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Taphonomy of Insects in Carbonates and Amber

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Taphonomy of Insects in Carbonates and Amber Palaeogeography, Palaeoclimatology, Palaeoecology 203 (2004) 19^64 www.elsevier.com/locate/palaeo Taphonomy of insects in carbonates and amber Xavier Mart|¤nez-Delclo's a, Derek E.G. Briggs b;Ã, Enrique Pen‹alver c a Dept. Estratigra¢a, Paleontologia i Geocie'ncies Marines, Fac. Geologia, Universitat de Barcelona, 08028 Barcelona, Spain b Department ofGeology and Geophysics, Yale University, P.O.Box 208109, New Haven, CT 06520-8109, USA c Institut Cavanilles de Biodiversitat i Biologia evolutiva, Universitat de Vale'ncia, Apart. 2085 Vale'ncia, Spain Received 15 July 2002; received in revised form 11 August 2003; accepted 16 September 2003 Abstract The major taphonomic processes that control insect preservation in carbonate rocks (limestones, travertines and nodules) are biological: insect size and wingspan, degree of decomposition, presence of microbial mats, predation and scavenging; environmental: water surface tension, water temperature, density and salinity, current activity; and diagenetic: authigenic mineralisation, flattening, deformation, carbonisation. The major taphonomic processes that control the preservation of insects in fossil resins (amber and copal) are different, but can be considered under the same headings ^ biological: presence of resin producers, size and behaviour of insects; environmental: latitude, climate, seasonality, resin viscosity, effects of storms and fires, soil composition; and diagenetic: resin composition, insect dehydration, pressure, carbonisation, thermal maturation, reworking, oxidation. These taphonomic processes are geographically and temporally restricted, and generate biases in the fossil record. Nevertheless, where insects occur they may be abundant and very diverse. Taphonomic processes may impact on phylogenetic and palaeobiogeographic studies, in determining the timing of the origin and extinction of insect groups, and in identifying radiations and major extinctions. Taphonomic studies are an essential prerequisite to the reconstruction of fossil insect assemblages, to interpreting the sedimentary and environmental conditions where insects lived and died, and to the investigation of interactions between insects and other organisms. ß 2003 Elsevier B.V. All rights reserved. Keywords: fossil insects; preservation; limestone; resin 1. Introduction their diversity through time have been reviewed (Carpenter, 1992; Labandeira and Sepkoski, Insects are by far the most diverse and success- 1993; Jarzembowski and Ross, 1996; Ross et ful group of macroscopic organisms and they play al., 2000; Jarzembowski, 2001b) as well as the an important role in all the terrestrial ecosystems palaeobiology of insect feeding (Labandeira et that they inhabit. The fossil record of insects and al., 1994; Labandeira, 1997). The earliest record of the Insecta is from the Lower Devonian of Gaspe¤ (Que¤bec) (Labandeira et al., 1988), but * Corresponding author. Tel./Fax: +1-203-432-8590. E-mail addresses: [email protected] the group is not evident in the fossil record in (X. Mart|¤nez-Delclo's), [email protected] (D.E.G. Briggs), signi¢cant numbers until the Upper Carbonifer- [email protected] (E. Pen‹alver). ous (Brauckmann et al., 1995). The number of 0031-0182 / 03 / $ ^ see front matter ß 2003 Elsevier B.V. All rights reserved. doi:10.1016/S0031-0182(03)00643-6 PALAEO 3225 5-1-04 20 X. Mart|¤nez-Delclo's et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 203 (2004) 19^64 insect orders present in the Permian was similar to sta«tten which preserve non-biomineralised tissues that of now; the record of families identi¢es ma- are a critical source of palaeobiological data that jor periods of origination in the Permo^Carbon- are not available from the ‘shelly’ fossil record iferous, Early Jurassic, Early Cretaceous and Pa- (Briggs, 1995a). The preservation of delicate laeocene (Jarzembowski and Ross, 1996). The structures allows for more detailed comparison degree to which the recorded diversity of insects with recent organisms. The controls on the pres- is a re£ection of taphonomic processes is very ervation of non-biomineralised tissues, so-called di⁄cult to determine. A knowledge of the condi- soft-bodied fossils, are more complex than those tions that led to the preservation of insect biotas, on shelly taxa, which are dominantly sedimento- and of the biases introduced by taphonomic pro- logical (Kidwell, 1991) ^ they include organic cesses, is also essential for interpreting the role of matter input, microbial activity, and environmen- insects in terrestrial ecosystems, such as organic tal geochemistry (Allison and Briggs, 1991a,b; matter recycling and the pollination and distribu- Briggs, 2003a). tion of plant taxa. Here we review the major ta- Insect body fossils occur in a variety of envi- phonomic processes that a¡ect insects. ronmental settings including peat deposits (Ken- Insects lack biomineralised tissues and are usu- ward, 1976; Hayashi, 1994; Lavoie et al., 1997), ally considered by palaeontologists as soft-bodied deserts (e.g. Chihuahuan Desert, USA, Quater- organisms. Exceptional conditions are normally nary: Elias, 1990; desert sands Mauritania, Qua- required to ensure their preservation, but where ternary, Azar pers. commun.), lakes and rivers taphonomic processes are favourable, insects may (e.g. Montsec and Las Hoyas, Spain, Lower be extremely abundant. However, there is clearly Cretaceous: Mart|¤nez-Delclo's, 1995; Mele¤ndez, a signi¢cant range in susceptibility to decay, for 1995; Se¤zanne and Auriol travertines, France, Eo- example between £ies and beetles, re£ecting con- cene: Nel and Blot, 1990; Papazian and Nel, trasts in the degree of sclerotisation of the cuticle. 1989), deltas (e.g. Vosges, France, Middle Trias- Nonetheless, laboratory experiments have shown sic: Gall, 1996; Marchal-Papier, 1998), lagoons that even £y carcasses may survive in quiet sedi- (e.g. Solnhofen, Germany, Upper Jurassic: mentary settings for more than a year without Malz, 1976; Frickhinger, 1994), open marine en- disarticulating (Mart|¤nez-Delclo's and Martinell, vironments (e.g. northern Switzerland, Lower Ju- 1993). rassic: Etter and Kuhn, 2000), and deeper marine The most exceptionally preserved insects in turbidites (e.g. Borreda', Spain, Eocene: Gaudant sedimentary rocks occur in ¢ne-grained laminated and Busquets, 1996). The spatiotemporal distribu- carbonates in lacustrine and shallow marine set- tion of these palaeoenvironments was a major tings, where conditions may be suitable for the control on the insect fossil record. preservation of truly labile soft-tissues (e.g. Soln- Where insects are preserved in calcium carbon- hofen, Germany: Frickhinger, 1994). They pro- ate, precipitation may occur as calcite mud (e.g. vide a basis for contrasting preservation with Green River, USA, Eocene: Ferber and Wells, that in amber. The di¡erent taphonomic processes 1995), aragonite mud (e.g. Rubielos de Mora, that control the preservation of insects in carbon- Spain, Miocene: Pen‹alver, 1998), and dolomite ate rocks and in amber result in samples of di¡er- mud (e.g. Karatau, Kazakhstan, Upper Jurassic: ent insect communities. Amber normally preserves Seilacher et al., 1985). Fossil insects also are pre- insects regardless of their susceptibility to decay, served in a variety of other sedimentary contexts: but selective preservation is often a feature of in- clays and marls (Nel, 1986), siltstones (Mart|¤nez- sect assemblages in carbonates. The occurrence of Delclo's and Nel, 1991), sandstones (Nel et al., insects is normally considered to identify a fossil 1993), lacustrine diatomites (Hong, 1985; Riou, deposit as a Konservat-Lagersta«tte or conserva- 1995), cherts (Whalley and Jarzembowski, 1981), tion deposit (sensu Seilacher, 1970) where the em- evaporites (Priesner and Quievreux, 1935; Schlu«- phasis is on the quality of preservation rather ter and Kohring, 2001), phosphates (Handschin, than the abundance of fossils. Konservat-Lager- 1944), coal measures (Bartram et al., 1987; Shear PALAEO 3225 5-1-04 X. Mart|¤nez-Delclo's et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 203 (2004) 19^64 21 and Kukalova¤-Peck, 1990; Jarzembowski, 2001a), Evidence of ecologic relationships between in- and asphalt (Miller, 1983, 1997; Iturralde-Vinent sects and other animals is sometimes preserved in et al., 2000). Here we focus on the two most im- limestones, where it is usually con¢ned to plant^ portant sources of fossil insects, carbonates and insect interactions (Labandeira, 1998; Waggoner, amber. 1999). Such behaviour is inferred on the basis of Repositories are abbreviated as follows: functional morphology, or gut contents (Schaal MNHN, Muse¤um National d’Histoire Naturelle, and Ziegler, 1992; Krassilov et al., 1997)or Paris, France; PIN, Palaeontological Institute, more commonly coprolites (Rothwell and Scott, Russian Academy of Sciences, Moscow, Russia; 1988), or an array of primary evidence for insect MCAM, Museu de la Cie'ncia, Fundacio¤ ‘La feeding (Ro«Mler, 2000; Labandeira and Phillips, Caixa’, Barcelona, Spain; EPGM, Dept. Estrati- 2002). Amber, on the other hand, commonly re- gra¢a, Paleontologia i Geocie'ncies Marines, Univ. veals interactions, such as reproduction (mating, Barcelona, Barcelona, Spain; MCNA, Museo egg laying), commensalism and parasitism be- Ciencias Naturales de AŁ lava, Vitoria^Gasteiz, tween di¡erent insects, and between insects and Spain; MCCM, Museo de Ciencias de Castilla^ other organisms such as nematodes, spiders, pseu- La Mancha, Cuenca, Spain; MPV,
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