The taxonomic and ecologic composition of Earth's biota has shifted dramatically through geologic time, with some clades going extinct while others diversified. Here, we derive a metric that quantifies the change in biotic composition due to extinction or origination and show that it equals the product of extinction/origination magnitude and selectivity (variation in magnitude among groups). We also define metrics that describe the extent to which a recovery (1) reinforced or reversed the effects of extinction on biotic composition and (2) changed composition in ways uncorrelated with the extinction. To demonstrate the approach, we analyzed an updated compilation of stratigraphic ranges of marine animal genera. We show that mass extinctions were not more selective than background intervals at the phylum level; rather, they tended to drive greater taxonomic change due to their higher magnitudes. Mass extinctions did not represent a separate class of events with respect to either strength of selectivity or effect. Similar observations apply to origination during recoveries from mass extinctions, and on average, extinction and origination were similarly selective and drove similar amounts of biotic change. Elevated origination during recoveries drove bursts of compositional change that varied considerably in effect. In some cases, origination partially reversed the effects of extinction, returning the biota toward the pre-extinction composition; in others, it reinforced the effects of the extinction, magnifying biotic change. Recoveries were as important as extinction events in shaping the marine biota, and their selectivity deserves systematic study alongside that of extinction.

Lanternfishes currently represent one of the dominant groups of mesopelagic fishes in terms of abundance, biomass, and diversity. Their otolith record dominates pelagic sediments below 200 m in dredges, especially during the entire Neogene. Here we provide an analysis of their diversity and rise to dominance primarily based on their otolith record. The earliest unambiguous fossil myctophids are known based on otoliths from the late Paleocene and early Eocene. During their early evolutionary history, myctophids were likely not adapted to a high oceanic lifestyle but occurred over shelf and upper-slope regions, where they were locally abundant during the middle Eocene. A distinct upscaling in otolith size is observed in the early Oligocene, which also marks their earliest occurrence in bathyal sediments. We interpret this transition to be related to the change from a halothermal deep-ocean circulation to a thermohaline regime and the associated cooling of the deep ocean and rearrangement of nutrient and silica supply. The early Oligocene myctophid size acme shows a remarkable congruence with diatom abundance, the main food resource for the zooplankton and thus for myctophids and whales. The warmer late Oligocene to early middle Miocene period was characterized by an increase in disparity of myctophids but with a reduction in their otolith sizes. A second and persisting secular pulse in myctophid diversity (particularly within the genus Diaphus) and increase in size begins with the “biogenic bloom” in the late Miocene, paralleled with diatom abundance and mysticete gigantism.

The highly elongated necks, and often tails, of sauropod dinosaurs were composed of concavo-convex vertebrae that provided stability without compromising mobility. Polarities of these concavo-convex joints in the neck and tail are anatomically opposite one another but mechanically equivalent. Opisthocoelous cervical vertebrae and procoelous caudal vertebrae have the convex articular face directed away from the body and the concave articular face directed toward the body. This “sauropod-type” polarity is hypothesized to be (1) more resistant to fracturing of the cotylar rim and (2) better stabilized against joint failure by rotation than the opposite polarity. We used physical models to test these two functional hypotheses. Photoelastic analysis of model centra loaded as cantilevers reveals that neither polarity better resists fracture of the cotylar rim; strain magnitude and localization are similar in both polarities. We assessed the rotational stability of concavo-convex joints using pairs of concavo-convex centra loaded near the joint. Sauropod-type joints withstood significantly greater weight before failure occurred, a pattern we interpret to be dependent on the position of the center of rotation, which is always within the convex part of the concavo-convex joint. In sauropod-type joints, the free centrum rotates about a center of rotation that lies within the more stable proximal centrum. In contrast, the opposite polarity results in a free centrum that rotates about an internal point; when the condyle rotates down and out of joint, the distal end rotates back toward the body, unopposed by ligamentous support. Sauropod-type joints remained stable with greater mobility, more mechanically advantageous tensile element insertions, and greater distal loads than the opposite polarity. The advantages conferred by this joint polarity would have facilitated the evolution of hyperelongated necks and tails by sauropods. Polarity of concavo-convex joints of the appendicular skeleton (e.g., hip, shoulder) is also consistent with the demands of rotational stability.

The application of molecular genetics, in particular comparative genomics, to the field of evolutionary biology is paving the way to an enhanced “New Synthesis.” Apart from their power to establish and refine phylogenies, understanding such genomic processes as the dynamics of change in genomes, even in hypothetical RNA-based genomes and the in vitro evolution of RNA molecules, helps to clarify evolutionary principles that are otherwise hidden among the nested hierarchies of evolutionary units. To this end, I outline the course of hereditary material and examine several issues including disparity, causation, or bookkeeping of genes, adaptation, and exaptation, as well as evolutionary contingency at the genomic level–issues at the heart of some of Stephen Jay Gould's intellectual battlegrounds. Interestingly, where relevant, the genomic perspective is consistent with Gould's agenda. Extensive documentation makes it particularly clear that exaptation plays a role in evolutionary processes that is at least as significant as–and perhaps more significant than–that played by adaptation.

Macrofossils from woodrat (Neotoma) middens serve as an important proxy for reconstructing past vegetation in arid and semiarid regions of North America. The presence/absence of plant macrofossils in middens can provide valuable information on temporal and spatial patterns of plant migration and range boundaries. The primary aim of this study was to determine how local plant abundance, distance of plant populations from midden sites, and species population density on the landscape affect the probability of occurrence of macrofossils in middens. The study was designed with the primary intent of determining the reliability of middens in detecting scattered populations of Pinus ponderosa. We analyzed macrofossil assemblages from 42 modern woodrat middens from West Carrizo Canyon in southeastern Colorado, near the current eastern range margin of Pinus ponderosa. We compared midden contents with composition of the surrounding vegetation, measuring distance from the midden to the nearest individual of selected plant species, and the percent cover of each species within 30 m of the midden. We used this information to model the probability of species presence in a midden across a range of population densities on the landscape. Macrofossils of Juniperus spp., Quercus gambelii, and Opuntia spp. were consistently found in middens regardless of their local abundance in vegetation, although populations occurred within 30 m of all middens. Pinus edulis and P. ponderosa occurred in nearly all middens within 20–30 m of individual trees. P. ponderosa was rare in middens >20–30 m away from individual trees. Results of a simple simulation model suggest that middens become absolutely reliable indicators of P. ponderosa presence on the landscape only when average tree density exceeds 50 stems ha−1. Woodrats reliably collected macrofossils of Pinus edulis, P. ponderosa, Juniperus spp., Quercus gambelii, and Opuntia spp. when populations of these taxa occur within 20–30 m of a midden site. Woodrats did not collect P. ponderosa when the nearest individuals were more than 30 m away. Low-density populations of these and other species may be difficult to detect in fossil woodrat-midden series owing to reduced probability that individuals grow within foraging distance of the middens. Data from this and similar studies can be used to construct and parameterize a forward model of macrofossil representation in woodrat middens.

The rise of durophagous predators during the Paleozoic represents an ecological constraint imposed on sessile marine fauna. In crinoids, it has been suggested that increasing predation pressure drove the spread of adaptations against predation. Damage to a crinoid's arms from nonlethal predation varies as a function of arm branching pattern. Here, using a metric for resilience to predation (“expected arm loss,” EAL), we test the hypothesis that the increase in predation led to more predation-resistant arm branching patterns (lower EAL) among Paleozoic crinoids. EAL was computed for 230 genera of Paleozoic crinoids and analyzed with respect to taxonomy and time. The results show significant variability among taxa. Camerates, especially monobathrids, display a pattern of increasingly convergent and predation-resistant arm morphologies from the Ordovician through the Devonian, with no significant change during the Mississippian. In contrast, the mean EAL among cladids follows no overall trend through the Paleozoic. Regenerating arms are known to be significantly more common in camerates than in other Paleozoic taxa; if regeneration is taken as a proxy for nonlethal interactions with durophagous predators, this indicates that nonlethal predation occurred more often among camerates throughout the Early and Middle Paleozoic. In addition, frequency of injury among camerates is inversely correlated with EAL and positively correlated with infestation by parasitic snails. From this we conclude that decreasing EAL signals a selective pressure in favor of resistance to grazing predation in camerates but not in other subclasses before the Mississippian, with an apparent relaxation in this constraint after the late Devonian extinctions.

Body mass is an important organism-level variable in mammalian biology, correlated with physiology, life history, and ecology. To analyze the dynamics of body size evolution, increases and decreases in body mass were tallied for ancestor-descendant (AD) species pairs for 519 terrestrial caniform taxa. To account for uncertainty phylogeny, a bootstrapping routine shuffled hypothesized AD pairs, and average proportions of increases were binned as a function of ancestral body mass. A set of models relating the rate of body size increase were evaluated with the Akaike Information Criterion (AIC). AIC selected three models of the candidate set as equivalent in support by the observed body mass data. These three models propose body size increase for small AD pairs and body size decrease for large AD pairs, although they differ in their treatment of taxa at intermediate sizes.

These results demonstrate the presence of constraints bounding the caniform distribution at large and small body sizes, stabilizing the distribution through time, which stands in contrast to a broader mammalian pattern. At a finer phylogenetic scale, subclades within intermediate size classes display proportions that are significantly different from unbiased, with several clades previously cited as examples of “Cope's Rule” showing biased increases in size, and basal mustelids (badgers, and allied genera), Mephitidae (skunks), and Vulpini (“foxes”) exhibiting biased decreases. The caniform pattern is therefore the result of superimposed, clade-specific trajectories, demonstrating that the inferred dynamics of body size evolution and even the direction of trends in body size evolution within the Caniformia, and for mammals in general, depend on the hierarchical scale of the analysis.

Studying the deep-time origins of macroecological phenomena can help us to understand their long-term drivers. Given the considerable spatiotemporal bias of the terrestrial fossil record, it behooves us to understand how much biological information is lost. The aim of this study is to establish whether latitudinal diversity gradients are detectable in a biased terrestrial fossil record. I develop a simulated fossilization approach, weighting the probability of terrestrial mammal species appearing in the fossil record based on body size and geographic-range size; larger species with larger range sizes are more likely to enter the fossil record. I create simulated fossil localities from the modern North American mammal record. I vary the percentage of species successfully fossilized and estimate the magnitude of the latitudinal diversity gradient (slope of the richness gradient and degree of species turnover). I find that estimates of the latitudinal diversity gradient are sensitive to the loss of species with small body size and geographic-range sizes. In some cases, simulated fossil-record bias completely obliterates evidence of declining richness with latitude, a phenomenon that is not ameliorated by the application of nonparametric richness estimation. However, if the rate of preservation is medium (50% of species) to high (75% of species), the magnitude of the latitudinal diversity gradient can be reliably estimated. Similarly, changes in the diversity gradient estimates are largely explained by differences in the diversity–climate relationship among iterations, suggesting that these relationships may be measurable in the fossil record.

Based on high-resolution palynological analysis of 680 samples from a core, short-term changes in plant diversity and floristic composition within the Paleogene greenhouse were detected in the lacustrine succession of a lower to middle Eocene maar lake at Messel (Federal State of Hesse, Germany). The microfloristic data show that taxonomic diversity increased rapidly within some decades during recolonization of a volcanically devastated area around the lake. With the establishment of a climax vegetation at the end of recolonization, the maximum in palynological diversity was reached within the crater area. During the following 640 Kyr the composition of the palynospectrum changed only gradually. However, different richness and evenness estimations show that alpha and gamma diversity decreased up to 35%, which can be related to the establishment of an equilibrium stage within the climax vegetation that led to the dominance of an assemblage of self-replacing species. Nevertheless, time-series analysis of alpha-diversity changes within the climax vegetation reveals that orbitally controlled climate change of Milankovitch and sub-Milankovitch order influenced the diversity of the vegetation, resulting in a rise of beta diversity. Based on the composition of the vegetation and comparison to modern analogues, our analysis proves that Eocene paratropical plant diversity increased during periods of slightly higher temperature and precipitation. Therefore, both composition and diversity of the vegetation was highly susceptible to minor-scale, short-term changes in climate, even during equable greenhouse conditions.

Muscle moment arms are important determinants of muscle function; however, it is challenging to determine moment arms by inspecting bone specimens alone, as muscles have curvilinear paths that change as joints rotate. The goals of this study were to (1) develop a three-dimensional graphics-based model of the musculoskeletal system of the Cretaceous theropod dinosaur Tyrannosaurus rex that predicts muscle-tendon unit paths, lengths, and moment arms for a range of limb positions; (2) use the model to determine how the T. rex hindlimb muscle moment arms varied between crouched and upright poses; (3) compare the predicted moment arms with previous assessments of muscle function in dinosaurs; (4) evaluate how the magnitudes of these moment arms compare with those in other animals; and (5) integrate these findings with previous biomechanical studies to produce a revised appraisal of stance, gait, and speed in T. rex. The musculoskeletal model includes ten degrees of joint freedom (flexion/extension, ab/adduction, or medial/lateral rotation) and 33 main muscle groups crossing the hip, knee, ankle, and toe joints of each hindlimb. The model was developed by acquiring and processing bone geometric data, defining joint rotation axes, justifying muscle attachment sites, and specifying muscle-tendon geometry and paths. Flexor and extensor muscle moment arms about all of the main limb joints were estimated, and limb orientation was statically varied to characterize how the muscle moment arms changed. We used sensitivity analysis of uncertain parameters, such as muscle origin and insertion centroids, to deterimine how much our conclusions depend on the muscle reconstruction we adopted. This shows that a specific amount of error in the reconstruction (e.g., position of muscle origins) can have a greater, lesser, similar, or no effect on the moment arms, depending on complex interactions between components of the musculoskeletal geometry. We found that more upright poses would have improved mechanical advantage of the muscles considerably. Our analysis shows that previously assumed moment arm values were generally conservatively high. Our results for muscle moment arms are generally lower than the values predicted by scaling data from extant taxa, suggesting that T. rex did not have the allometrically large muscle moment arms that might be expected in a proficient runner. The information provided by the model is important for determining how T. rex stood and walked, and how the muscles of a 4000–7000 kg biped might have worked in comparison with extant bipeds such as birds and humans. Our model thus strengthens the conclusion that T. rex was not an exceptionally fast runner, and supports the inference that more upright (although not completely columnar) poses are more plausible for T. rex. These results confirm general principles about the relationship between size, limb orientation, and locomotor mechanics: exceptionally big animals have a more limited range of locomotor abilities and tend to adopt more upright poses that improve extensor muscle effective mechanical advantage. This model builds on previous phylogenetically based muscle reconstructions and so moves closer to a fully dynamic, three-dimensional model of stance, gait, and speed in T. rex.

The most popular theoretical propositions are often those which spell the final doom to a notion that has already fallen into disrepute. This is as true today as it was when August Weismann (1892) published his seminal volume, Der Keimplasma, which far more than Cuvier's vituperative attacks, put the ideas of Lamarck to final rest.

Fusion occurred between pairs of conspecific colonies of Permian fenestrate bryozoans that were of approximately equal radius at time of contact. In contrast, no fusion occurred in pairs of conspecific colonies that were of unequal radius at time of contact. It is suggested that histologically compatible colonies developed from larvae that settled essentially simultaneously and that the larvae were siblings, lecithotrophic with low mobility, and of the same genotype (i.e., polyembryonic). Thus polyembryony appears to have existed as a reproductive “strategy” for at least 275 Myr, and its scarcity in modern taxa cannot be due to recentness of evolution.

Comprising Holostei and Teleostei, the ~32,000 species of neopterygian fishes are anatomically disparate and represent the dominant group of aquatic vertebrates today. However, the pattern by which teleosts rose to represent almost all of this diversity, while their holostean sister-group dwindled to eight extant species and two broad morphologies, is poorly constrained. A geometric morphometric approach was taken to generate a morphospace from more than 400 fossil taxa, representing almost all articulated neopterygian taxa known from the first 150 million years—roughly 60%—of their history (Triassic‒Early Cretaceous). Patterns of morphospace occupancy and disparity are examined to: (1) assess evidence for a phenotypically “dominant” holostean phase; (2) evaluate whether expansions in teleost phenotypic variety are predominantly abrupt or gradual, including assessment of whether early apomorphy-defined teleosts are as morphologically conservative as typically assumed; and (3) compare diversification in crown and stem teleosts. The systematic affinities of dapediiforms and pycnodontiforms, two extinct neopterygian clades of uncertain phylogenetic placement, significantly impact patterns of morphological diversification. For instance, alternative placements dictate whether or not holosteans possessed statistically higher disparity than teleosts in the Late Triassic and Jurassic. Despite this ambiguity, all scenarios agree that holosteans do not exhibit a decline in disparity during the Early Triassic‒Early Cretaceous interval, but instead maintain their Toarcian‒Callovian variety until the end of the Early Cretaceous without substantial further expansions. After a conservative Induan‒Carnian phase, teleosts colonize (and persistently occupy) novel regions of morphospace in a predominantly gradual manner until the Hauterivian, after which expansions are rare. Furthermore, apomorphy-defined teleosts possess greater phenotypic variety than typically assumed. Comparison of crown and stem teleost partial disparity indicates that, despite a statistically significant increase in crown teleost disparity between the Late Jurassic and earliest Cretaceous, stem teleosts remained important long-term contributors to overall teleost disparity during this time.

The end-Permian mass extinction occurred alongside a large swath of environmental changes that are often invoked as extinction mechanisms, even when a direct link is lacking. One way to elucidate the cause(s) of a mass extinction is to investigate extinction selectivity, as it can reveal critical information on organismic traits as key determinants of extinction and survival. Here we show that machine learning algorithms, specifically gradient boosted decision trees, can be used to identify determinants of extinction as well as to predict extinction risk. To understand which factors led to the end-Permian mass extinction during an extreme global warming event, we quantified the ecological selectivity of marine extinctions in the well-studied South China region. We find that extinction selectivity varies between different groups of organisms and that a synergy of multiple environmental stressors best explains the overall end-Permian extinction selectivity pattern. Extinction risk was greater for genera that had a low species richness, narrow bathymetric ranges limited to deep-water habitats, a stationary mode of life, a siliceous skeleton, or, less critically, calcitic skeletons. These selective losses directly link the extinctions to the environmental effects of rapid injections of carbon dioxide into the ocean–atmosphere system, specifically the combined effects of expanded oxygen minimum zones, rapid warming, and potentially ocean acidification.

Carbon and nitrogen stable isotopic compositions of fossil materials from Lake Messel (47 Ma) in Germany are used to investigate Eocene ecosystem dynamics. Autolithified soft tissues of terrestrial and aquatic vertebrate organisms, as well as plant compression fossils, contain organic material (20–50 wt% C, 1–6 wt% N), which appears to retain precursor compositions. Stable isotopic compositions (δ13C and δ15N) of Messel fossils are similar to those reported for components in modern lacustrine ecosystems. These data show trophically sensible enrichments relative to food sources, reflect multiple feeding strategies for each organism (e.g., omnivory, planctivory, piscivory), and differentiate between benthic and pelagic organic carbon sources. These chemical data broadly confirm existing Messel food web models based on coprolite and gut content analyses. δ13C values for the lacustrine shale range from −30.3 to −26.3‰, pointing to mixed terrestrial and aquatic origins for primary producers in the food web. δ13C values for primary consumers such as insects overlap with those for primary producers but are comparatively enriched in 15N. Secondary and higher consumers (fish, crocodiles, and frogs) are associated with even more positive δ15N values and show a more constrained range of δ13C values. Omnivory appears widespread in both low and high trophic level consumers. Hence, the stable isotopic compositions of Messel fossils are complex and overlap, and must be combined with paleontological investigations in order to be conclusive. This study represents the first comprehensive isotopic reconstruction, featuring tens of components, of an ecosystem of Eocene age. A thorough understanding of trophic structure in Eocene Lake Messel contributes to the global databank of ecological history.

A computer model of background and mass extinctions in a taxonomic hierarchy has been used to study the effects of different extinction patterns in a search for clues as to the causes of actual extinction events. Model taxa at four levels were built up from speciation events in adaptive space according to rules of origination which seem plausible biologically. The frequency distribution of species among the three higher taxonomic levels in the model is similar to that in living marine taxa which have good fossil records. Three mass extinction patterns were imposed on the model after species diversity had attained equilibrium (i.e., when speciation = background extinction): random; bloc (contiguous niches were cleared); and clade (all members of selected higher taxa were removed). Effects on the taxonomic profile varied with pattern. Four of the five historical mass extinctions resemble the effects of the random pattern. End-Permian families were harder hit than those in the random model, but this may be a result of an extremely high species extinction level. It is concluded that the effect of extinctions on the taxonomic hierarchy provides a tool to help in understanding extinction causes.

During land plant evolution, a change in stelar architecture, i.e., in the geometric arrangement of the water-conducting tissue inside the plant axis, can be observed. In the most primitive stele type, the protostele, the vascular tissue is organized as a simple central strand. Further evolutionary changes led to more peripherally arranged vascular tissues. In the siphonostele, for example, the vascular tissue forms a hollow cylinder filled with pith. A functional explanation of this early stelar evolution is provided in the present paper. Using a numerical simulation approach, we analyze the water transport properties of various protostelic and siphonostelic model axes. The results indicate that several geometric parameters are relevant for understanding the water transport properties of various stele types and for explaining the early stelar evolution: the parenchymatic path lengths (i.e., the distance between the xylem surface and the transpiring plant surface), the ratio of xylem surface over transpiring surface, and the ratio of cross-sectional area of xylem to cross-sectional area of the parenchyma outside of the xylem. As a whole, the evolution of early stele types may be viewed as a size-related multi-criteria optimization process in which the xylem volume as well as the fluid pressure gradients in the parenchyma and in the xylem are minimized. For slender plant axes, a protostele appears to be the optimal stelar architecture. In wider plant axes, however, other stelar architectures (such as a siphonostele) prove to be more efficient than a protostele.

High levels of biodiversity and endemism in ancient lakes have motivated research on evolutionary processes in these systems. Drill-core records from Lake Titicaca (Bolivia, Peru), an ancient lake in the high-elevation Altiplano, record the history of climate, landscape dynamics, and diatom evolution. That record was used to examine the patterns and drivers of morphological evolution of an endemic species complex of diatoms in the lake, the Cyclostephanos andinus complex. In an attempt to delineate species within the complex based on morphology, no discernible evidence was found for species separation based on an ordination analysis of multiple characters, but multiple populations were detected based on the distribution of valve size in individual samples. Likelihood modeling of phyletic evolution showed that size evolved through punctuated change. Correlation of size trends with environmental variables indicates that C. andinus size responded to regional environmental change driven by global processes that influenced Lake Titicaca by affecting lake level and thermal stratification.

The founders of North American vertebrate paleontology, F. V. Hayden, Joseph Leidy, E. D. Cope, O. C. Marsh, and their colleagues, collected and described the first suites of fossil mammals obtained from the rich Tertiary successions of the western United States. Among them were remains of fossil horses, and subsequent study of these resulted in an interpretation that supported the concept of Darwinian gradualism as the major mode of evolution. The fossil record of horses also contributed importantly to the demise of orthogenesis as an evolutionary pattern, and to the evaluation of evolutionary rates and long-term evolutionary trends in a major phyletic group of organisms.

Within ancient ecosystems, it is generally difficult to determine the specific diets of species from higher trophic levels, which in turn hinders our understanding of trophic relationships and energy flow through these systems. To better understand the ecology of taxa at higher trophic levels, we used analysis of tooth enamel stable carbon isotope values to infer the dietary preferences of Canis edwardii and Smilodon gracilis from the Leisey Shell Pit 1A (LSP 1A) and Inglis 1A, two Pleistocene localities in Florida. The goals of the analyses were to (1) determine whether these carnivorans specialized in particular prey types or maintained a generalist diet; (2) ascertain whether carbon isotope values support what was previously suggested about the ecology of these species; and (3) establish what ecological details of ancient food webs can be discovered by carbon isotope analyses at higher trophic levels. Results show that the sampled carnivoran carbon isotope values are distributed among suspected prey isotope values, suggesting that varied prey were taken at the study localities. Prey compositions were modeled for each carnivoran species by using Stable Isotope Analysis in R (SIAR). The modeled diets indicate that each studied carnivoran had a generalist diet; however, there are differences in how these taxa achieved dietary generalization. At the glacial Inglis 1A locality, sampled individuals of C. edwardii and S. gracilis show similar isotope values and modeled dietary prey proportions, although both carnivorans do show a preference for grazing prey species. The similar isotopic values, and calculated prey proportions, observed between these species may imply greater interspecific competition for food. At the interglacial LSP 1A locality, C. edwardii shows values similar to those observed at Inglis 1A. In contrast, the data for S. gracilis shows a preference for consuming browsing prey species. Further, its restricted range of carbon isotope values suggests that S. gracilis may have concentrated its feeding within a particular habitat. Examination of stable carbon isotope values among species at higher trophic levels reveals that some intricacies of ancient food webs can be discerned.