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The Cenozoic succession of the East Pisco Basin preserves the sedimentary record of several episodes of deformation of the forearc crust along the Peruvian margin. The 1:50,000 scale geological map presented here encompasses an area of about 1,000 km2 lying astride the Ica River, and contributes to our understanding of the timing and mode of basin filling and deformation. Our novel two-fold megasequence framework provides a sound basis for establishing a first-order tectono-stratigraphic setting of the mid-Eocene–upper Miocene succession exposed in the study area. We interpret that the mid-Eocene to lower Oligocene succession studied in this work (megasequence P) was deposited in a single forearc basin, which was dissected into the present-day West and East Pisco basins by a fault-bounded basement high during the late Oligocene, and subsequently overlain by the Miocene fill of the East Pisco basin (megasequence N).

We report 130 vertebrate fossils preserved as bony elements and the co-occurring assemblage of fish teeth and spines from the lower strata of the Pisco Formation exposed along the western side of the lower Ica Valley (East Pisco Basin, Peru). Geological mapping at 1:10,000 scale reveals that all these fossils originate from the Langhian–Serravallian P0 allomember. In the study area, P0 is up to ∼40 m thick and features a sandy lower portion, reflecting shoreface deposition, that fines upwards into a package of offshore silts. Marine vertebrates only occur in the lower sandy layers and include whales, dolphins, reptiles, birds, and bony and cartilaginous fishes. The reconstructed paleoenvironment is consistent with a warm-water, marginal-marine setting with a strong connection to the open ocean. This work helps to elucidate the rich yet still poorly understood middle Miocene portions of the Pisco Formation, and highlights the need to conserve this outstanding Fossil-Lagerstätte.

Bioapatite of fossil bone and teeth is susceptible to alteration and ion exchange during burial and diagenesis, varying its Sr content through the geological time. Nevertheless, fossil shark teeth are a powerful proxy for both chronostratigraphic and paleoecological reconstructions, thanks to the presence of the enameloid, a hard outer layer consisting of resistant fluorapatite crystallites. Here, we analyze fossil shark teeth from the Miocene sediments of the Chilcatay Formation of the Pisco Basin (southwestern Peru) with the aim of dating poorly constrained strata in this region. (Ultra)structural and compositional analyses of fossil lamniform and carcharhiniform teeth are performed through macroscopical observations, optical microscopy and SEM-EDS for evaluating the preservation state of the collected teeth. Shark teeth display a compact and well preserved outer enameloid layer formed by highly ordered bundles of crystallites that is distinctly separated by a more porous and heterogeneous inner core of dentine featuring diagenetic artefacts and microborings. Compositional mapping highlights differences in the distribution of Ca, P, F, and S in the enameloid and dentine, and chemical results show a Sr content that is consistent with the range reported for extant shark teeth. The best preserved teeth were selected for Strontium Isotope Stratigraphy (SIS), measuring the 87Sr/86Sr values in the enameloid and obtaining numerical (absolute) age estimates. In the Ica River Valley, SIS dates the Chilcatay strata to the Burdigalian (between 19.1 and 18.1 Ma), in agreement with previous radiometric, isotopic and biostratigraphic ages obtained in the same region. At Media Luna, the Chilcatay strata are dated herein for the first time, resulting in a slightly older age of 21.8–20.1 Ma (late Aquitanian–early Burdigalian). These results strengthen the notion that the Sr-ratio of shark teeth can be successfully applied for obtaining reliable age estimates via SIS.

Based on mapping of laterally traceable stratigraphic discontinuities, we propose a high-resolution allostratigraphic scheme for one of the world’s foremost fossil marine vertebrate Lagerstätten: the lower Miocene strata of the Chilcatay Formation exposed along the Ica River near Zamaca, southern Peru. Measured sections combined with 1:10,000 scale mapping of a 24 km2 area provide an overview of the stratal architecture, as well as a general facies framework and interpretation of the various depositional settings. As a whole, the Chilcatay alloformation is bounded by the CE0.1 unconformity at the base and the PE0.0 unconformity at the top. An internal Chilcatay surface, termed CE0.2, splits the alloformation into two distinct allomembers (Ct1 and Ct2). The Ct1 allomember comprises three facies associations recording deposition in shoreface, offshore, and subaqueous delta settings. The Ct2 allomember comprises two facies associations, recording deposition in shoreface and offshore settings. Using these data, we place the rich marine vertebrate assemblage in a precise spatial and stratigraphic framework. The well-diversified vertebrate assemblage is dominated by cetaceans (mostly odontocetes) and sharks (mostly lamniforms and carcharhiniforms); rays, bony fish, and turtles are also present. Taxonomic novelties include the first records of baleen whales, platanistids, and eurhinodelphinids from the Chilcatay Formation.

The northward-flowing Humboldt Current hosts perpetually high levels of productivity along the western coast of South America. Here, we aim to elucidate the deep-time history of this globally important ecosystem based on a detailed palaeoecological analysis of the exceptionally preserved middle–upper Miocene vertebrate assemblages of the Pisco Formation of the East Pisco Basin, southern Peru. We summarise observations on hundreds of fossil whales, dolphins, seals, seabirds, turtles, crocodiles, sharks, rays, and bony fishes to reconstruct ecological relationships in the wake of the Middle Miocene Climatic Optimum, and the marked cooling that followed it. The lowermost, middle Miocene Pisco sequence (P0) and its vertebrate assemblage testify to a warm, semi-enclosed, near-shore palaeoenvironment. During the first part of the Tortonian (P1), high productivity within a prominent upwelling system supported a diverse assemblage of mesopredators, at least some of which permanently resided in the Pisco embayment and used it as a nursery or breeding/calving area. Younger portions of the Pisco Formation (P2) reveal a more open setting, with wide-ranging species like rorquals increasingly dominating the vertebrate assemblage, but also local differences reflecting distance from the coast. Like today, these ancient precursors of the modern Humboldt Current Ecosystem were based on sardines, but notably differed from their present-day equivalent in being dominated by extremely large-bodied apex predators like Livyatan melvillei and Carcharocles megalodon.

As other marine minerals, phosphates - and in particular bioapatite - incorporate Sr from seawater during formation (Staudigel et al., 1985). Due to long-term variations in the relative abundance of Sr isotopes in the global ocean, 87Sr/86Sr values can be measured in minerals and compared to calibration curves for obtaining age estimates via Strontium Isotope Stratigraphy (SIS) (McArthur et al., 2020). Although hydroxyapatite is susceptible to alteration and Sr concentration in fish teeth can change during burial and diagenesis (Martin & Scher, 2004), fossil shark teeth have been successfully used for Sr-dating, especially when enameloid is analyzed (Schmitz et al., 1997; Harrell et al., 2016). In order to test the feasibility of this method in the fish tooth-rich marine sediments of the East Pisco Basin (Peru), and aiming to date some poorly-constrained strata of this region, we analyzed the ultrastructure and composition of fossil shark teeth from the Chilcatay Formation. This Miocene formation consists of massive sandstones and basement boulders overlain by bioclastic sandstones and diatomaceous and tuffaceous siltstones (Di Celma et al., 2019). It is characterized by an abundant marine vertebrate assemblage, among which elasmobranchs are present (Landini et al., 2019). Teeth of Isurus sp., Cosmopolitodus hastalis, Isurus oxyrinchus, Megalolamna paradoxodon and Physogaleus contortus were collected from Chilcatay beds at the localities of Zamaca, Media Luna and near Cerro Colorado. Teeth were investigated through an optical microscopy and SEM. After taphonomic observations, 11 teeth underwent ICP-OES and 87Sr/86Sr analyses. Shark teeth show a compact and non-porous outer enameloid layer that is distinctly separated from the more porous and heterogeneous inner core of dentine. Ultrastructure analysis shows that the enameloid is formed by highly-ordered bundles of fluoroapatite crystallites, which are often well-preserved, whereas the dentine displays a bone-lik

This paper is the first integrated account of the sedimentology, stratigraphy, and vertebrate paleontology for the marine strata of the Chilcatay Formation exposed at Ullujaya, Pisco basin (southern Peru). An allostratigraphic framework for the investigated strata was established using geological mapping (1:4000 scale) and conventional sedimentary facies analysis and resulted in recognition of two unconformity-bounded allomembers (designated Ct1 and Ct2 in ascending order). The chronostratigraphic framework is well constrained by integration of micropaleontological data and isotope geochronology and indicates deposition during the early Miocene. The marine vertebrate fossil assemblage is largely dominated by cetaceans (odontocetes), whereas isolated teeth and spines indicate a welldiversified elasmobranch assemblage. Our field surveys, conducted to evaluate the paleontological sensitivity of the investigated strata, indicate that vertebrate remains only came from a rather restricted stratigraphic interval of the Ct1 allomember and reveal the high potential for these sediments to yield abundant and scientifically significant fossil assemblages.

A newly discovered elasmobranch assemblage from the fossil-bearing area of Zamaca (Chilcatay Formation, southern Peru) is described herein, providing a first comprehensive view on the early Miocene shark and ray paleocommunities of the East Pisco Basin, whose sedimentary infill represents one of the most important Cenozoic Fossil-Lagerstätten worldwide. The studied assemblage includes at least twenty-two species attributed to twelve families and five orders. Thirteen taxa are recorded for the first time from the Chilcatay Formation, and four of them are recorded for the first time from the Pacific coast of South America. The reconstructed paleoenvironmental scenario is consistent with a shallow-marine coastal area, representative of a sheltered shelfal setting, influenced by both brackish and open-ocean waters. This paleoenvironment was inhabited by a community of small mesopredator elasmobranchs that exploited the Zamaca area as a nursery ground and recruitment area. The structure of the Zamaca assemblage is mainly explained by three key-features: 1) a taxonomic composition dominated by two shark lineages, Lamniformes and Carcharhiniformes, the former being predominant; 2) the leading role played by two species, Carcharhinus brachyurus and †Cosmopolitodus hastalis, accounting for about 60% of the analyzed specimens; 3) the distinctly juvenile imprint of the entire assemblage. Striking similarities emerge between the elasmobranch assemblage from Zamaca and the late Miocene one from Cerro Colorado (Pisco Formation, East Pisco Basin), thus suggesting the persistence of a peculiar “biological enclave” driven by the concurrence of the ecological, environmental, and oceanographic factors that characterized the coast of present-day Peru during the Neogene

This paper describes a ∼200 m-thick section of the Pisco Formation exposed at Cerro Colorado, an important fossiliferous site in the Ica desert. In order to properly place the fauna in its correct relative position, this study establishes the stratigraphic framework within which the different fossil-bearing intervals of this site can be compared and may prove invaluable in future high-resolution studies on the faunal change. Most of the Pisco Formation deposits exposed at Cerro Colorado consist of gently dipping fine-grained sandstones, diatomaceous siltstones and diatomites with minor ash layers and dolomites deposited within nearshore and offshore settings. To facilitate detailed stratigraphic correlations within the Pisco strata for a 30 km2area, eight marker beds have been defined and large-scale (1:10,000 scale) geological mapping conducted to determine fault positions, styles and offsets. The geological map shows that there are two important angular unconformities in the study area. The first one is the interformational basal unconformity of the Pisco Formation against folded, faulted, and planated Oligo-Miocene rocks of the Chilcatay Formation. The second is a low-angle intraformational erosional discontinuity of up to 4° angular discordance that allows the subdivision of the Pisco stratigraphy exposed in the study area into two informal allomembers. Dating of the exposed succession by diatom biostratigraphy suggests that the age of the lower allomember is late Miocene, whereas the upper allomember is late Miocene or younger.

Field mapping and sedimentological study of outcrop sections exposed along the Ica River valley permitted the establishment of a regional allostratigraphic framework for the upper Miocene portion of the Pisco Formation. The stratigraphy of the studied interval is illustrated using a new 1:20,000-scale geological map which reveals that this formation is a cyclical sedimentary succession composed of three fining-upward allomembers. The bounding surfaces defining each allomember are transgressively modified subaerial unconformities. They converge and merge landward into a single composite surface representing the time-transgressive lower boundary of the Pisco Formation. Accordingly, the extent of the stratigraphic gap associated with the basal unconformity varies significantly throughout the basin and increases toward the basin margins. The timing of allomember-bounding surfaces coincides with that of major oxygen-isotope maxima in the deep-sea oxygen isotopic record and matches the ages of eustatic sequence boundaries identified elsewhere, indicating glacio-eustatic falls due to the growth of Antarctica ice sheets as a viable mechanism for their development

The taphonomy and palaeoecology of the early Miocene (Burdigalian) vertebrate assemblage of Ullujaya (East Pisco Basin, Peru) is here described. Vertebrate remains are concentrated in marine facies (Ct1a association) of the exposed Chilcatay Formation (dated 19–18 Ma) deposited within a 30–40 m deep, semi-enclosed, offshore environment. Coupled with ichnological observations, the size distribution of pyrite framboid relics reveals fluctuation of euxinic and oxic-dysoxic conditions at the seafloor. The assemblage is dominated by toothed cetaceans (kentriodontids, squalodelphinids, physeteroids, and the eurhinodelphinid-like Chilcacetus), together with a large dermochelyid turtle, some bony fish, and diverse elasmobranchs, mostly juveniles of Carcharhinus brachyurus and Cosmopolitodus hastalis. The vertebrate assemblage comprises a coastal community, dominated by mesopredators, representative of a warm-temperate, sheltered embayment connected with riverine and open-ocean environments. Vertebrate skeletons are typically disarticulated and incomplete, and some bone elements display shark bite marks. Microborings are observed at the bone surface. Bones exhibit a good degree of apatite mineralisation and bone cavities are locally filled by Ca-Mg carbonates. Our taphonomic observations suggest prolonged flotation of carcasses during which they were subject to biogenic and physical processes of partial destruction (including scavenging by sharks), before final deposition on a soft compact substrate. Preservation was favoured by the oxygen-deficient bottom conditions that inhibited the action of benthic macro-scavengers

We report on bite marks incising fossil mammal bones collected from upper Miocene deposits of the Pisco Formation exposed at Aguada de Lomas (southern Peru) and attributed to the giant megatooth shark Carcharocles megalodon. The bitten material includes skull remains referred to small-sized baleen whales as well as fragmentary cetacean and pinniped postcrania. These occurrences, the first in their kind from the Southern Hemisphere, significantly expand the still scarce record of bite marks for C. megalodon; moreover, for the first time a prey (or scavenging item) of C. megalodon is identified at the species level (as Piscobalaena nana, a diminutive member of the extinct mysticete family Cetotheriidae). Due to the fragmentary nature of the studied material, the exact origin of the detected marks (i.e., by scavenging or by active predation) cannot be ascertained. Nevertheless, relying on actualistic observations and size-based considerations, we propose that diminutive mysticetes (e.g., cetotheriids) were some of the target prey of adult C. megalodon, at least along the coast of present-day Peru. C. megalodon is thus here interpreted as an apex predator whose trophic spectrum was focused on relatively small-sized prey. Lastly, we propose a link between the recent collapse of various lineages of diminutive mysticetes (observed around 3 Ma) and the extinction of C. megalodon (occurring around the end of the Pliocene).

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 on40Ar/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.

The enormous concentration of marine vertebrates documented within the Pisco Formation is unique for Peru and South America and places this unit among the prime fossil Lagerstätten for Miocene to Pliocene marine mammals worldwide. In order to provide a robust stratigraphic framework for the fossil-bearing locality of Cerro Los Quesos, this study presents a 1:10,000 scale geological map covering an area of about 21 km2, a detailed measured section spanning 290 m of strata, and a refined chronostratigraphy for the studied succession well constrained by diatom biostratigraphy and high-resolution40Ar/39Ar isotopic dating of three interbedded ash layers. Within the apparently monotonous, diatomite-dominated sedimentary section, the Pisco Formation has been subdivided into six local members, with stratigraphic control over the different outcrops facilitated by the establishment of a detailed marker bed stratigraphy based on 15 readily distinguishable sediment layers of different nature.

The Mio-Pliocene Pisco Formation (Peru) is a world-famous marine vertebrate Lagerstätte. Several fossil specimens are wrapped up in dolomitic nodules. Some others lie in the sediment displaying dolomite only in bone cavities (e.g., mesorostral canal and endocranium). With the aim to understand whether the precipitation of the dolomitic nodules influenced the formation of the Lagerstätte, we collected field data on a high number of fossil vertebrates and conducted petrographic and mineralogical analyses on samples representative of the variable development of concretions. Our results revealed positive relationships between size, completeness and articulation of skeletons and the presence of an external nodule. Clear evidence of chemoautotrophic communities that thrived on the carcasses is scarce. Microborings are often found in the cortical bone tissues together with iron oxides; the former are left by microorganisms feeding on the carcass, the latter are traces of former Fe sulphides, a product of organic matter degradation. We suggest that an early burial of the skeletons was a determinant factor in the development of dolomite concretions, since it allowed methanogenesis and anaerobic sulphate reduction exploiting the lipids in the bones and the organic matter dispersed in the sediments. Dolomite precipitation was driven by the same bacteria operating during the suphophilic stage of whale-fall communities. Textural observations imply that dolomite precipitated shortly after the burial of carcasses. The increase of alkalinity generated by sulphate reduction and methanogenesis caused a rapid precipitation of the dolomite within skeletal cavities and prevented the degradation of the bones and diagenetic compression of skeletons; the nodules themselves prevented erosion of fossils after exhumation. Therefore, nodule formation had a crucial role in the development of the Pisco Lagerstätte.

Laminated diatomaceous mudstones are the most characteristic lithology of the Mio- Pliocene Pisco Formation, Ica desert, Peru; discrete tuff layers intercalate to them. The Pisco Fm. deposited during the Neogene in the East Pisco Basin, one of the Andean forearc basins which run eastward of the Peru-Chile trench. Nowadays, the Pisco Fm. is one of the most famous Fossillagerstätte in the world. Its popularity is due to the great abundance of exceptionally well preserved marine vertebrates, especially cetaceans, lying in the diatomaceous mudstone. Many specimens are indeed fully articulated and often concentrated in horizontal accumulation layers. This study is meant to understand how diatom deposition may have positively interfered in the genesis of this Fossillagerstätte, by: 1) analyzing diatom laminae and their different assemblages and obtaining information on the seasonality affecting the water column during deposition of the Pisco Fm.; 2) recognizing and evaluating any evidence of exceptional harmful algal blooms (HABs) coinciding with the fossil vertebrate accumulation levels; HABs were recently evoked to explain four discrete horizons showing fossil marine vertebrates accumulation in the late Miocene of Atacama desert, Chile (Pyenson et al., 2014); 3) investigating the hypothesis that diatom and diatom mats deposition may have favored the preservation of vertebrates by lowering the level of dissolved oxygen on the sea floor; indeed, these conditions would have prevented the development of a whale-fall community that generally leads to the fast decay of the carcasses and the final disarticulation of skeletons; 4) evaluating the sedimentation rates of the diatomaceous mudstones to understand how long it would have taken for a carcass to be completely buried, thus, verifying whether fossil preservation is mainly due to the high sedimentation rates of diatomaceous mudstones as suggested by Brand et al. (2004); Moreover, we are investigating diatom assemblages c

The Mio-Pliocene Pisco Formation (Ica Desert, Peru) is known for its vast outcrops, which preserve one of the richest concentrations of fossil marine vertebrates on Earth. During the last thirty years, punctual studies carried out on several exceptionally well preserved individual specimens highlighted the extraordinary diversity of vertebrates that populated the area: cetaceans, pinnipeds, turtles, sea birds, and fishes are just a few examples. Through our successive fieldwork campaigns, for the first time a detailed taphonomic and stratigraphic study of the fossil assemblages from localities of the oldest strata (middlelate Miocene) of the Pisco Fm. was undertaken. Stratigraphy was used to correctly assign specimens to well-defined strata and, consequently, to evaluate the temporal variation in the composition and concentration of the fossils along the investigated stratigraphical sequence. GPS data were collected for all the fossil vertebrates and a preliminary systematic determination was carried out in the field for all the specimens, while the most interesting and well-preserved ones were taken in laboratory for preparation and detailed study. Taphonomic observations were also carried out: orientation of the skeletons, skeletal completeness, bone articulation, associated invertebrates, associated shark teeth, and evidence of bioerosion. Sediment samples enclosing the fossils were collected along the continuous vertical diatomaceous mudstone sequences and were examined through different analyses (grain size analyses, micropalaeontological analyses, major and minor elements analyses, etc.) to understand the condition of burial and to obtain a more precise dating of the fossiliferous layers, using calcareous nannoplankton (where present), diatoms, silicoflagellates, radiolarians, and dinoflagellates as biostratigraphic markers. Some of these microfossils were also used to measure as proxies to infer on paleo-depth and water paleo-temperature. Samples for radiomet