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.

We highlight the historical and contemporary policies that govern paleontological research on federally recognized Native American lands. The United States has a long history of fossil dispossession from Indigenous Peoples, and federal policies surrounding the management of Native American lands (i.e., reservations), and the geological resources therein, have changed through time. These changes reflect shifting popular and political ideologies regarding Native American nations’ sovereignty and self-governance. As of 2022, the United States has a government-to-government relationship with federally recognized Tribal entities, but that has not always been the case. Historians have divided post-contact Native American federal policy into distinct eras: Colonial Times to 1820, Native American Removal and Reservation (1820–1887), Allotments and Attempted Assimilation (1887–1934), Reorganization and Preservation (1934–1953), Termination and Relocation (1953–1968), and Tribal Self-Determination (1968–present). Documentation of how the federal policies from each of these eras continue to impact current paleontological research is limited. We summarize major legislative actions, court cases, and historical events that have affected paleontological resource management in Native American territory. We use this historical context to identify federal policy gaps and highlight legal nuances associated with fossil collection and ownership, particularly given the importance of fossils to some Native Americans’ cultural patrimony. Finally, we explore how these gaps affect scientific research and highlight best practices for conducting paleontological research on vertebrate, invertebrate, and paleobotanical body and trace fossils using the CARE (Collective Benefit, Authority to Control, Responsibility, Ethics) Principles for Indigenous Data Governance (https://www.gida-global.org/care).

Over the last 50 years, access to new data and analytical tools has expanded the study of analytical paleobiology, contributing to innovative analyses of biodiversity dynamics over Earth's history. Despite—or even spurred by—this growing availability of resources, analytical paleobiology faces deep-rooted obstacles that stem from the need for more equitable access to data and best practices to guide analyses of the fossil record. Recent progress has been accelerated by a collective push toward more collaborative, interdisciplinary, and open science, especially by early-career researchers. Here, we survey four challenges facing analytical paleobiology from an early-career perspective: (1) accounting for biases when interpreting the fossil record; (2) integrating fossil and modern biodiversity data; (3) building data science skills; and (4) increasing data accessibility and equity. We discuss recent efforts to address each challenge, highlight persisting barriers, and identify tools that have advanced analytical work. Given the inherent linkages between these challenges, we encourage discourse across disciplines to find common solutions. We also affirm the need for systemic changes that reevaluate how we conduct and share paleobiological research.

The body armor of ankylosaurians is a unique morphological feature among dinosaurs. While ankylosaurian body armor has been studied for decades, paleohistological analyses have only started to uncover the details of its function. Yet there has been an overall bias toward sampling ankylosaurian remains from the Northern Hemisphere, with limited quantitative studies on the morphological and functional evolution of the osteoderms composing their body armor. Here, we describe new ankylosaurian materials recovered from the Late Cretaceous of Antarctica that, in combination with data compiled from the literature, reveal new insights into the evolution of the ankylosaurian body armor. Based on histological microstructure and phylogenetic results, the new Antarctic material can be assigned to Nodosauridae. This group shares the absence/poor development of their osteodermal basal cortex and highly ordered sets of orthogonal structural fibers in the superficial cortex. Our morphospace analyses indicate that large morphological diversity is observed among both nodosaurids and ankylosaurids, but osteoderms became more functionally specialized in late-diverging nodosaurids. Besides acting as effective protection against predation, osteoderms also exhibit highly ordered structural fibers in nodosaurids, enabling a decrease in cortical bone thickness (as in titanosaurs), which could have been co-opted for secondary functions, such as calcium remobilization for physiological balance. The latter may have played a key role in nodosaurid colonization of high-latitude environments, such as Antarctica and the Arctic Circle.

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.

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.

Geographically explicit, taxonomically resolved fossil occurrences are necessary for reconstructing macroevolutionary patterns and for testing a wide range of hypotheses in the Earth and life sciences. Heterogeneity in the spatial and temporal distribution of fossil occurrences in the Paleobiology Database (PBDB) is attributable to several different factors, including turnover among biological communities, socioeconomic disparities in the intensity of paleontological research, and geological controls on the distribution and fossil yield of sedimentary deposits. Here we use the intersection of global geological map data from Macrostrat and fossil collections in the PBDB to assess the extent to which the potentially fossil-bearing, surface-expressed sedimentary record has yielded fossil occurrences. We find a significant and moderately strong positive correlation between geological map area and the number of fossil occurrences. This correlation is consistent regardless of map unit age and binning protocol, except at period level; the Neogene and Quaternary have non-marine map units covering large areas and yielding fewer occurrences than expected. The sedimentary record of North America and Europe yields significantly more fossil occurrences per sedimentary area than similarly aged deposits in most of the rest of the world. However, geographic differences in area and age of sedimentary deposits lead to regionally different expectations for fossil occurrences. Using the sampling of surface-expressed sedimentary units in North America and Europe as a predictor for what might be recoverable from the surface-expressed sedimentary deposits of other regions, we find that the rest of the globe is approximately 45% as well sampled in the PBDB. Using age and area of bedrock and sampling in North America and Europe as a basis for prediction, we estimate that more than 639,000 occurrences from outside these regions would need to be added to the PBDB to achieve global geological parity in sampling. In general, new terrestrial fossil occurrences are expected to have the greatest impact on our understanding of macroevolutionary patterns.

The spatial distribution of in situ sessile organisms, including those from the fossil record, provides information about life histories, such as possible dispersal and/or settlement mechanisms, and how taxa interact with one another and their local environments. At Nilpena Ediacara National Park (NENP), South Australia, the exquisite preservation and excavation of 33 fossiliferous bedding planes from the Ediacara Member of the Rawnsley Quartzite reveals in situ communities of the Ediacara Biota. Here, the spatial distributions of three relatively common taxa, Tribrachidium, Rugoconites, and Obamus, occurring on excavated surfaces were analyzed using spatial point pattern analysis. Tribrachidium have a variable spatial distribution, implying that settlement or post-settlement conditions/preferences had an effect on populations. Rugoconites display aggregation, possibly related to their reproductive methods in combination with settlement location availability at the time of dispersal and/or settlement. Additionally, post-settlement environmental controls could have affected Rugoconites on other surfaces, resulting in lower populations and densities. Both Tribrachidium and Rugoconites also commonly occur as individuals or in low numbers on a number of beds, thus constraining possible reproductive strategies and environmental/substrate preferences. The distribution of Obamus is consistent with selective settlement, aggregating near conspecifics and on substrates of mature microbial mat. This dispersal process is the first example of substrate-selective dispersal among the Ediacara Biota, thus making Obamus similar to numerous modern sessile invertebrates with similar dispersal and settlement strategies.

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.

Grass-dominated ecosystems cover ~40% of Earth's terrestrial surface, with tropical grasses accounting for ~20% of global net primary productivity. C3 (cool/temperate) and C4 (tropical and subtropical) grass distribution is driven primarily by temperature. In this work, we used phytolith assemblages collected from vegetation plots along an elevation and temperature gradient in the northern Andes (Colombia and Ecuador) to develop a paleothermometer for the region. To accomplish this, we created a transfer function based on the inverse relationship between mean annual temperature (MAT) and the phytolith-based climatic index (Ic), which is the proportion of C3 over C4 grass phytoliths (GSSCP). To evaluate how accurately the index reflects C4–C3 grass abundance in vegetation plots, we compared it with semiquantitative floristic estimates of C4–C3 grass abundance. To further evaluate the 1 − Ic index as a proxy for C4–C3 grass abundance, we compared it with corresponding δ13C values (an independent proxy for C4–C3 vegetation). Results indicate that (1) GSSCP assemblages correctly estimate C4–C3 grass abundance in vegetation plots; (2) the Ic index outperforms the δ13C record in estimating C4–C3 grass abundance, even in open-vegetation types; and (3) our Ic index–based model accurately predicts MAT. This new calibrated proxy will help improve paleotemperature reconstructions in the northern Andes since at least the emergence and spread of C4 grasses in the region during the late Miocene.

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.

Body size is an important trait with implications for energy use and ecology as well as generation time and evolutionary rates. Turritelline gastropods are widely distributed through geologic time and space, making them an excellent group for evaluating macroevolutionary patterns. To evaluate the pattern of body-size change in turritelline gastropods, we compiled a dataset of shell lengths of 316 species of turritelline gastropods spanning the Jurassic to Recent. Type specimens were almost always significantly larger than specimen distributions from the same species. We found that turritelline gastropod size was inversely correlated with latitude, a trend likely driven by the Neogene–Recent diversification of small-bodied Southern Hemisphere taxa. A time series model was applied to distinguish among three possible macroevolutionary patterns: unbiased random walk (no directional trend), biased random walk (directional trend), and stasis (no net change). We determined that turritelline gastropods have experienced stasis in body size throughout their evolutionary history, adding to the growing literature documenting directionless body-size trends in marine invertebrate clades. Stasis of geographically widespread clades may be the result of ecological variability across the environmental range occupied by the group or differential diversification into opposing environments. Turritelline life-history patterns, especially their reproductive strategy that combines a short life span and decline in growth rate around 1 year of age to reallocate energy to reproduction, might circumvent selection for longevity and larger size, while further decrease in minimum size is likely limited by feeding efficiency and anti-predatory defense. The expectation that species or clades should continue to evolve to occupy larger size classes conflicts with the evolutionary advantages of small size, which in turritelline gastropods include high generational turnover and larger population sizes that yield opportunities for genetic variance.

The ecogeographic rule known as Bergmann's rule suggests that there is a positive relationship between body size and latitude when comparing closely related taxa. The underlying mechanism or mechanisms to explain this pattern vary as widely as the taxa that seem to follow it, which has led to skepticism over whether Bergmann's rule should be considered a rule at all. Despite this, Bergmann's rule is widespread among modern birds, mammals, beetles, and some amphibians, but far fewer extinct taxa have been subjected to tests of Bergmann's rule. To examine whether Bergmann's rule is detected in extinct taxa, we compared body-size proxies in Lystrosaurus recovered from Early Triassic–aged strata in Antarctica, South Africa, India, and China. Our results reveal that average body size is largest at mid-northern paleolatitudes (~45°N) instead of the highest southern paleolatitudes (~70°S). Additionally, maximum body size is consistent across the Northern and Southern Hemispheres, indicating that Bergmann's rule did not apply for Lystrosaurus during the Early Triassic. To test potential sample size biases in our results, we used rarefaction and subsampling to show that only the Karoo Basin is well sampled and that large individuals are exceedingly rare, except in the Turpan-Junggar Basin of Xinjiang, China. Taken together, our results suggest that Lystrosaurus had the potential to reach large body sizes in each of the latitudinally widespread geologic basins studied here, but that local conditions may have allowed individuals at mid-northern paleolatitudes a greater chance of reaching a large size compared with southern congeners that suffered increased mortality when young or at a small size.

Drilling predation is a common reason for mortality of benthic mollusks but did not become common until the late Mesozoic. The scarcity of drill holes in the early Mesozoic fossil record limits our understanding of the evolution of drilling behavior and its role on shaping early Mesozoic marine communities. Here, we use drilling traces on several bivalve taxa from the Lower Jurassic (Pliensbachian) marine soft-bottom deposits in northern Germany to explore behavioral patterns of the predator (e.g., site selectivity, change in site-selective behavior with age). Although none of the known drilling gastropod groups existed in the Pliensbachian, including the studied localities, the drill-hole morphology suggests that the predator was probably a gastropod. The ecology and identity of the target prey changes from a diverse array of epifaunal to infaunal taxa in older deposits to focus on a single, large, deep infaunal taxon, Gresslya intermedia, in younger deposits, suggesting a potential trend in prey selectivity over time. Spatial point pattern analysis of traces (SPPAT) reveals an aggregated pattern of drill holes on Gresslya, suggesting strong selectivity in drill-hole location. Drilling on a single large infaunal taxon and site selectivity are common patterns also inferred previously from the drilled deep infaunal Eothyasira from the Pliensbachian of southern Germany. In addition to the scarcity of predators, the highly specialized behavior of the early drilling predators, including strong prey selectivity in terms of prey identity and life habit, can partly explain the rarity of the early Mesozoic drill holes.

The impact of global climate events on local ecosystems can vary spatially. Understanding this potential heterogeneity can illuminate which environments will be most impacted and the proximal drivers of ecosystem responses. Cenomanian–Turonian marine deposits of the Western Interior Seaway (WIS) record paleoceanographic changes associated with the Greenhorn transgression and the onset of Oceanic Anoxic Event 2 (OAE2). They provide an ideal setting to study basin-wide paleoecological responses during a global perturbation. Here, we integrate benthic foraminiferal assemblages from before, during, and after OAE2 via multivariate ordination analysis to examine spatial patterns in faunal responses across the western United States on a common scale and to interrogate a previously defined faunal marker commonly used for basin-wide correlation, the Benthonic Zone (BZ). We identify oxygenation and organic matter quality as primary and secondary controls of faunal variation across the 10 stratigraphic records and use this variation to infer paleoenvironmental changes. Stratigraphic trends reveal, in contrast to previous studies, deoxygenation at the onset of OAE2. They also reveal temporal patterns in oxygenation and productivity consistent with the gradual northward migration of a southern water mass into the WIS. This spatial heterogeneity hinders the use of the BZ as a temporal marker, because assemblages change in response to diachronous environmental change, and thus timing of the BZ with respect to OAE2 is not consistent across the basin. Our study demonstrates that regional processes can overshadow ecosystem responses to global events and underscores the importance of considering how changes in the position of water masses impact the expression of global biogeochemical perturbations.

Studies of insect herbivory on fossilized leaves tend to focus on a few, relatively simple metrics that are agnostic to the distribution of insect damage types among host plants. More complex metrics that link particular damage types to particular host plants have the potential to address additional ecological questions, but such metrics can be biased by sampling incompleteness due to the difficulty of distinguishing the true absence of a particular interaction from the failure to detect it—a challenge that has been raised in the ecological literature. We evaluate a range of methods for characterizing the relationships between damage types and host plants by performing resampling and subsampling exercises on a variety of datasets. We found that the components of beta diversity provide a more valid, reliable, and interpretable method for comparing component communities than do bipartite network metrics and that the rarefaction of interactions represent a valid, reliable, and interpretable method for comparing compound communities. Both beta diversity and rarefaction of interactions avoid the potential pitfalls of multiple comparisons. Finally, we found that the host specificity of individual damage types is challenging to assess. Whereas bipartite network metrics are sufficiently biased by sampling incompleteness to be inappropriate for fossil herbivory data, alternatives exist that are perfectly suitable for fossil datasets with sufficient sample coverage.

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.

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.

Conodont elements have high rates of morphological evolution, but the drivers of this disparity are debated. Positive allometric relationships between dimensions of food-processing surfaces and entire P1 elements have been used to argue that these elements performed mechanical digestion. If involved in food processing, the surface of the element should grow at a rate proportional to the increase in energy requirements of the animal. This inference of function relies on the assumption that the energy requirements of the animal grew faster (≅ mass0.75) than the tooth area (≅ mass0.67). We reevaluate this assumption based on metabolic rates across animals and calculate the allometry in platform-bearing P1 elements of Late Triassic co-occurring taxa, Metapolygnathus communisti and Epigondolella rigoi, using 3D models of ontogenetic series. Positive allometry is found in platform and element dimensions in both species, supporting a grasping-tooth hypothesis, based on the assumption that metabolic rate in conodonts scaled with body mass similarly to that in fish and ectotherms. We also calculate the curvature of the P1 platform surface using the Dirichlet normal energy (DNE) as a proxy for diet. DNE values increase with body mass, supporting the assumption that conodont metabolic rates increased faster than mass0.67. We finally find that adults in both taxa differ in their food bases, which supports trophic diversification as an important driver of the remarkable disparity of conodont elements.

The spherical encompassing final chamber of the planktonic foraminifera Orbulina universa is a prime example of a complex character whose evolution has been documented by a sequence of intermediate forms. However, the mechanism that induced evolution of the spherical chamber remain unclear. Here we show that shortly after the emergence of Orbulina, documented throughout the oceans, a convergent evolutionary transition occurred in the semi-isolated Paratethys, leading to the emergence of the endemic Velapertina, which occupied a similar niche to Orbulina in the surface waters. Using X-ray computed tomography scanning, we show that the evolution of the encompassing final chamber involved the same sequence of steps in both lineages, combining a progressively spherical shell shape with changes in the position, number, and sizes of apertures. The similarity in the sequence of character acquisitions suggests structural determinism in the way foraminiferal shells are constructed and the presence of natural selection favoring a spherical morphology. Collectively, the emergence of spherical chambers in the two lineages at a similar time suggests that the evolution of this spectacular complex character occurred in response to a singular environmental driver.