The Ichnologic Record of the Continental Invertebrate Invasion: Evolutionary Trends in Environmental Expansion, Ecospace Utilization, and Behavioral Complexity

The combined study of continental trace fossils and associated sedimentary facies provides valuable evidence of colonization trends and events throughout the Phanerozoic. Colonization of continental environments was linked to the exploitation of empty or under-utilized ecospace. Although the nonmarine trace fossil record probably begins during the Late Ordovician, significant invasion of nonmarine biotopes began close to the Silurian-Devonian transition with the establishment of a mobile arthropod epifauna (Diplichnites ichnoguild) in coastal marine to alluvial plain settings. Additionally, the presence of vertical burrows in Devonian high-energy fluvial deposits reflects the establishment of a stationary, deep suspension-feeding infauna of the Skolithos ichnoguild. The earliest evidence of plant-arthropod interaction occurred close to the Silurian-Devonian boundary, but widespread and varied feeding patterns are known from the Carboniferous. During the Carboniferous, permanent subaqueous lacustrine settings were colonized by a diverse, mobile detritus-feeding epifauna of the Mermia ichnoguild, which reflects a significant paleoenvironmental expansion of trace fossils. Paleozoic ichnologic evidence supports direct routes to the land from marginal marine environments, and migration to lakes from land settings. All nonmarine sedimentary environments were colonized by the Carboniferous, and subsequent patterns indicate an increase in ecospace utilization within already colonized depositional settings. During the Permian, backfilled traces of the Scoyenia ichnoguild record the establishment of a mobile, intermediate-depth, deposit-feeding infauna in alluvial and transitional alluvial-lacustrine sediments. Diversification of land plants and the establishment of ecologically diverse plant communities through time provided new niches to be exploited by arthropods. Nevertheless, most of the evolutionary feeding innovations took place relatively early, during the Late Paleozoic or early Mesozoic. A stationary deep infauna, the Camborygma ichnoguild, was developed in Triassic transitional alluvial-lacustrine deposits. Terrestrial environments hosted the rise of complex social behavioral patterns, as suggested by the probable presence of hymenopteran and isopteran nests in Triassic paleosols. An increase in diversity of trace fossils is detected in Triassic-Jurassic eolian deposits, where the ichnofauna displays more varied behavioral patterns than their Paleozoic counterparts. Also, a mobile, intermediate- depth, depositfeeding infauna, the Vagorichnus ichnoguild, was established in deep lake environments during the Jurassic. In contrast to Paleozoic permanent subaqueous assemblages typified by surface trails, Jurassic ichnocoenoses are dominated by infaunal burrows. High density of infaunal deposit-feeding traces of the Planolites ichnoguild caused major disruption of lacustrine sedimentary fabrics during the Cretaceous. Most insect mouthpart classes, functional feeding groups, and dietary guilds were established by the end of the Cretaceous. Diversification of modern insects is recorded by the abundance and complexity of structures produced by wasps, bees, dung-beetles, and termites in Cretaceous-Tertiary paleosols. The increase in bioturbation migrated from fluvial and lake-margin settings to permanent subaqueous lacustrine environments through time.