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Shell Repair As a Response to Attempted Predation in Some Palaeozoic and Younger Gastropods

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Shell Repair As a Response to Attempted Predation in Some Palaeozoic and Younger Gastropods Shell Repair as a Response to Attempted Predation in some Palaeozoic and Younger Gastropods ANNA LINDSTRÖM ISBN 91-506-1829-6 urn:nbn:se:uu:diva-6033 ! " ##" $#%## & & & ' ( ) ' * +' ##"' , - ' . ' ' /,0 1$2"#32$4 123' & & 5 & & 2 6 & & ' 6 ( ' 7 && & - & 8 ( & & & ' ( & 9 ' / & & & & 2 ' / &9 & & - & && ' , ( &9 &$#:( ;: & ' - & ( ( & ' , 2 & ( &9 ' - ( &9 & $.'$ : ;' :' &9 < - ( 4'4 : ;3'3 : ( & = ( 6' 6 & ( & & & ' > & 6 ( - ( ' & 5 ' 7 - !"#$ %% & & # '()%*+ & ? + * ##" /,0 1$2"#32$4 123 % %%% 23#@@ A %BB '5'B C D % %%% 23#@@E Till Henning, Selma & Agnes List of Papers I Lindström, A. and Peel, J. S. 1997. Failed predation and shell repair in the gastropod Poleumita from the Silurian of Gotland, Sweden. Bulletin of the Czech Geological Survey 72, 115-126. II Lindström, A. and Peel, J. S. 2003. Shell repair and mode of life of Praenatica gregaria (Gastropoda) from the Devonian of Bohemia (Czech Republic). Palaeontology 46, 623-633. III Lindström, A. 2003. Shell breakage in two pleurotomarioid gastropods from the Upper Carboniferous of Texas, and its relation to shell morphology. GFF 125, 39- 46. IV Lindström, A. and Peel, J. S. (in press). Repaired injuries and shell form in some Palaeozoic pleurotomarioid gastropods. Acta Palaeontologica Polonica 50. V Lindström, A., Ebbestad, J. O. R. and Peel, J. S. (manuscript). Predation on bellerophontiform mollusks in the Paleozoic VI Lindström, A. (manuscript). Shell repair and shell form in Jurassic pleurotomarioidean gastropods from England. VII Lindström, A. (manuscript). Predator-prey interaction through the Phanerozoic: the history of shell repair in pleurotomarioid gastropods. Reproduction of papers I-IV was made with permission of the copyright holder. Paper I © 1997 Czech Geological Survey Paper II © 2003 The Palaeontological Association Paper III © 2003 Geologiska Föreningen Paper IV © 2005 Instytut Paleobiologii PAN Papers V-VII © by the authour(s) Contents Introduction ............................................................................................................................... 9 Aims of this study................................................................................................................. 9 Predation in the fossil record................................................................................................... 11 Detection of predation in fossils ......................................................................................... 11 Drilling ........................................................................................................................... 11 Shell breakage ................................................................................................................ 12 Unsuccessful predatory attacks........................................................................................... 12 Shell repair frequency .................................................................................................... 12 Predators past and present.............................................................................................. 13 Other causes of breakage................................................................................................ 14 Adaptations in gastropods against predation ...................................................................... 14 The papers in this study........................................................................................................... 16 Poleumita, Praenatica and predation ................................................................................. 16 Predation on bellerophontiform molluscs........................................................................... 17 Shell repair in pleurotomarioid gastropods......................................................................... 17 Conclusions......................................................................................................................... 19 Svensk sammanfattning...........................................................................................................20 Bakgrund............................................................................................................................. 20 Hur upptäcker man predation bland fossil? ........................................................................ 20 Artiklarna i avhandlingen ................................................................................................... 21 Acknowledgements ................................................................................................................. 23 References ............................................................................................................................... 24 Introduction Organisms are well adapted to the environments in which they live, but the interaction of environmental change and natural selection has prompted the evolution of new morphologies throughout geologic time. In modern, as well as ancient seas the growth, size and shape of, for example, a gastropod’s shell are controlled by several factors, such as temperature, salinity, climate, metabolic rate, calcification and physical stress (Vermeij, 1978). One other factor is extremely important in ‘shaping’ communities and the organisms that live in them: predation. Predation is an every day phenomenon in modern animal communities and it has led to a great variety of adaptations among prey organisms in their quest to avoid being eaten, such as escape responses, camouflage, living in a cryptic habitat, chemical defence and the widespread development of armour. Gastropods afford an excellent opportunity for study since their shells are easily recognized from the Cambrian onwards, and usually contain a full record of their life history in their shells. The importance of predation as an evolutionary factor throughout the Phanerozoic, in a coevolution through geologic time between the prey and their predators, was emphasized by Vermeij (1977) in his paper on the ‘Mesozoic Marine Revolution’. Vermeij (1977, 1987) stated that predation increased with time from the Palaeozoic to Recent and that this development was seen in the evolution of shell morphology. Gastropod shell morphologies changed from simple unornamented forms in the Palaeozoic, to more sturdy forms with thick shells and protecting ornamentation in the Mesozoic. During the same time period an increase in the incidence of shell repair was also detected (Vermeij et al., 1981; Vermeij et al. 1982), coinciding with the evolution of new groups of predators (Vermeij, 1977, 1987). Increased predation pressure seemed to be one of the factors that fueled this radical evolution in shell morphology. Morphological changes induced by increased predation pressure have also been found in other groups of invertebrates (Meyer and Macurda, 1977; Signor and Brett, 1984; Aronson, 1991). As is frequently the case, increased awareness has stimulated description of more cases of repaired shell injuries after presumably failed predatory attacks in fossils other than gastropods (Strimple and Beane, 1966; Vorwald, 1982; Mapes and Hansen, 1984; Conway Morris and Jenkins, 1985; Stridsberg, 1985; Alexander, 1986; Bond and Saunders, 1989; Ebbestad and Högström, 2000, Kröger, 2004). A marked increase noted in the study of the relationships between predators and prey in fossil organisms, resulting in several publications, although this aspect is still in its infancy (Harper et al. 1998; Kowalewski and Kelley, 2002; Leighton, 2002; Kelley et al. 2003). Aims of this study In this project, Vermeij’s prediction concerning low rates of predation has been examined through studies of the frequency of shell repair in assemblages of Palaeozoic gastropods from different geological periods (Fig. 1). Comparative studies also explored the relationship between shell morphology and the frequency and form of shell breaks. Similar morphologies were examined through geological time to investigate potential variation in predatory strategies and responses. Studies of this kind usually require access to relatively large 9 collections of exquisitely preserved gastropods, which in itself has been a limiting factor in terms of the age and composition of the available fossil samples. Figure 1. Geological time table with the papers in the thesis indicated by an arrow at their approximate position in time. 10 Predation in the fossil record Detection of predation in fossils Widespread as predation is in every ecological community today, it is not a process which is easily measurable when it comes to fossils. In recent faunas, animals can be studied both in the laboratory and in the field, but how is predatory behaviour detected in fossils? Direct evidence of a predator caught ‘in action’ is unambiguous but such cases are extremely rare in the fossil record, for example the Ordovician asteroid humped over a bivalve in the extraoral feeding posture described by Blake and Guensburg (1994). Other unambiguous evidence of predatory
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