Oichnus Bromley Borings in the Irregular Echinoid Hemipneustes Agassiz from the Type Maastrichtian (Upper Cretaceous,The Netherlands and Belgium)

Three ichnospecies of Oichnus Bromley occur in tests of the large holasteroid echinoid Hemipneustes striatoradiatus (Leske) in the type area of the Maastrichtian (Upper Cretaceous) in The Netherlands and Belgium; Oichnus simplex Bromley (penetrative), Oichnus paraboloides Bromley (nonpenetrative and showing two distinct morphologies), and Oichnus excavatus isp. nov. (nonpenetrative). The two distinct morphologies of O. paraboloides (both shallow, one with a central boss) are gregarious, but do not occur together on the same specimens, suggesting they were generated by different taxa. Oichnus paraboloides with a central boss occurs on H. striatoradiatus from the upper Nekum Member, Maastricht Formation (Maastrichtian, Upper Cretaceous). Tests of the host echinoid are smaller in the overlying Meerssen Member, Maastricht Formation, where they are infested by O. excavatus, the largest borings considered herein, which have concave walls and a large central boss. Blisters inside tests from the Meerssen Member show that this infestation occurred when the echinoid was alive. It is postulated that producers of these borings in H. striatoradiatus may have been genetically related and increased in size during the Maastrichtian even as the host echinoids showed a size decrease. This size increase in H. striatoradiatus was genetic and cannot be related to increase in size of borings.     

Oichnus simplex Bromley infesting Hemipneustes striatoradiatus (Leske) (Echinoidea) from the Maastrichtian type area (Upper Cretaceous,The Netherlands)

Abstract

The large holasteroid echinoid Hemipneustes striatoradiatus (Leske) was exploited by diverse invertebrate encrusters and borers during the Maastrichtian, both pre- and post-mortem. In life, the specimen described herein was perforated by multiple Oichnus simplex Bromley borings close to the apical system. Each engendered a growth reaction from the echinoid, a mound-like swelling on the external surface of the test with the boring at the centre. These would have been moved away from the apical system as the echinoid grew and inserted new plates apically. Whether this infestation was the product of numerous individual organisms or, less likely, just one organism (gastropod?) that relocated when discouraged by each mound-like swelling is uncertain. Similar growth reactions are known from other echinoderms, but associated with non-penetrative Oichnus paraboloides Bromley.     

Tithonia oxfordiana,a new irregular echinoid associated with Jurassic seep deposits in south‐east France

Abstract: The infaunal irregular echinoid, Tithonia oxfordiana, is described and compared to congeneric species previously described from Upper Jurassic and lowermost Cretaceous strata. This new species characterizes a monospecific echinoid assemblage, which occurs only in some places where deep‐marine middle Oxfordian deposits are exposed in south‐east France. Specimens are closely packed and clearly concentrated at the top of small carbonate chemoherms; a close connection of the echinoids with the emission of reduced chemicals, which were oxidized by chemoautotrophic bacteria, is highly probable. Based on general test shape and plate architecture, T. oxfordiana probably was a deposit feeder on chemosynthetic organic matter produced by such bacteria. In view of the fact that T. oxfordiana is the sole species of the Jurassic genus Tithonia known from Oxfordian strata, it is postulated that chemoherms possibly acted as refugia for these peculiar echinoids, which have an episodic record between the Callovian and Valanginian.     

Internal Disorder: Post-Mortem Burrows within the Tests of the Holasteroid Echinoid Echinocorys Leske (Upper Cretaceous to Paleocene)

A test of the Lower Paleocene holasteroid echinoid Echinocorys ex. gr. scutata Leske from Göynük, Turkey, preserves a Planolites beverleyensis (Billings) burrow close to the internal surface. Identification of such a specimen is fortuitous; only where the test has broken cleanly away from the internal mould can such burrows be exposed, if present at all. A second specimen, from the Upper Cretaceous of Norfolk, England, bears a sinuous boring identified as Planolites montanus Richter.     

A Dense Infestation of Round Pits in the Irregular Echinoid Hemipneustes striatoradiatus (Leske) from the Maastrichtian of the Netherlands

A test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the upper Meerssen Member (subunit IVf-6; Maastricht Formation) in the type area of the Maastrichtian Stage (Late Cretaceous) was infested by 170 non-penetrative pits assigned to Oichnus excavatus (Donovan and Jagt). The echinoid is assumed to have been alive at that time. The distribution of pits is approximately bilaterally symmetrical and mainly supraambital; pits on the oral surface may have been carried subambitally by echinoid plate growth. Many pits are comparatively shallow; they probably were partly filled by the echinoid after death of the pit-forming organisms. The area around the periproct is devoid of pits as is the anterior ambulacrum (amb III). The latter was probably kept free of larval pit-formers by the tube feet; other ambulacra, with tube feet adapted for respiration, lacked such defense.     

An Intersection in Time and Space: Significance of Modern Invertebrate Borings in Upper Cretaceous Echinoids

Reworked fossils may contribute unique data to ichnology, stratigraphy, and paleoecology. Reworked Upper Cretaceous echinoids are found on Ocean Isle Beach, North Carolina, United States. Tests of Hardouinia sp. cf. H. mortonis were intensely bored by clionaid sponges (Entobia) and bivalves (Gastrochaenolites), with rare polychaete annelids (Caulostrepsis). Evidence for a high-energy environment in the Late Cretaceous is provided by the sandstone infill and a geopetal infill. Hardouinia were bored on all surfaces, largely lack surface detail and have been corraded, exposing the internal structure of some borings, indicating that these specimens were tumbled in a modern, high energy marine environment. Borers showed substrate preferences: Entobia is largely limited to the test, but do penetrate the sandstone infill in some specimens; Caulostrepsis mainly infested the crystalline calcite of the echinoid test; and Gastrochaenolites penetrated both tests and lithified sandstone infill. This Entobia-Gastrochaenolites assemblage in these echinoids differs from the Entobia-Caulostrepsis assemblage recognized from the approximately coeval, but spatially distant, tests of Upper Cretaceous Echinocorys ex gr. scutata found reworked on the North Sea coast of Norfolk, England. Unlike the Entobia-Gastrochaenolites association of North Carolina, the boring association in Norfolk is between Entobia and Caulostrepsis, which are found in both tests and chalk infill. Thus, the lithology of the infill is an important factor in determining these differences.     

Site Selectivity of the Pit Oichnus excavatus Donovan and Jagt Infesting Hemipneustes striatoradiatus (Leske) (Echinoidea) in the Type Maastrichtian (Upper Cretaceous,The Netherlands)

Two tests of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the type area of the Maastrichtian Stage (Upper Cretaceous) bear a varied infestation of episkeletozoans (oysters, bryozoan colony, and serpulids), borings (probable Caulostrepsis isp., Oichnus simplex Bromley), surface abrasion (Gnathichnus? isp.), and pits (O. excavatus Donovan and Jagt). Only O. excavatus represents a premortem infestation. In one specimen, the four individual pits of this ichnospecies are each associated with a different ambulacrum and pore pairs that, in life, bore respiratory tube feet; the anterior ambulacrum, of different gross morphology, is not infested. In the second test, three out of four of the same ambulacra are infested, although there are also O. excavatus in the interambulacra. The association between O. excavatus and the ambulacra of the echinoid, and thus its tube feet, is open to several plausible explanations, but most likely provided some form of feeding or protective advantage to the pit-forming organism.     

Extremes of Pit Infestation and Growth Deformity in a Crinoid Column,Permian of Timor

Crinoids are diverse and well-known from the Permian of Timor, but the literature has failed to document the numerous specimens of crinoid pluricolumnals from the fauna, many showing unusual morphology or yielding palaeoecological information. A curious and instructive specimen demonstrates the relationship between a living Permian crinoid and coeval invasive, pit-forming, invertebrates in detail. The pit-former is not preserved; most likely it was unmineralized or, if mineralized, then the shell simply dropped out. The infesting organism made pits assigned to the ichnospecies Oichnus paraboloides Bromley. The pit-former was unusually site selective. Either (1) one spatfall attached to just one side of the elevated (either up-current or down-current) or recumbent column and each individual centered their pits on the sutures between adjacent columnals; or (2) a single individual migrated along the column. The living crinoid showed an extreme reaction to this infestation. Excess stereom growth on the side of the pits transformed what was a circular column by addition of a thick, triangular ridge on the pitted side.     

Rogerella isp. Infesting the Pore Pairs of Hemipneustes striatoradiatus (Leske) (Echinoidea: Upper Cretaceous,Belgium)

An otherwise well-preserved test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the Emael Member, Maastricht Formation (Maastrichtian, Upper Cretaceous) of Belgium, is infested by encrusting bivalves and foraminiferans and the boring Rogerella isp. In this specimen, Rogerella preferentially infested and modified the ambulacral pore pairs of the echinoid close to the apex. This was not a commensal association. The echinoid test shows no growth deformations in response to this invasion; pore pairs are locally strongly infested; and encrusting invertebrates testify to the long post-mortem residence time of the test on the sea floor. Rather, the pore pairs of the dead echinoid were crannies attractive to settling larvae of acrothoracian barnacles, the producers of Rogerella.     

Experimental morphology of coronar growth in regular echinoids

Summary The coronar growth of a cidaroid and an aulodont echinoid are investigated by means of tetracycline labelling. The results are compared with earlier investigations on a stirodont and on a camarodont echinoid in order to evaluate the general features of coronar growth. In all echinoids new coronar plates are added at the apical end of the corona throughout the life cycle. The plates are shifted towards the peristome and they grow peripherically.In cidaroids ambulacral (A-) plates are detached from the firm corona. They are transformed into scales covering the peristomial field. The interambulacral (IA-) plates, however, are partially reabsorbed at the peristomial margin. In this manner the oldest solitary interambulacral plates are lost. The subsequent plates are arranged in pairs. The cidaroids thus show interradial growth even at their peristomial margin. This is unique to echinoids.In non-cidaroids there is a perignathic girdle made up of paired ambulacral auricles with interambulacral ridges in between. In some species the ridge is a solitary element. Therefore interradial growth cannot occur in the peristomial margin. In other species the ridge consists of several elements, but it also grows as a whole. Slight resorption of calcite occurs in places at the peristomial margin. In other places, however, calcite is added onto the peristomial edge. In non-cidaroids, therefore, the widening of the peristome is achieved solely by means of lateral growth in the plates bordering the peristome. The shift of the coronar plates from apicad to orad in noncidaroids is a relative shift only.In all echinoids the coronar plates are arranged in meridional columns. All plates grow up to the peristome. Their growth rates are relatively uniform towards the adambulacral sutures (which run between A- and IA-columns). Their growth rates towards the perradius and the interradius respectively are high in younger plates which are positioned above the ambitus, and decrease rapidly in plates located below the ambitus. Near the peristome the interradiad and perradiad growth rates are always considerably lower than adradiad growth rates. Perradial and interradial growth serve to adjust the plates in size and shape to their respective position in the corona.     

The infaunal echinoid Micraster: Taphonomic pathways indicated by sclerozoan trace and body fossils from the Upper Cretaceous of northern Spain

Micraster echinoid fossils are common in the Upper Cretaceous Olazagutía Formation of northern Spain. Tests frequently record sclerobiont signatures (including bioerosion and encrustation), left by reaction and/or coaction phenomena. Among bioerosion structures, Oichnus simplex, O. paraboloides, O. ichnosp. A, Trypanites solitarius, Rogerella ichnosp. indet., Centrichnus cf. eccentricus, Maeandropolydora ichnosp. indet. and fungal microborings are found, as well as pits and fractures. As for sclerozoan body fossils, bivalves (Dimyidae, Anomiidae, Plicatulidae and other Pectinacea), polychaete annelids (serpulids and spirorbids), lituolid foraminiferans (Haddoniidae and Coscinophragmatidae) and bryozoans (cheilostomate ones and others undetermined), as well as other less common groups, have been identified. Taphonomic paths followed by Micraster tests are analysed, based on conservation state and degree of colonisation and sedimentary filling. This allows to distinguish between accumulated fossils and non accumulated ones (including resedimented and reelaborated ones).     

IRREGULAR ECHINOIDS—AN INSUFFICIENTLY KNOWN GROUP

Some aspects of echinoid evolution from an endocyclic towards an exocyclic apical system, in the light of embryological and palaeontological data, are discussed. A primitive feature—imbrication of interamhulacral plates in Jurassic irregular echinoids, recently described—proves once more that this group is insufficiently known. Every revision of taxonomic or phylogenetic concepts concerning this group of echinoids should be preceded by careful morphological analysis of the material.     

Development of the nervous system in the brittle star Amphipholis kochii

There are several studies of neural development in various echinoderms, but few on ophiuroids, which develop indirectly via the production of pluteus larvae, as do echinoids. To determine the extent of similarity of neuroanatomy and neural development in the ophiuroids with other echinoderm larvae, we investigated the development of the nervous system in the brittle star Amphipholis kochii (Echinodermata: Ophiuroidea) by immunohistochemistry. Immunoreactive cells first appeared bilaterally in the animal pole at the late gastrula stage, and there was little migration of the neural precursors during A. kochii ontogeny, as is also the case in echinoids and holothuroids. On the other hand, neural specification in the presumptive ciliary band near the base of the arms does occur in ophiuroid larvae and is a feature they share with echinoids and ophiuroids. The ophiopluteus larval nervous system is similar to that of auricularia larvae on the whole, including the lack of a fine network of neurites in the epidermis and the presence of neural connections across the oral epidermis. Ophioplutei possess a pair of bilateral apical organs that differ from those of echinoid echinoplutei in terms of relative position. They also possess coiled cilia, which may possess a sensory function, but in the same location as the serotonergic apical ganglia. These coiled cilia are thought to be a derived structure in pluteus-like larvae. Our results suggest that the neural specification in the animal plate in ophiuroids, holothuroids, and echinoids is a plesiomorphic feature of the Ambulacraria, whereas neural specification at the base of the larval arms may be a more derived state restricted to pluteus-like larvae.     

Loriolella,a key taxon for understanding the early evolution of irregular echinoids

Loriolella Fucini is an enigmatic and poorly understood Early Jurassic sea-urchin that has been interpreted both as a regular and an irregular echinoid. The discovery of new and well-preserved material of the type species L. ludovicii (Meneghini) from the Upper Pliensbachian (Lower Jurassic) of Lombardy, Italy, has clarified the morphology of this genus for the first time. Loriolella resembles regular echinoids in having a large (almost certainly monocyclic) apical disc enclosing the periproct, and large primary interambulacral tubercles and spines. However, its peristome is extremely small with only vestigial buccal notches and auricles, and the ambulacra expand adorally forming cassiduloid-like phyllodes. Cladistic analysis suggests that Loriolella is the earliest known microstomatid irregular echinoid. Its unique mixture of primitive and derived characters is important for interpreting the initial steps by which irregular echinoids arose from diadematoid-like regular ancestors.     

Domes,arches and urchins: The skeletal architecture of echinoids (Echinodermata)

Summary A combination of simple membrane theory and statical analysis has been used to determine how stresses are carried in echinoid skeletons. Sutures oriented circumferentially are subject principally to compression. Those forming radial zig-zags are subject to compression near the apex and tension near the ambitus. Radial and circumferential sutures in Eucidaris are equally bound with collagen fibers but in Diadema, Tripneustes, Psammechinus, Arbacia and other regular echinoids, most radial sutures are more heavily bound, and thus stronger in tension. Psammechinus, Tripneustes and several other echinoids have radial sutures thickened by ribs which increase the area of interlocking trabeculae. Ribs also increase flexural stiffness and carry a greater proportion of the stress. Further, ribs effectively draw stress from weaker areas pierced by podial pores, and increase the total load which can be sustained.Allometry indicates that regular echinoids become relatively higher at the apex as size increases, thus reducing ambital stresses. Some spatangoids with very high domes (eg Agassizia) maintain isometry, but others (eg Meoma) become flatter with size. Both holectypoids (Echinoneus) and cassiduloids (Apatopygus) maintain a constant height to diameter relationship. Flattening, and consequently ambital tensile stress, is greatest in the clypeasteroids. In this group the formation of internal buttresses which preferentially carry stress, reaches maximum development. A notable exception, however, is the high domed Clypeaster rosaceus.In this analysis it was assumed that local buckling or bending does not occur. The test of some echinoids (e.g. Diadematoida) have relatively wide sutures swathed in collagen, which allows local deformation. Others (e.g. Arbacia) have rigid sutures with reduced collagen. In Psammechinus and other members of the Order Echinoida, in addition to rib formation, inner and outer surface trabeculae are thickened so that the individual plates are stiffened. Some spatangoids (Meoma, Paleopneustes) have extensive sutural collagen, but the cassiduloid Apatopygus has collagen confined to junctions of sutures, and elsewhere the joints are strengthened and stiffened by fusion of trabeculae. Fusion of surface trabeculae is almost complete in the holectypoid, Echinoneus, and the sutures are obscured.     

Revision of the family Idelinellidae,with a review of the Permian Eoblattida (Insecta)

Permian Eoblattida, which include the families Tillyardembiidae, Protembiidae (=Sylvardembiidae), Soyanopteridae, and Idelinellidae, are reviewed. The latter family is revised; it includes Idelinella macroptera Storozhenko, 1997 from the Middle Permian (Lower Kazanian Substage) of northern European Russia, Permostriga augustalis Novokshonov, 1999, Sylvastriga miranda Aristov, 2004, Strigulla cuculiophora (Aristov, 2002) comb. nov., Cucullistriga cucullata Aristov et Rasnitsyn, gen. et sp. nov., Scutistriga scutata Aristov et Rasnitsyn, gen. et sp. nov. from the Lower Permian (Kungurian Stage) of the Urals, Permeoblatta borealis Rasnitsyn et Aristov, 2010 from the Upper Permian (Upper Severodvinian Substage) of northern European Russia, and probably also Rasstriga americana Aristov et Rasnitsyn, gen. et sp. nov. from the Upper Carboniferous (Desmoinesian Stage) of Illinois.     

Ecophenotypic Variation and Developmental Instability in the Late Cretaceous Echinoid Micraster brevis (Irregularia; Spatangoida)

The Late Cretaceous echinoid genus Micraster (irregular echinoids, Spatangoida) is one of the most famous examples of a continuous evolutionary lineage in invertebrate palaeontology. The influence of the environment on the phenotype, however, was not tested so far. This study analyses differences in phenotypical variations within three populations of Micraster (Gibbaster) brevis from the early Coniacian, two from the Münsterland Cretaceous Basin (Germany) and one from the North Cantabrian Basin (Spain). The environments of the Spanish and the German sites differed by their sedimentary characteristics, which are generally a crucial factor for morphological adaptations in echinoids. Most of the major phenotypical variations (position of the ambitus, periproct and development of the subanal fasciole) among the populations can be linked to differences in their host sediments. These phenotypic variations are presumed to be an expression of phenotpic plasticiy, which has not been considered in Micraster in previous studies. Two populations (Erwitte area, Germany; Liencres area, Spain) were tested for stochastic variation (fluctuating asymmetry) due to developmental instability, which was present in all studied traits. However, differences in the amount of fluctuating asymmetry between both populations were recognised only in one trait (amount of pore pairs in the anterior paired petals). The results strengthen previous assumptions on ecophenotypic variations in Micraster.     

Entobia exogyrarum (Frič, 1883) from the Upper Cretaceous of the Bohemian Cretaceous Basin

The validity of the little-known ichnospecies Entobia exogyrarum (Fri?) is verified. Shells of an oyster, Rhynchostreon suborbiculatum (Lam.), represent the substrate for the sponge borings. The borings occur in many Upper Cenomanian to Middle Turonian localities of the Bohemian Cretaceous Basin. This study revealed that the sponges attacked shells of living oysters. Entobia exogyrarum (Fri?) also represents one of the shallowest occurrences of Entobia borings in the Upper Cretaceous rocks.     

Ontogeny and origin of the brooding system in Antarctic urechinid sea urchins (Echinodermata,Holasteroida)

Summary Echinoids usually broadcast gametes, and do not generally engage in a high degree of parental care. However, when they do, juveniles are typically maintained among the spines, or in shallow, external depressions in the test itself. The brooding Antarctic holasteroids Urechinus mortenseni and Plexechinus nordenskjoldi are bizarre exceptions: females develop an elaborate brooding system in which a small number of direct developing young are protected. Ontogeny of post-natal brooding urechinids is marked by profound divergence in the growth trajectories of male and female apical systems. In females, this leads to dramatic departures from the patterns found in all other echinoids. Otherwise, coronal skeleton allometry of males and females is almost identical. Juveniles in brood pouches grow larger than the diameter of the apical aperture through which they must pass to reach the external environment. The apical plates, from which the brooding system is suspended, hinge downward to enlarge the aperture, allowing the young to emerge from the female. A possible origin for the brooding system suggests derivation by centripetal plate addition from the ocular plates in the coronal skeleton. We develop a contrasting model for the origin of the brooding system that relies on a proposed homology between genital and periproctal elements of the apical system of echinoids and the more highly developed dorsal skeleton of other echinoderm classes.