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Pisco Formation
Pisco Formation
Pisco Formation
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The Pisco Formation is a geologic formation located in Peru, on the southern coastal desert of Ica and Arequipa. The approximately 640 metres (2,100 ft) thick formation was deposited in the Pisco Basin, spanning an age from the Late Miocene up to the Early Pliocene, roughly from 9.6 to 4.5 Ma. The tuffaceous sandstones, diatomaceous siltstones, conglomerates and dolomites were deposited in a lagoonal to near-shore environment, in bays similar to other Pacific South American formations as the Bahía Inglesa and Coquimbo Formations of Chile.

Key Information

The Pisco Formation is considered one of the most important Lagerstätten sites,[2][3] based on the large amount of exceptionally preserved marine fossils, including sharks (most notably megalodon), birds including penguins, whales and dolphins, marine crocodiles, and Thalassocnus, a marine giant sloths.[4]

Other famous fossils from this site include the giant raptorial sperm whale Livyatan,[5] the sperm whale relative Acrophyseter, and the walrus-like dolphin Odobenocetops.[6]

Description

[edit]

The Pisco Formation of the Pisco Basin consists of tuffaceous sandstones, diatomaceous yellow to gray siltstones and a basal conglomerate.[7] The formation is deposited from Pisco in the north to Yauca in the south. The northern portion is known as the Ocucaje Area and the southern part as the Sacaco Area.[8] The total thickness of the formation is estimated at 640 metres (2,100 ft).[9] The formation unconformably overlies the Chilcatay and Caballas Formations.

Paleobiota of the Pisco Formation

[edit]

The Pisco Formation has provided a rich resource of marine fauna, including marine mammals like cetaceans and seals, large fishes, reptiles, and penguins.[10] It is also one of the richest sites in the world for fossil cetaceans, with close to 500 examples being found in the formation.[11]

The oldest fossils of the aquatic sloth Thalassocnus (T. antiquus) come from the Aguada de Lomas horizon of the Pisco Formation and were dated at roughly 7 Ma. The youngest specimen (T. carolomartini) was found in the Sacaco horizon and dated to approximately 3 Ma.[12] Thalassocnus was preyed upon by the probable apex predators of the environment, Livyatan and megalodon.[13][14] The youngest strata belonging to the formation have been dated at 2 Ma, corresponding to the Early Pleistocene (Uquian). Fossils of the modern Humboldt penguin were found in these deposits at the Yauca locality.[15]

Birds

[edit]
Taxa Species Locality Materials Description Images Notes
Ciconiidae indet. Gen. et sp. indet. A stork
[16]
Fulmarus Indet. sp. A petrel
[17]
Morus M. peruvianus Sud-Sacaco A set of limb elements A relative of living gannets (Sulidae)
[18]
Perugyps P. diazi Sud-Sacaco A limb element (right carpometacarpus). A New World vulture (Cathartidae)
[18]
Pelagornis Pelagornis sp. Aguada de Lomas Proximal carpometacarpus and right humerus ends A pseudotoothed bird (Pelagornithidae)
[19][20]
Pelecanus Indet. sp. A pelican
[21]
Phalacrocorax P. aff. bougainvillii A relative of the Guanay cormorant
[17][22]
cf. Phalacrocorax sp. Probable cormorant remains
Rhamphastosula R. aguierrei Sud-Sacaco West A cranium skull A sulid bird with an enlarged beak
[17][23]
R. ramirezi Sud-Sacaco West A skull
Spheniscus S. humbodti Sud Sacaco Archaic Humboldt penguin
[15]
S. megarhampus Sud Sacaco A partial skeleton Large-beaked banded penguin
[15]
S. muizoni Cerro la Bruja A partial skeleton. The oldest banded penguin
[24][25]
S. urbinai Sud-Sacaco West A partial skeleton (partial skeleton with skull) A larger banded penguin than S. muizoni
[15]
Sula S. brandi Cerro Colorado A neurocranium lacking interorbital septum, lacrimals, palatines, pterygoids, jugals, quadrates, and right quadratojugal.Proximal portion of the beak, including most of the right dentary and angular. Relatives of living boobies (Sulidae)
[18][26]
S. magna Sud Sacaco A set of limb elements.
S. sulita Sud Sacaco A limb element (coracoid)

Fish

[edit]

Bony fish

[edit]
Taxa Species Description Images Notes
Alosinae indet. sp. A type of herring
[10]
Centropomidae C. aff. Psamoperca A snook
[10]
Triglidae indet. sp. A type of sea robin
[10]
Xiphiidae indet. sp. A swordfish
[10]
Sardinops S. humboldti A sardine
[27]
[28]

Rays

[edit]
Taxa Species Description Images Notes
Myliobatis indet. sp. A species of eagle ray
[10]

Sharks

[edit]
Taxa Species Locality Materials Description Images Notes
Carcharias C. taurus Archaic sand tiger shark
[17]
Carcharhinus indet. sp. A requiem shark
[10]
Carcharodon C. carcharias Archaic great white shark and close relative, respectively
[29][17]
C. hubbelli
Cosmopolitodus C. hastalis The broad-toothed "mako"
[10]
Hexanchus H. gigas A cow shark
[29]
Isurus I. oxyrhinchus Archaic shortfin mako
[19]
Otodus O. megalodon The largest of the megatoothed sharks (Otodontidae) and of all fishes
[10]

Mammals

[edit]

Cetaceans

[edit]
Taxa Species Locality Material Description Images Notes
Acrophyseter A. deinodon Sud-Sacaco A skull A small raptorial physeteroid (sperm whale relative)
[18][30]
A. robustus Cerro la Bruja A skull
Atocetus A. iquensis Cero la Bruja A skull A small kentriodontid whale
[31][32]
Australithax A. intermedia El Jahuay A long-snouted porpoise (Phocoenidae)
[33]
Balaenoptera B. siberi Aguada de Lomas A partial skeleton Archaic rorqual (Balaenopteridae)
[19][34]
Belonodelphis B. peruanus Cerro la Bruja A skull An elongated oceanic dolphin (Delphinidae)
[19][31][35]
Brachydelphis B. jahuayensis El Jahuay A partial skull An early oceanic dolphin (Delphinidae)
[19][31][36][37]
B. mazeasi El Jahuay A partial skull
Brujadelphis B. ankylorostris Cerro la Bruja A skull with complete jaw A toothed whale of uncertain relation (incertae sedis)
[38]
Hemisyntrachelus H. oligodon Sud-Sacaco A skull An orca relative
[17]
Incakujira I. anillodefuego Aguada de Lomas A preserved skeleton A small rorqual (Balaenopteridae)
[19][39]
I. fordycei Aguada de Lomas A preserved skeleton
Kogia K. danomurai Sacaco A partial skull consisting of partial cranium, preserving the facial and dorsolateral regions of the cranial vault, but lacking most of the rostrum and basicranium Basal member of Kogia, the genus of pygmy and dwarf sperm whale
[40]
Koristocetus K. pescei Aguada de Lomas A partial skull A small kogiid
[41]
Livyatan L. melvillei Cerro Colorado A partially preserved skull with teeth and lower jaw A very large raptorial physeteroid with 36 centimetres (1.18 ft) teeth
[5]
Lomacetus L. ginsburgi Aguada de Lomas A porpoise relative (Phocoenidae)
[19][42]
Mamaziphius M. reyesi Cerros la Mama y la Hija A partial skull consisting of partial cranium lacking most of the rostrum, the zygomatic processes of the squamosal, the occipital condyles and the ear bones An early beaked whale (Ziphiidae)
[43]
Messapicetus M. gregarius Cerro Colorado A partial skeleton consists of a skull, mandibles, humeri, vertebrae, and scapula An early beaked whale (Ziphiidae)
[44]
Miocaperea M. pulchra Aguada de Lomas A partial skull A cetothere whale
[45]
Ninoziphius N. platyrostris Sacaco A partial skeleton A giant beaked whale-relative
[17]
Odobenocetops O. leptodon Sacaco A preserved skull A tusked odontocete belonging to its unique family
[6]
O. peruvianus Sacaco A partial skull
Piscobalaena P. nana Sud-Sacaco A skull A small cetothere
[17][27]
Piscocetus P. sacaco Sacaco A partial skeleton An extinct cetacean
[29]
Piscolithax P. aenigmaticus Aguada de Lomas A partial skeleton A porpoise relative (Phocoenidae)
[17][19]
Platyscaphokogia P. landinii Cerro Hueco la Zorra, An incomplete skull lacking the tip of the rostrum and the basicranium. An early beaked whale (Ziphiidae)
[46]
Pliopontos P. littoralis Sud-Sacaco A partial skull An early oceanic dolphin (Delphinidae)
[17]
Scaphokogia S. cochlearis Aguada de Lomas An extinct kogiid
[19][47]

Pinnipeds

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Seals
Taxa Species Locality Material Description Images Notes
Acrophoca A. longirostirus Sub-Sacaco A partial skull. Archaic southern seal (Monachinae)
[17]
Australophoca A. changorum Aguada de Lomas A partial skeleton consists of incomplete right ulna, right radius, right and left humeri, and other unidentified remains. A phocidae seal.
[19]
Hadrokirus H. martini Sub-Sacaco A partial skull Archaic southern seal (Monachinae)
[17][48]
Hydrarctos H. lomasiensis Sub-Sacaco A skull. A sea lion and fur seal relative (Otariidae)
[17]
Icaphoca I. choristodon Cerro La Bruja A subcomplete skull with associated left and right mandibles, atlas, axis, third, fourth, and fifth cervical vertebrae. Archaic southern seal (Monachinae) [49]
Magophoca M. brevirostris Cerro la Bruja A partial skeleton of a male. Archaic southern seal (Monachinae)
[50]
Piscophoca P. pacifica Sub-Sacaco A partial skull Archaic southern seal (Monachinae)
[27]

Xenarthrans

[edit]
Sloths
Taxa Species Locality Materials Description Images Notes
Thalassocnus T. antiquus Aguada de Lomas A partial skeleton including the skull, mandible, and most of the postcranial skeleton. a semi-aquatic giant sloth inhabiting marine environments
[4][17]
[18][29]
[51]
T. carolomartini Sacaco An associated skull and mandible and two articulated hands, probably belonging to the same individual.
T. littoralis Sud-Sacaco A skull with missing jugals.
T. natans Sud-Sacaco A skull, mandible, and partial skeleton.

Mollusks

[edit]

Bivalves

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Taxa Species Description Images Notes
Dosinia indet. sp.
[17]

Polychaetes

[edit]
Taxa Species Description Images Notes
Diplochaetetes D. mexicanus A cirratulid bristle worm
[52]

Gastropods

[edit]
Taxa Species Description Images Notes
Acanthina A. obesa
[17]
A. triangularis
[17]
Concholepas C. kieneri
[17]
Herminespina indet. sp.
[17]

Reptiles

[edit]

Crocodilians

[edit]
Taxa Species Locality Materials Description Images Notes
Eusuchia indet. sp.
[17]
Piscogavialis P. jugaliperforatus Sacaco A partial skeleton and a skull. A gryposuchine (gharial relative)
[18][27]
P. laberintoensis Laberinto area, Ladera de Lisson Hill A complete skull and mandible, along with postcranial elements
[53]
Sacacosuchus S. cordovai Sacaco A nearly complete skull A gharial relative (Gavialidae)
[54]

Turtles

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Taxa Species Locality Materials Description Images Notes
Pacifichelys P. urbinai A sea turtle (Cheloniidae)
[13][55]
Chelonia indet. sp. Green sea turtle relative
[10]

Correlations

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Laventan

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Laventan correlations in South America
Formation Honda Honda Aisol Cura-Mallín Pisco Ipururo Pebas Capadare Urumaco Inés Paraná Map
Basin VSM Honda San Rafael Caldera Pisco Ucayali Amazon Falcón Venezuela Paraná
Pisco Formation is located in South America
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation
Pisco Formation (South America)
Country  Colombia  Bolivia  Argentina  Chile  Peru  Venezuela  Argentina
Boreostemma
Hapalops
Miocochilius
Theosodon
Xenastrapotherium
Mylodontidae
Sparassodonta
Primates
Rodents
Birds
Terror birds
Reptiles
megalodon
Flora
Insects
Environments Fluvial Fluvio-deltaic Fluvio-lacustrine Fluvio-deltaic Fluvial
Laventan volcanoclastics

Laventan fauna

Laventan flora
Volcanic Yes

See also

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References

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Further reading

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Pisco Formation

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The Pisco Formation is a prominent geological unit exposed in the East Pisco Basin along the southern coast of , spanning from approximately 14.8 to 6.7 million years ago (Ma) and renowned worldwide as an exceptional Fossil-Lagerstätte for well-preserved marine vertebrates. This formation, with a thickness varying between 200 and over 1,000 meters, consists primarily of diatom-rich sediments deposited in a basin influenced by the ancient , providing critical insights into marine ecosystems. Geologically, the Pisco Formation is subdivided into three main allomembers—P0 (14.8–12.4 Ma), P1 (9.5–8.9 Ma), and P2 (8.5–6.7 Ma)—separated by unconformities and reflecting cyclic changes in and depositional environments from shallow nearshore to deeper offshore shelf settings. Its lithology is dominated by fine-grained marine mudstones, siltstones, and diatomites, interspersed with sandstones, conglomerates, layers, and phosphatic intervals that indicate episodes of and low-oxygen conditions conducive to preservation. The basin's formation resulted from along the Peru , leading to a progression from semi-enclosed bays in the early stages to more open marine conditions later, with evidence of rocky shores and storm-influenced deposition. Paleontologically, the Pisco Formation stands out for its extraordinary concentration of articulated skeletons and soft-tissue remains, including over 890 documented specimens from 16 localities, with cetaceans comprising about 85% of the assemblage (61% mysticetes and 24% odontocetes). Notable fossils include the giant predatory Livyatan melvillei, the Carcharocles megalodon, diverse dolphins, seals, seabirds, turtles, crocodiles, sharks, rays, and bony fishes such as Sardinops cf. sagax, offering glimpses into a productive upwelling-driven that supported early marine . Taphonomic features like rapid burial in soupy sediments, phosphatization, and rare preservations of and stomach contents further highlight its status as one of the most significant marine fossil sites globally.

Geographic and tectonic setting

Location and extent

The Pisco Formation is primarily exposed in the East Pisco Basin along the southern coastal plain of , extending discontinuously approximately 200 km from the Ica region near the Pisco River southward to the Yauca area. This span corresponds to latitudes roughly between 13.5°S and 15.6°S. The formation's thickness varies significantly across the basin, ranging from 200 to 1000 m. Major outcrop areas include the Ocucaje region in the north, the Sacaco area in the central portion, and the Yauca locality in the south, where the formation is well-exposed in hills and valleys of the arid coastal desert. The northern boundary lies near the Pisco River valley, while the southern limit is marked by the vicinity of the Yauca River; inland, the extent is constrained by the rising terrain of the Andean foothills to the east. The Pisco Formation unconformably overlies older units such as the Chilcatay Formation in various parts of the basin. These exposures provide accessible sites for geological fieldwork, with prominent localities including Cerro Colorado in the Ica Desert and Agua Blanca, where detailed stratigraphic sections have been documented.

Regional geological context

The Formation is situated within the East Pisco Basin, a component of the broader basin that parallels the -Chile off the coast of southern . This tectonic regime arises from the ongoing oblique subduction of the Nazca Plate beneath the South American Plate, at a convergence rate of approximately 7-8 cm per year, which has driven the development of these basins since the late . The East Pisco Basin's evolution commenced in the middle Eocene with a major marine transgression, marked by the deposition of the Paracas Formation, reflecting initial and inundation of the region amid early Andean margin . By the , uplift of the Coastal —a structural high linked to the overriding plate's deformation—interrupted this , creating a barrier that influenced routing and led to localized erosional unconformities within the basin fill. This uplift phase, tied to intensified dynamics, transitioned the basin toward renewed accommodation space in the late , facilitating the accumulation of the Pisco Formation's thick marine sequences. The profoundly shaped the basin's sedimentary record, with detrital inputs primarily sourced from volcanic in the western , as evidenced by the abundance of tuffaceous components and populations dated to 23-1.2 Ma from the Huaylillas-Calipuy and Barroso . These volcaniclastic materials, including syndepositional ash layers, underscore the proximal influence of arc during Pisco Formation deposition. Basin geometry and patterns were further controlled by an array of high-angle normal faults, striking NNW-NNE, which bound depocenters and induce significant thickness variations across the East Pisco Basin, with offsets up to several hundred meters accommodating differential loading and extension.

Stratigraphy

Lithology

The Pisco Formation is predominantly composed of tuffaceous sandstones, diatomaceous siltstones, conglomerates, and dolomitic limestones. These lithologies reflect a marine sedimentary succession with significant biogenic and volcanic inputs. The sandstones are typically medium- to fine-grained, incorporating components such as glass shards, , and from tuffaceous layers. Shell fragments, including bivalves and gastropods, are common in beds, while phosphatic nodules and intraclasts occur throughout, particularly in coarser units. Conglomerates feature well-rounded pebbles to boulders in a sandy or silty matrix, often phosphate-enriched. Diatomaceous siltstones and limestones are fine-grained, with the former dominated by biogenic silica from coastal diatoms and the latter by microcrystalline dolomite. Vertically, the formation exhibits a fining-upward trend, beginning with basal conglomerates that transition to sandstones and siltstones, interspersed with intermittent shell beds. In the East Pisco Basin, the formation reaches thicknesses up to 1000 m. Diagenetic alterations include widespread dolomitization, forming nodular horizons and cementing siltstones and sandstones in upper units, as well as localized silicification evident in chert layers and replacement.

Internal subdivisions

The Pisco Formation exhibits a cyclical nature, composed of three depositional sequences or allomembers designated P0 (basal), P1 (middle), and P2 (upper), each characterized by transgressive-regressive cycles bounded by intraformational unconformities. These subdivisions are defined through sequence stratigraphic analysis, with erosional surfaces marking the bases of each allomember and reflecting episodic sea-level fluctuations. Thicknesses and lithological details vary by locality within the basin. The basal P0 allomember comprises diatomaceous mudstones, fine- to medium-grained sandstones, phosphatic nodule beds, and dolomite-cemented mudstones, attaining a thickness of approximately 40 m in studied sections, and represents the lowermost depositional phase overlying the Chilcatay Formation via a major . The middle P1 allomember consists of diatomaceous mudstones, siltstones, sandstones, and coquinas, is notably rich in fossils, and reaches about 100 m in thickness, separated from P0 by an erosional . The upper P2 allomember includes sandstones, silts, conglomerates, and coquinas, with a thickness of over 200 m, transitioning upward into units, and is bounded below by another that onlaps previous strata.

Age and biostratigraphy

Dating methods

The primary dating method for the Pisco Formation involves radiometric techniques, particularly 40Ar/39Ar dating applied to sanidine and crystals from interbedded layers (tuffs). This approach provides direct numerical age constraints by measuring the decay of 40K to 39Ar in potassium-bearing minerals, with samples typically processed through stepwise heating in a mass spectrometer to isolate the argon release spectrum and calculate plateau ages. Magnetostratigraphy has been employed to refine relative age assignments through the analysis of remanent magnetization in sedimentary rocks, correlating observed normal and reversed polarity zones to the global geomagnetic polarity timescale. Paleomagnetic samples are collected, demagnetized using or alternating field methods, and analyzed for characteristic directions, enabling to specific chrons and assessment of tectonic rotations or rates. Biostratigraphic integration utilizes microfossils such as nannofossils and planktonic to establish zonal schemes, with nannofossil assemblages assigned to standard biozones based on first and last appearances of marker species like Triquetrorhabdulus rugosus. Foraminiferal complements this by identifying zones through species like Globorotalia menardii, providing tied to the international chronostratigraphic framework. biostratigraphy is also key, particularly for the upper allomembers. Recent post-2020 studies have incorporated U-Pb dating on grains from tuffaceous layers and detrital sediments, using ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS) to determine crystallization ages and maximum depositional constraints, often combined with analysis to link sources to Andean . For instance, Ochoa et al. (2022) applied this method to refine ages in the Sacaco area, enhancing integration with prior radiometric data. has been used to constrain ages in the basal P0 allomember.

Temporal framework

The Pisco Formation spans the , from approximately 14.8 to 6.7 million years ago (Ma). This chronological range is established through integrated , , and isotopic analyses, reflecting a period of significant marine in the East Pisco Basin. Some regional exposures in the Sacaco area suggest the upper part extends to about 4.5 Ma into the early , but this is not representative of the entire formation and may include overlying units like the Caracoles Formation. Internally, the formation is subdivided into three unconformity-bounded depositional sequences or allomembers: P0 (14.8–12.4 Ma), P1 (9.5–8.9 Ma), and P2 (8.5–6.7 Ma). These subdivisions represent transgressive-regressive cycles, with ages constrained by diatom biostratigraphy, strontium isotope stratigraphy, and 40Ar/39Ar dating of intercalated volcanic ashes. 40Ar/39Ar dating results, in particular, have refined the timing of P1 and P2, confirming their placement within the late Tortonian to stages. Early uncertainties about temporal overlap between the and underlying Chilcatay formations, particularly regarding the nature of their bounding , have been resolved through recent correlations, which demonstrate a clear separation with the Chilcatay ending around 18 Ma.

Sedimentary

The Pisco Formation exhibits a range of sedimentary reflecting shallow-marine depositional environments along the Peruvian coast. Key associations include shoreface sands composed of fine- to coarse-grained, cross-stratified calcarenites and sandstones, characterized by unidirectional cross-lamination, swaley cross-stratification, and wave ripples indicative of storm-influenced wave-dominated settings. These sands are often moderately to highly bioturbated and transition upward into finer-grained deposits. Open marine silts occur as finely laminated, gray to white diatomaceous siltstones and mudstones, rich in planktic and benthic diatoms, deposited in low-energy, highly productive offshore environments. Within these silts, phosphate-rich layers form as transgressive lags, consisting of phosphatic clasts, nodules, and fragments, resulting from phosphogenesis in suboxic conditions with low rates. occurs as veins and casts in the formation, associated with later diagenetic processes. The formation's cycle architecture is organized into three main depositional sequences (P0, P1, P2), each comprising transgressive systems tracts with deepening-upward successions from shoreface sands to offshore diatomaceous silts, bounded by ravinement surfaces. Regressive phases show shallowing-upward trends, with coarser conglomeratic and sandy intervals at sequence bases marking progradational episodes. These cycles reflect eustatic sea-level fluctuations and local tectonic in a forearc basin setting. Trace fossils are abundant, particularly in shoreface and transition zones, including pre-lithification burrows and borings that indicate softground conditions during deposition. The Glossifungites ichnofacies dominates at sequence boundaries, featuring firmground burrows such as vertical shafts and horizontal networks that colonized omission surfaces. Post-depositional alterations include incipient cementation by dolomite and , forming nodules within siltstones and around clasts, often associated with microbial activity and early . Basin-wide erosion surfaces, represented by unconformities separating the sequences, show evidence of subaerial exposure and ravinement, with irregular and lag deposits. Diatomaceous siltstones, a dominant , preserve these features with minimal compaction due to high silica content.

Paleoecological interpretations

The paleoecological interpretations of the Pisco Formation reveal a dynamic influenced by coastal , sea-level variations, and fluctuating salinity in nearshore to lagoonal settings. The lower P0 allomember (middle ) records a warm, semi-enclosed shallow-water environment with restricted circulation, as evidenced by nearshore calcarenites and coastal assemblages indicating subtropical conditions during the Climatic Optimum. This setting supported initial nutrient enrichment from early activity associated with the precursor to the modern , fostering high primary productivity in a protected embayment. In the P1 allomember (, Tortonian), depositional conditions shifted to shallow marine nearshore environments with evidence of restricted circulation and localized hypersaline episodes, inferred from evaporitic features such as beds and bedded evaporites in lagoonal . These indicators suggest periodic in semi-protected coastal lagoons, interspersed with nutrient-rich incursions that enhanced productivity. Prominent , driven by strengthened coastal currents, is documented by abundant microfossils (e.g., Thalassionema and Chaetoceros) and phosphatic lags at sequence bases, which formed during transgressive ravinement and concentrated organic phosphates from high-biomass decay. Sea-level fluctuations, linked to eustatic cycles and tectonic in the forearc basin, produced cyclical transgressive-regressive patterns, with unconformities marking lowstands that exposed phosphatic hardgrounds. Cooler temperatures relative to P0 reflect post-Middle Climate Transition cooling, yet overall warm temperate marine conditions prevailed with seasonal pulses. The upper P2 allomember () indicates a transition to more open marginal marine settings with rocky shores, where intensified in distal areas sustained elevated productivity and a complex trophic structure. Transgressive trends throughout the formation, culminating in deeper offshore diatomaceous siltstones, underscore eustatic control on shifts, while persistent warm with El Niño-like variability maintained overall stability. These interpretations highlight the Pisco Formation as a record of an ancient Humboldt Current-influenced system, where upwelling-driven productivity and sea-level dynamics shaped coastal paleoecologies.

Paleobiota

Invertebrates

The fossil assemblage of the Pisco Formation is remarkably diverse, with mollusks representing the most prominent group and serving as key indicators of the paleoenvironmental conditions in the ancient ecosystem. Numerous species of mollusks have been documented, encompassing a wide array of bivalves and gastropods that inhabited shallow marine to lagoonal settings. Bivalves such as Ostrea, Pecten, Dosinia, Chionopsis, and Panopea are particularly abundant, often occurring in dense shell concentrations that reflect episodes of elevated biological productivity and rapid sedimentation. Gastropods, including Conus, distans, Miltha, Cancellaria, Northia, and various buccinids, further diversify the record, with their distributions aiding in biostratigraphic correlations across the formation's sequences. These molluscan assemblages, preserved primarily as molds and casts in diatomaceous shales, highlight thriving benthic communities adapted to the nutrient-rich coastal waters. Polychaetes contribute to the invertebrate paleobiota through the presence of cirratulid , which were first reported in 2021 from the strata of the Pisco Formation and associated Chilcatay Formation. These tube-building annelids formed organomineralized structures in subtidal environments, indicating stable, soft-bottom habitats conducive to reef development. Other notable invertebrate groups include , echinoids, and , which, though less dominant than mollusks, provide insights into the formation's and macrofossil dynamics. Benthic , such as Lepidocyclina, Miogypsina, Operculina, Amphistegina, and Cycloclypeus, occur in shallow-water carbonates and are valuable for , spanning assemblages from the late to late that reflect shifts in water depth and nutrient availability. Echinoid fragments appear subordinately in bioclastic deposits, alongside , contributing to skeletal grainstones formed by reworking in offshore settings. , often identified as balanids similar to Austromegabalanus, form dense accumulations in , signaling high-energy, nearshore conditions with encrusting growth on substrates. Shell beds dominated by bivalves, echinoids, and underscore the high productivity of the benthic realm, likely driven by in the proto-Humboldt Current. Recent post-2020 investigations have highlighted additional components, including ostracods from the P2 allomember, such as Cytheridea sp., Xestoleberis sp., Hemicytherura sp., Costa edwardsii, and Aurila sp., utilized in geochemical analyses to reconstruct paleoenvironmental conditions. These findings emphasize the role of in lagoonal ecosystems, where size and abundance patterns in shell assemblages likely responded to fluctuations in benthic oxygenation and nutrient flux.

Vertebrates

The record of the Pisco Formation is exceptionally rich and diverse, dominated by marine and coastal species that reflect adaptations to the productive systems of the ancient . This assemblage includes over 500 cetacean specimens alone, alongside numerous remains of elasmobranchs, reptiles, birds, and other mammals, highlighting a complex trophic web in nearshore to open-marine environments. Fish represent a foundational component of the , with elasmobranchs and bony showing specialized marine predatory and schooling behaviors. , such as Carcharocles megalodon, reached lengths of up to 17 m and occupied niches, preying on large marine vertebrates in open-water settings. Other like Cosmopolitodus hastalis and Carcharhinus brachyurus utilized nursery areas in shallower bays, indicating reproductive adaptations to coastal habitats. Rays, including myliobatids such as Myliobatis spp., were durophagous bottom-feeders, crushing shelled prey on the seafloor with reinforced jaws suited to benthic marine life. Bony , particularly teleosts like clupeids (Sardinops cf. sagax), formed massive schools that served as keystone prey, with hundreds of specimens preserving evidence of high productivity in zones. A new species, Sardinops humboldti, was described in 2025 based on articulated skeletons from the upper . Reptiles in the Pisco Formation exhibit clear transitions to fully marine lifestyles, particularly among cheloniids and crocodilians inhabiting coastal lagoons and shelves. Crocodilians, represented by longirostrine forms like Piscogavialis sp., adapted to piscivory in brackish to marine waters, with slender snouts for capturing fish in shallow embayments. Sea turtles dominated open-marine , including cheloniids such as Pacifichelys urbinai, which displayed durophagous for crushing mollusks and crustaceans, underscoring their role as mid-level consumers in productive nearshore ecosystems. Birds are primarily seabirds with diving and flighted adaptations for exploiting surface and mid-water resources. Penguins, such as Spheniscus muizoni, were pursuit divers specialized for underwater on schools in coastal areas. Procellariids and sulids, including Sula brandii and Sula figueroae, represented flighted forms that foraged over open seas, with remains indicating aerial piscivory and scavenging behaviors tied to nutrient-rich surface waters. Mammals, especially cetaceans, pinnipeds, and xenarthrans, showcase peak diversity and specialization for fully aquatic or semi-aquatic niches. The cetacean assemblage includes approximately 50 species, spanning mysticetes like Piscobalaena nana (filter-feeders adapted to krill-rich waters) and odontocetes such as the macroraptorial Livyatan melvillei (up to 17 m long, with conical teeth for tearing large prey) and suction-feeding Odobenocetops spp. (walrus-like tusks for benthic foraging). Recent discoveries, including the 2024 beaked whale Nazcacetus urbinai (a small-bodied ziphiid with reduced dentition for deep suction feeding, dated to ~7.45 Ma) and a 12-million-year-old porpoise skeleton from 2025, further illustrate odontocete evolutionary radiations in productive marine settings. Pinnipeds like Piscophoca sp. were monachine seals adapted for bottom-feeding in coastal zones, while xenarthrans such as semi-aquatic sloths (Thalassocnus spp.) grazed on seagrasses in protected lagoonal habitats, bridging terrestrial and marine realms.

Taphonomy and preservation

The Pisco Formation is recognized as an exceptional marine Fossil-Lagerstätte, renowned for preserving articulated skeletons, contents, and soft tissues in remarkable detail. Approximately 27% of specimens exhibit full articulation, including complete mysticete whales with associated plates phosphatized for enhanced preservation. contents, such as masses of remains in the abdominal regions of whales like Piscobalaena nana, provide direct evidence of diet and are preserved through rapid mineralization processes. Soft tissues, including and possible integumentary structures, are often found within nodules, highlighting the formation's capacity for Konservat-Lagerstätte conditions. Taphonomic processes in the Pisco Formation are dominated by rapid in anoxic or dysoxic bottom waters, which minimized scavenging and . Carcasses experienced scour-induced self- during storms, penetrating soupy diatomaceous sediments, followed by early diagenetic mineralization via recrystallization and dolomite formation around skeletons. Phosphatization particularly aids in preserving delicate features, such as fringes and scales in gut contents. Bonebeds, formed from mass mortality events possibly linked to algal blooms or seasonal anoxia, occur in concentrations at sites like Cerro , where hundreds of individuals accumulated in low-energy settings. Preservation biases in the Pisco Formation favor large vertebrates, with cetaceans comprising 85% of the assemblage and mysticetes overrepresented at 61% due to their durable, dense bones resisting post-mortem and decay. Size-sorting is evident in current-influenced deposits, where larger skeletons (>10 m) show better articulation than smaller odontocetes, which are more prone to dispersal. These biases reflect hydrodynamic and durability factors rather than original abundance, leading to an incomplete record skewed toward . The formation has yielded over 500 cetacean specimens, underscoring these patterns. Recent research from 2021 to 2025 has elucidated cyclical taphonomic variations tied to the formation's three allomembers (P0, P1, P2), which represent transgressive-regressive cycles influencing sediment type and oxygen levels. Studies highlight how dolomite layers in these allomembers record fluctuating diagenetic environments, enhancing preservation during anoxic phases. For instance, analyses of assemblages across localities reveal periodic mass accumulations linked to boundaries, providing a framework for understanding temporal biases in the record.

Stratigraphic correlations

Regional comparisons

The Pisco Formation exhibits stratigraphic equivalence with other marine sequences along the western margin of , particularly the Bahía Inglesa Formation in northern and the Urumaco Formation in northwestern . These units represent contemporaneous depositional episodes in and marginal marine settings influenced by Andean tectonics and volcanism during the Tortonian to stages. The Pisco and Bahía Inglesa formations, with their upper sections spanning the (approximately 9.5–6.1 Ma), share lithofacies dominated by sandstones, siltstones, and tuffaceous deposits, reflecting nearshore to inner shelf environments shaped by the proto-Humboldt Current. Biostratigraphic correlations highlight shared marine vertebrate assemblages across these Pacific and Caribbean coast formations, including odontocete cetaceans such as kentriodontids (e.g., Kentriodon sp.) and other delphinidans, alongside sharks like Carcharodon hastalis and aquatic sloths (Thalassocnus natans). Mass mortality events, evidenced by bonebeds of rorqual whales and pinnipeds in the Pisco and Bahía Inglesa formations, suggest analogous paleoecological dynamics, including harmful algal blooms and rapid burial in supratidal settings. The Urumaco Formation, dated to the late Miocene (ca. 11.6–5.3 Ma), complements this regional pattern with similar cetacean diversity, though its more marginal marine to estuarine facies distinguish it from the fully open-marine Pisco sequence. Lithostratigraphically, the Formation's tuffaceous sandstones and diatom-rich siltstones find analogs in the of the Bahía Inglesa Formation, where layers and bioclastic sands indicate comparable supply from Andean sources. Marginal equivalents in the Paraná Basin, such as tuffaceous intervals in the Entrerriano Group, reflect broader dispersal of volcanogenic sediments across southern during this period. These matches underscore a unified regional response to enhanced arc magmatism and . Prominent disconformities within the Pisco Formation, such as those bounding its internal allomembers (P1 and P2), record erosional hiatuses tied to eustatic sea-level falls, correlated to glacio-eustatic events such as Mi6 (ca. 10.4 Ma) and Mi7 (ca. 8.7 Ma). These unconformities, marked by conglomerate lags and angular discordances, mirror similar sequence boundaries in the Bahía Inglesa Formation, where sea-level lowstands facilitated exposure and ravinement surfaces. Such features highlight the Pisco Formation's sensitivity to far-field tectonic and climatic forcings across the American margin.

Global equivalents

The Pisco Formation as a whole spans the middle to (ca. 14.8–6.7 Ma), with upper sections in areas like Sacaco extending into the early Zanclean stage (to ca. 4.5 Ma). This temporal framework aligns it with global marine sequences, including those of the European , where Tortonian deposits reflect similar shallow to outer shelf environments during a period of tectonic and climatic transition. The Zanclean portion corresponds to the onset of the , marked by renewed marine flooding and faunal turnover in Paratethyan basins. Calcareous nannofossil supports correlations for parts of the formation, with sections encompassing zones NN9 to NN11 in low-latitude Tethyan sequences, characterized by marker species such as Catinaster coalitus in NN9 and quinqueramus in NN11, along with co-occurring taxa like Amaurolithus tricorniculatus. Tectonically, the Pisco Formation represents a forearc basin setting analogous to the Monterey Formation of , both developed along convergent margins with organic-rich, siliceous mudstones formed under intense coastal . Shared features include dolomite precipitation in anoxic muds and abundant phosphatic nodules, reflecting nutrient-rich waters that supported prolific . The depositional record of the Pisco Formation also aligns with major global climate events, including the Climatic Optimum (ca. 17–14 Ma) in its lower sections and the subsequent cooling trends toward the . This is evidenced by shifts from warm, semi-enclosed conditions in the middle to cooler, open-marine regimes in the upper , mirroring broader oceanic reorganization.

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