
4.4  Bedding-Plane Markings 
103 
 as ichnofosss, or lebensspuren. Study of trace fossils constitutes the discipline of 
ichnology, which has become increasingly complex since the  mid-1950s and has 
spawned  a  massive  body  of literature.  Several  of these  books  are  listed  under 
"Further Reading" at the end of this chapter. 
nds  ace  Fossils.  Trace fossils are not true bodily  preserved fossils;  that is, 
ey 
do not form by conversion of a skeleton into a body fossil.  They are  simply 
structures that originated through the activities of organisms. Interpreted broadly, 
biogenic  structures  can be  considered to  clude the  following:  (1) bioturbation 
stctures (burrows, tracks, trails, root penetration structures), (2) biostratification 
suctures  (algal  stromatolites,  graded  bedding of biogenic  origin),  (3) bioero
sion structures (borings, scrapings, bitings), and (4) excrement (coprolites, such as 
fe cal pellets or fecal castings). Not all geologists regard biostratification structures 
as ace fossils, and these structures are not commonly included in published dis
ssions of trace fossils. 
Trace  fossils are  classified  into  ichnogenera  on  the basis of characteristics 
at relate  to major behavioral  traits  of organisms  and are given generic names 
 as  Ophiomorpha.  Distinctive  but  less  important characteristics  are  used  to 
identify  ichnospecies, e.g.,  Ophiomorpha  nodosa.  Trace fossils  are  produced by a 
host of mine organisms such as crabs, flatfish, clams, molluscs, worms, shrimp, 
d eel.  nonmarine environments,  organisms such as insects,  spiders,  worms, 
llipedes, snails, and lizards can produce a variety of burrows and tunnels; ver
brate  organisms leave tracks;  and plants leave root traces. The organisms that 
puce traces are rarely preserved with the traces; thus, the trace maker is  com
monly 
not known. Therefore, the names applied to ichnogenera and ichnospecies 
geray do not refer to the trace makers themselves. 
ace Fossil Assemblages.  From a sedimentological standpoint, study of trace-fossil 
assemblages  has  commonly  proven  to  be more  useful  than  study  of individual 
icogenera  or ichnospecies. A trace-fossil assemblage is a basic collective term 
at embraces all of the trace fossils present within a single unit of rock Although 
vaous kinds of trace-fossil assemblages are recognized, groupg of trace fossils 
to  ichnofacies  has  particular  significance  in  paleoenvironmental  studies. 
Seilacher  (1964,  1967) introduced the concept of ichnofacies to describe associa
 of trace fossils that are recurrent in time and space and that reect environ
mental conditions such as water  depth  (bathymetry),  salinity,  and the nature of 
e  substrate in or on which they formed (e.g., mud vs. sand bottom). Fundamen
tly,  ichnofacies are sedimentary facies  defined  on the  basis of trace fossils, and 
each inofacies may include several ichnogenera. 
Seilacher (1967) established six ichnofacies, which he named after character
istic inogenera. Four of these (Skolithos,  Cruziana, Zoophycos, and Nereites) were 
based 
on  e marine water depth at which they were interpreted to occur (Table 
4.3; Fig. 4.38). The Glossungites  ichnofacies was established for  traces  that occur 
  to hard marine  surfaces, and  the  Scoyenia  ichnofacies characterized non
marine  environments.  Subsequently,  Frey  and  Seilacher  (1980)  established  the 
Tpanites  ichnofacies  for  hardgrounds  and  rockgrounds;  Bromley,  Pemberton, 
d  Rahmani  (1984)  proposed  the  Te redolites  ichnofacies  for  borgs  in  wood 
(woodgrou
nds); and Frey and Pemberton (1987) established the Psilonichus ichno
cies for  softgrounds in the marine to nonmarine environment. Several addition
 icofacies have also been proposed (e.g., Bromley, 1996, p. 241); however, the 
ne  inofacies shown in Ta ble 4.3  are  most commonly used.  Sedimentologists 
 parcularly interested in  the Skolithos, Cruziana, Nereites, and Zoophycos  ichno
facies,  which have the greatest potential for interpreting ancient marine environ
mtal conditions.