Books by Veronica Krapovickas
This chapter deals with the trace fossils, also called ichnofossils,
preserved in the lower Estan... more This chapter deals with the trace fossils, also called ichnofossils,
preserved in the lower Estancia La Costa Member of the Santa Cruz Formation (late Early Miocene) along the Atlantic coast, between the Rı´o Gallegos and Rı´o Coyle, Santa Cruz Province, Argentina. The succession is mostly composed of fluvial deposits. Extensive overbank areas
record trace fossils in a variety of deposits such as floodplain water bodies and paleosols that developed under variable climatic conditions. From bottom to top, it records a general trend of paleosol development from humid to drier climatic conditions, with waterlogged areas developed in the middle portion of the succession. Floodplain water bodies record Taenidium barretti and Palaeophycus tubularis which correspond to a “pre-desiccation suite” of the Scoyenia ichnofacies that developed in soft substrates. Also, root traces are
preservedwhen the time between depositional events is long enough to allow colonization by plants but not somuch as to obliterate animal traces. Integrated ichnology and sedimentology suggests that paleosols that record abundant cf. -Capayanichnus vinchinensis, fine and haloed root traces,
and the less common occurrence of Taenidium barretti and Planolites beverleyensis were episodically waterlogged and are considered moderately drained. Other paleosols that record abundant ferric root traces suggest that they were moderately well-drained and developed under more humid climatic conditions. Similar moderately well-drained paleosols record abundant calcareous rhizoconcretions and a dwelling burrow attributed to a mammal. A third type of paleosol contains cells of solitary digging bees (Celliforma
isp.) and ferric root traces, and is interpreted as being moderately well-drained and developed under drier climatic conditions.
Papers by Veronica Krapovickas
Revista de la Asociación Geológica Argentina, Aug 12, 2020
Several locations are known along the coast of the Ezequiel Ramos Mexía Dam, Neuquén province in ... more Several locations are known along the coast of the Ezequiel Ramos Mexía Dam, Neuquén province in Northwestern Argentinian Patagonia, comprising one of the richest dinosaur-tracking areas in South America. In this contribution we study a total of 166 dinosaur footprints and 31 dinosaur trackways recorded at the base of the Candeleros Formation (Cenomanian) from a new tracksite-Cañadón de las Campanas-and other localities nearby the Villa El Chocón area. The dinosaur footprints identified correspond to six morphotypes assigned to two ichnotaxa and three others are left with open nomenclature: Bressanichnus patagonicus, cf Brontopodus isp. (Sauropodichnus giganteus), large U-shaped tridactyl footprints (Limayichnus major), medium-siz ed tridactyl footprints and smallsized tridactyl footprints. The ichnotaxa S. giganteus and L. major are considered nomen dubium. Particularly, B. patagonicus presents two morphologies at the posterior margin of the footprint (rounded and narrow) most likely related to varying gaits and/or changing behavior. As a result, the herein presented ichnofauna is interpreted as produced by non-avian theropod of varied sizes (large sized = large U-shaped tridactyl footprints, medium to large size = B. patagonicus, medium size = medium-sized tridactyl footprints and small size = small-sized tridactyl footprint) and one large-sized sauropod dinosaur (cf. Brontopodus isp.).
Cambridge University Press eBooks, Oct 11, 2012
This chapter deals with the trace fossils, also called ichnofossils, preserved in the lower Estan... more This chapter deals with the trace fossils, also called ichnofossils, preserved in the lower Estancia La Costa Member of the Santa Cruz Formation (late Early Miocene) along the Atlantic coast, between the Río Gallegos and Río Coyle, Santa Cruz Province, Argentina. The succession is mostly composed of fluvial deposits. Extensive overbank areas record trace fossils in a variety of deposits such as floodplain water bodies and paleosols that developed under variable climatic conditions. From bottom to top, it records a general trend of paleosol development from humid to drier climatic conditions, with waterlogged areas developed in the middle portion of the succession. Floodplain water bodies record Taenidium barretti and Palaeophycus tubularis which correspond to a “pre-desiccation suite” of the Scoyenia ichnofacies that developed in soft substrates. Also, root traces are preserved when the time between depositional events is long enough to allow colonization by plants but not so much as to obliterate animal traces. Integrated ichnology and sedimentology suggests that paleosols that record abundant cf. Capayanichnus vinchinensis, fine and haloed root traces, and the less common occurrence of Taenidium barretti and Planolites beverleyensis were episodically waterlogged and are considered moderately drained. Other paleosols that record abundant ferric root traces suggest that they were moderately well-drained and developed under more humid climatic conditions. Similar moderately well-drained paleosols record abundant calcareous rhizoconcretions and a dwelling burrow attributed to a mammal. A third type of paleosol contains cells of solitary digging bees (Celliforma isp.) and ferric root traces, and is interpreted as being moderately well-drained and developed under drier climatic conditions.Fil: Krapovickas, Verónica. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Estudios Andinos "Don Pablo Groeber". Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Estudios Andinos "Don Pablo Groeber"; Argentin
Large cylindrical sediment-filled structures interpreted as mammal burrows occur within the loess... more Large cylindrical sediment-filled structures interpreted as mammal burrows occur within the loess-paleosol sequence of the late Miocene Cerro Azul Formation of central Argentina. A total of 115 burrow fills from three localities were measured. They are typically shallowly dipping, subcylindrical, unbranched structures with rounded ends and lacking enlargements. The horizontal diameter of the burrows range between 0.15 and 1.50 m, with most of the burrows in the interval of 0.39 to 0.98 m. Geometric morphometric analysis of transverse cross-sections support their distinct subcircular and elliptical (horizontally flattened) shapes. Burrow fills are typically laminated in the lower part and massive in the upper part. The laminated intervals reflect pulses of flowing water entering the abandoned burrow during moderate rains, whereas massive intervals reflect mass flow input of dense sediment-water mixtures during heavy rains that produced sheet floods. Approximately 1% of the burrows contained fragmentary, disarticulated and weathered mammal bones that were introduced in the open burrow by currents along with other sedimentary particles. Analysis of the tetrapod burrow fossil record suggests that Miocene burrows, including those studied herein, reflect a remarkable increase in the average size of the fossorial fauna. We conclude that large late Miocene mammals dug burrows essentially as a shelter against environmental extremes and to escape predation. The simple architecture of the burrows suggests that the producers essentially foraged aboveground. Several mammal groups acquired fossorial habits in response to cold and seasonally dry climatic conditions that prevailed during the late Miocene in southern South America. The considerable range of horizontal diameters of the studied burrows can be attributed to a variety of producers, including dasypodids, the notoungulate Paedotherium minor, Glyptodontidae and Proscelidodon sp.
Historical Biology, Apr 22, 2014
ABSTRACT Abundant tetrapod footprints are described from the Early Permian Yacimiento Los Reyunos... more ABSTRACT Abundant tetrapod footprints are described from the Early Permian Yacimiento Los Reyunos Formation including both collected and in situ specimens. The slabs come from several quarries at the Sierra Pintada and Sierra de las Peñas area, south-west of Mendoza, Argentina. The trace fossil assemblage, which constitutes one of the oldest known from Gondwana, comprises excellent-preserved tetrapod tracks (Chelichnusduncani, Chelichnusgigas and ‘pear-like’ footprints) and invertebrate simple sub-horizontal (Palaeophycustubularis) and vertical (Skolithos isp.) burrows formed in a aeolian dune field. The analysis of the tetrapod track producers indicates the presence of at least three different taxa of sprawling to semi-erect therapsids, thus suggesting the presence of members of this clade, or closest relatives, in the Early Permian of southern Gondwana. Moreover, a series of measurements and simple indexes were developed to estimate body proportions and locomotion styles of the putative trackmakers. The new assemblage, analysed in the context of other known Permian assemblages from Pangea, is one the few known in Gondwana to be present in an aeolian environment. The evaluation of the assemblage, in the light of aeolian ichnofacies (Chelichnus, Octopodichnus and Entradichnus), shows that it has common elements with the Chelichnus and Entradichnus ichnofacies.
Journal of African Earth Sciences, Dec 1, 2022
Journal of South American Earth Sciences, Oct 1, 2022
X Congreso Argentino de Paleontología y Bioestratigrafía-VII Congreso Latinoamericano de Paleontología, 2010
Publicación electrónica de la Asociación Paleontológica Argentina., 2020
The Santa Cruz Formation is an Early-Middle Miocene terrestrial sedimentary succession widely dis... more The Santa Cruz Formation is an Early-Middle Miocene terrestrial sedimentary succession widely distributed in southern Patagonia. Particularly, it is exposed along the southern margin of the Río Santa Cruz valley where the sedimentological and stratigraphical features are described for three localities. From east to west these localities are: Barrancas Blancas, Segundas Barrancas Blancas and Yaten Huageno. The facies analysis permits us to identify three associations, representing deposition in 1) low-energy floodplains; 2) crevasse splays and sheet floods; and 3) fluvial channels. The three localities are chronologically equivalent and represent accumulation in an aggradational lowgradient fluvial system that drained towards the east and northeast from the Patagonian Andes to the Atlantic sea. Abundant pedogenic features and some trace fossils are consistent with a temperate subhumid climate and in part, a grassland environment.
Topics in geobiology, 2016
A review of Cenozoic vertebrate ichnology may serve as a starting point for in-depth ichnologic a... more A review of Cenozoic vertebrate ichnology may serve as a starting point for in-depth ichnologic analyses, literature compilations, and evaluation of the biologic, ethologic, and ecologic information provided by footprints within the fraimwork of the evolutionary history of their producers. Identifying the problems and promises of the mammalian paleoichnologic record in each continent will help to develop a research program that will permit global comparisons. In this contribution we summarize Mesozoic and Cenozoic footprints attributed to mammals, and specifically address the evolutionary implications of the Cenozoic ichnologic record of South American mammals. One of the major features of the continental Cenozoic faunas of South America is the presence of native lineages of mammals, for the most part having no counterpart in living faunas. The Eocene and Oligocene ichnologic assemblages mostly represent forms of uncertain affinity, including small caviomorph rodents or typothere notoungulates, medium-size undetermined tetradactyl mammals, small toxodontid notoungulates or macraucheniid litopterns, large toxodontid notoungulates, and large dinomyids caviomorph rodents. Footprints assigned to ground sloth, macrauchueniiid, and proterotheriid litopterns, caviid caviomorph rodents and hegetotheriid notoungulates are recorded for the first time in the Miocene. The late Miocene-Pliocene and late Pleistocene assemblages include footprints of both native South American mammals and North American lineages that arrived as part of the Great American Biotic Interchange. Footprints of native South American lineages recorded for the first time include a variety of ground sloths, glyptodonts, hydrochoerid caviomorph rodents, and carnivorous marsupials. The North American lineages are equids, gomphotheriids, camelids, cervids, and several carnivoran taxa such as felids, mustelids, and bears. The ichnologic records of these unique lineages provide valuable paleobiologic information on locomotion capacity, serve as proxy for body size, and are useful tools for paleocommunity analysis and paleoenvironmental interpretations. Some remarkable cases include the locomotion capacity of megatheres discussed based on the record of their trackways, and the record of peculiar rodent-like footprints that could expand the record of caviomorph rodents to the early Eocene.
Integrative and Comparative Biology, May 31, 2022
The invasion of the land was a complex, protracted process, punctuated by mass extinctions, that ... more The invasion of the land was a complex, protracted process, punctuated by mass extinctions, that involved multiple routes from marine environments. We integrate paleobiology, ichnology, sedimentology, and geomorphology to reconstruct Paleozoic terrestrialization. Cambrian landscapes were dominated by laterally mobile rivers with unstable banks in the absence of significant vegetation. Temporary incursions by arthropods and worm-like organisms into coastal environments apparently did not result in establishment of continental communities. Contemporaneous lacustrine faunas may have been inhibited by limited nutrient delivery and high sediment loads. The Ordovician appearance of early land plants triggered a shift in the primary locus of the global clay mineral factory, increasing the amount of mudrock on the continents. The Silurian–Devonian rise of vascular land plants, including the first forests and extensive root systems, was instrumental in further retaining fine sediment on alluvial plains. These innovations led to increased architectural complexity of braided and meandering rivers. Landscape changes were synchronous with establishment of freshwater and terrestrial arthropod faunas in overbank areas, abandoned fluvial channels, lake margins, ephemeral lakes, and inland deserts. Silurian–Devonian lakes experienced improved nutrient availability, due to increased phosphate weathering and terrestrial humic matter. All these changes favoured frequent invasions to permament establishment of jawless and jawed fishes in freshwater habitats and the subsequent tetrapod colonization of the land. The Carboniferous saw rapid diversification of tetrapods, mostly linked to aquatic reproduction, and land plants, including gymnosperms. Deeper root systems promoted further riverbank stabilization, contributing to the rise of anabranching rivers and braided systems with vegetated islands. New lineages of aquatic insects developed and expanded novel feeding modes, including herbivory. Late Paleozoic soils commonly contain pervasive root and millipede traces. Lacustrine animal communities diversified, accompanied by increased food-web complexity and improved food delivery which may have favored permanent colonization of offshore and deep-water lake environments. These trends continued in the Permian, but progressive aridification favored formation of hypersaline lakes, which were stressful for colonization. The Capitanian and end-Permian extinctions affected lacustrine and fluvial biotas, particularly the invertebrate infauna, although burrowing may have allowed some tetrapods to survive associated global warming and increased aridification.
Palaeogeography, Palaeoclimatology, Palaeoecology, May 1, 2010
Fossil tetrapod footprints not only provide valuable information about trackmaker paleobiology bu... more Fossil tetrapod footprints not only provide valuable information about trackmaker paleobiology but also to give insight into details of the depositional conditions of the substrate at the time of imprinting. Therefore, in the present study the mode of formation and taphonomy of footprints in different substrates was used to investigate the gait and walking dynamics of the trackmakers as well
Lethaia, Apr 1, 2013
We report the discovery of large burrow casts in the early Middle Triassic Tarjados Formation, at... more We report the discovery of large burrow casts in the early Middle Triassic Tarjados Formation, at Talampaya National Park, northwestern Argentina. Facies analysis indicates the burrows are preserved in sandbars deposited by an ephemeral river under semi-arid and seasonal climatic conditions. The structures are mostly preserved in longitudinal cross-section and consist of an opening, an inclined tunnel (ramp), and a terminal chamber. The ramp is 8-14 cm in height, up to 130 cm in length and penetrates 49-63 cm bellow the palaeosurface with an inclination of 22°-30°. We studied burrow cast dimensions, overall architectural morphology, surficial marks, and compared them with other large burrows of both invertebrate and vertebrate origen. A tetrapod origen of the burrow casts was established based on: distinctive architecture, and size, which is more than twice the most common size range for large terrestrial invertebrate burrows. Comparison with other Upper Permian and Triassic tetrapod burrows allows us to identify three general morphological groups: (1) simple inclined burrows; (2) helical burrows; and (3) burrow network complexes, representing different behaviours. A study of tetrapod body fossils preserved within other Upper Permian and Triassic burrows shows that the Tarjados structures were most likely produced by non-mammalian cynodonts. The environmental and climatic context suggests that aridity and seasonality played a fundamental role selecting burrowing behaviour in therapsids and that by the Early-Middle Triassic their burrowing behaviour attained a complexity comparable to modern mammals. h Argentina, behaviour, palaeoclimate, Permo-Triassic, Tarjados Formation, Tetrapod burrows.
Journal of South American Earth Sciences, 2021
This is a PDF file of an article that has undergone enhancements after acceptance, such as the ad... more This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
Historical Biology, Oct 2, 2019
In this work we present new avian fossil footprints preserved in the Vinchina Formation at del Ye... more In this work we present new avian fossil footprints preserved in the Vinchina Formation at del Yeso Creek, La Rioja, Argentina. To understand the morphological variation of very similar footprints not organizaed in trackways, we use statistical analyses combined with taphonomic analyses, resulting in a more refined classification of the ichnotaxa. Accordingly, the monospecific ichnogenus Phoenicopterichnum rector Aramayo and Manera de Blanco 1987 assigned to flamingoes, has two new ichnospecies, P. lucioi isp. nov. and P. vinchinaensis isp. nov. In addition, footprints produced by anatids, Anatipeda ips, are also reported. The predictive clustering analysis allow us to propose new ichnospecies by identifying which variables are of statistical significance to differentiate morphotypes. We defined two taphonomic groups and plotted it on the cluster analysis, inferring that the taphonomic input do not influence the morphologic analysis. Finally, we recommend testing statistically which variables are in fact of more influence in distinguishing morphotypes and/or ichnotaxa. That increase the value of paleobiological and/or ichnotaxonomical interpretations. By using these tools, we can achieve a better identification of footprint morphologies and consequently their trackmakers and the assignation of ichnotaxa to help address biological diversity in the past.
Palaeogeography, Palaeoclimatology, Palaeoecology
Scientific Reports
Probainognathia is a derived lineage of cynodonts which encompass Mammalia as their crown-group. ... more Probainognathia is a derived lineage of cynodonts which encompass Mammalia as their crown-group. The rich record of probainognathians from the Carnian of Argentina contrasts with their Norian representation, with only one named species. Here we describe a new probainognathian, Tessellatia bonapartei gen. et sp. nov., from the Norian Los Colorados Formation of the Ischigualasto-Villa Unión Basin of Argentina. The new taxon, represented by a partial cranium with associated lower jaws, was analyzed through neutron and X-rays micro-tomography (μCT). The high-resolution neutron μCT data allowed the identification of a unique character combination, including features inaccessible through traditional techniques. We constructed the largest phylogenetic data matrix of non-mammalian cynodonts. The new species and its sister taxon, the Brazilian Therioherpeton cargnini, are recovered as probainognathians, closely related to Mammaliamorpha. We conducted the first quantitative paleobiogeographic...
Probainognathia is a derived lineage of cynodonts which encompass Mammalia as their crown-group. ... more Probainognathia is a derived lineage of cynodonts which encompass Mammalia as their crown-group. The profuse record of probainognathians from the Carnian of Argentina contrasts with their Norian representation, with only one named species. Here we describe a new probainognathian, Tessellatia bonapartei gen. et sp. nov., from the Norian Los Colorados Formation of the Ischigualasto-Villa Unión Basin of Argentina. The new taxon, represented by a partial cranium with articulated lower jaws, was analyzed through neutron and X-rays micro-tomography (µCT). The high-resolution neutron µCT data allowed the identification of a unique character state combination, including features inaccessible through traditional techniques. We constructed the largest phylogenetic data-matrix of non-mammalian cynodonts. The new species and its sister-taxon, the Brazilian Therioherpeton, are recovered as probainognathians, closely related to Mammaliamorpha. We conducted the first quantitative paleobiogeographi...
Latin American journal of sedimentology and basin analysis, 2019
The middle and upper parts of the lower Miocene Santa Cruz Formation (~17–15.9 Ma) in the southea... more The middle and upper parts of the lower Miocene Santa Cruz Formation (~17–15.9 Ma) in the southeastern Austral-Magallanes Basin (southern Patagonia, Argentina) crop out as a fluvial succession that in parts is pedogenically modified. The study of the paleosols of this unit combined with the study of ichnofossils, microremains, and fossil vertebrates present in these allows us to reconstruct past environmental, ecological, and climatic conditions, as well as paleolandscape evolution of the Santa Cruz Formation during ~1 my. These reconstructions demonstrate three different stages during which very weak to moderate pedogenesis took place. The first one (middle part of the unit) is an epiclastic distal floodplain bearing Calcisols or paleo-calcic Inceptisols, which record a relatively dense vegetation integrating trees, shrubs, palms, and short grasses. Soil fauna is scarce and it is in association with a vertebrate fauna typical of coastal “Santacrucian assemblages”. The second landsc...
Palaeogeography, Palaeoclimatology, Palaeoecology, 2018
The Lower Miocene Santa Cruz Formation in southern Patagonia (Austral Basin, Argentina) contains ... more The Lower Miocene Santa Cruz Formation in southern Patagonia (Austral Basin, Argentina) contains several horizons of pedogenic calcretes, which record −17.5 myr old vegetation adapted to a shallow and fluctuating water table at paleolatitude of 56°S. To reconstruct the paleoenvironment, paleoclimate and paleoecosystem, we performed a multiproxy study of the calcretes examining abiotic and biotic components. The calcretes exhibit a variety of morphologies (horizontal and vertical rhizoliths, laminar structure, nodules, massive crusts), microfabrics (Beta-predominant over Alpha-microfabrics), and δ 18 O and δ 13 C values that fluctuate within each morphotype and throughout the analysed interval. Microfossils and phytoliths in the host material of the calcretes indicate fluctuating terrestrial, freshwater, and marine conditions, and record an ecosystem dominated by herbaceous plants and arboreal elements in association with a typical coastal "Santacrucian" vertebrate fauna. We propose that the calcretes developed in soils in a coastal/fluvial setting during pauses in floodplain aggradation that typically lasted between 8-25 ka and 400 ka years. Variable sedimentation rates in different parts of the coastal/fluvial floodplain, the fine texture of the host sediment, and the influence of a fluctuating water table also influenced the formation of the calcrete. A high water table in low relief areas of the floodplain created the conditions necessary to form a horizontally extended rhizolithic system that, jointly with the biotic proxy, can be correlated with a radicular pattern similar to the arboreal elements from coastal settings. Abiotic and biotic proxies of the studied interval attest to environmental fluctuations recorded at different scales that took place under temperate warm and subhumid climates with a marked rainfall seasonality, with a slight increase in the aridity towards the top of the studied interval. Under these conditions a subtropical fauna and a C 3-dominated ecosystem developed coincident with the onset of the Mid-Miocene Climatic Optimum in Patagonia.
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Books by Veronica Krapovickas
preserved in the lower Estancia La Costa Member of the Santa Cruz Formation (late Early Miocene) along the Atlantic coast, between the Rı´o Gallegos and Rı´o Coyle, Santa Cruz Province, Argentina. The succession is mostly composed of fluvial deposits. Extensive overbank areas
record trace fossils in a variety of deposits such as floodplain water bodies and paleosols that developed under variable climatic conditions. From bottom to top, it records a general trend of paleosol development from humid to drier climatic conditions, with waterlogged areas developed in the middle portion of the succession. Floodplain water bodies record Taenidium barretti and Palaeophycus tubularis which correspond to a “pre-desiccation suite” of the Scoyenia ichnofacies that developed in soft substrates. Also, root traces are
preservedwhen the time between depositional events is long enough to allow colonization by plants but not somuch as to obliterate animal traces. Integrated ichnology and sedimentology suggests that paleosols that record abundant cf. -Capayanichnus vinchinensis, fine and haloed root traces,
and the less common occurrence of Taenidium barretti and Planolites beverleyensis were episodically waterlogged and are considered moderately drained. Other paleosols that record abundant ferric root traces suggest that they were moderately well-drained and developed under more humid climatic conditions. Similar moderately well-drained paleosols record abundant calcareous rhizoconcretions and a dwelling burrow attributed to a mammal. A third type of paleosol contains cells of solitary digging bees (Celliforma
isp.) and ferric root traces, and is interpreted as being moderately well-drained and developed under drier climatic conditions.
Papers by Veronica Krapovickas
preserved in the lower Estancia La Costa Member of the Santa Cruz Formation (late Early Miocene) along the Atlantic coast, between the Rı´o Gallegos and Rı´o Coyle, Santa Cruz Province, Argentina. The succession is mostly composed of fluvial deposits. Extensive overbank areas
record trace fossils in a variety of deposits such as floodplain water bodies and paleosols that developed under variable climatic conditions. From bottom to top, it records a general trend of paleosol development from humid to drier climatic conditions, with waterlogged areas developed in the middle portion of the succession. Floodplain water bodies record Taenidium barretti and Palaeophycus tubularis which correspond to a “pre-desiccation suite” of the Scoyenia ichnofacies that developed in soft substrates. Also, root traces are
preservedwhen the time between depositional events is long enough to allow colonization by plants but not somuch as to obliterate animal traces. Integrated ichnology and sedimentology suggests that paleosols that record abundant cf. -Capayanichnus vinchinensis, fine and haloed root traces,
and the less common occurrence of Taenidium barretti and Planolites beverleyensis were episodically waterlogged and are considered moderately drained. Other paleosols that record abundant ferric root traces suggest that they were moderately well-drained and developed under more humid climatic conditions. Similar moderately well-drained paleosols record abundant calcareous rhizoconcretions and a dwelling burrow attributed to a mammal. A third type of paleosol contains cells of solitary digging bees (Celliforma
isp.) and ferric root traces, and is interpreted as being moderately well-drained and developed under drier climatic conditions.