History of Ichnology: The Origins of Trace Fossil Taxonomy and the Contributions of Joseph F. James and Walter H. Häntzschel

The taxonomy of trace fossils has had a somewhat controversial history because they do not represent the actual animal remains but rather their work on and in the substrate. As such, traditional palaeontologists and zoologists have viewed them with some skepticism. Ichnologists owe a great debt to two geologists: Joseph F. James of Cincinnati and Walter H. Häntzschel of Hamburg, who took it upon themselves to impose some order on the chaos that constituted trace fossil taxonomy at the time. James, working independently and in ignorance of Alfred Nathorst, arrived at and utilized many of the same criteria his Swedish counterpart employed to criticize the fucoid origins of many trace fossils in the late 19th century. With his restudy of the systematics of Fucoides, Skolithos, and Arthrophycus, James brought to light many of the taxonomical nightmares that faced—and are still facing—the fledging science and can be rightfully considered the first trace fossil taxonomist. During the 1940s and 1950s, Häntzschel collected the widely scattered pertinent data from the literature, an immense task that, when published in 1962 (and later revised and expanded in 1975), made trace fossils accessible to further research and started a worldwide boom in trace fossil research.     

History of Ichnology: Early Ichnology Poems and Their Poets

In the early nineteenth century, the discovery of trace fossils fired the imagination of the discoverers and inspired them to compose unique poems. John Joly of Ireland wrote a poem about the invertebrate trace fossil Oldhamia that had been discovered in rocks around Bray Head. In the United States, the Reverend Edward Hitchcock discovered what he felt were giant bird tracks that later turned out to be reptile tracks. The traces inspired Hitchcock, who published a poem “The Sandstone Bird” in The Knickerbocker magazine under the pseudonym Poetaster. The poem is about a sorceress bringing the great sandstone bird back to life and probably represents the first ichnological poem.     

Ulisse Aldrovandi (1522–1605): The Study of Trace Fossils During the Renaissance

The Renaissance was a time of flourishing for literature, art and science. A prominent figure in Renaissance science was the Italian naturalist Ulisse Aldrovandi, considered today to be one of the founding fathers of modern paleontology. Aldrovandi is known to have widely studied body fossils, but his work on trace fossils has yet to be explored by science historians. This paper proposes to critically analyze Aldrovandi's approach to trace fossils, based on his writings (most of which are still unpublished) and on the illustrations that accompany them. In his studies, the Bologna-born naturalist accurately describes such ichnological celebrities as Gastrochaenolites and Cosmorhaphe and discusses a few theoretic principles on the trace-making process. The study of the works of Aldrovandi brings to light a fascinating and little-known source of the history of ichnology.     

Sir William Jardine and the Ichnology of Annandale

ABSTRACT

Sir William Jardine was a polymath; he was one of the world's foremost ornithologists, and ichthyologists, as well as a knowledgeable geologist, entomologist and botanist. He was a talented and skilled artist and engraver and a keen hunter and fisherman. He was also the author of the book The Ichnology of Annandale that represents the first book ever published on the subject of ichnology. It was here in this volume that Jardine coined the word ichnology. It is also one of the rarest ichnology publications, as it is believed that only 135–140 copies were ever printed. This volume remains the most lavishly illustrated book in vertebrate ichnology literature. This paper discusses the work of Jardine, the finds from his estate and his role in defining ichnology as a science.     

A Multifaceted Approach to Ichnology

Trace fossils link paleontology and sedimentology in ways that most body fossils cannot achieve. A multifaceted approach to ichnology helps to bridge the gap the biologic and geologic standpoints, to connect different levels of analysis (conceptual framework, explanatory schemata and applications), and to reconcile dichotomic views (e.g., adaptationist vs. interactionist approaches, time's arrow vs, time's cycle perspectives) by integrating the multiple aspects of the science of animal-substrate interactions. In this context, ichnologic investigations provide important links among numerous different fields. Analysis of specific ichnofaunas using this integrated approach may result in important and meaningful contributions to our understanding of paleoecology, sedimentology, sequence stratigraphy, reservoir characterization, biostratigraphy, evolutionary paleoecology, paleoceanography, and paleoclimatology.     

Ichnological analysis: The common ground between ichnofacies workers and ichnofabric analysts

This work compares the approaches taken by the two schools of applied ichnology (ichnofacies and ichnofabric analysts) and proposes an integrated approach for future improvement. There are many similarities between the two approaches, which mainly differ in the resolution at which ichnological analysis is undertaken. Ichnofabric work is by necessity performed on a bed-by-bed basis and interpreted at a scale consistent with that of the sedimentologist's facies. Ichnofacies studies, in contrast, seldom report bed-by-bed changes in ichnological content, instead highlighting bedset to facies association-scale changes in ichnology through comparison with summary models of ichnological trends.This work highlights the fact that workers from both schools utilize the same tools, with slightly different emphases, to reach their ichnological conclusions. Contention between the two schools has in recent years centred upon which is the better method for studying trace fossils—an artificial construct that detracts from the fact that both are extremely useful to sedimentological and palaeoenvironmental studies. Ichnofabric analysis is in addition, a highly effective means of documenting bioturbated sedimentary fabrics, as the basis for petrophysical studies. Ichnofacies and ichnofabric analysts use the same toolbox of ichnological methods to determine ancient palaeoenvironments—the scale of observation is generally different, but the results closely comparable. The recognition of key stratal surfaces on ichnological grounds is also possible with both methodologies. Ichnofacies analysts look to recognise Glossifungites ichnofacies firmgrounds in offshore mudstones. Ichnofabric studies would look to recognise overprinting of fabrics created by groups of organisms from different palaeoenvironments, but associated with the same key surface.It is proposed that the terms “ichnofacies method” and “ichnofabric analysis” are abandoned in favour of a neutral Ichnological Analysis Method, encompassing the approaches and terminology of both to produce a unified, scientifically rigorous, user-friendly methodology.     

HISTORY OF ICHNOLOGY—Ichnological Eccentrics: The Curious Case of Dr. Joseph Barratt of Middletown,Connecticut

Joseph Barratt was a British-born and educated physician who settled in the United States in 1819. He had a great interest in natural history, collecting both plants and insects and studying geology, mycology, ornithology, chemistry, meteorology, Native Americans, and local history. He was apparently a man of great energy and ambition but one who could not focus to see a project to completion. Barratt was active in the early history of the discovery of vertebrate footprints in the Newark Supergroup in the eastern United States but latter developed some very strange theories regarding the age and significance of these deposits. In his latter years, Barratt's mental state deteriorated and he became even more obsessed with trying to publicize his outlandish theories. Dr. Joseph Barratt is remembered in ichnology for basically two things; he sold Edward Hitchcock a superb specimen of vertebrate tracks; and he may have the most elaborate ichnological tombstone ever constructed.     

Birding by Foot: A Critical Look at the Synapomorphy- and Phenetic-Based Approaches to Trackmaker Identification of Enigmatic Tridactyl Mesozoic Traces

One of the goals of vertebrate ichnology is to use trace fossils as an additional source of data to determine the palaeoecological makeup of vertebrate paleoecosystems. The features in both the synapomorphy-based and phenetic-based methods of attributing a trace to an osteologic trackmaker are those that are affected by preservational conditions, convergent due to size and/or habitat of the trackmaker, and are morphologically variable within taxa. Despite the drawbacks, the phenetic-based, or “gestalt” method, is still the most comprehensive, if not always synapomorphy-supported, means of using the largest amount of data (morphologic and behavioral) preserved for identifying tracks as avian. To date there are too few synapomorphies that are both pedes specific and are consistently preserved in footprints to be a practical method for attributing tracks to an avian trackmaker. There is still much more comparative ichnological and statistical work to be done to discern novel traits that can be used to delineate between the traces of large avian and small nonavian Mesozoic theropods.     

Ichnology and sedimentology of a tide‐influenced delta in the Ordovician from the northeastern Alborz range of Iran (Kopet Dagh region)

The Middle–Late Ordovician Ghelli Formation in the Kopet Dagh region northeastern Alborz range of Iran is composed of siliciclastic rocks deposited in a variety of turbidite to marginal marine environments (deep marine clastic fan and related environments and prodelta, distal delta front, proximal delta front and distributary channels). The ichnology of the middle member of the Ghelli Formation is here reported. Combined sedimentological and ichnological analysis allows recognition of a tide‐influenced deltaic succession in the middle member of the Ghelli Formation consisting of three main facies associations: lower distributary channels, delta front and prodelta facies. Twenty‐two ichnogenera have been identified in this marginal marine succession: Arenicolites, Arthrophycus, Bergaueria, Chondrites, Conichnus, Cruziana, Cylindrichnus, Didymaulichnus, Diplichnites, Diplocraterion, Helminthopsis, Gyrochorte, Lockeia, Macaronichnus, Monomorphichnus, Palaeophycus, Planolites, Rosselia, Rusophcus, Skolithos, Teichichnus and ?Scolicia. Their distribution is clearly linked with lithofacies and depositional palaeoenvironments. The ichnological analysis reveals that the trace fossil suites developed in these environments indicate low diversity and low to moderate abundance of burrows, poor development of tiering and sporadic distribution. Low ichnodiversity and low bioturbation intensities with higher bioturbation clusters consist of facies‐crossing ichnogenera, and the impoverishment of suspension‐feeding trophic types indicates ‘stressed’, non‐archetypal expression of the Cruziana ichnofacies. The periodic higher intensities of bioturbation due to variations in hydrodynamic regimes of tidal currents reflect the archetypal of the Cruziana ichnofacies (and rare Skolithos ichnofacies).     

Trace fossils from lower Cambrian Hongjingshao Formation,Yunnan, China: Taxonomy,palaeoecology, palaeoenvironment

The Hongjingshao Formation (Cambrian Series 2, Stage 3) yields abundant, complex trace fossils that have not been systematically investigated up till now. Our detailed ichnological study of the traces from this formation in Malong area, Yunnan, China, recognizes four groups: (1) simple horizontal or sub-horizontal burrows; (2) complex branched burrow systems; (3) arthropod trails and; (4) simple vertical burrows belonging to the Cruziana ichnofacies. A trophic web is reconstructed for the community in Hongjingshao Formation based on both trace and body fossils. The ichnological and sedimentological features of the formation indicate an intertidal setting. The trace fossil assemblage demonstrates that Early Cambrian organisms were able to colonize very shallow marine environments, which further supports the landward expansion of the Early Cambrian ecosystems.     

Making Tracks

The science of ichnology has grown out of its early phase of infancy, but has not yet entered into the stage of senility. As long as we ichnologists continue playing around with trace fossils just for the fun of it while at the same time employing trace fossils as serious tools in solving geologic problems, our science will continue to be a dynamic and exciting pursuit for many years to come.     

Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca,Spain

Gibert, J.M. de, Mas, G. & Ekdale, A.A. 2012: Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca, Spain. Lethaia, Vol. 45, pp. 574–585. Unusual helical trace fossils occur in Tortonian shelfal calcarenites in the island of Mallorca. Their morphology may be either simple (ichnogenus Gyrolithes) or, more commonly, consist of two concentric helical burrows (ichnogenus Lapispira). They bear a very characteristic pelleted lining and are associated and probably connected to Thalassinoides and Ophiomorpha burrow systems very abundant in the same unit. These features allow the interpretation that the tracemaker was a thalassinidean shrimp. The complex and compound nature of these trace fossils is comparable to that seen in other modern and fossil crustacean burrow systems, and it reflects the behavioural plasticity of the architects. □Trace fossils, ichnology, ichnofabrics, crustacean burrows, Miocene, shallow marine, Lapispira.     

A LEAFCUTTER BEE TRACE FOSSIL FROM THE MIDDLE EOCENE OF PATAGONIA, ARGENTINA, AND A REVIEW OF MEGACHILID (HYMENOPTERA) ICHNOLOGY

Abstract:  The ichnospecies Phagophytichnus pseudocircus isp. nov. is described to include trace fossils characterized by leaf-margin excisions showing eccentricity values of 0.35–0.65 and more than 270 degrees of an arc, a non cuspate margin and vein stringers or necrotic flaps of tissue along the margin. A method for determining ellipse eccentricity was performed on leaf discs obtained from the nests of the modern leafcutter bee Megachile rotundata (Hymenoptera: Megachilidae), which provided objectively obtained values comparable to the trace fossil from the middle Eocene of Argentina and other world-wide ichnological records, historically and subjectively considered to be 'circular' trace fossils and attributed to leafcutter bees. The material described herein represents the first evidence for fossil Megachilidae from the Southern Hemisphere.     

Bioerosion trace fossils on bones of the Cretaceous South American theropod Buitreraptor gonzalezorum Makovicky,Apesteguía and Agnolín, 2005 (Deinonychosauria)

The ichnological record provides valuable information on the lifestyle, behaviour, and other palaeobiological and palaeoecological aspects of the biota. Here, we describe an interesting case of bioerosion trace fossils in bones of Buitreraptor gonzalezorum Makovicky, Apesteguía and Agnolín, 2005, a deinonychosaurian theropod from the fossiliferous locality of La Buitrera, Río Negro, Patagonia, Argentina. The trace fossils are morphologically diverse and preserved in a great percentage of the skeleton, including the jaw, vertebrae and limbs. Four main groups of trace fossils have been informally named as Parallel-Edge Furrows, Overlapped Grooves, Punctures and Lined. Parallel-Edge Furrows are in turn subdivided into four subgroups: isolated furrows, parallel pairs, opposed pairs and a combination of parallel and opposed pairs. The bioerosion trace fossils were probably generated by scavenging activities, and the semi-articulated preservation of the skeleton and the small size of each individual trace indicate small-sized tracemakers. Mammals are the main candidates although some traces may have been generated by crocodyliforms and insects such as dermestids and termites. This evidence provides additional information about palaeoenvironmental conditions, taphonomic processes, taxonomic diversity and ecological relationships that characterised this part of northern Patagonia at Early Cretaceous times.     

Fossil footprints and what they mean for hominin paleobiology

Hominin footprints have not traditionally played prominent roles in paleoanthropological studies, aside from the famous 3.66 Ma footprints discovered at Laetoli, Tanzania in the late 1970s. This contrasts with the importance of trace fossils (ichnology) in the broader field of paleontology. Lack of attention to hominin footprints can probably be explained by perceptions that these are exceptionally rare and “curiosities” rather than sources of data that yield insights on par with skeletal fossils or artifacts. In recent years, however, discoveries of hominin footprints have surged in frequency, shining important new light on anatomy, locomotion, behaviors, and environments from a wide variety of times and places. Here, we discuss why these data are often overlooked and consider whether they are as “rare” as previously assumed. We review new ways footprint data are being used to address questions about hominin paleobiology, and we outline key opportunities for future research in hominin ichnology.     

Ichnodiversity and ichnodisparity: significance and caveats

Ichnodiversity has been used as a proxy for environmental stress and stability in facies interpretations and to reconstruct evolutionary radiations and colonization histories in evolutionary palaeoecology. The three components of global diversity are exported from ecology and adapted for ichnology. Alpha ichnodiversity is used for palaeoenvironmental characterization, being assessed for individual facies. Beta ichnodiversity is commonly overlooked, although it may provide information about degree of similarity between ichnofaunas formed along environmental gradients. Gamma ichnodiversity may provide clues to detect ichnofossil provincialism. The concept of disparity may also prove to be significant in ichnological studies. Whereas ichnodiversity refers to ichnotaxonomic richness, ichnodisparity provides a measure of the variability of morphological plans in biogenic structures. Changes in global ichnodiversity do not necessarily parallel changes in ichnodisparity. For example, while the Cambrian explosion involved a dramatic increase in both, the Ordovician radiation essentially reflects an increase only in the former. Ichnodiversity and ichnodisparity should be used with caution because they are both affected by taphonomic processes. High diversity of superficial to shallow‐tier trace fossils may result from enhanced preservation due a poorly developed mixed layer, rather than a true reflection of ecosystem performance, as shown by underexploited infaunal ecospace after biotic crises (e.g. end‐Permian mass extinction).     

Digital image treatment applied to ichnological analysis of marine core sediments

Characterization of trace fossils in marine core sediments is, most times, difficult due to the weak differentiation between biogenic structures and the host sediment, especially in pelagic and hemipelagic facies. This problem is accentuated where a high degree of bioturbation is associated with composite ichnofabrics. Simple methods are presented here based on modifications to image features such as contrast, brightness, vibrance, saturation, exposure, lightness, and color balance using the software Adobe Photoshop CS6 (Adobe Systems, San Jose, CA, USA) to enhance visibility and thus allow for a better identification of the trace fossils. Adjustments involving brightness, levels and vibrance generally give better results. This approach was applied to marine cores of pelagic and hemipelagic sediments obtained from the Integrated Ocean Drilling Program Expedition 339, Site U1385. Enhancing the digital images facilitates ichnological analysis through improving the visibility of weakly observed trace fossils, and in some cases revealing traces not detected previously.     

Cretaceous small scavengers: feeding traces in tetrapod bones from Patagonia, Argentina

Ecological relationships among fossil vertebrate groups are interpreted based on evidence of modification features and paleopathologies on fossil bones. Here we describe an ichnological assemblage composed of trace fossils on reptile bones, mainly sphenodontids, crocodyliforms and maniraptoran theropods. They all come from La Buitrera, an early Late Cretaceous locality in the Candeleros Formation of northwestern Patagonia, Argentina. This locality is significant because of the abundance of small to medium-sized vertebrates. The abundant ichnological record includes traces on bones, most of them attributable to tetrapods. These latter traces include tooth marks that provde evidence of feeding activities made during the sub-aerial exposure of tetrapod carcasses. Other traces are attributable to arthropods or roots. The totality of evidence provides an uncommon insight into paleoecological aspects of a Late Cretaceous southern ecosystem.     

Origin of the Mckean Complex:Evidence from Timberline

Abstract

The Fourth of July valley site (5BL 120), a singlecomponent hunting camp at timberline in the Colorado Front Range, was occupied approximately 6000 radiocarbon years ago. Projectile points from the occupation area are typologically intermediate between James Allen and McKean Lanceolate points, and between Pryor Stemmed and Duncan points. The age and timberline location of the site, together with the transitional nature of the projectile points, support Husted’s (1968) hypothesis that the McKean complex developed from Plano progenitors in high-mountain Altithermal refuge areas.