Claudio Di Celma

Distribution of fossil marine vertebrates in Cerro Colorado, the type locality of the giant raptorial sperm whale Livyatan melvillei (Miocene, Pisco Formation, Peru)

Hundreds of fossil marine vertebrates cropping out at Cerro Colorado (Pisco Basin, Peru) are identified and reported on a 1:6500 scale geological map and in a joined stratigraphic section. All the fossils are from the lower strata of the Pisco Formation, dated in this area to the late middle or early late Miocene. They are particularly concentrated (88%) in the stratigraphic interval from 40 to 75 m above the unconformity with the underlying Chilcatay Formation. The impressive fossil assemblage includes more than 300 specimens preserved as bone elements belonging mostly to cetaceans (81%), represented by mysticetes (cetotheriids and balaenopteroids) and odontocetes (kentriodontid-like delphinidans, pontoporiids, ziphiids, and physeteroids, including the giant raptorial sperm whale Livyatan melvillei). Seals, crocodiles, sea turtles, seabirds, bony fish, and sharks are also reported. Isolated large teeth of Carcharocles and Cosmopolitodus are common throughout the investigated stratigraphical interval, whereas other shark teeth, mostly of carcharinids, are concentrated in one sandy interval. This work represents a first detailed census of the extraordinary paleontological heritage of the Pisco Basin and the basis for future taphonomic, paleoecological, and systematic studies, as well as a much needed conservation effort for this extremely rich paleontological site.

Basin physiography and tectonic influence on sequence architecture and stacking pattern: Pleistocene succession of the Canoa Basin (central Ecuador)

Facies, shell bed features, and sequence stratigraphic framework for the shallow-marine Pleistocene upper Canoa and Tablazo Formations are presented, based on outcrop data from the southern coast of Cabo San Lorenzo, Ecuador. Sediments of this succession exhibit a distinct cyclic pattern, consisting of a stack of eight depositional sequences (cyclothems) likely developed under the main control of orbitally induced sea-level changes. As a rule, within the studied interval an idealized cyclothem is composed of a transgressive systems tract (TST) and a highstand systems tract (HST), whereas deposits attributable to the lowstand and falling-stage systems tracts are not present. Transgressive lithosomes may be defined by estuarine deposits interposed between the sequence boundary and the ravinement surface (back-barrier wedge) and by upward fining shoreface to inner-shelf facies successions above the ravinement (backstepping shelf wedge). Separated by an expanded siliciclastic core, hiatal shell concentrations occur at the base (onlap shell beds) and the top (backlap shell beds) of the transgressive shelf wedges, and some occur at the base of highstand systems tracts (downlap shell beds). On the basis of sedimentary facies, geometry, taphonomy, and paleoecology of shell beds, and the nature of the transition between siliciclastic and mollusk-bearing sediments, cyclothems were classified into two main types that show dependence upon paleoshoreline morphological configuration: sheltered (in the upper Canoa Formation) and exposed (in the Tablazo Formation). Notwithstanding the different synsedimentary tectonic and climatic regimes, the Ecuadorian cyclothems share basic patterns of condensation and facies assemblages with other roughly coeval cyclothemic successions around the world. This suggests that (1) hiatal shell bed development is not just a temperate-latitude phenomenon; (2) a global process, such as glacio-eustatic sea-level change, is the primary mechanism of control for the general architecture of sequences; and (3) specific paleogeographic settings play an important role by determining the taphonomic and paleoecologic characteristics of key shell beds, the nature of their contacts with the encasing sediments, and the type of the component set of facies. At a multicycle time scale, tectonics influenced the long-term trend of the relative sea-level changes and consequently the large-scale stratigraphic organization. Owing to the continued tectonic uplift of the area, successive high-frequency depositional sequences are nested to form a longer-order falling-stage sequence set.

No deep diving: evidence of predation on epipelagic fish for a stem beaked whale from the Late Miocene of Peru.

Although modern beaked whales (Ziphiidae) are known to be highly specialized toothed whales that predominantly feed at great depths upon benthic and benthopelagic prey, only limited palaeontological data document this major ecological shift. We report on a ziphiid–fish assemblage from the Late Miocene of Peru that we interpret as the first direct evidence of a predator–prey relationship between a ziphiid and epipelagic fish. Preserved in a dolomite concretion, a skeleton of the stem ziphiid Messapicetus gregarius was discovered together with numerous skeletons of a clupeiform fish closely related to the epipelagic extant Pacific sardine (Sardinops sagax). Based on the position of fish individuals along the head and chest regions of the ziphiid, the lack of digestion marks on fish remains and the homogeneous size of individuals, we propose that this assemblage results from the death of the whale (possibly via toxin poisoning) shortly after the capture of prey from a single school. Together with morphological data and the frequent discovery of fossil crown ziphiids in deep-sea deposits, this exceptional record supports the hypothesis that only more derived ziphiids were regular deep divers and that the extinction of epipelagic forms may coincide with the radiation of true dolphins.

Piscivory in a Miocene Cetotheriidae of Peru: first record of fossilized stomach content for an extinct baleen-bearing whale

Instead of teeth, modern mysticetes bear hair-fringed keratinous baleen plates that permit various bulk-filtering predation techniques (from subsurface skimming to lateral benthic suction and engulfment) devoted to various target prey (from small invertebrates to schooling fish). Current knowledge about the feeding ecology of extant cetaceans is revealed by stomach content analyses and observations of behavior. Unfortunately, no fossil stomach contents of ancient mysticetes have been described so far; the investigation of the diet of fossil baleen whales, including the Neogene family Cetotheriidae, remains thus largely speculative. We report on an aggregate of fossil fish remains found within a mysticete skeleton belonging to an undescribed late Miocene (Tortonian) cetotheriid from the Pisco Formation (Peru). Micro-computed tomography allowed us to interpret it as the fossilized content of the forestomach of the host whale and to identify the prey as belonging to the extant clupeiform genus Sardinops. Our discovery represents the first direct evidence of piscivory in an ancient edentulous mysticete. Since among modern mysticetes only Balaenopteridae are known to ordinarily consume fish, this fossil record may indicate that part of the cetotheriids experimented some degree of balaenopterid-like engulfment feeding. Moreover, this report corresponds to one of the geologically oldest records of Sardinops worldwide, occurring near the Tortonian peak of oceanic primary productivity and cooling phase. Therefore, our discovery evokes a link between the rise of Cetotheriidae; the setup of modern coastal upwelling systems; and the radiation of epipelagic, small-sized, schooling clupeiform fish in such highly productive environments.

Fossil marine vertebrates of Cerro Los Quesos: Distribution of cetaceans, seals, crocodiles, seabirds, sharks, and bony fish in a late Miocene locality of the Pisco Basin, Peru

ABSTRACT One-hundred and ninety-two fossil marine vertebrate specimens, preserved as bone elements cropping out at Cerro Los Quesos (Pisco Basin, Peru), are identified and reported on a 1:4,000 scale geological map and in the corresponding stratigraphic section. All the fossils originate from the Pisco Formation, which is dated in this area to the late Miocene (from 7.55 Ma to ≥6.71 Ma, based on 40Ar/39Ar analyses of three volcanic ash layers along the section). Specimens are particularly concentrated near the top of the two main hills, where the geologically youngest portion of the examined section crops out. The impressive fossil assemblage includes cetaceans (91.6%), represented by mysticetes (balaenopteroids and cetotheriids) and odontocetes (phocoenids, physeteroids, and ziphiids, including the holotype of Nazcacetus urbinai). Seals, a crocodile, a seabird, bony fish, and sharks are also reported. Isolated large teeth of Carcharocles and Cosmopolitodus are common and, in several instances, associated to mysticete skeletons. Together with a similar work recently published for the other late Miocene locality of Cerro Colorado, this work represents a case study for the detailed inventory of the extraordinary paleontological heritage of the Pisco Basin. As such, it constitutes the basis for future taphonomic, paleoecological, and systematic studies, as well as for a much-needed conservation effort.

Inside baleen: Exceptional microstructure preservation in a late Miocene whale skeleton from Peru

Exceptionally preserved delicate baleen microstructures have been found in association with the skeleton of a late Miocene balaenopteroid whale in a dolomite concretion of the Pisco Formation, Peru. Microanalytical data (scanning electron microscopy, electron probe microanalysis, X-ray diffraction) on fossil baleen are provided and the results are discussed in terms of their taphonomic and paleoecological implications. Baleen fossilization modes at this site include molding of plates and tubules, and phosphatization. A rapid formation of the concretion was fundamental for fossilization. We suggest that the whale foundered in a soft sediment chemically favorable to rapid dolomite precipitation, allowing the preservation of delicate structures. Morphometric considerations on the baleen plates and bristles coupled with the reconstructed calcification of the latter permit speculation on the trophic preferences of this balaenopteroid whale: the densely spaced plates and the fine and calcified bristles provide evidence for feeding on small-sized plankton, as does the modern sei whale.

Sedimentologic and diagenetic controls on pore-network characteristics of Oligocene–Miocene ramp carbonates (Majella Mountain, central Italy)

This article addresses the controls exerted by sedimentologic and diagenetic factors on the preservation and modification of pore-network characteristics (porosity, pore types, sizes, shapes, and distribution) of carbonates belonging to the Bolognano Formation. This formation, exposed at the Majella Mountain, Italy, is composed of Oligocene–Miocene carbonates deposited in middle- to outer-ramp settings. The carbonates consist of (1) grainstones predominantly composed of either larger benthic foraminifera, especially Lepidocyclina, or bryozoans; (2) grainstones to packstones with abundant echinoid plates and spines; and (3) marly wackestones to mudstones with planktonic foraminifera. The results of this field- and laboratory-based study are consistent with skeletal grain assemblages, grain sizes, sorting, and shapes, all representing the sedimentologic factors responsible for high values of connected primary macroporosity in grainstones deposited on the high-energy, middle to proximal outer ramp. Cementation, responsible for porosity reduction and overall macropore shape and distribution in grainstones to packstones deposited on the intermediate outer ramp, was mainly dependent on the following factors: (1) amount of echinoid plates and spines, (2) grain size, (3) grain sorting and shapes, and (4) clay amount. Differently, in the wackestones to mudstones, laid down on the low-energy, distal outer ramp, matrix is the key sedimentologic factor responsible for low values of scattered macroporosity and dominance of microporosity. The aforementioned results may be useful to improve the prediction of reservoir quality by means of mapping, simulating, and assessing individual carbonate facies with peculiar pore-network characteristics.

Sedimentology and high-resolution sequence stratigraphy of the late middle to late Miocene Angostura Formation (western Borbón Basin, northwestern Ecuador)

An integrated analysis of facies and palaeontological content of the middle to late Miocene (c. 12.4–9.1 Ma) Angostura Formation led to the identification of facies associations indicative of shoreface and inner-shelf settings. The Angostura Formation comprises eight high-frequency sequences that are stacked to form a tectonically driven lowstand sequence set. The most complete examples of sequences are bounded by transgressively modified unconformities and exhibit a threefold subdivision: (1) a basal suite of deepening-upward shoreface sediments (transgressive systems tract), including a base-of-cycle shell concentration; (2) a mid-cycle shell bed, consisting of molluscan shells dispersed in a matrix of inner-shelf muddy fine-grained sandstones; (3) a shallowing-upward unit of inner-shelf to lower shoreface sediments almost barren of mollusc fossils (highstand systems tract). Biostratigraphic constraints allowed a reasonable correlation between sequence bounding unconformities and the late middle to late Miocene high-frequency glacio-eustatic changes derived from recent δ18O studies. This correlation has far-reaching implications and leads to the following conclusions: (1) glacio-eustasy in tune with oxygen isotope changes at fourth-order frequency (200 ka–1 Ma duration) may have been the principal factor regulating stratigraphic packaging in the Angostura Formation; (2) these sequences provide an excellent shallow-marine outcrop record of late middle to late Miocene Antarctic glaciations.

TAPHONOMIC AND PALEOECOLOGICAL ANALYSES (MOLLUSKS AND FISHES) OF THE SÚA MEMBER CONDENSED SHELLBED, UPPER ONZOLE FORMATION (EARLY PLIOCENE, ECUADOR)

Abstract The early Pliocene sediments of the Súa Member (Onzole Formation) are spectacularly exposed along coastal cliffs in the surroundings of Súa (northwestern Ecuador). The shellbed at the base of these strata contains rich mollusk and fish assemblages and provides a rare opportunity to document the virtually unknown Pliocene shallow-water faunas of Ecuador. Stratigraphic context, faunal composition (both mollusks and fishes), biofabric trend, ichnological signatures, and taphonomic features presented here, all indicate that the shellbed is a stratigraphically condensed (hiatal) skeletal concentration generated primarily by low rates of net background sedimentation. Shellbeds of this type, immediately overlying a Glossifungites-demarcated ravinement surface, are regarded as onlap shellbeds and record landward stratal convergence and attenuation during the earliest phases of marine transgression. Although composed of largely autochthonous-parautochthonous specimens, the fossil assemblage contains mollusk species from a range of water depths and both soft- and hard-bottom habitats, implying in situ time averaging and admixture of different assemblages as transgression proceeded on a sediment-starved shoreline. The mollusk assemblage is composed of 54 taxa (26 bivalves, 26 gastropods, 2 scaphopods), and the fish assemblage of 31 taxa belonging to 16 families. Both are indicative of a well-oxygenated marine biotope swept by currents and waves, and devoid of a benthic macrophyte cover. With the exception of a few fish species and a single gastropod, the largest number of the mollusk and fish taxa are presently distributed throughout the Tropical Eastern Pacific Biogeographic Region.

SEQUENCE STRATIGRAPHY AND PALEONTOLOGY OF THE UPPER MIOCENE PISCO FORMATION ALONG THE WESTERN SIDE OF THE LOWER ICA VALLEY (ICA DESERT, PERU)

The sequence stratigraphic framework and a summary of the fossil fauna of the upper Miocene portion of the Pisco Formation exposed along the western side of the Ica River (southern Peru) is presented through a new geological map encompassing an area of about 200 km 2 and detailed chronostratigraphic analyses. Extensive field mapping and sedimentological study of outcrop sections have shown that the Pisco Formation is a cyclical sediment unit composed of at least three fining-upward, unconformity-bounded depositional sequences, designated P0, P1, and P2 from oldest to youngest. In the study area, these sequences progressively onlap a composite basal unconformity from southwest to northeast. Integration of biostratigraphic and tephrochronologic age determinations constrains the ages of the three Pisco sequences within the study area. Based on the age of surrounding sediments, a conservative estimate of the age of P0 suggests deposition of these strata between 17.99 ± 0.10 Ma and 9.00 ± 0.02 Ma, whereas diatom biostratigraphy and calculated 40 Ar/ 39 Ar ages converge to indicate that strata of the P1 sequence were deposited sometime between 9.5 Ma and 8.9 Ma and that those of the P2 sequence are younger than 8.5 Ma and older than 6.71 ± 0.02 Ma. Our survey for both vertebrate and macro-invertebrate remains in the three sequences confirms the outstanding paleontological value of the Pisco Formation and contributes to depict regional faunal shifts in the fossil assemblage.