Mass mortality of an asteriid starfish (Forcipulatida,Asteroidea, Echinodermata) from the late Maastrichtian (Late Cretaceous) of Morocco

Abstract: A mass occurrence, numbering tens of thousands of individuals in a single lens, belonging to a single species of asteroid, is described from the late Maastrichtian (Late Cetaceous) of Morocco. The lens of partially silicified asteroidal limestone is made up largely of fully articulated specimens of similar size and probably represents the mass mortality of a single recruitment. By comparison with mass strandings of the present‐day species Asterias rubens (Linnaeus), it can be inferred that a feeding swarm of individuals was swept into a submarine channel by either a storm or an exceptionally strong tidal current, and permanently buried. The genus and species are herein described as Cretasterias reticulatus gen. et sp. nov. The exceptional preservation of the material enables the identification of wreath organs (clusters of crossed pedicellariae set in a dermal pad around spines) for the first time in the fossil record. Comparison between extant Asteriidae, putative fossil asteriids and C. reticulatus provides ambiguous evidence of its affinities; it appears to display a combination of plesiomorphic and derived characters. It is shown that all Mesozoic forcipulatid asteroids described so far share a very simple arm construction (single row of adradial ossicles) unknown in adult extant Forcipulatida.     

First evidence for the early Aptian Oceanic Anoxic Event (OAE1a) from the Western margin of the Pindos Ocean (NW Greece)

In this paper, new biostratigraphic, stable isotope (C, O) and organic geochemical data are presented for the Pindos Zone in NW Greece (SE Epirus region) in order to investigate whether the organic carbon rich strata of the Kalarrytes sections A and B correspond to a local expression of the early Aptian Oceanic Anoxic Event (OAE 1a or Selli Event) or not. The Pindos Zone Mesozoic to Tertiary sedimentary sequence constitutes the deep-sea sedimentary cover of the Pindos Ocean, which was separated from its oceanic basement as an accretionary prism during the complete closure of this ocean, and was emplaced westwards onto the adjacent Gavrovo-Tripolis carbonate platform. Stable carbon and oxygen isotope data from the Kalarrytes sections reveal an isotopic composition compatible with the characteristic features of the OAE 1a. Calcareous nannofossil and radiolarian biostratigraphy indicates an early Aptian age for both sequences. Biomarker analysis on the organic-rich intervals reveals the primary marine origin of the organic matter, with substantial contribution from bacteria, cyanobacteria and dinoflagellates, as well as a significant terrigenous input. Findings of authigenic framboidal pyrite provide evidence for the prevalence of sulphidic conditions during deposition. Furthermore, similarities between the biomarker signatures of the Pindos organic-rich strata and coeval strata of early Aptian age where the impact of OAE 1a has previously been recorded, are identified. The presented data show that the organic-rich intervals of both Kalarrytes sections constitute the first records of the OAE 1a in oceanic deposits of Greece.     

Comptoniaster adamsi nov. sp. (Echinodermata, Asteroidea) from the middle Cretaceous of Texas and its phylogenetic position

Comptoniaster adamsi nov. sp. (Asteroidea, Valvatida, Goniasteridae) is described from the middle Cretaceous (Cenomanian-Turonian) Britton Formation of northcentral Texas, USA. The new species provides the focus for an exploratory cladistic analysis of Mesozoic asteroids of the Comptoniaster-Tylasteria Group sensu G. Breton. The systematics of Mesozoic goniasterids has relied heavily on the morphology of marginal ossicles, which generally are the best-preserved elements of the skeleton. Unfortunately, marginal ossicular data are scanty for most species because ossicles tend to be morphologically simple yet varied even within individuals, and few even partially articulated specimens are available to provide more comprehensive information. Further, both plesiomorphy and homeomorphy have been important. Because of limited available data, phylogenetic reconstruction here is preliminary. Nevertheless, a number of taxon groupings recognized in the literature are recovered, and stratigraphic distribution provides some support for results. Data unfortunately are particularly incomplete for species of Comptoniaster, the focus of the study. Comptoniaster adamsi nov. sp. clustered with three other species assigned to the genus, including the type, all Cretaceous in age. Jurassic species of Comptoniaster are more widely distributed in the analysis, perhaps reflecting an early stage in diversification or perhaps suggesting the need for species assignment reassessment, but also reflecting the limited available data.     

Ultrastructure of the gametogenesis of the common Mediterranean starfish,Echinaster (Echinaster) sepositus

Echinaster (Echinaster) sepositus is one of the most abundant sea stars in western Mediterranean rocky bottoms, yet its reproductive biology remains virtually unknown. Here we report the ultrastructure of its gametogenesis over 2 consecutive years. It is a gonochoric species with an annual reproductive cycle spawning gametes in late summer and early autumn. Each arm of every individual contained two gonads (dark red in females and yellow in males). In both sexes, the gonad was a single, large sac composed of several smaller sacs. The gonad wall consisted of two multilayered sacs, outer and inner, separated by the genital haemal sinus. The histology of the gonad wall was consistent with that found in other asteroids. Oogenesis was continuous during the year, but eggs were spawned only in late summer. Oocytes were in close relationship with follicular cells that are suggested to transfer nutrients to the oocytes. Spermatogenesis was restricted to 5–6 months in spring-summer. It occurred in columns with an axial interstitial cell supporting each column, and producing processes towards the lumen that remained connected to spermatogenic cells by intercellular junctions. Developing sperm cells were found along the length of the column, while spermatozoa were found free in the testis lumen. Spermatogenesis followed the pattern described for echinoderms, to give rise to an acrosome-bearing, round-shaped spermatozoon. The histology and cytology of the reproductive process in E. sepositus followed the general pattern found for asteroids.     

Toarcian carbon isotope shifts and nutrient changes from the Northern margin of Gondwana (High Atlas, Morocco, Jurassic): Palaeoenvironmental implications

The Early Toarcian is marked by a global perturbation of the carbon cycle and major marine biological changes. These coincide with a general decrease in calcium carbonate production and an increase in organic carbon burial, and culminate in the so-called Toarcian Oceanic Anoxic Event. It is believed that the environmental crisis was triggered by the activity of the Karoo-Ferrar large igneous province. In order to further document the Early Toarcian palaeoenvironmental perturbations, carbon isotope, total organic matter, calcareous nannofossils and phosphorus content of the Amellago section in the High Atlas rift basin of Morocco were investigated. This section is extremely expanded compared to the well-studied European sections. Its position along the northern margin of the Gondwana continent is of critical importance because it enables an assessment of changes of river nutrient input into the western Tethyan realm. The carbon isotope curve shows two negative excursions of equal thickness and amplitude, at the Pliensbachian-Toarcian boundary and at the transition from the Polymorphum to the Levisoni Zone. This confirms the supra-regional nature of these shifts and highlights the possible condensation of the first “boundary” shift in European sections. Phosphorus content is used to trace palaeo-nutrient changes and shows that the two negative carbon isotope shifts are associated with increased nutrient levels, confirming that these episodes are related to enhanced continental weathering, probably due to elevated greenhouse gases in the atmosphere. In the High Atlas Basin, the increase in nutrient levels at the Pliensbachian-Toarcian boundary is moreover likely to be the main factor responsible for the coeval demise of the Saharan carbonate platform. A middle Toarcian event, centered on the boundary between the Bifrons and Gradata Zones, characterized by a positive carbon isotope excursion and nutrient level rise, is documented in the Amellago section.     

Paleoenvironmental changes across the Cenomanian/Turonian Boundary Event (Oceanic Anoxic Event 2) as indicated by benthic foraminifera from the Demerara Rise (ODP Leg 207)

This study is based on Cenomanian to lower Turonian sediments of Ocean Drilling Program (ODP) Sites 1258, 1259, 1260, and 1261 from the Demerara Rise (Leg 207, western tropical Atlantic, off Suriname) that are oriented along a paleodepth transect. Studied sediments include the Cenomanian/Turonian Boundary Event (CTBE) or Oceanic Anoxic Event 2 (OAE 2) and consist of laminated black shales with TOC values between 5% and 10% below and above OAE 2 and up to 29% within the OAE 2 interval. Benthic foraminiferal assemblages in this eutrophic environment are generally characterized by low diversities and strong fluctuations of abundances, indicating oxygen depletion and high organic matter fluxes. Dominant taxa at all sites are Bolivina anambra, Gavelinella dakotensis, Tappanina sp., Praebulimina prolixa, and Neobulimina albertensis. Based on the positive stable carbon isotope excursion characteristics of OAE 2 we subdivided the studied successions into three intervals: (1) the interval below OAE 2; (2) the carbon isotope excursion reflecting OAE 2; and (3) the interval above OAE 2. In the bathymetrically shallower Sites 1260 and 1261 benthic foraminiferal assemblages indicate anoxic to sometimes slightly dysoxic bottom-water conditions below the OAE 2 interval. The bathymetrically deepest Site 1258, in contrast, reflects more oxygenated bottom waters with an almost continuous occurrence of benthic foraminifera. It is therefore suggested that the shallower sites were located within the oxygen minimum zone (OMZ), whereas Site 1258 was below the OMZ. During OAE 2 anoxic conditions prevailed at the shallower sites. At Site 1258 benthic foraminifera indicate severe dysoxic but not anoxic conditions. This pattern is proposed to reflect a strengthening of the OMZ possibly related to increased primary production during OAE 2. A short-term repopulation event of benthic foraminifera in the lower third of the OAE 2 interval was observed at all sites, reflecting a brief bottom-water oxygenation event. This short-lived event parallels a surface-water cooling and is probably equivalent to the “Plenus Cool Event” in Europe and the “benthonic zone” in the U.S. Western Interior. The benthic foraminifera of a ~0.5 Ma interval following OAE 2 still indicate oxygen depletion of bottom waters. Subsequently, however, a strong increase in benthic foraminiferal abundance and diversity reflects better oxygenation of the bottom-water masses, probably related to a weakening of the OMZ due to decreasing organic matter flux.     

Larger foraminifera as a substratum for encrusting bryozoans (Late Oligocene,Tethyan Seaway,Iran)

Considering the diversity and abundance of larger foraminifera examined from a wide range of Late Oligocene to Early Miocene palaeoenvironments in the Tethyan Seaway, encrusting bryozoans make extremely little use of their tests as substratum. Significant encrustations by bryozoans were exclusively found on large (ø c. 6 cm), undulating tests of Lepidocyclina spp., on which, however, a remarkable 34 taxa of encrusting bryozoans were recorded. This shallow-water fauna of Chattian age was analyzed in respect of the bryozoan taxa present, colony growth type, and mode of budding, colony size, as well as onset of reproduction. Taxic and morphological similarities between the fossil assemblage and modern faunas encrusting mobile substrata indicate a long history of bryozoans as part of the interstitial habitat, while the tests of certain larger foraminifera may have played a significant role in the evolution of shallow-water bryozoans by providing substrata for encrusting species in otherwise unfavorable environments.     

First record of Mugil capurrii (Mugilidae, Perciformes) in the Gulf of Guinea

Leaping African mullet Mugil capurrii was caught along the Togolese coast in the Gulf of Guinea. This is the first record of this species which usually occurs from Morocco to Guinea Bissau and the southernmost point of its known distribution.     

Green algae from the Lower Cretaceous carbonate of northern Shiraz (Zagros Mountains,SW Iran)

In northern Shiraz (SW Iran), Lower Cretaceous carbonate was studied in detail. In this study, nine species of dasycladacea algae were classified. There are different species of dasycladacea algae which belong to seven different genera: Actinoporella, Cylindroporella, Dissocladela, Heteroporella, Neomeris, Salpingoporella, Trinocladus; one species of udoteaceae belongs to Bouenia; one species of acetabulariaceae belongs to Clypeina and the microproblematicum Coptocampylodon was also seen. Among the green algae, dasycladaceae and acetabulariceae are the most frequent and udoteaceans are rare in Zagros Mountains. The genus of Trinocladus is a new record for Lower Cretaceous (Upper Albian) in SW Iran.     

First giraffid skulls (Bohlinia attica) from the late Miocene Maragheh fauna,Northwest Iran

An almost complete skull and a second partial skull of Bohlinia attica (Artiodactyla: Giraffidae) from the late Miocene of Maragheh in northwestern Iran is described along with a complete upper dentition from Samos in Greece. These specimens enrich what is known of this species. The skull has massive bent ossicones. The braincase is horizontal to the face in lateral view. The upper premolars have strongly curved styles. The metapodials of this species are elongate with a deep plantar trough. The braincase is short and the occipital does not extend caudally. The type material of Bohlinia is from Pikermi but this taxon is also known from localities in F.Y.R.O. Macedonia, Greece, Turkey, Bulgaria, Iraq, and Iran. The new specimens are similar to others except that the ossicone terminates in a knob and the palatine choanae are positioned caudally. This taxon along with Honanotherium, which is most similar but with shorter metapodials and simpler premolars, can be placed in the subfamily Bohlininae.     

Pollen morphology of the genus Gagea (Liliaceae) in Iran

Pollen grains of 26 species of the genus Gagea distributed in Iran were examined by light and scanning electron microscopy. Detailed pollen morphological characteristics are given for these species. Among the studied species, the newly described G. iranica together with G. olgae and G. graminifolia possess the smallest pollen grains, and the widely distributed G. lutea the largest ones. Our studies show that the sculpturing of exine provides valuable characters for separating the species, sometimes even for closely related ones, and delimitation of natural groups within the genus. The exine of the genus Gagea is in most cases perforated upon tectum or rarely tectate-columellate. The muri are solid or structured, compound, simpli-, dupli- or pluricolumellate. It seems that the structure of muri is very important in recognizing natural groups within the genus. Tectal perforations vary from <0.2 to 2.0 μm in diameter among the studied species. Regarding sculpturing of the exine in proximal face, four basic types of pollen grains can be distinguished: reticulate, microreticulate, foveolate and perforate. Within the reticulate type there is sufficient variation in exine structure at distal face to describe three subtypes: reticulate, microreticulate and perforate. A diagnostic key is given for all studied taxa based on palynomorphological characters. For a limited number of populations of selected Iranian species of Gagea, further aspects of pollen biology were studied. It seems that populations with ploidy levels other than diploidy, show a low percentage of pollen fertility. Moreover, the rate of pollen fertility is correlated with the manner of the reproduction in certain species.     

Track analysis of the marine palaeofauna from the Turonian (Late Cretaceous)

Aim  To analyse the worldwide distribution patterns of Turonian marine biotas using a panbiogeographical approach.
Location  Turonian localities of southern and north-eastern Brazil, Mexico, Canada, central Europe, England and Morocco.
Method  Panbiogeographical track analysis.
Results  Nine generalized tracks and six nodes were found. The generalized tracks comprise two vicariant track patterns (one northern and one mid-southern) across the Atlantic.
Main conclusions  The generalized tracks show clearly two separate marine biotas, which were associated with the proto-South Atlantic and the proto-North Atlantic oceans. These generalized tracks, as well as the two vicariant track clusters between the north and south Atlantic, are identified by vicariant relationships shared by most of the taxa analysed, and illustrate the final break up of the Gondwana and Laurasia supercontinents and the consequences of vicariant events for the biogeography of the Atlantic Ocean.     

Depositional environment of the Upper Cretaceous orbitolinid-rich microfacies in the Kuh-e Mazar anticline (Kerman Province,Central Iran)

This study is focused on the depositional model and paleoenvironmental distribution patterns of orbitolinids-rich microfacies in an Upper Cretaceous carbonate succession in the Kuh-e Mazar anticline in Kerman Province, Central Iran. Twelve microfacies indicating a tidal flat (including intertidal and supratidal) and an inner ramp (consisting of subtidal lagoon and orbitolinid–rudist barrier shoal) belonging to a shallow-water ramp-type carbonate platform were recognised. Generally, the orbitolinid associations in the carbonate ramp are indicative of shallow warm waters with normal salinity. The occurrence and abundance of thick or conical orbitolinids in both shoal and lagoon show that such associations were well adapted to different environmental conditions including different depth, substrate stability and water energy. The slightly discoidal, almost discoidal and discoidal forms were only present in the deeper depths of the environment including the lagoon with muddy and more stable substrates and lower water energy. However, the orbitolinids in the inner part of the carbonate ramp were scarce or almost absent in the deep inner to middle lagoon due to the deep restricted environmental conditions unfavourable for benthic marine life.     

Fruit anatomy of the genus Pimpinella L. (Apiaceae) in Iran

The genus Pimpinella L., with about 170–180 species in the world, is one of the largest genera of the family Apiaceae (Umbelliferae). Based on the Flora Iranica treatment, this genus has 25 species in the Iranian plateau, including 19 species in Iran, and six of those (P. tragioides, P. deverroides, P. pastinacifolia, P. anisactis, P. khorasanica and P. khayyamii) are endemic.     

First record of the Late Cretaceous giant bird Gargantuavis philoinos from the Iberian Peninsula

Gargantuavis philoinos is a large terrestrial bird, initially described from several bones (synsacrum, cervical vertebra, femora) discovered in the Late Cretaceous deposits of southern France. The synsacrum described here comes from the late Campanian (Late Cretaceous) site of Laño, in north-western Spain, and is similar in all respects to the other synsacra of Gargantuavis philoinos described from France. This is therefore the first specimen attributed to this species to be reported outside France. This find increases the diversity of the fossil assemblage from Laño and the geographical distribution of this bird, which existed on the Ibero-Armorican island, which included southern France and the Iberian Peninsula, during the Late Cretaceous.     

Taxonomy and paleobiogeographic implication of Glabrobournonia Morris and Skelton (Hippuritida,Radiolitidae) from the Late Cretaceous Yigeziya Formation,southwestern Tarim Basin

A re-examination of the specimens that were identified as Biradiolites minor Pojarkova from the late Campanian to early Maastrichtian middle member of the Yigeziya Formation of southwestern Tarim Basin reveals that they should be assigned to the genus Glabrobournonia Morris and Skelton. Glabrobournonia is a genus of radiolitids characterized by indented radial bands, salient ridges on the shell margins and absence of fine ribs on the surface of the right valve. Apart from southwestern Tarim Basin, Glabrobournonia minor (Pojarkova) has also been recorded from the late Campanian of Fergana and Alai basins. The central Asian, late Campanian to early Maastrichtian G. minor differs from the late Campanian to Maastrichtian, eastern Arabian type species Glabrobournonia arabica Morris and Skelton in the flat left valve and an additional fourth ridge on the junction of the dorsal and posterior sides of the right valve. Biradiolites ingens (Des Moulins) could be the direct ancestor of Glabrobournonia. The paleogeographic distribution of Glabrobournonia suggests that this genus dispersed to central Asia from the late Campanian time, becoming widely distributed in the eastern Tethyan region rather than endemic to eastern Arabia. Correspondingly, specimens belonging to Gyropleura yielded from the same bed as G. minor in southwestern Tarim Basin, are similar to the specimens which were attributed to the eastern Arabian Gyropleura sp.; Campanian to early Maastrichtian Osculigera specimens described from the Yigeziya Formation are comparable with those known from the Campanian–Maastrichtian of Iran, Afghanistan and eastern Arabia. The similarity of the rudist assemblages between central Asia and eastern Arabia suggests a faunal connection and affinity between the north and south margins of the eastern Tethyan realm during Campanian to early Maastrichtian times.     

First record of the Olive Ridley Turtle,Lepidochelys olivacea,in Iranian coastal waters (Testudines,Cheloniidae)

Abstract

Three specimens of the Olive Ridley Turtle (Lepidochelys olivacea) were collected in Guader (Gwatar) Bay in the Sea of Oman on 6 January, 1997. Two of them have been preserved for the Zoological Museum of Gorgan University. This is the first report of the Olive Ridley Turtle in Iranian coastal waters.     

Mosasaurus beaugei Arambourg, 1952 (Squamata, Mosasauridae) from the Late Cretaceous phosphates of Morocco

Mosasaurus beaugei Arambourg, 1952 was based on isolated teeth from the Maastrichtian phosphatic deposits of Morocco. The recent discovery of new material, including skull and mandibular remains, improves our knowledge of this species. M. beaugei shares the following synapomorphies with the genus Mosasaurus: large teeth bearing two prominent carinae and with asymmetrical labial and lingual surfaces, the labial one being flattened and strongly facetted and the lingual one being convex; premaxillae with a small pointed rostrum and dentary without rostrum; palatal elements closely united; coronoid with very large ventromedial process overlying the prearticular. M. beaugei is characterised by the following autapomorphies: 12-13 maxillary teeth; marginal teeth bearing 3-5 prisms on the labial surface and 8-9 on the lingual one; palatine with posterior border concave and perpendicular to the long axis of the skull; splenial visible laterally on half of the dentary ventral surface; coronoid with anterior wing well developed and bearing two notches. M. beaugei is only known to date in the Maastrichtian phosphates of Morocco.