A taphonomic approach to the genetic interpretation of clypeasteroid accumulations from the Miocene of Tarragona,NE Spain

Belaústegui, Z., Nebelsick, J.H., Gibert, J.M. de, Domènech, R. & Martinell, J. 2012: A taphonomic approach to the genetic interpretation of clypeasteroid accumulations from the Miocene of Tarragona, NE Spain. Lethaia, Vol. 45, pp. 548–565. Clypeasteroid accumulations are common in Cenozoic shallow marine sediments, particularly in the Neogene, as they are also in analogue modern environments. In this article, four clypeasteroid accumulations from Miocene (Serravallian) shallow marine carbonates of the El Camp de Tarragona Basin (NE Spain) are studied. Two of them are dominated by flat‐shaped Parascutella, whereas the other two consist exclusively of bell‐shaped Clypeaster. The combination of the taphonomic analysis of the tests, the palaeoecological and taphonomic information provided by the associated fossils, and the stratigraphical and sedimentological context of each one of these beds allows interpreting them as autochthonous/parautochthonous accumulations resulting from in situ reworking in moderate energy settings, except for one that constitutes a true sand dollar coquina deposited by a storm event. This contribution explores the potential of thorough taphonomic analysis of marine invertebrate skeletal concentrations and substantiates the importance of clypeasteroid echinoids as producers of shell beds in the Neogene. □ Clypeasteroids, Echinoids, Miocene, Taphonomy, Tarragona.     

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.     

Gut microbiota in the burying beetle,Nicrophorus vespilloides,provide colonization resistance against larval bacterial pathogens

Carrion beetles, Nicrophorus vespilloides, are reared on decomposing carrion where larvae are exposed to high populations of carcass‐derived bacteria. Larvae do not become colonized with these bacteria but instead are colonized with the gut microbiome of their parents, suggesting that bacteria in the beetle microbiome outcompete the carcass‐derived species for larval colonization. Here, we test this hypothesis and quantify the fitness consequences of colonization with different bacterial symbionts. First, we show that beetles colonized by their endogenous microbiome produce heavier broods than those colonized with carcass‐bacteria. Next, we show that bacteria from the endogenous microbiome, including Providencia rettgeri and Morganella morganii, are better colonizers of the beetle gut and can outcompete nonendogenous species, including Serratia marcescens and Escherichia coli, during in vivo competition. Finally, we find that Providencia and Morganella provide beetles with colonization resistance against Serratia and thereby reduce Serratia‐induced larval mortality. This effect is eliminated in larvae first colonized by Serratia, suggesting that while competition within the larval gut is determined by priority effects, these effects are less important for Serratia‐induced mortality. Our work suggests that an unappreciated benefit of parental care in N. vespilloides is the social transmission of the microbiome from parents to offspring.     

Bioerosive structures from Miocene marine mobile‐substrate communities in southern Spain,and description of a new sponge boring

Abstract: Neogene palaeoshore sediments are abundantly represented along the Mediterranean coast of Iberia. An outcrop north of the Sierra Tejeda, named La Resinera, exposes concentrations of pebbles and boulders of marble, comprising an upper Miocene marine beach deposit. The high diversity of bioerosion trace fossils present in these boulders includes structures produced by polychaete annelids, demosponges, echinoids and endolithic bivalves, which indicate a shallow shoreface environment. The ichnotaxa represented are Maeandropolydora sulcans, Caulostrepsis taeniola, Entobia geometrica, Entobia ovula, Circolites kotoncensis, Gastrochaenolites torpedo, Gastrochaenolites lapidicus, Gastrochaenolites ornatus and Gastrochaenolites turbinatus. The borings are Tortonian (late Miocene) in age. Also present, and particularly abundant, are large sponge borings that have a single chamber from which radiating canals emerge. This trace fossil is designated as Entobia resinensis isp. nov.     

Regional‐scale dominance of non‐framework building corals on Caribbean reefs affects carbonate production and future reef growth

Coral cover on Caribbean reefs has declined rapidly since the early 1980's. Diseases have been a major driver, decimating communities of framework building Acropora and Orbicella coral species, and reportedly leading to the emergence of novel coral assemblages often dominated by domed and plating species of the genera Agaricia, Porites and Siderastrea. These corals were not historically important Caribbean framework builders, and typically have much smaller stature and lower calcification rates, fuelling concerns over reef carbonate production and growth potential. Using data from 75 reefs from across the Caribbean we quantify: (i) the magnitude of non‐framework building coral dominance throughout the region and (ii) the contribution of these corals to contemporary carbonate production. Our data show that live coral cover averages 18.2% across our sites and coral carbonate production 4.1 kg CaCO₃ m^?2 yr^?1. However, non‐framework building coral species dominate and are major carbonate producers at a high proportion of sites; they are more abundant than Acropora and Orbicella at 73% of sites; contribute an average 68% of the carbonate produced; and produce more than half the carbonate at 79% of sites. Coral cover and carbonate production rate are strongly correlated but, as relative abundance of non‐framework building corals increases, average carbonate production rates decline. Consequently, the use of coral cover as a predictor of carbonate budget status, without species level production rate data, needs to be treated with caution. Our findings provide compelling evidence for the Caribbean‐wide dominance of non‐framework building coral taxa, and that these species are now major regional carbonate producers. However, because these species typically have lower calcification rates, continued transitions to states dominated by non‐framework building coral species will further reduce carbonate production rates below ‘predecline’ levels, resulting in shifts towards negative carbonate budget states and reducing reef growth potential.     

Pathologies of non‐marine bivalve shells from the Late Triassic of Poland

Gorzelak, P., Nied?wiedzki, G. & Skawina, A. 2010: Pathologies of non‐marine bivalve shells from the Late Triassic of Poland. Lethaia, Vol. 43, pp. 285–289. Shells of Late Triassic non‐marine bivalves from Lisowice (Lipie ?l?skie clay pit, southern Poland), which co‐occur with remains of several vertebrate taxa (mammal‐like reptiles, carnivorous dinosaurs, pterosaurs, temnospondyl amphibians, hybodont sharks, dipnoan and ganoid fish), bear evidence of pathologies. Distribution, dimension and shape of some of these injuries (radiate tooth marks) closely match the dental morphology of lungfish (here probably represented by the genus Ceratodus). Thus, we interpret these pathologies as evidence of unsuccessful predatory attack on bivalves by this fish. This interpretation is also consistent with modern examples of such behaviour among lungfish. The feasibility that other culprits caused other pathologies (shell scarring and wedges) on the bivalves analysed is also discussed. Discovery of these traces constitutes important evidence of predator–prey interaction, which provides ‘fingerprints’ of trophic structure within this Late Triassic freshwater ecosystem. □Freshwater bivalves, lungfish, pathologies, predation, Triassic.     

Comparative phylogeography of six herpetofauna species in Cyprus: late Miocene to Pleistocene colonization routes

The colonization patterns of oceanic islands are often interpreted through transmarine dispersal. However, in islands with intense human activities and unclear geological history, this inference may be inappropriate. Cyprus is such an island, whose geotectonic evolution has not been clarified yet to the desired level for biogeographical reconstructions, leaving the questions of ‘how the Cypriote biota arrived’ and ‘does the dispersal have the formative role in patterns of its diversification’ unanswered. Here, we address these issues through a reconstruction of the evolutionary history of six herptiles (Ablepharus budaki, Ophisops elegans, Acanthodactylus schreiberi, Telescopus fallax, Pelophylax cf. bedriagae, and Hyla savignyi) by means of mitochondrial DNA (cytochrome b and 16S rRNA), applying a Bayesian phylogenetic, biogeographical, and chronophylogenetic analyses. The phylogeographical analyses show that the colonization history of those species in Cyprus started in the late Miocene and extended into the Pliocene and Pleistocene, with geodispersal, transmarine dispersal, and human‐mediated dispersal having their share in shaping the diversification of Cypriote herptiles. The revealed patterns could be divided into three biogeographical categories: old colonizers that arrived in Cyprus during the late Miocene or early Pliocene either by a land bridge (geodispersal) which connected Cyprus with the mainland or by transmarine dispersal, younger colonizers that reached the island through transmarine dispersal from the Middle East, and new settlers that arrived through human‐induced (voluntary or not) introductions. This work advances our knowledge of the biogeography of Cyprus and highlights the need to consider both geo‐ and transmarine dispersal when dealing with islands whose associations do not have a straightforward interpretation. © 2013 The Linnean Society of London     

Periostracal mineralization in the gastrochaenid bivalve Spengleria

We investigated the spikes on the outer shell surface of the endolithic gastrochaenid bivalve genus Spengleria with a view to understand the mechanism by which they form and evaluate their homology with spikes in other heterodont and palaeoheterodont bivalves. We discovered that spike formation varied in mechanism between different parts of the valve. In the posterior region, spikes form within the translucent layer of the periostracum but separated from the calcareous part of the shell. By contrast those spikes in the anterior and ventral region, despite also forming within the translucent periostracal layer, become incorporated into the outer shell layer. Spikes in the posterior area of Spengleria mytiloides form only on the outer surface of the periostracum and are therefore, not encased in periostracal material. Despite differences in construction between these gastrochaenid spikes and those of other heterodont and palaeoheterodont bivalves, all involve calcification of the inner translucent periostracal layer which may indicate a deeper homology.     

Bored and Burrowed: An Unusual Echinoid Steinkern from the Type Maastrichtian (Upper Cretaceous,Belgium)

A test of the irregular echinoid Hemipneustes striatoradiatus (Leske) was dissolved in dilute hydrochloric acid to expose the flint steinkern. This revealed slender, string-like borings that were mainly limited to ambulacra (Talpina isp. cf. T. canna (Price)); less common pouch-like borings limited to part of one ambulacrum only (Rogerella isp. cf. R. pattei (Saint Seine)); and slender, sinuous to contorted burrows, Planolites montanus Richter. Ingress to the test (borers) and sediment infill (burrowers) was presumably through the ambulacral pore pairs of the dead test. Borings and burrows were probably not coeval. Such infestations of fossil echinoids may be locally common, but their identification is controlled by the difficulty of seeing features of both the test and the infill.     

Differential colonization of wheat cultivars by two biotypes of Russian wheat aphid （Homoptera： Aphididae）

Susceptible and resistance wheat cultivars, Triticum aestivum L, were presented to two biotypes of Russian wheat aphid, Diuraphis noxia （Mordvilko）, in multiple choice tests to assay their relative acceptability as host plants. Both apterae （third and fourth instars） and alate adults were offered plants at the two-leaf stage in different cultivar combinations at 22±1℃ and 16：8 （L： D） hour photoperiod. Apterae were released from Petri dishes in the center of a circle of test plants, whereas alatae dispersed from a mature aphid colony to settle on plants arranged in rows. Both alatae and apterous nymphs of both biotypes readily colonized all cultivars tested：‘2137＇, ‘Akron＇,‘Ankor’,‘ Halt’ ,‘ Jagger’ ,‘ Prairie Red’ , ‘Stanton＇,‘TAM 107＇,‘TAM 110＇,‘Trego＇, ‘ Yuma＇, and ‘Yumar＇. Fewer biotype I apterae responded （settled and fed） in the combination containing more resistant （Dn4- and Dny-expressing） cultivars, compared to the combinations that had fewer. The reverse was true for biotype 2 apterae; more aphids responded in the combination containing the largest number of Dn4 expressing cultivars. Differential colonization of cultivars was observed in only one combination, in which biotype 2 apterae colonized Akron and Yumar in larger numbers than they did Stanton and Yuma. A separate experiment confirmed that, 48 hours after infestation, more biotype 2 apterae abandoned plants of Yuma than plants of Yumar. This differential response was likely due to genetic differences between the two ＇ near isogenic＇ lines that include the lack of Dn4 expression in Yuma. Choice tests with alatae did not result in differential rates of cultivar colonization by either biotype in any combination tested. These results suggest that young wheat plants appear to lack any meaningful antixenosis toward D. noxia, even though the aphids appear to perceive, and sometimes respond to, certain differences in cultivar suitability.     

Histochemical Studies of the Non‐Host Resistance and the Role of Actin Cytoskeleton in Pepper Against Colletotrichum orbiculare

The penetration process and defence reactions (hypersensitive response, oxidative burst and cell wall fortification) of Colletotrichum orbiculare were studied histochemically on pepper cultivar ‘A11’ (non‐host) and susceptible cucumber cultivar ‘Changchun Thorn’ (host). The results indicate that C. orbiculare could hardly penetrate the non‐host pepper leaves. It was papillae rather than hypersensitive response and H₂O₂ that played an important role in resisting the colonization and development of C. orbiculare on the non‐host pepper. The depolymerization of the actin microfilament weakened the papilla deposition of pepper and allowed successful penetration of the non‐adapted C. orbiculare, suggesting that the actin cytoskeleton of pepper is significant in preventing the invasion of the non‐host pathogen C. orbiculare.     

Epi- and endobiontic organisms on Late Jurassic crinoid columns from the Negev Desert, Israel: Implications for co-evolution

Columns of the articulate crinoids Millericrinus and Apiocrinites from the Upper Jurassic (Upper Callovian) Zohar and Matmor formations of the Negev Desert of Israel display abundant encrusting organisms of about ten species, as well as diverse trace fossils produced by endobionts. Pluricolumnals were colonized by epi- and endobiontic organisms both during life and post-mortem. Skeletonized encrusting organisms include abundant ostreid bivalves (which evidently colonized both live and dead crinoid columnals), two types of serpulid worms, encrusting foraminifera, three species of bryozoans, and small encrusting sclerosponges. Several types of borings are present: Trypanites (possibly produced by sipunculids), Gastrochaenolites (crypts of boring lithophagid bivalves), elliptical barnacle? borings, and channel-like annelid? borings. In addition, approximately 16% of the pluricolumnals display circular parabolic embedment pits assignable to the ichnogenus Tremichnus. They are associated with substantial deformation of the containing columnals and were probably the work of host-specific ectoparasitic organisms. Discovery of Tremichnus on Jurassic crinoids extends the range of this trace by almost 100 million years, providing evidence for one of the longest-ranging host-parasite interactions documented thus far (over 200 million years). The relationship of epibionts to the Jurassic crinoids thus ranged from simple utilization of dead hard substrate to probable opportunistic commensalism in forms that colonized the live upright stems, as in some oysters, through host-specific parasitism in the case of Tremichnus.     

A New Ichnospecies of Gastrochaenolites Leymerie from the Pleistocene Port Morant Formation of Southeast Jamaica and the Taphonomy of Calcareous Linings in Clavate Borings

The ichnospecies Gastrochaenolites pickerilli isp. nov. is based on ten borings found in a shell of the gastropod Strombus gigas Linné from the Pleistocene (Sangamonian) Port Morant Formation of southeast Jamaica. These borings bear morphological similarities to Gastrochaenolites torpedo Kelly and Bromley but differ from all other Gastrochaenolites ispp. in having prominent and numerous calcareous meniscate structures arrayed adjacent to one side of the boring. These menisci are concave towards the center of the boring and are the remnants of calcareous tubes that lined earlier boreholes, that the boring bivalve treated as part of the lithified substrate when relocating. They are thus evidence of the former positions of borings that, unusually, were breached as the bivalve migrated sideways. Although this was a common behavior for Gastrochaenolites-producing bivalves within this substrate, the reason for it occurring is uncertain.     

Patterns of sclerobiont colonization on the rugose coral Schlotheimophyllum patellatum (Schlotheim, 1820) from the Silurian of Gotland,Sweden

Analysis of mushroom-shaped rugose corals Schlotheimophyllum patellatum (Schlotheim, 1820) from the Silurian (Upper Visby Beds, Lower Wenlock, Sheinwoodian) of Gotland, Sweden, showed that they were colonized on both the upper (exposed) and lower (cryptic) sides by a variety of encrusting and boring (sclerobiont) biotas, represented by 10 taxa and at least 23 species. Bryozoans and microconchid tubeworms, the most abundant encrusters, dominated on the cryptic undersides of the corals, while the dominant endobionts responsible for Trypanites borings overwhelmingly dominated the exposed surfaces. Except for cnidarian sphenothallids, which were exclusive colonizers of the underside of only one coral host, no other encrusters could be referred to as obligate cryptobionts. Because the upper surface of these corals was likely covered by soft-tissues during life, in specimens lifted off the sea-floor sclerobionts must have settled on the cryptic sides first. They could colonize the upper side only after the coral’s death, unless it was covered by sediment as could be the case in some flat specimens. With time, the space on the underside of the coral skeleton may have progressively been filled by sediment as well, precluding further colonization by sclerobionts. In that respect, the colonization patterns of these corals by encrusters and borers were controlled by the complex interplay of environmental factors, sclerobiont dynamics and coral growth in a given Silurian habitat. Compared with Silurian stromatoporoid hosts, the sclerobiont diversity and abundance noted on the Schlotheimophyllum corals may be regarded as representative for the Silurian as a whole.     

Towards a balanced view of pike in Ireland: a reply to Ensing

In our recent study of the population genetics of pike (Esox lucius) in Ireland (Pedreschi et al., 2014 , Journal of Biogeography, 41 , 548–560), we reported the existence of two main demographic units and showed that these may correspond to two independent and temporally staggered colonization events, the first of which may have been too old to be caused or assisted by human translocations. Ensing (2015, Journal of Biogeography, doi: 10.1111/jbi.12410 ) first used our genotypic data to explore alternative historical scenarios, then attempted to reconcile the ‘two‐wave’ colonization process of Ireland by pike with translocation activities by humans in Neolithic/Bronze age times. Here we illustrate why the evidence base for Ensing's reconstruction is weak and we outline a realistic strategy to better understand the role of pike in Irish freshwater ecosystems.     

‘Becoming a species by becoming a pest’ or how two maize pests of the genus Ostrinia possibly evolved through parallel ecological speciation events

New agricultural pest species attacking introduced crops may evolve from pre‐existing local herbivores by ecological speciation, thereby becoming a species by becoming a pest. We compare the evolutionary pathways by which two maize pests (the Asian and the European corn borers, ACB and ECB) in the genus Ostrinia (Lepidoptera, Crambidae) probably diverged from an ancestral species close to the current Adzuki bean borer (ABB). We typed larval Ostrinia populations collected on maize and dicotyledons across China and eastern Siberia, at microsatellite and mitochondrial loci. We found only two clusters: one on maize (as expected) and a single one on dicotyledons despite differences in male mid‐tibia morphology, suggesting that all individuals from dicotyledons belonged to the ABB. We found evidence for migrants and hybrids on both host plant types. Hybrids suggest that field reproductive isolation is incomplete between ACB and ABB. Interestingly, a few individuals with an ‘ABB‐like’ microsatellite profile collected on dicotyledons had ‘ACB’ mtDNA rather than ‘ABB‐like’ mtDNA, whereas the reverse was never found on maize. This suggests asymmetrical gene flow directed from the ACB towards the ABB. Hybrids and backcrosses in all directions were obtained in no‐choice tests. In laboratory conditions, they survived as well as parental strain individuals. In Xinjiang, we found ACB and ECB in sympatry, but no hybrids. Altogether, our results suggest that reproductive isolation between ACB and ABB is incomplete and mostly prezygotic. This points to ecological speciation as a possible evolutionary scenario, as previously found for ECB and ABB in Europe.     

Dark septate endophytes colonizing the roots of ‘non-mycorrhizal’ plants in a mine tailing pond and in a relatively undisturbed environment,Southwest China

Dark septate endophytes (DSEs), one of the most common fungal colonizers of roots, are considered to overlap in function with mycorrhizal fungi. However, there is little knowledge on the distribution and identity of DSEs in ‘non-mycorrhizal’ plants. In the current study, colonization and diversity of DSEs colonizing the roots of eight typically ‘non-mycorrhizal’ families were assessed. In total, 120 root samples of 31 plant species were all colonized by DSEs. Intensity of DSE colonization varied greatly among different plant species, with a range of 0.56–47.56%, 8.13% on average. Cladosporium, Cyphellophora and Phialophora were the dominant genera, with a relative abundance of more than 60% over a total of 90 isolates. Our results showed that diverse DSE species colonized the roots of ‘non-mycorrhizal’ plants, especially they were more common in degraded mine tailings than in the undisturbed site, but their integral roles to the functional roots are in need of further experimental demonstration.     

Influence of cabbage resistance and colour upon the diamondback moth and its parasitoid Oomyzus sokolowskii

Host plant resistance and biological control are vital integrated pest management tools against the diamondback moth (DBM), Plutella xylostella (L.) (Lepidoptera: Plutellidae), but to date no study has investigated this system including the DBM parasitoid Oomyzus sokolowskii (Kurdjumov) (Hymenoptera: Eulophidae). We examined oviposition and development of P. xylostella exposed to two commercial cabbage cultivars (green ‘Chato de quintal’ and red ‘Roxo’) and possible effects upon O. sokolowskii. Under free‐choice tests, DBM females laid significantly more eggs on plants of the green cabbage, even though several population growth parameters showed that DBM developed better on the red cabbage. Furthermore, a laboratory free‐choice test with artificially green‐ and red‐painted kale leaf discs demonstrated a similar oviposition preference pattern, with green colour being preferred over red colour. The preference was apparently visually mediated; olfactometer tests showed similar attraction of moths to both green and red cultivars in choice and non‐choice tests. Host plant cultivar had no statistically significant effect on female parasitoid behaviour towards DBM larvae, nor on parasitoid numbers or longevity. Moreover, wasps parasitizing DBM larvae reared on the green cultivar developed more quickly and in larger numbers per parasitized larva. Thus, feeding on green cabbage rather than red does not hinder, and potentially even enhances, control of DBM by O. sokolowskii. On a practical level, these results suggest that intercalating green cabbage cultivars as a trap crop might help protect more profitable red cultivars in growing fields.     

Ichnofacies and microbial build-ups on Late Miocene rocky shores from Menorca (Balearic Islands), Spain

Angular unconformites between Jurassic and Miocene strata are exposed in the sea cliffs of Cala Cigonya on the northwest coast of Menorca in the Balearic Islands of Spain. The geological discontinuities represent rocky shores on opposite sides of a former headland with 15 m of topographic relief. On the east flank, Jurassic dolomite is overlain by Upper Miocene (Tortonian to Messinian) breccia and laminated limestone. Here, a partially exhumed dolomite surface records Miocene bivalve borings of the ichnospecies Gastrochaenolites torpedo and G. lapidicus that achieved a density of >1,000 borings/m². Other associated traces include sponge borings (Entobia isp.) and polychaete borings (Caulostrepsis isp.). A breccia deposit 0.8 m thick was derived from the underlying dolomite and angular clasts still retain evidence of bivalve borings. Above follows a succession of laminated limestone beds more than 5 m thick, including some levels with dome-shaped stromatolites and other horizons with reworked dolomite clasts. Thin-section analysis of the laminated limestone reveals dark and light couplets 0.2 mm thick consistent with microbial origins. In contrast, the west flank was buried by coarse sandstone and laminated sediments. Here, dwelling structures of regular echinoids (Circolites kotoncensis) are the dominant traces preserved on the dolomite surface, reaching a maximum density of 66 borings/m². Associated borings include Entobia geometrica as well as rare traces of Gastrochaenolites isp. and Trypanites isp. Notable for the absence of a basal Miocene breccia, the west flank is interpreted as a sheltered rocky shore coeval with an exposed rocky shore on the east flank. Today, heavy surf on the north coast of Menorca is related to the Tramontana winds that blow out of Spain during winter months. Similar atmospheric circulation patterns must have prevailed during the Late Miocene, but the replacement of ichnofacies by microbial build-ups resulted from increased salinity during the Messinian crisis.     

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.