Selectivity of Perinereis aibuhitensis (Polychaeta,Nereididae) feeding on sediment

Deposit feeding polychaetes play an important role in the acceleration of the biogeochemical processes of the sediment through bioturbation. Feeding is one of the important factors of bioturbation. However, knowledge of the feeding biology of polychaetes, especially the subsurface deposit feeder, is limited. The objective of this study is to characterize the feeding selection of Perinereis aibuhitensis with different body weights. The animals were fed with natural sediment from their original habitat in the lab. The feed intake and particle size of sediment were measured to find any evidence of feeding selection. A two-way ANOVA showed that the particle size class significantly affected the particle size frequency of the ingested and the remaining sediment (P?P?>?0.05). Bivariate correlation analysis showed that the particle size frequency of sediment, ingested and remaining sediment in different size classes were linearly related (P?P?P?P. aibuhitensis preferred smaller particles in the sediment, which was limited by the particle size distribution of the sediment in which they live. The nitrogen and organic carbon contents in the faeces might be the thresholds when P. aibuhitensis selects sediment particles as food. These results demonstrated the particle selectivity of P. aibuhitensis, and may be applicable to other subsurface deposit feeding polychaetes.     

Palaeobotanical remains associated with dinosaur fossils from the Camarillas Formation (Barremian) of Galve (Teruel,Spain)

A well-preserved macroflora and rich palynological assemblages corresponding to the Camarillas Formation (early-middle Barremian) in the San Cristóbal and Galve Mine sites from the Galve sub-basin in northeastern Spain are presented here. These remains represent the first fossil plant evidence from these deposits. Within plant macroremains, the cheirolepidiacean Pseudofrenelopsis aff. varians (Fontaine) Watson has been found. The palynological assemblage yielded well-preserved spores and pollen grains, mainly dominated by the genus Classopollis. Spores are also abundantly represented by schizaeacean spores (Cicatricosisporites and Plicatella). This spore assemblage supports an early-middle Barremian age for these localities. It is noteworthy that small basal angiosperm pollen grains of the genera Crassipollis and Retimonocolpites, together with other indeterminate grains, have been reported here. This flora constitutes the primary food producer for dinosaur at that time, and thus the information of the flora is important for the understanding of the ecological background for the dinosaur evolution during the Early Cretaceous.     

The effect of suspended clay on ciliate population growth rates

• 1 Renewed batch cultures of three algivorous and one bacterivorous ciliate showed that ciliate reproductive rates increase with increasing food concentration up to an asymptote. Maximum r_m values for Strobilidium gyrans, Bursaridium difficile and Euplotes eurystomus feeding on the cryptomonad alga Rhodomonas minuta were 0.87, 0.86 and 0.45 day-¹, respectively, while Paramecium aurelia feeding on the bacterium Enterobactor aerogenes had an r_m of 0.40 day-¹. B. difficile had a food threshold (r_m= 0) of 0.38 μg Cml-¹ and P. aurelia had a threshold of 1.6μg Cml-¹.
• 2 The presence of suspended fine (particle size <1 μm) and coarse (particle size 1–2μm) clay suppressed the half maximum reproductive rate of the ciliate Strobilidium gyrans by up to 69% but had no effect on B. difficile, E. eurystomus or P. aurelia. Populations of ciliates unaffected by clay may be released in situations where suspended particles negatively affect their metazooplankton predators and competitors.

     

Palynofloras from the upper Barremian-Aptian Nishihiro Formation (Outer Zone of southwest Japan) and the appearance of angiosperms in Japan

Palynomorphs are reported for the first time from the Nishihiro Formation (Wakayama Prefecture, Outer Zone of southwest Japan). The Nishihiro Formation consists of brackish to shallow marine deposits, dated as late Barremian to Aptian from geological correlations. Spores prevail in the assemblage, representing Filicopsida (mainly Cyatheaceae and Anemiaceae), Marchantiopsida and Lycopsida. The pollen assemblage is dominated by Coniferales, whereas Gnetales and Bennettitales/Cycadales are only rarely observed. Moreover, we report angiosperm pollen grains of the genus Retimonocolpites for the first time in the Early Cretaceous sediments of Japan. Pollen grains of the Retimonocolpites Group are typical of early angiosperms and commonly found in assemblages from the early to mid-Cretaceous of all paleofloristic provinces. Until this paper, the oldest angiosperm fossils in Japan were represented by a single seed and a wood reported from the Albian of Hokkaido. The oldest reliable angiosperm pollen grains were reported in Hokkaido from the Cenomanian, and in Honshu from the Coniacian. Thus, Retimonocolpites pollen grains reported in the present study represent the oldest record of angiosperms in Japan. They indicate an appearance of the angiosperms in Japan older than thought until now, which is consistent with that proposed elsewhere in eastern Asia.     

Preliminary palaeobotanical notes on the Early Cretaceous Cintura Formation (Sonora,Mexico)

The Cintura Formation of Albian–Cenomanian age in the Cabullona Basin yielded an important fossil flora, including palynomorphs, leaf impressions and fossil trunks. At the base of the Marquechi Member a poorly preserved palynomorph assemblage is recognised. The palynological assemblage is dominated by gymnosperms, mainly by cheirolepidiacean genus Classopollis. A noteworthy feature is the presence of angiosperm pollen grains of the genera Clavatipollenites, Retimonocolpites and Tucanopollis. A rich, diverse and well-preserved macroflora of leaf impressions has been observed in the uppermost part of this formation belonging to the San Juan Member. There are at least eight morphospecies of leaves including an abundance of taxa with possible botanical affinities to the family Sapindaceae. The angiosperm pollen grains together with the sapindalean leaves constitute the oldest record of such remains in Mexico.     

Scaphohippus, A New Genus of Horse (Mammalia: Equidae) from the Barstow Formation of California

A new genus of hypsodont equid, Scaphohippus, is recognized from the Green Hills Division (Ba1), Second Division (Ba1), and Barstow Faunas (Ba2) of the medial Miocene Barstow Formation, as well as from Ba1 and Ba2 faunas in the Punchbowl Formation of California, the Tesuque Formation of New Mexico, the Pawnee Creek Formation of Colorado, and the Olcott Formation of Nebraska. The genus includes two species, Scaphohippus sumani n. sp. and Scaphohippus intermontanus n. sp. Scaphohippus is a member of the “protohippine” clade, but is differentiated from its more derived sister taxon, Protohippus, by an arc-shaped incisor battery, a shorter I3-P1 diastema, and a more rounded protocone. The upper cheek teeth of S. sumani n. sp. have an unworn crown height of 40 mm. The protocone remains isolated until as much as 60% of wear, and the enamel fossettes are more complex than those of S. intermontanus n. sp. S. intermontanus n. sp. has an unworn crown height of 48–50 mm, the protocone opens by 30% of wear, and the enamel fossettes are simpler than those of S. sumani n. sp. Phyletic analysis shows S. sumani n. sp. to be the more primitive species. Similar facial and dental characteristics reveal a likely common ancestor with Merychippus insignis. The wider geographic and temporal range of S. sumani n. sp. reported here allows for useful phyletic and biogeographic interpretations. S. sumani n. sp. and M. insignis diverged during the late Hemingfordian (He2) of the Great Plains and both taxa dispersed throughout the western United States. S. intermontanus n. sp. is only recognized from the Barstow Formation, where it was likely directly descended from S. sumani n. sp.     

Soil composition affects the nesting behavior of blue-tailed bee-eaters (Merops philippinus) on Kinmen Island

The blue-tailed bee-eater (Merops philippinus) is a summer migrant that breeds on Kinmen Island, located off the west coast of Taiwan, about 5 km from the southern coast of mainland China. The aim of this study was to investigate why blue-tailed bee-eaters build their nests in sandy loam and sandy clay loam, but not in clay loam. Soil chemical and physical properties, and mineralogical composition were measured for the different soil types. Clay loam had a significantly lower pH, Na, and base saturation than did sandy loam or sandy clay loam. Clay loam had a significantly higher N, cation-exchange capacity (CEC), K, and free iron (Fe_d) and aluminum oxide (Al_d) contents than the other soil types. Clay loam had significantly lower sand and higher clay content, and higher bulk density and penetration resistance than the other soil types. The correlation coefficients (r ²) between penetration resistance and Fe_d, Al_d, and clay contents were 0.997, 0.848, and 0.779, respectively. Soil strength and compaction are important criteria for bee-eaters nesting-site selection. The lower pH of clay loam would enhance the exchangeable Al and acidity, further increasing the soil aggregation. Thus, it might prevent the bee-eaters from excavating nesting burrows.     

Taxonomic review of Agraphydrus from Japan (Coleoptera: Hydrophilidae: Acidocerinae)

The Japanese species of the genus Agraphydrus are revised. Five species are recognized, including Agraphydrus ogatai sp. nov. from Honshu, Shikoku and Kyushu. The new species can be distinguished by the coarse ground punctures on the pronotum and elytra, and by the shape of parameres. All species except A. luteilateralis are redescribed. All species are diagnosed, relevant morphological characters are illustrated, distributional records are summarized and an updated key is provided.     

Oldest known Dicoelosia and Epitomyonia,deep water brachiopods from the Beiguoshan Formation (Middle Katian,Upper Ordovician), Shaanxi,north China

Abstract: A diverse brachiopod fauna from a relatively deep water carbonate facies of the Upper Ordovician Beiguoshan Formation (uppermost Caradoc – lower Ashgill, middle Katian) is characterized by small shells and contains the oldest known Dicoelosia and Epitomyonia, two diagnostic taxa of deep water brachiopod palaeocommunities during the Late Ordovician and Silurian. Three new species are recognized: Dicoelosia cordiformis sp. nov., Dicoelosia perbrevis sp. nov. and Epitomyonia fui sp. nov. These pioneer forms of the family Dicoelosiidae show a relatively high degree of morphological plasticity. The shells of Dicoelosia from the Beiguoshan Formation range from the typical slender‐lobed form with a concavoconvex profile to the strongly equibiconvex, fat‐lobed morphotype that was not known previously until the late Silurian. The Beiguoshan dicoelosiids point to an important attribute of the deep water brachiopods: small generalists with high morphological plasticity, which make them ideal candidates as progenitors for the evolution of shallow water brachiopod faunas in shelf and platform depositional environments.     

Biodegradation of pyrene in sand,silt and clay fractions of sediment

Microbial degradation is the dominant pathway for natural attenuation of PAHs in environmental compartments such as sediments, which in turn depends on the bioavailability of PAHs. The bioavailability of PAHs has seldom been studied at the sediment particle size scale. We evaluated biodegradation of pyrene by Mycobacterium vanbaalenii PYR-1 as a function of sediment particle sizes, and investigated the relationship between the rate of degradation on sand, silt and clay particles with their individual desorption kinetics measured with the Tenax extraction method. Regression analysis showed that the total organic carbon (TOC), black carbon (BC), and specific surface area (SSA) of the specific particle size fractions, instead of the particle size scale itself, were closely related (P < 0.01) with the mineralization rate. While the fraction in the rapid desorption pool (F _rapid) ranged from 0.11 to 0.38 for the whole sediments and different size groups, the fractions mineralized after 336-h incubation (0.52 to 0.72) greatly surpassed the F _rapid values, suggesting utilization of pyrene in the slow desorption pool (F _slow). A biodegradation model was modified by imbedding a two-phase desorption relationship describing sequential Tenax extractions. Model analysis showed that pyrene sorbed on silt and clay aggregates was directly utilized by the degrading bacteria. The enhanced bioavailability may be attributed to the higher chemical concentration, higher TOC or larger SSA in the silt and clay fractions, which appeared to overcome the reduced bioavailability of pyrene due to sorption, making pyrene on the silt and clay particles readily available to degrading microbes. This conjecture merits further investigation.     

Angiosperm pollen grains of San Just site (Escucha Formation) from the Albian of the Iberian Range (north-eastern Spain)

The most complete angiospermous palynological assemblage for the middle-earliest late Albian of north-eastern Spain is reported from the Middle Member of the Escucha Formation at the San Just site. Nine palynological samples yielded mainly monosulcates (Clavatipollenites, Dichastopollenites, Hammenia, Jusinghipollis, Liliacidites, Pennipollis including proposed new species P. escuchensis, Retimonocolpites, and Transitoripollis) and a few tricolpates (Phimopollenites, Rousea, Striatopollis, and Tricolpites). These assemblages, mainly composed of angiosperm pollen grains, suggest that significantly diversified flowering plants flourished both in brackish and freshwater swamp plain environments represented by the Escucha Formation in Albian times, although they were less diverse than angiosperms in the Late Cretaceous. The palynological assemblages share greatest similarities with those of other southern Laurasian localities, although there are some taxa more typical of palynofloras of northern Gondwana.     

Life-history traits of the six Panonychus species from Japan (Acari: Tetranychidae)

Six species of the genus Panonychus are known from Japan. Life-history parameters of all six species were investigated at 25 °C, and for three species two strains of different geographical origin were included. The intrinsic rate of natural increase (r _m ) ranged from 0.172/day for the P. osmanthi albino strain to 0.209/day for P. citri, while the net reproductive rate (R ₀) varied from 23.98 in the thelytokous species P. thelytokus to 46.61 in P. citri. Both values were higher in the polyphagous species (P. ulmi, P. mori and P. citri), which are considered crop pests, than those in the oligophagous species (P. thelytokus and P. osmanthi), considered non-pests. The only exception was P. bambusicola, an oligophagous non-pest species with R ₀- and r _m -values closely resembling those of the three polyphagous species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Calcineurin Plays an Important Role in the Shell Formation of Pearl Oyster (Pinctada fucata)

Calcineurin (CN) is a multifunctional protein involved in many important physiological processes in mammalians, but the function of CN in mollusks is still largely unknown. In the present study, through the shell regeneration system, the changes of enzymatic activity of CN were determined in the process of shell regeneration in pearl oyster Pinctada fucata. CN was activated immediately and continuously in the shell regeneration process. The speed of shell regeneration was measured and the ultrastructure of inner shell surface was observed by scanning electron microscopy after inhibiting CN by intramuscular injection of immunosuppresant cyclosporine A (CsA). The results showed that the speed of shell regeneration was delayed and the morphology of calcite and aragonite in the inner shell surface became abnormal when CN was inhibited by CsA. Meanwhile, RT-PCR analysis revealed that the expression of P. fucata BMP-2 in mantle tissue decreased with CsA injection. In vitro secretion level of proteoglycans (PGs) in primary cultures of mantle cells was also decreased when mantle cells were exposed to CsA. Taken together, our results, for the first time, show that CN is involved in the shell formation through regulating the expression of Pf-BMP-2 in mantle tissue, which controls the secretion of PGs/GAGs of the mantle epithelial cells.     

Structure and histochemistry of mycorrhizae synthesized between Arbutus menziesii (Ericaceae) and two basidiomycetes,Pisolithus tinctorius (Pisolithaceae) and Piloderma bicolor (Corticiaceae)

Arbutoid mycorrhizae were synthesized in growth pouches between Arbutus menziesii Pursch. (Pacific madrone) and two broad host range basidiomycete fungi, Pisolithus tinctorius (Pers.) Coker and Couch and Piloderma bicolor (Peck) Jülich. P. tinctorius induced the formation of dense, pinnate mycorrhizal root clusters enveloped by a thick fungal mantle. P. bicolor mycorrhizae were usually unbranched, and had a thin or non-existent mantle. Both associations had the well-developed para-epidermal Hartig nets and intracellular penetration of host epidermal cells by hyphae typical of arbutoid interactions. A. menziesii roots developed a suberized exodermis which acted as a barrier to cortical cell penetration by the fungi. Ultrastructurally, the suberin appeared non-lamellar, but this may have been due to the imbedding resin. Histochemical analyses indicated that phenolic substances present in epidermal cells may be an important factor in mycorrhiza establishment. Analyses with X-ray energy dispersive spectroscopy showed that some of the granular inclusions present in fungal hyphae of the mantle and Hartig net were polyphosphate. Other inclusions were either protein or polysaccharides.     

Sequestration and turnover of plant- and microbially derived sugars in a temperate grassland soil during 7 years exposed to elevated atmospheric pCO2

Temperate grasslands contribute about 20% to the global terrestrial carbon (C) budget with sugars contributing 10–50% to this soil C pool. Whether the observed increase of the atmospheric CO₂ concentration (pCO₂) leads to additional C sequestration into these ecosystems or enhanced mineralization of soil organic matter (SOM) is still unclear. Therefore, the aim of the presented study was to investigate the impact of elevated atmospheric pCO₂ on C sequestration and turnover of plant‐ (arabinose and xylose) and microbially derived (fucose, rhamnose, galactose, mannose) sugars in soil, representing a labile SOM pool. The study was carried out at the Swiss Free Air Carbon Dioxide Enrichment (FACE) experiment near Zurich. For 7 years, Lolium perenne swards were exposed to ambient and elevated pCO₂ (36 and 60 Pa, respectively). The additional CO₂ in the FACE plots was depleted in ¹³C compared with ambient plots, so that ‘new’ (<7 years) C inputs could be determined by means of compound‐specific stable isotope analysis (¹³C : ¹²C). Samples were fractionated into clay, silt, fine sand and coarse sand, which yielded relatively stable and labile SOM pools with different turnover rates. Total sugar sequestration into bulk soil after 7 years of exposure to elevated pCO₂ was about 28% compared with the control plots. In both ambient and elevated plots, total sugar concentrations in particle size fractions increased in the order sand2 for coarse sand, fine sand and silt (about 274%, 17% and 96%, respectively) but about 14% lower for clay compared with the control plots, corroborating that sugars belong to the labile SOM pool. The fraction of newly produced sugars gradually increased by up to 50% in bulk soil samples after 7 years under elevated pCO₂. In the ambient plots, sugars were enriched in ¹³C by up to 10‰ when compared with bulk soil samples from the same plots. The enrichment of ¹³C in plant‐derived sugars was up to 13.4‰ when compared with parent plant material. After 7 years, the δ¹³C values of individual sugars decreased under elevated (¹³C‐depleted) CO₂ in bulk soil and particle size fractions, varying between −13.7‰ and −37.8‰ under elevated pCO₂. In coarse and fine sand, silt and clay fractions newly produced sugars made up 106%, 63%, 60% and 45%, respectively, of the total sugars present after 7 years. Mean residence time (MRT) of the sugars were calculated according to two models revealing a few decades, mean values increasing in the order coarse sand2 led to a net sequestration of about 30% of labile SOM (sugars) while no increase of total organic C was observed at the same plots. The additional labile SOM is gradually incorporated into more stable SOM pools such as silt and clay fractions in the medium term (<7 years). MRT of labile (sugar) SOM under elevated pCO₂ is in the same order of magnitude when compared with studies under ambient pCO₂ though no direct comparison of elevated and ambient plots was possible.     

Early Triassic vertebrate burrows from the Katberg Formation of the south‐western Karoo Basin,South Africa

Bordy, E.M., Sztanó, O., Rubidge, B.S. & Bumby, A. 2010: Early Triassic vertebrate burrows from the Katberg Formation of the south‐western Karoo Basin, South Africa. Lethaia, Vol. 44, pp. 33–45. Very large (～30–35 cm), uniform diameter cylindrical burrows were found at two localities, ～100–110 m above Permo‐Triassic boundary in the fluvial Katberg Formation (main Karoo Basin, South Africa). Analysis of their morphology and stratigraphical distribution allows us to improve both the understanding of the ethology of burrowing, and also the reconstruction of the earliest Triassic ecosystems. These burrows have a single opening that leads, via a large, uniform diameter, semi‐horizontal tunnel, to a rounded terminus. These 3‐m‐long structures descend at angles of ～30° to a maximum of 1.5 m depth. They are devoid of chambers, branching, cross‐cutting, coiling or spiralling. Filled with coarse sediments, some have a <5‐mm clay lining, and most have subtle indentations and various scratch marks. These burrows were possibly excavated as resting, hiding or aestivating shelters, and are tentatively attributed to dicynodonts (i.e. Lystrosaurus murrayi and L. declivis). Data suggest that burrowing was widespread after the P/Tr boundary event, when in this part of Gondwana, dryland fluvial systems had large fluctuations in flow with extended low‐flow periods or drought punctuated by high‐discharge periods. We hypothesize that these constructed refuges played a role in the biodiversity recovery and maintenance in the Early Triassic (Induan) ecosystem. □Early Triassic, Karoo Basin, Katberg Formation, South Africa, Tetrapod Burrows.     

A new basal ornithopod dinosaur (Frenchman Formation,Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America

A small, articulated basal ornithopod skeleton from the Frenchman Formation (late Maastrichtian) of Saskatchewan (RSM P 1225.1), previously referred to the taxon Thescelosaurus, differs from both recognized species of this taxon (Thescelosaurus neglectus and Thescelosaurus garbanii). The differences are taxonomically informative and we recognize this specimen as the holotype of a new species, Thescelosaurus assiniboiensis sp. nov. , diagnosed by the presence of two autapomorphies, and displaying plesiomorphic traits more similar to those of Parksosaurus, than to those of the other Thescelosaurus species. The Frenchman Formation also harbours an intriguing faunal assemblage in which Thescelosaurus represents one of the most abundant dinosaur taxa, and preserves a relatively high proportion of small (putatively juvenile and subadult) specimens of many dinosaur taxa. Further work that increases the faunal sample from this formation, and that permits quantitative comparisons with contemporary formations, will determine whether or not these differences are well supported, and will determine their ultimate palaeobiological significance. Identification of a third species of Thescelosaurus from the late Maastrichtian of North America suggests that this taxon was more diverse than previously recognized, and shows an increase in diversity from the Campanian through the late Maastrichtian, contrasting the trends seen in most other ornithischian clades. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1157–1198.     

Facies of a Lower Ordovician carbonate shelf (Mungok Formation: Taebaeksan Basin,Korea)

Summary The lithologic associations within the Lower Ordovician Mungok Formation in Korea define four depositional facies that formed across a continental margin fringing the Sino-Korean block: these facies represent lagoonal/restricted marine, shoal, inner shelf, and outer shelf environments. The stacking pattern of these facies reveals two systems tracts composed of five depositional sequences. The lower highstand systems tract consists of the lagoonal/restricted marine and shoal facies, whereas the upper lowstand systems tract comprises, in ascending order, inner shelf, outer shelf, and inner shelf facies. Three trilobite biofacies are recognized in the Mungok Formation: i.e.,Yosimuraspis, Kainella, andShumardia biofacies in ascending order. TheYosimuraspis Biofacies is dominated byYosimuraspis but also containsJujuyaspis andElkanaspis. The predominance of the endemic eponymous taxon suggests a lagoonal/restricted marine environment. The nearly monotaxicKainella Biofacies, which comprises pandemic genera such asKainella and occasionallyLeiostegium, may represent a less restricted environment than theYosimuraspis Biofacies. TheShumardia Biofacies occurs in the marlstone/shale lithofacies through relatively thick stratigraphic interval and is dominated by cosmopolitan trilobite taxa with some endemic species. The lithofacies and cosmopolitan trilobite assemblage of theShumardia Biofacies indicate that it occupied an outer shelf environment. The vertical succession of lithofacies and trilobite biofacies in the Mungok Formation records in general a shift from a restricted, shallow water environment to deeper-water environment.     

The short-term effects of fresh kelp (Macrocystis integrifolia) on physical properties of a fine-textured soil

In a two year field investigation fresh kelp (M. integrifolia) was broadcast and incorporated into plots of silty clay loam soil in the lower Fraser Valley of British Columbia. The effects of this amendment on bulk density, particle density, total porosity and aeration (volume of pores occupied by air after a saturated core was allowed to freely drain for 12 h at a soil water potential of −0.60 m) were measured. Soil aeration was increased in the first year with 30 and 60 t ha^-1 kelp application but decreased with the 120 t ha^-1 application. This soil aeration effect did not persist into the second year. Soil bulk density, particle density and total porosity were not significantly affected.     

A new ichthyodectiform (Actinopterygii,Teleostei) from the Lower Cretaceous Marizal Formation,north‐east Brazil

Abstract: A new genus and species, Ogunichthys triangularis, from the Early Cretaceous Marizal Formation in the Tucano Basin, Bahia State, Brazil, can be assigned to the teleost order Ichthyodectiformes and to the suborder Ichthyodectoidei by the presence of ten synapomorphies of these clades. Ogunichthys gen. nov. cannot be placed into any of the currently recognized families within Ichthyodectoidei and differs from all other ichthyodectoid genera in the ratio between the depth of the dentary symphysis and the depth of the coronoid process, the relationships between the length of the coronoid process and the alveolar border of the dentary, and a supraoccipital crest with posterior border fringed. The new taxon occurs in Early Cretaceous (Aptian) strata of the Marizal Formation, which are related to periods during epicontinental seas extended from the Caribbean Tethys into the South American continent. Rather than representing a single continuous widespread seaway, these epicontinental seas reflect local tectonic events, sometimes including locally distinctive faunas.