Evolutionary events may impact the geological carbon cycle via transient imbalances in silicate weathering, and such events have been implicated as causes of glaciations, mass extinctions, and oceanic anoxia. However, suggested evolutionary causes often substantially predate the environmental effects to which they are linked—problematic when carbon cycle perturbations must be resolved in less than a million years to maintain Earth's habitability. What is more, the geochemical signatures of such perturbations are recorded as they occur in widely distributed marine sedimentary rocks that have been densely sampled for important intervals in Earth history, whereas the fossil record—particularly on land—is governed by the availability of sedimentary basins that are patchy in both space and time, necessitating lags between the origination of an evolutionary lineage and its earliest occurrence in the fossil record. Here, we present a simple model of the impact of preservational filtering on sampling to show that an evolutionary event that causes an environmental perturbation via weathering imbalance should not appear earlier in the rock record than the perturbation itself and, if anything, should appear later rather than simultaneously. The Devonian Hangenberg glaciation provides an example of how evolutionary events might be more fruitfully considered as potential causes of environmental perturbations. Just as the last samplings of species lost in mass extinction are expected to come before the true environmental event, first appearance should be expected to postdate the geological expression of a lineage's environmental impact with important implications for our reading of Earth history.

Notosuchia is a group of mostly terrestrial crocodyliforms. The presence of a prominent crest overhanging the acetabulum, slender straight-shafted long bones with muscular insertions close to the joints, and a stable knee joint suggests that they had an erect posture. This stance has been proposed to be linked to endothermy, because it is present in mammals and birds and contributes to the efficiency of their respiratory systems. However, a bone paleohistological study unexpectedly suggested that Notosuchia were ectothermic organisms. The thermophysiological status of Notosuchia deserves further analysis, because the methodology of the previous study can be improved. First, it was based on a relationship between red blood cell size and bone vascular canal diameter tested using 14 extant tetrapod species. Here we present evidence for this relationship using a more comprehensive sample of extant tetrapods (31 species). Moreover, contrary to previous results, bone cross-sectional area appears to be a significant explanatory variable (in addition to vascular canal diameter). Second, red blood cell size estimations were performed using phylogenetic eigenvector maps, and this method excludes a fraction of the phylogenetic information. This is because it generates a high number of eigenvectors requiring a selection procedure to compile a subset of them to avoid model overfitting. Here we inferred the thermophysiology of Notosuchia using phylogenetic logistic regressions, a method that overcomes this problem by including all of the phylogenetic information and a sample of 46 tetrapods. These analyses suggest that Araripesuchus wegeneri, Armadillosuchus arrudai, Baurusuchus sp., Iberosuchus macrodon, and Stratiotosuchus maxhechti were ectothermic organisms.

The rise of jawed vertebrates (gnathostomes) and extinction of nearly all jawless vertebrates (agnathans) is one of the most important transitions in vertebrate evolution, but the causes are poorly understood. Competition between agnathans and gnathostomes during the Devonian period is the most commonly hypothesized cause; however, no formal attempts to test this hypothesis have been made. Generally, competition between species increases as morphological similarity increases; therefore, this study uses the largest to date morphometric comparison of Silurian and Devonian agnathan and gnathostome groups to determine which groups were most and least likely to have competed. Five agnathan groups (Anaspida, Heterostraci, Osteostraci, Thelodonti, and Furcacaudiformes) were compared with five gnathostome groups (Acanthodii, Actinopterygii, Chondrichthyes, Placodermi, and Sarcopterygii) including taxa from most major orders. Morphological dissimilarity was measured by Gower's dissimilarity coefficient, and the differences between agnathan and gnathostome body forms across early vertebrate morphospace were compared using principal coordinate analysis. Our results indicate competition between some agnathans and gnathostomes is plausible, but not all agnathan groups were similar to gnathostomes. Furcacaudiformes (fork-tailed thelodonts) are distinct from other early vertebrate groups and the least likely to have competed with other groups.

Bryozoans are active non-phototrophic biomineralizers that precipitate their calcareous skeletons in seawater. Carbonate saturation states varied temporally and spatially in Paleozoic oceans, and we used the Bryozoan Skeletal Index (BSI) to investigate whether bryozoan calcification was controlled by seawater chemistry in Paleozoic trepostome and cryptostome bryozoans. Our results show that cryptostome bryozoan genera were influenced by ocean chemistry throughout the Paleozoic and precipitated the most calcite at lower latitudes, where carbonate saturation states are generally higher, and less in midlatitudes, where carbonate will be relatively undersaturated. Trepostome bryozoan genera show a similar but weaker trend for the Ordovician to Devonian, suggesting that, like the cryptostomes, they were unable to metabolically overcome falling saturation states and simply precipitated less robust skeletons at higher latitudes. Carboniferous to Triassic trepostomes differ, however, and show a trend toward increased calcification at higher latitudes, indicating an ability to overcome unfavorable carbonate saturation states. Analysis of Permian trepostomes at the species level indicates this is most pronounced in the Southern Hemisphere, where calcification is matched by increased feeding capacity. It is proposed that this increased feeding capacity allowed trepostomes to metabolically overcome unfavorable carbonate saturation states. The differing responses of trepostome and cryptostome bryozoans to carbonate saturation states suggest that bryozoans should not be considered as a single group in marine extinctions linked to ocean chemistry changes. Likewise, it would suggest that modern stenolaemate and gymnolaemate bryozoans should be treated separately when considering their response to modern ocean chemistry changes.

The repeated return of tetrapods to aquatic life provides some of the best-known examples of convergent evolution. One comparison that has received relatively little focus is that of mosasaurids (a group of Late Cretaceous squamates) and archaic cetaceans (the ancestors of modern whales and dolphins), both of which show high levels of craniodental disparity, similar initial trends in locomotory evolution, and global distributions. Here we investigate convergence in skull ecomorphology during the initial aquatic radiations of these groups. A series of functionally informative ratios were calculated from 38 species, with ordination techniques used to reconstruct patterns of functional ecomorphospace occupation. The earliest fully aquatic members of each clade occupied different regions of ecomorphospace, with basilosaurids and early russellosaurines exhibiting marked differences in cranial functional morphology. Subsequent ecomorphological trajectories notably diverge: mosasaurids radiated across ecomorphospace with no clear pattern and numerous reversals, whereas cetaceans notably evolved toward shallower, more elongated snouts, perhaps as an adaptation for capturing smaller prey. Incomplete convergence between the two groups is present among megapredatory and longirostrine forms, suggesting stronger selection on cranial function in these two ecomorphologies. Our study highlights both the similarities and divergences in craniodental evolutionary trajectories between archaic cetaceans and mosasaurids, with convergences transcending their deeply divergent phylogenetic affinities.

Understanding current and future biodiversity responses to changing climate is pivotal as anthropogenic climate change continues. This understanding is complicated by the multitude of available metrics to quantify dynamics and by biased sampling protocols. Here, we investigate the impact of sampling protocol strategies using a data-rich fossil record to calculate effective diversity using Hill numbers for the first time on Paleogene planktonic foraminifera. We sample 22,830 individual tests, in two different size classes, across a 7 Myr time slice of the middle Eocene featuring a major transient warming event, the middle Eocene climatic optimum (MECO; ~40 Ma), at study sites in the midlatitude North Atlantic. Using generalized additive models, we investigate community responses to climatic fluctuations. After correcting for any effects of fossil fragmentation, we show a peak in generic diversity in the early and middle stages of the MECO as well as divergent trajectories between the typical size-selected community (>180 μm) and a broader assemblage, including smaller genera (>63 μm). Assemblages featuring smaller genera are more resilient to the climatic fluctuations of the MECO than those assemblages that feature only larger genera, maintaining their community structure at the reference Hill numbers for Shannon's and Simpson's indices. These results raise fundamental questions about how communities respond to climate excursions. In addition, our results emphasize the need to design studies with the aim of collecting the most inclusive data possible to allow detection of community changes and determine which species are likely to dominate future environments.

Developmental plasticity, where traits change state in response to environmental cues, is well studied in modern populations. It is also suspected to play a role in macroevolutionary dynamics, but due to a lack of long-term records, the frequency of plasticity-led evolution in deep time remains unknown. Populations are dynamic entities, yet their representation in the fossil record is a static snapshot of often isolated individuals. Here, we apply for the first time contemporary integral projection models (IPMs) to fossil data to link individual development with expected population variation. IPMs describe the effects of individual growth in discrete steps on long-term population dynamics. We parameterize the models using modern and fossil data of the planktonic foraminifer Trilobatus sacculifer. Foraminifera grow by adding chambers in discrete stages and die at reproduction, making them excellent case studies for IPMs. Our results predict that somatic growth rates have almost twice as much influence on population dynamics than survival and more than eight times more influence than reproduction, suggesting that selection would primarily target somatic growth as the major determinant of fitness. As numerous paleobiological systems record growth rate increments in single genetic individuals and imaging technologies are increasingly available, our results open up the possibility of evidence-based inference of developmental plasticity spanning macroevolutionary dynamics. Given the centrality of ecology in paleobiological thinking, our model is one approach to help bridge eco-evolutionary scales while directing attention toward the most relevant life-history traits to measure.

Conspicuous centers of biodiversity are frequently attributed to local conditions that promote speciation or resistance to extinction, but recent diversification studies indicate this mode of explanation might not be very general, so it may be fruitful to revisit the role of dispersal in concentrating biodiversity. Here we consider the processes underlying the marine diversity hotspot in the Indo-West Pacific among comatulid crinoids, suspension-feeding echinoderms conspicuous on modern tropical reefs. We used ancestral-range reconstruction on a phylogeny of extant crinoids, assembled a new occurrence database of fossil comatulids and interrogated it with probabilistic preservational models, and developed a morphological character matrix to estimate the relationships among living and fossil comatulids. Ancestral-range reconstruction on a phylogeny of extant comatulids recovers an origin outside the Indo-Pacific and elevated dispersal into it. A new occurrence database records the comatulid clade spreading out gradually from origin in the Early Jurassic of the West Tethys. Comatulids do not appear in their modern hotspot until the Oligocene, and taphonomic analyses show these results cannot be explained solely as a result of inadequate sampling in Asia and Oceania. Finally, phylogenetic analyses demonstrate that deeply nested crown-group comatulids had originated before the clade became well established in the East Tethys, implying many independent dispersals into the modern hotspot. These consilient results suggest a biodiversity hotspot that owes its existence to dispersals out of the ancient West Tethys rather than to elevated in situ diversification.

A length–frequency sample (n = 295) from a fossil population of the Ordovician trilobite Triarthrus eatoni Hall, 1838, assembled and analyzed by J. L. Cisne in 1973 is here reexamined using methods of length–frequency analysis commonly used in fishery science and marine biology. Theoretical considerations and the empirical data at hand suggest that the growth of T. eatoni was not “linear,” but asymptotic, as is the growth of most Recent marine invertebrates. The parameters of the von Bertalanffy growth function (L∞ = 41 mm, K = 0.29 yr−1) suggest that T. eatoni, which apparently lived in a challenging environment, grew somewhat more slowly than the extant marine isopod Ceratoserolis trilobitoides (Eights, 1833), used here as Recent analogue to T. eatoni. This trilobite probably lived up to 10 years, rather than the suggested 4 years, and its mortality rate was 15%–20% per year rather than 30%–40% per year. These represent the first estimates of trilobite absolute growth characteristics using methods known to accurately model growth in extant water-breathing ectotherms. These provide a baseline for trilobite growth that can be used to make inferences about growth in other species. The approach used here may also be applied to other trilobites for which suitable length–frequency data exist.

Dinosaur embryos cause a lot of excitement in the scientific literature and are often widely reported because of the general public's interest in dinosaur biology. Well-preserved, articulated oviraptorosaur embryos in eggs are usually interpreted as representing a stage of development close to hatching because of their large size and good level of skeletal ossification. Based on this evidence, a recent report suggested that the position of the one embryo's head was reminiscent of an avian-like hatching position. Here we explore how the developmental stage of well-preserved oviraptorosaur embryos can be estimated, rather than assumed. This will help in our understanding of their developmental biology and its evolutionary consequences. Using quantitative methods and comparison with modern crocodilian embryos, we show that all articulated oviraptorosaur embryos are small relative to the egg and most likely at a stage of development equivalent to around 50%–60% of the developmental period, that is, not even close to hatching. This conclusion is supported by the fact that many elements of the crocodilian skeleton are well ossified many weeks before hatching and the position of oviraptorosaur embryos’ heads was also comparable to a crocodilian embryo many days before hatching. Misunderstandings about the stage of the developmental biology of these well-preserved oviraptorosaur embryos hampers our understanding of the true nature of their reproductive biology. We urge a more conservative approach to their interpretation. This is important, because misunderstandings in the minds of the public about dinosaur biology are hard to counter once poorly evidenced ideas have been reported around the world.

The number of individuals of species varies, but estimating abundance, given incomplete and biased sampling in both contemporary and fossilized communities, is challenging. Here, we describe a new occupancy model in a hierarchical Bayesian framework with random effects, in which multispecies occupancy and detection are modeled as a means to estimate relative species abundance and relative population densities. The modeling framework is suited for temporal samples of fossil communities with repeated sampling including multiple species with similar preservation potential. We demonstrate our modeling framework using a fossil community of benthic organisms to estimate relative species abundance dynamics and changing relative population densities of focal species in nine (geological) time intervals over 2.3 Myr. We also explore potential explanatory factors (paleoenvironmental proxies) and temporal autocorrelation that could provide extra information on unsampled time intervals. The modeling framework is applicable across a wide range of questions on species-level dynamics in paleoecological community settings.

The rapid and accurate taxonomic identification of fossils is of great significance in paleontology, biostratigraphy, and other fields. However, taxonomic identification is often labor-intensive and tedious, and the requisition of extensive prior knowledge about a taxonomic group also requires long-term training. Moreover, identification results are often inconsistent across researchers and communities. Accordingly, in this study, we used deep learning to support taxonomic identification. We used web crawlers to collect the Fossil Image Dataset (FID) via the Internet, obtaining 415,339 images belonging to 50 fossil clades. Then we trained three powerful convolutional neural networks on a high-performance workstation. The Inception-ResNet-v2 architecture achieved an average accuracy of 0.90 in the test dataset when transfer learning was applied. The clades of microfossils and vertebrate fossils exhibited the highest identification accuracies of 0.95 and 0.90, respectively. In contrast, clades of sponges, bryozoans, and trace fossils with various morphologies or with few samples in the dataset exhibited a performance below 0.80. Visual explanation methods further highlighted the discrepancies among different fossil clades and suggested similarities between the identifications made by machine classifiers and taxonomists. Collecting large paleontological datasets from various sources, such as the literature, digitization of dark data, citizen-science data, and public data from the Internet may further enhance deep learning methods and their adoption. Such developments will also possibly lead to image-based systematic taxonomy to be replaced by machine-aided classification in the future. Pioneering studies can include microfossils and some invertebrate fossils. To contribute to this development, we deployed our model on a server for public access at www.ai-fossil.com.

A dramatic shift from carbonate-rich to chert-rich marine strata occurred during the Permian and is frequently attributed to the increased activity of siliceous sponges and their biosiliceous sedimentation. The first-order ecologic consequences of this transition, if any, remain opaque. We analyze fossil occurrence data from the Phosphoria Basin (western North America) to test whether the presence of siliceous sponges, which are correlated with basin-wide chert strata, influenced the recruitment of benthic fauna. Using published lithologic descriptions, we categorized fossil collections by formation, facies, and lithology and used these data to code detrended correspondence analysis and nonmetric multidimensional scaling ordinations. We also analyzed the clustering of taxa into faunal units termed biofacies.

Results from these analyses indicate that fossil collections occurring in chert and carbonate are closely associated in faunal composition and community structure. These collections preferentially occur in the inner- to mid-ramp facies, in agreement with previous studies. Although largely similar in composition, collections of chert and carbonate lithology exhibit differences in the frequency and abundance of accessory brachiopod taxa (e.g., Composita and Hustedia), possibly a result of greater biosiliceous sedimentary input.

This study compares the δ15N values and the trophic position of two seabird species throughout the late Holocene in three regions in the southwestern Atlantic Ocean to assess the hypothesis that the decimation of megafauna led to changes in the trophic position of mesopredators. Modern and ancient mollusk shells were also analyzed to account for changes in the isotopic baseline through time. Results revealed that modern Magellanic penguins have higher δ15N values than their ancient conspecifics in the three regions, after controlling for changes in the isotopic baseline. This was also true for modern Imperial shags compared with ancient unidentified cormorants/shags from the two areas where ancient specimens were recovered (southern Patagonia and the Beagle Channel). Such temporal variability might be caused by three non–mutually exclusive processes: decreased availability of pelagic squat lobster resulting from decreasing primary productivity through the late Holocene, increased availability of small fishes resulting from the sequential depletion of other piscivores (South American fur seal and sea lion and Argentine hake) since the late eighteenth century, and modification of the migratory patterns of Magellanic penguins. Although disentangling the relative contribution of all those processes is impossible at this time, the results reported here demonstrate that the ecology of Magellanic penguins and Imperial shags has undergone major changes since the late Holocene.

We examine temporal and spatial variation in morphology of the ammonoid cephalopod Discoscaphites iris using a large dataset from multiple localities in the Late Cretaceous (Maastrichtian) of the U.S. Gulf and Atlantic Coastal Plains, spanning a distance of 2000 km along the paleoshoreline. Our results suggest that the fossil record of D. iris is consistent with no within-species net accumulation of phyletic evolutionary change across morphological traits or the lifetime of this species. Correlations between some traits and paleoenvironmental conditions as well as changes in the coefficient of variation may support limited population-scale ecophenotypic plasticity; however, where stratigraphic data are available, no directional changes in morphology occur before the Cretaceous/Paleogene (K/Pg) boundary. This is consistent with models of “dynamic” evolutionary stasis. Combined with knowledge of life-history traits and paleoecology of scaphitid ammonoids, specifically a short planktonic phase after hatching followed by transition to a nektobenthic adult stage, these data suggest that scaphitids had significant potential for rapid morphological change in conjunction with limited dispersal capacity. It is therefore likely that evolutionary mode in the Scaphitidae (and potentially across the broader ammonoid clade) follows a model of cladogenesis wherein a dynamic morphological stasis is periodically interrupted by more substantial evolutionary change at speciation events. Finally, the lack of temporal changes in our data suggest that global environmental changes had a limited effect on the morphology of ammonoid faunas during the latest Cretaceous.

The Cambrian evolutionary radiations are marked by spectacular biotic turnovers and the establishment of increasingly tiered food chains. At the base of these food chains are primary producers, which in the Cambrian fossil record are chiefly represented among organic-walled microfossils. The majority of these microfossil remains have traditionally been attributed to an informal category of incertae sedis called “acritarchs,” based entirely on form taxonomy. Acritarch form taxa have been intensely used for biostratigraphy and in large-scale studies of phytoplankton diversity. However, both prospects have been challenged by cases of taxonomic inconsistencies and oversplitting arising from the large phenotypic plasticity seen among these microfossils. The acritarch form genus Skiagia stands as an ideal case study to explore these taxonomic challenges, because it encompasses a number of form species widely used in lower Cambrian biostratigraphy. Moreover, subtle morphological differences among Skiagia species were suggested to underlie key evolutionary innovations toward complex reproductive strategies. Here we apply a multivariate morphometric approach to investigate the morphological variation of Skiagia-plexus acritarchs using an assemblage sourced from the Buen Formation (Cambrian Series 2, Stages 3–4) of North Greenland. Our analysis showed that the species-level classification of Skiagia discretizes a continuous spectrum of morphologies. While these findings bring important taxonomic and biostratigraphic hurdles to light, the unequal frequency distribution of life cycle stages among Skiagia species suggests that certain elements of phytoplankton paleobiology are nonetheless captured by Skiagia form taxonomy. These results demonstrate the value of using morphometric tools to explore acritarch phenotypic plasticity and its potential ontogenetic and paleoecological drivers in Cambrian ecosystems.

Trace fossils record foraging behaviors, the search for resources in patchy environments, of animals in the rock record. Quantification of the strength, density, and nature of foraging behaviors enables the investigation of how these may have changed through time. Here, we present a novel approach to explore such patterns using spatial point process analyses to quantify the scale and strength of ichnofossil spatial distributions on horizontal bedding planes. To demonstrate the utility of this approach, we use two samples from the terminal Ediacaran Shibantan Member in South China (between 551 and 543 Ma) and the early Cambrian Nagaur Sandstone in northwestern India (between 539 and 509 Ma). We find that ichnotaxa on both surfaces exhibited significant nonhomogeneous lateral patterns, with distinct levels of heterogeneity exhibited by different types of trace fossils. In the Shibantan, two ichnotaxa show evidence for mutual positive aggregation over a shared resource, suggesting the ability to focus on optimal resource areas. Trace fossils from the Nagaur Sandstone exhibit more sophisticated foraging behavior, with greater niche differentiation. Critically, mark correlation functions highlight significant spatial autocorrelation of trace fossil orientations, demonstrating the greater ability of these Cambrian tracemakers to focus on optimal patches. Despite potential limitations, these analyses hint at changes in the development and optimization of foraging at the Ediacaran/Cambrian transition and highlight the potential of spatial point process analysis to tease apart subtle differences in behavior in the trace fossil record.

Pteridinium simplex is an iconic erniettomorph taxon best known from late Ediacaran successions in South Australia, Russia, and Namibia. Despite nearly 100 years of study, there remain fundamental questions surrounding the paleobiology and paleoecology of this organism, including its life position relative to the sediment–water interface, and how it fed and functioned within benthic communities. Here, we combine a redescription of specimens housed at the Senckenberg Forschungsinstitut und Naturmuseum Frankfurt with field observations of fossiliferous surfaces, to constrain the life habit of Pteridinium and gain insights into the character of benthic ecosystems shortly before the beginning of the Cambrian. We present paleontological and sedimentological evidence suggesting that Pteridinium was semi-infaunal and lived gregariously in aggregated communities, preferentially adopting an orientation with the long axis perpendicular to the prevailing current direction. Using computational fluid dynamics simulations, we demonstrate that this life habit could plausibly have led to suspended food particles settling within the organism's central cavity. This supports interpretation of Pteridinium as a macroscopic suspension feeder that functioned similarly to the coeval erniettomorph Ernietta, emblematic of a broader paleoecological shift toward benthic suspension-feeding strategies over the course of the latest Ediacaran. Finally, we discuss how this new reconstruction of Pteridinium provides information concerning its potential relationships with extant animal groups and state a case for reconstructing Pteridinium as a colonial metazoan.

Neogene and Pleistocene African suids displayed convergent evolutionary trends in the third molar (M3) morphology, with increasingly elongated and higher crowns through time. While these features can prevent premature loss of masticatory functionality and potentially increase long-term reproductive success, changes in dental occlusal traits such as enamel complexity and thickness can also improve chewing efficiency and increase short-term energetic return. While both long-term and short-term benefits can contribute to the thriving of a lineage, the selective pressures associated with each category can be different. To examine how crown elongation correlates with these functional occlusal traits, we selected M3s of Kolpochoerus, Notochoerus, and Metridiochoerus from Kenya and South Africa, dated between 3.0 Ma and 0.4 Ma. To account for dental wear, we used micro-computed tomography imaging of unworn/slightly worn M3s to simulate wear progression within each tooth. We compared morphometric representatives of occlusal enamel complexity and thickness among the specimens following their respective wear trajectories. We found that M3 elongation correlates with higher occlusal complexity and thinner enamel in Notochoerus and Metridiochoerus lineages through time. In Kolpochoerus, enamel complexity and thickness were generally maintained through time, despite M3 elongation. The differences in M3 morphometric trends suggest that Kolpochoerus likely experienced a different set of selective pressures on functional occlusal traits compared with Notochoerus and Metridiochoerus. The shared evolutionary trends of M3 specialization among Notochoerus and Metridiochoerus suggest similar selective pressures on their chewing efficiency and the possibility of a dietary niche overlap in more xeric habitats.

Pollen malformations have been proposed as a paleoenvironmental stress proxy. However, the frequency and variability of pollen malformations under near-optimal conditions and environmental stress, as well as their developmental origins, remain unclear. To bridge these gaps, we compared pollen malformation frequencies and assemblages of 14 extant conifer genera of Pinaceae and Podocarpaceae producing saccate (winged) grains grown under near-optimal conditions. These baseline pollen yields were compared with those produced by Pinus mugo ‘Columnaris’ cultured under an abiotic stress—three experimentally heightened ultraviolet-B radiation (UV-B) regimes proposed for the end-Permian crisis. We additionally reviewed previous cytological literature of abnormal microsporogenesis in conifers. Under near-optimal conditions, malformations comprise <3% of pollen yields in 12 out of 13 bisaccate genera and >10% of yields in the naturally trisaccate Dacrycarpus dacrydioides. We detected no phylogenetic pattern in malformation assemblages of the baseline comparisons. UV-B–irradiated P. mugo produced significantly higher malformation frequencies and different assemblage compositions when compared with baseline bisaccate lineages. We propose that pollen malformations originate during the meiotic and tetrad stages of microsporogenesis and present a framework for the ontogeny of different malformation types seen in the fossil record. Malformations comprising >3% of bisaccate pollen yields can be used as a paleoenvironmental stress proxy, but rare, naturally trisaccate lineages are not suitable for such assessments. Furthermore, heightened UV-B not only increases pollen malformation production, but also alters the types of abnormalities trees produce. Different environmental stresses may therefore leave behind distinct fingerprints in the fossil record.