Experimental neoichnology of crawling stalked crinoids

Stalked crinoids have long been considered sessile. In the 1980s, however, observations both in the field and of laboratory experiments proved that some of them (isocrinids) can actively relocate by crawling with their arms on the substrate, and dragging the stalk behind them. Although it has been argued that this activity may leave traces on the sediment surface, no photographs or images of the traces produced by crawling crinoids have been available. Herein, we present results of neoichnological experiments using the shallowest species of living stalked crinoid, Metacrinus rotundus, dredged from Suruga Bay (near the town of Numazu, Shizuoka Prefecture, ~ 140 m depth). Our results demonstrate that isocrinids produce characteristic locomotion traces, which have some preservation potential. They are composed of rather deep and wide, sometimes weakly sinuous, central drag marks left by the stalk and cirri, and short, shallow scratch marks made by the arms. Based on the functional morphology and taphonomy, it has been argued that the ability to autotomize the stalk and relocate had already evolved in the oldest stem-group isocrinids (holocrinids), likely in response to increased benthic predation pressure during the so-called Mesozoic marine revolution. Our data show that this hypothesis may be corroborated in the future by ichnological findings, which may provide more direct proof of active locomotion in Triassic holocrinids

The paleobiogeographical significance of the Silurian and Devonian trilobites of Japan

Six major groups of trilobites from the Silurian and Devonian of Japan are evaluated for their paleobiogeographical signature. Silurian illaenids and scutelluids show four generic-level and at least two species-level links with the Australian segment of the Gondwana paleocontinent; encrinurids also indicate two generic-level links with Australia and also the South China paleocontinent; whilst Devonian phacopids, and possibly proetids, suggest at least two generic-level links with the North China paleocontinent. These different patterns may reflect the fragmentary biostratigraphical record of Japanese trilobites, but they also appear to reflect paleoenvironmental parameters associated with lithofacies, and paleoecology. Thus, Japanese assemblages of proetids and phacopids occurring in deep-water clastic lithofacies have counterparts in similar settings in North China, and Japanese scutelluids and illaenids are strongly associated with shallow marine carbonate lithofacies that are similar to those of their occurrences in Australia. Japanese encrinurids occur in carbonate rocks indicative of shallow marine settings in the Kurosegawa Terrane, and they demonstrate a consistent paleobiogeographical affinity with Australia and South China. Larval ecology cannot be directly assessed for Japanese trilobite groups. However, proetids have consistently been shown to have planktonic protaspides, whereas illaenids, scutelluids, and encrinurids have benthic protaspides. Planktonic protaspides would have a greater propensity for distribution in ocean currents than benthic ones, and therefore may be of more limited paleobiogeographical utility. The combined data from the six different groups indicates that the complex paleobiogeographical patterns of the Japanese trilobite assemblages need to be interpreted with caution, and similarity of taxa does not necessarily denote paleogeographical proximity to other regions

Fixed, Free, and Fixed: The Fickle Phylogeny of Extant Crinoidea (Echinodermata) and Their Permian-Triassic Origin

Although the status of Crinoidea (sea lilies and featherstars) as sister group to all other living echinoderms is well-established, relationships among crinoids, particularly extant forms, are debated. All living species are currently placed in Articulata, which is generally accepted as the only crinoid group to survive the Permian–Triassic extinction event. Recent classifications have recognized five major extant taxa: Isocrinida, Hyocrinida, Bourgueticrinina, Comatulidina and Cyrtocrinida, plus several smaller groups with uncertain taxonomic status, e.g., Guillecrinus, Proisocrinus and Caledonicrinus. Here we infer the phylogeny of extant Crinoidea using three mitochondrial genes and two nuclear genes from 59 crinoid terminals that span the majority of extant crinoid diversity. Although there is poor support for some of the more basal nodes, and some tree topologies varied with the data used and mode of analysis, we obtain several robust results. Cyrtocrinida, Hyocrinida, Isocrinida are all recovered as clades, but two stalked crinoid groups, Bourgueticrinina and Guillecrinina, nest among the featherstars, lending support to an argument that they are paedomorphic forms. Hence, they are reduced to families within Comatulida. Proisocrinus is clearly shown to be part of Isocrinida, and Caledonicrinus may not be a bourgueticrinid. Among comatulids, tree topologies show little congruence with current taxonomy, indicating that much systematic revision is required. Relaxed molecular clock analyses with eight fossil calibration points recover Articulata with a median date to the most recent common ancestor at 231–252 mya in the Middle to Upper Triassic. These analyses tend to support the hypothesis that the group is a radiation from a small clade that passed through the Permian–Triassic extinction event rather than several lineages that survived. Our tree topologies show various scenarios for the evolution of stalks and cirri in Articulata, so it is clear that further data and taxon sampling are needed to recover a more robust phylogeny of the group

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

Early Cretaceous Isocrinus from northeast Japan

Volume: 28Start Page: 629End Page: 64

Arm autotomy and arm branching pattern as anti-predatory adaptations in stalked and stalkless crinoids

Arm autotomy was induced in a living specimen of Metacrinus rotundus (Echinodermata: Crinoidea). An arm was autotomized at a ligamentary articulation known as a cryptosyzygy, following incision by scissors distal to the break point. Although sessile stalked crinoids cannot entirely escape from a predatory attack by arm autotomy and they do not have an active defense, arm autotomy at cryptosyzygies reduces damage and arm loss by effective distribution, and by minimizing trauma and facilitating subsequent regeneration. The paradigmatic distribution of cryptosyzygies in which arm loss is set at a minimum, compared with the actual distribution, shows that these two patterns are similar and that actual specimens successfully reduce arm loss by the effective distribution of cryptosyzygies. The crinoid branching pattern also affects arm loss, and two different paradigms are discussed: anti-predatory and harvesting. Arm branching patterns of various isocrinids have tended toward the anti-predatory configuration from the Jurassic to the Recent, suggesting that the isocrinids have coped with increased predation. Shallow-water comatulids generally adopt the anti-predatory paradigm in their branching pattern, whereas many deep-water, stalked crinoids adopt a harvesting paradigm, reflecting that shallow-water comatulids receive more predatory attacks than do deep-water crinoids

Survival of crinoid stalk fragments and its taphonomic implications: additional discussion

Volume: 2Start Page: 285End Page: 28

La reconquête triasique par les échinodermes

L\u27intervalle de temps correspondant aux extinctions massives du Permo-Trias fut une période importante dans l\u27histoire évolutive des échinodermes. Le détail des modalités de ces extinctions, en particulier de celles de la reconquête se produisant immédiatement après les extinctions, est rarement abordé, car les données paléontologiques sur les échinodermes du Permien et du Trias sont estimées par trop insuffisantes. Pourtant, seuls les Holothuroidea et les Asteroidea souffrent d\u27un manque de représentants fossiles au début du Trias. Mais, même dans ces deux groupes, les modalités des extinctions et de la reconquête peuvent être déduites des analyses cladistiques récentes. Le cas des Holothuroidea, qui ne présentent pas d\u27extinctions au niveau des familles au cours de l\u27intervalle du Permo-Trias, vraisemblablement en raison de leur mode de nutrition détritivore, se révèle unique parmi les échinodermes. En revanche, les Echinoidea, les Crinoidea et probablement aussi les Asteroidea subirent à cette époque des évolutions en « goulot ». Parmi les échinides, la radiation post-permienne n\u27a véritablement eu lieu qu\u27à partir du Trias supérieur (Carnien), quoiqu\u27elle ait pu commencer au début du Trias. Chez les crinoïdes, la diversité taxonomique augmente considérablement à partir du Ladinien supérieur, alors que des analyses cladistiques suggèrent une diversification plus précoce, au début du Trias (Indusien). Bon nombre de restes de crinoïdes, non encore décrits, provenant du Trias inférieur de différentes régions du globe, indiquent que la radiation post-permienne du groupe a dû être plus rapide qu\u27il n\u27est couramment admis. Dans le Spathien, les ossicules sont parfois tellement abondants qu\u27ils participent à l\u27édification de roches. La rapidité de la radiation post-permienne des Ophiuroidea est démontrée par l\u27existence au Trias inférieur d\u27au moins sept espèces d\u27ophiures, même si les affinités à un niveau supérieur de ces taxons demeurent pour l\u27instant encore problématiques et si les données du Permien supérieur sont mal connues. Au sein des échinodermes du Trias inférieur, les ophiures constituent le groupe affichant la plus grande diversité, à la fois sous la forme d\u27individus complets et sous celle d\u27ossicules désarticulés. Les holothuries ont probablement engagé leur radiation post-permienne au début du Trias, mais les données de l\u27analyse cladistique militent en faveur d\u27un âge largement Anisien. Tous les échinodermes connus du Trias inférieur étaient des animaux de petite taille peuplant les eaux très peu profondes, oxygénées, dans la zone d\u27activité des vagues des basses latitudes.The Permian–Triassic mass extinction interval was an important time in the evolutionary history of the echinoderms. Details of the extinction and, in particular the immediate post-extinction recovery in the Early Triassic, are seldom addressed because of a perception that the Permian–Triassic echinoderm fossil record is too poor. However, only the Holothuroidea and Asteroidea lack any Early Triassic fossil representatives. Even in these groups, details of the extinction and recovery can be inferred from recent cladistic analyses. The Holothuroidea are unique amongst the echinoderms in showing no family level extinction through the Permian–Triassic interval, possibly due to their deposit-feeding lifestyle. In contrast, the Echinoidea, Crinoidea and probably the Asteroidea underwent severe evolutionary bottlenecks during that time. In the echinoids, significant post-Permian radiation occurred from the Late Triassic (Carnian), although it may have begun in the Early Triassic. In the Crinoidea, fossil diversity increases dramatically from the Late Ladinian, although cladistic analyses suggest that initial diversification took place in the Earliest Triassic (Induan). Many undescribed crinoid remains from Lower Triassic strata worldwide also imply that the post-Permian radiation in this group may have been more rapid than currently thought. Locally in the Spathian, crinoid ossicles may approach rock-building densities. The presence of at least seven Early Triassic fossil ophiuroid species may indicate rapid post-Permian radiation in the Ophiuroidea, although the higher level affinities of these taxa are presently unresolved and the Late Permian record is poorly known. Ophiuroid remains are the most diverse echinoderm fossils during the Early Triassic, comprising both complete body fossils and disarticulated ossicles. Holothuroids possibly radiated in the Early Triassic, but current evidence from cladistic analysis favours a largely Anisian age for the post-Permian radiation in this group. All known Early Triassic echinoderms were small-sized animals that inhabited very shallow, oxygenated, low palaeolatitude environments within wave base.</p

Supplemental figure 2

SUPPLEMENTAL DATA 2—Columnar section of the Rusophycus avalonensis Zone of the Chapel Island Formation in Fortune Head, Newfoundland