A Devonian crinoid with a diamond microlattice.

Owing to their remarkable physical properties, cellular structures, such as triply periodic minimal surfaces (TPMS), have multidisciplinary and multifunctional applications. Although these structures are observed in nature, examples of TPMS with large length scales in living organisms are exceedingly rare. Recently, microstructure reminiscent of the diamond-type TPMS was documented in the skeleton of the modern knobby starfish Protoreaster nodosus. Here we report a similar microlattice in a 385 Myr old crinoid Haplocrinites, which pushes back the origins of this highly ordered microstructure in echinoderms into the Devonian. Despite the low Mg2+/Ca2+ ratio of the 'calcite' Devonian sea, the skeleton of these crinoids has high-Mg content, which indicates strong biological control over biomineralogy. We suggest that such an optimization of trabecular arrangement additionally enriched in magnesium, which enhances the mechanical properties, might have evolved in these crinoids in response to increased predation pressure during the Middle Palaeozoic Marine Revolution. This discovery illustrates the remarkable ability of echinoderms, through the process of evolutionary optimization, to form a lightweight, stiff and damage-tolerant skeleton, which serves as an inspiration for biomimetic materials.

Comparative actualistic study hints at origins of alleged Miocene coprolites of Poland.

Excrement-shaped ferruginous masses have been recovered from the Miocene of Turów mine in south-western Poland. These siderite masses have been the subject of much controversy, having been interpreted either as being coprolites, cololithes or pseudofossils created by mechanical deformation of plastic sediment. Here we present the results of mineralogical, geochemical, petrographic and microtomographical analyses. Our data indicate that these masses consist of siderite and iron oxide rather than phosphate, and rarely contain recognizable food residues, which may suggest abiotic origins of these structures. On the other hand, evidence in support of a fecal origin include: (i) the presence of two distinct morphotypes differing in size and shape, (ii) the presence of rare hair-like structures or coalified inclusions and (iii) the presence of rare fine striations on the surface. Importantly, comparative actualistic study of recent vertebrate feces shows overall resemblance of the first morphotype (sausage-shaped with rare coalified debris) to excrements of testudinoid turtles (Testudinoidea), whose shell fragment was found in the investigated locality. The second morphotype (rounded to oval-shaped with hair-like structures), in turn, is similar to the feces of some snakes (Serpentes), the remains of which were noted in the Miocene of the neighborhood areas. Other potential producers (such as lizards and crocodiles) and even abiotic origins cannot be fully excluded but are less likely.

Ausichicrinites zelenskyyi gen. et sp. nov., a first nearly complete feather star (Crinoidea) from the Upper Jurassic of Africa.

Fossil comatulids, referred to as feather stars, are mostly known from highly disarticulated specimens. A single isolated element (centrodorsal) has been the basis for taxonomic description of a vast majority of fossil comatulids. Here, we report a nearly complete, and thus extremely rare, comatulid from the Upper Jurassic (Tithonian) of the Blue Nile Basin in central western Ethiopia that provides a unique insight into the morphology of comatulid arms and cirri. It is assigned to Ausichicrinites zelenskyyi gen. et sp. nov. and is the first Jurassic comatulid from the African continent. The new taxon shows some similarities with representatives of the Mesozoic Solanocrinitidae but also has close resemblance with the modern family Zygometridae, exclusively known from the Holocene of western Pacific and eastern Indian Oceans. This morphologic similarity is considered to be due to convergence. The first example of pinnule regeneration in a fossil feather star is reported, which reinforces the hypothesis about the importance of predation in the evolution of these crinoids.

Unraveling the hidden paleobiodiversity of the Middle Devonian (Emsian) crinoids (Crinoidea, Echinodermata) from Poland.

Most previous publications on Devonian crinoids from the Holy Cross Mountains in Poland have concentrated on crinoid columns, and until now, little has been published about crinoid cups and calyxes. Herein, five crinoid taxa are described from an abundant occurrence of aboral cups and partial crowns from the Bukowa Góra Member (Emsian) in the Holy Cross Mountains of southern Poland. The following taxa are described: Bactrocrinites sp., Codiacrinus sevastopuloi sp. nov., Halocrinites geminatus (Bohatý, 2005), Halocrinites schlotheimii Steininger, 1831, and a single brachial plate from a flexible crinoid placed in Flexibilia incertae sedis. Simple discoid holdfasts are also present encrusted to cylindrical stromatoporoids. These taxa are the first crinoids described from the remains of partial crowns from Emsian strata of Poland.

Shared patterns in body size declines among crinoids during the Palaeozoic extinction events.

Crinoids were among the most abundant marine benthic animals throughout the Palaeozoic, but their body size evolution has received little attention. Here, we compiled a comprehensive database on crinoid calyx biovolumes throughout the Palaeozoic. A model comparison approach revealed contrasting and complex patterns in body size dynamics between the two major crinoid clades (Camerata and Pentacrinoidea). Interestingly, two major drops in mean body size at around two mass extinction events (during the late Ordovician and the late Devonian respectively) are observed, which is reminiscent of current patterns of shrinking body size of a wide range of organisms as a result of climate change. The context of some trends (marked declines during extinctions) suggests the cardinal role of abiotic factors (dramatic climate change associated with extinctions) on crinoid body size evolution; however, other patterns (two intervals with either relative stability or steady size increase in periods between mass extinctions) are more consistent with biotic drivers.

Substrate type and palaeodepth do not affect the Middle Jurassic taxonomic diversity of crinoids.

Crinoids are largely considered as good indicators for determining environmental conditions. They are robust proxies for inferring changes in salinity and sedimentation rate and for inferring substrate type. Some crinoid groups (e.g., certain comatulids, cyrtocrinids, millericrinids) have a depth preference, thus, making them useful for palaeodepth estimation. The hypotheses that crinoid distribution is substrate-dependent (rock type) or palaeodepth-dependent is tested here based on (a) archival Bathonian-Callovian (Middle Jurassic) crinoid occurrences from Poland and (b) newer finds from five boreholes from eastern Poland. Qualitative data suggests that isocrinids and cyclocrinids occur in both carbonate and siliciclastic rocks. The cyrtocrinids and roveacrinids occur within carbonate rocks, whereas the comatulids are exclusive to siliciclastics. In terms of palaeodepth, most crinoid groups dominate in shallow environments with the sole exception of cyrtocrinids, that are ubiquitous and occur in both shallow (near shore and shallow marine) and slightly deeper (deeper sublittoral to open shelf) settings. The occurrences of the cosmopolitan taxa, Chariocrinus andreae and Balanocrinus subteres (isocrinids), is independent of both substrate type and palaeodepth. Quantitative analyses (Analysis Of Variance; ANOVA) based on substrate type, i.e., substrate-dependency (claystones, sandstones and limestones), and palaeodepth i.e., palaeodepth-dependency (near shore, shallow-marine, mid-ramp and offshore), corroborate qualitative results. Statistical analysis suggest that the distribution of crinoids shows a strong substrate-dependency but not for palaeodepth, although very weak significance (low p value) is noted for near shore and shallow marine settings and crinoid distribution.

Bromalites from the Upper Triassic Polzberg section (Austria); insights into trophic interactions and food chains of the Polzberg palaeobiota.

A rich assemblage of various types of bromalites from the lower Carnian "Konservat-Lagerstätte" from the Reingraben Shales in Polzberg (Northern Calcareous Alps, Lower Austria) is described for the first time in detail. They comprise large regurgitalites consisting of numerous entire shells of ammonoid Austrotrachyceras or their fragments and rare teuthid arm hooks, and buccal cartilage of Phragmoteuthis. Small coprolites composed mainly of fish remains were also found. The size, shape and co-occurrence with vertebrate skeletal remains imply that regurgitalites were likely produced by large durophagous fish (most likely by cartilaginous fish Acrodus). Coprolites, in turn, were likely produced by medium-sized piscivorous actinopterygians. Our findings are consistent with other lines of evidence suggesting that durophagous predation has been intense during the Triassic and that the so-called Mesozoic marine revolution has already started in the early Mesozoic.

Experimental neoichnology of post-autotomy arm movements of sea lilies and possible evidence of thrashing behaviour in Triassic holocrinids.

Echinoderms exhibit remarkable powers of autotomy. For instance, crinoids can shed arm and stalk portions when attacked by predators. In some species, it has been reported that the autotomized arms display vigorous movements, which are thought to divert the attention of predators. This phenomenon, however, has not been well explored. Here we present results of experiments using the shallowest water species of living stalked crinoid (Metacrinus rotundus) collected at 140 m depth. A wide range of movements of detached arms, from sluggish writhing to violent flicks, was observed. Interestingly, autotomized arms produce distinct traces on the sediment surface. They are composed of straight or arched grooves usually arranged in radiating groups and shallow furrows. Similar traces were found associated with detached arms of the oldest (Early Triassic) stem-group isocrinid (Holocrinus). This finding may suggest that the origins of autotomy-related thrashing behaviour in crinoids could be traced back to at least the Early Triassic, underscoring the magnitude of anti-predatory traits that occurred during the Mesozoic Marine Revolution. A new ethological category, autotomichnia, is proposed for the traces produced by thrashing movements of shed appendages.

Uncovering the hidden diversity of Mississippian crinoids (Crinoidea, Echinodermata) from Poland.

Partial crinoid crowns and aboral cups are reported from the Mississippian of Poland for the first time. Most specimens are partially disarticulated or isolated plates, which prevent identification to genus and species, but regardless these remains indicate a rich diversity of Mississippian crinoids in Poland during the Mississippian, especially during the late Viséan. Lanecrinus? sp. is described from the late Tournaisian of the Debnik Anticline region. A high crinoid biodiversity occurred during late Viséan of the Holy Cross Mountains, including the camerate crinoids Gilbertsocrinus? sp., Platycrinitidae Indeterminate; one flexible crinoid; and numerous eucladid crinoids, including Cyathocrinites mammillaris (Phillips), three taxa represented by partial cups left in open nomenclature, and numerous additional taxa known only from isolated radial plates, brachial plates, and columnals. To date, the youngest occurrence of Gilbertsocrinus was the early Viséan of the United States, thus the present finding in upper Viséan extends this genus range. Furthermore, the occurrence of Lanecrinus? sp. expands the Western European range of this genus into the Tournaisian. A single partially disarticulated crown, Crinoidea Indeterminate B, is described from the Serpukhovian of the Upper Silesian Coal Basin. In addition, several echinoid test plates and spines are also reported.

Body-size increase in crinoids following the end-Devonian mass extinction.

The Devonian period ended with one of the largest mass extinctions in the Earth history. It comprised a series of separate events, which eliminated many marine species and led to long-term post-extinction reduction in body size in some groups. Surprisingly, crinoids were largely unaffected by these extinction events in terms of diversity. To date, however, no study examined the long-term body-size trends of crinoids over this crucial time interval. Here we compiled the first comprehensive data sets of sizes of calyces for 262 crinoid genera from the Frasnian-Visean. We found that crinoids have not experienced long-term reduction in body size after the so-called Hangenberg event. Instead, size distributions of calyces show temporal heterogeneity in the variance, with an increase in both the mean and maximum biovolumes between the Famennian and Tournaisian. The minimum biovolume, in turn, has remained constant over the study interval. Thus, the observed pattern seems to fit a Brownian motion-like diffusion model. Intriguingly, the same model has been recently invoked to explain morphologic diversification within the eucladid subclade during the Devonian-early Carboniferous. We suggest that the complex interplay between abiotic and biotic factors (i.e., expansion of carbonate ramps and increased primary productivity, in conjunction with predatory release after extinction of Devonian-style durophagous fishes) might have been involved not only in the early Mississippian diversity peak of crinoids, but possibly also in their overall passive expansion into larger body-size niches.

Late Cretaceous crinoids (Echinodermata) from the southwestern margin of the Holy Cross Mts. (southern Poland) and phylogenetic relationships among bourgueticrinids.

A systematic account of crinoids from the Upper Coniacian-Lower Campanian of the southwestern margin of the Holy Cross Mountains in southern Poland is presented. Seven crinoid taxa [Marsupites testudinarius (von Schlotheim), Bourgueticrinus ellipticus (Miller), Bourgueticrinus sp., I.? granosus Valette, Isocrinus? sp., Nielsenicrinus carinatus Roemer and Austinocrinus bicoronatus (von Hagenow)] are described and illustrated. The new material from Poland extends down the stratigraphic range of Austinocrinus bicoronatus to the Lower Campanian. Morphometric data support that Bourgueticrinus ellipicus and B.? suedicus are conspecific. Taphonomy and paleoecology of recorded crinoid assemblages are discussed. Phylogeny of Cretaceous bourgueticrinids is also revisited.

Troglomorphism in the middle Triassic crinoids from Poland.

In this paper, we document the Middle Triassic marine fauna recovered from the fissure/cave system of Stare Gliny (southern Poland) developed in the Devonian host dolomite. The fossils are mostly represented by in situ preserved and small-sized holdfasts of crinoids (Crinoidea) that are attached to the cave walls. Other fossils found in the cave infills include articulated brittle stars and brachiopods. Our findings constitute the oldest Mesozoic evidence for troglophile crinoids. We suggest that troglomorphism in these echinoderms was likely related to protection against predation, which underscores the magnitude of anti-predatory adaptations to increased predation pressure that occurred during the Early Mesozoic Marine Revolution.

Reassessing the improbability of a muscular crinoid stem.

Muscular articulations in modern stalked crinoids are only present in the arms. Although it has been suggested that certain coiled-stemmed fossil taxa may have been functionally adapted to utilize muscles, evidence supporting this interpretation is lacking. Here, we use cathodoluminescence and SEM to reveal the skeletal microstructure of the enigmatic coiled-stemmed taxon Ammonicrinus (Flexibilia). Based on the well-established link between skeletal microstructure and the nature of infilling soft tissues in modern echinoderms, we reconstructed the palaeoanatomy of the Middle Devonian ammonicrinids. We show that their median columnals with elongated lateral columnal enclosure extensions (LCEE) have stereom microstructure unexpectedly resembling that in the crinoid muscular arm plates. In particular, large ligamentary facets, that are present on each side of a transverse ridge, are mainly comprised of fine galleried stereom that is indicative of the mutable collagenous tissues. In contrast, fine labyrinthic stereom, commonly associated with muscles, is situated in the periphery on each side of the surface of elongated LCEE. Our findings thus strongly suggest that the muscles may have also been present in the stem of ammonicrinids. These results reassess the previous hypotheses about evolution of muscles in crinoids and provide new insights into the mode of life of Ammonicrinus.

Microlens arrays in the complex visual system of Cretaceous echinoderms.

It has long been assumed that photosensitivity in echinoderms is mainly related to diffuse photoreception mediated by photosensitive regions embedded within the dermis. Recent studies, however, have shown that some extant echinoderms may also display modified ossicles with microlenses acting as sophisticated photosensory organs. Thanks to their remarkable properties, these calcitic microlenses serve as an inspiration for scientists across various disciplines among which bio-inspired engineering. However, the evolutionary origins of these microlenses remain obscure. Here we provide microstructural evidence showing that analogous spherical calcitic lenses had been acquired in some brittle stars and starfish of Poland by the Late Cretaceous (Campanian, ~79 Ma). Specimens from Poland described here had a highly developed visual system similar to that of modern forms. We suggest that such an optimization of echinoderm skeletons for both mechanical and optical purposes reflects escalation-related adaptation to increased predation pressure during the so-called Mesozoic Marine Revolution.

Drill holes and predation traces versus abrasion-induced artifacts revealed by tumbling experiments.

Drill holes made by predators in prey shells are widely considered to be the most unambiguous bodies of evidence of predator-prey interactions in the fossil record. However, recognition of traces of predatory origin from those formed by abiotic factors still waits for a rigorous evaluation as a prerequisite to ascertain predation intensity through geologic time and to test macroevolutionary patterns. New experimental data from tumbling various extant shells demonstrate that abrasion may leave holes strongly resembling the traces produced by drilling predators. They typically represent singular, circular to oval penetrations perpendicular to the shell surface. These data provide an alternative explanation to the drilling predation hypothesis for the origin of holes recorded in fossil shells. Although various non-morphological criteria (evaluation of holes for non-random distribution) and morphometric studies (quantification of the drill hole shape) have been employed to separate biological from abiotic traces, these are probably insufficient to exclude abrasion artifacts, consequently leading to overestimate predation intensity. As a result, from now on, we must adopt more rigorous criteria to appropriately distinguish abrasion artifacts from drill holes, such as microstructural identification of micro-rasping traces.

Ophiuroids discovered in the middle triassic hypersaline environment.

Echinoderms have long been considered to be one of the animal phyla that is strictly marine. However, there is growing evidence that some recent species may live in either brackish or hypersaline environments. Surprisingly, discoveries of fossil echinoderms in non-(open)marine paleoenvironments are lacking. In Wojkowice Quarry (Southern Poland), sediments of lowermost part of the Middle Triassic are exposed. In limestone layer with cellular structures and pseudomorphs after gypsum, two dense accumulations of articulated ophiuroids (Aspiduriella similis (Eck)) were documented. The sediments with ophiuroids were formed in environment of increased salinity waters as suggested by paleontological, sedimentological, petrographical and geochemical data. Discovery of Triassic hypersaline ophiuroids invalidates the paleontological assumption that fossil echinoderms are indicators of fully marine conditions. Thus caution needs to be taken when using fossil echinoderms in paleoenvironmental reconstructions.

Predator-induced macroevolutionary trends in Mesozoic crinoids.

Sea urchins are a major component of recent marine communities where they exert a key role as grazers and benthic predators. However, their impact on past marine organisms, such as crinoids, is hard to infer in the fossil record. Analysis of bite mark frequencies on crinoid columnals and comprehensive genus-level diversity data provide unique insights into the importance of sea urchin predation through geologic time. These data show that over the Mesozoic, predation intensity on crinoids, as measured by bite mark frequencies on columnals, changed in step with diversity of sea urchins. Moreover, Mesozoic diversity changes in the predatory sea urchins show a positive correlation with diversity of motile crinoids and a negative correlation with diversity of sessile crinoids, consistent with a crinoid motility representing an effective escape strategy. We contend that the Mesozoic diversity history of crinoids likely represents a macroevolutionary response to changes in sea urchin predation pressure and that it may have set the stage for the recent pattern of crinoid diversity in which motile forms greatly predominate and sessile forms are restricted to deep-water refugia.

Pelagic crinoids (Roveacrinida, Crinoidea) discovered in the Neogene of Poland.

Until recently, it has been assumed that pelagic crinoids, the roveacrinids (Roveacrinida, Crinoidea), became extinct during the Cretaceous-Paleogene boundary event. Recent finds of well-preserved roveacrinidal remains (brachials and radials) in the Danian (Early Paleogene) of Poland showed that they survived into the earliest Cenozoic. This group was thus characterized as a "dead clade walking". Here, we present fossil evidence that these pelagic crinoids survived in Poland until at least the Middle Miocene (Badenian, ca. 14 Myr ago)-more than 50 Myr after their supposed extinction. These Miocene roveacrinids constitute the first documented evidence of Roveacrinida in strata of Neogene age, thus prolonging the stratigraphic range of pelagic crinoids. This find characterizes the order as a "Lazarus taxon" rather than a "dead clade walking" group.

Post-Paleozoic crinoid radiation in response to benthic predation preceded the Mesozoic marine revolution.

It has been argued that increases in predation over geological time should result in increases in defensive adaptations in prey taxa. Recent in situ and laboratory observations indicate that cidaroid sea urchins feed on live stalked crinoids, leaving distinct bite marks on their skeletal elements. Similar bite marks on fossil crinoids from Poland strongly suggest that these animals have been subject to echinoid predation since the Triassic. Following their near-demise during the end-Permian extinction, crinoids underwent a major evolutionary radiation during the Middle-Late Triassic that produced distinct morphological and behavioral novelties, particularly motile taxa that contrasted strongly with the predominantly sessile Paleozoic crinoid faunas. We suggest that the appearance and subsequent evolutionary success of motile crinoids were related to benthic predation by post-Paleozoic echinoids with their stronger and more active feeding apparatus and that, in the case of crinoids, the predation-driven Mesozoic marine revolution started earlier than in other groups, perhaps soon after the end-Permian extinction.