, with the aragonite compensation depth     related, brachiopod-dominated communities
             (ACD) placed at 1–2 km. Both the CCD and        in the Lower Silurian rocks of Wales and the
             ACD vary with latitude, being shallower at      Welsh borderlands (see Chapter 12). These
             higher latitudes, and both parameters have      communities stretched from the intertidal
             varied throughout geological time. Neverthe-    zone in the east to the deep shelf and conti-
             less, depth alone probably has little effect on   nental slope towards the west, matching per-
             biotic distribution, rather the many depth-     fectly the ancient paleogeography. This whole
             related factors can be used to reconstruct      system has been revamped and is now more
             the water depths of ancient marine              widely known as the benthic assemblage zones

             communities.                                    (Fig. 4.15). These zones are defined on a wide
               Finally, the state of the substrate, rates of   range of faunal and sedimentological criteria
             sedimentation and turbidity dramatically        and may be subdivided, internally, on the
             affect the distributions of benthic organisms   basis of, for example, substrate type and the
             (Brenchley & Pickerill 1993). Organisms have    degree of turbulence (Brett et al. 1993).
             complex ecological requirements, some pre-
             ferring a particular grain size, a certain type   Describing fossil communities
             of organic material or they even respond to
             chemical signals (chemotaxic). There are also   Sometimes the simplest jobs are the hardest
             complex taphonomic feedback processes,          to do properly. For over a hundred years,
             where biogenic substrates such as shell         paleontologists have provided lists of species
             pavements can form attachment sites for new     from particular localities, but these are not
             communities. In general terms, within near-     helpful for ecological work unless the relative
             shore environments, there is a broad correla-   abundances of the different species are docu-
             tion between community distribution and         mented as well. We need to know which
             grain size. Diversity tends to be highest in    species dominate (sometimes one species
             muddy sands, moderate in sandy muds,            makes up more than 50% of the sample) and
             low in pure sands and virtually zero in soft    which are rare (i.e. less than 5% of the col-
             muds. Moreover whether the sediments            lection). Now it is more common to docu-
             form soupy muds, loose sands, fi rmgrounds       ment the absolute and relative abundance of
             or hardgrounds will infl uence  faunal           each organism, illustrated graphically with
             distributions.                                  frequency histograms, and based on data
                                                             derived from line transects, quadrats or more
                                                             commonly now from bed-by-bed collecting or
             Paleocommunities
                                                             bulk samples.
             Paleocommunities are recurrent groups of          Counting conventions remain a problem.
             organisms related to some specifi c set of envi-  With many organisms it is relatively simple to
             ronmental conditions or limiting factors.       calculate how many individuals were actually
             Many of the concepts and techniques applied     represented in a given assemblage: univalved
             to marine fossil communities are based on the   species (e.g. gastropods) count as one, whereas
             work of biologists such as the Danish scientist   twin-valved species (e.g. bivalves and brachio-
             Carl Petersen, researching in the late 1800s    pods) may be assessed by adding the most
             and early 1900s. Petersen recognized a series   common valve (right or left, dorsal or ventral)
             of level-bottom benthic communities around      to the number of articulated or conjoined
             the Scandinavian coasts; the major control on   shells. Animals that molt, such as ostracods
             community distribution was water depth,         and trilobites, colonial organisms and those
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