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Meridiolestida

Meridiolestida is an extinct clade of dryolestoid cladotherian mammals endemic to , primarily , spanning from the stage of the to the early . Known for their distinctive dental morphology featuring a reversed triangular occlusal pattern in the cheek teeth—lacking the protocone and basined talonid of tribosphenic therians—they represent a non-therian lineage that achieved ecological diversity as insectivores, carnivores, and possibly herbivores. Fossils are concentrated in Patagonia ( and ), with potential Eocene occurrences in , highlighting their role in the isolated and early faunas of the region. Within Dryolestoidea, Meridiolestida forms a monophyletic group distinct from dryolestids, encompassing several families such as Mesungulatidae (e.g., Mesungulatum ferox, one of the largest known members reaching up to 1 meter in length) and Reigitheriidae (e.g., Reigitherium, featuring specialized crushing ). Recent reassessments have expanded recognized diversity, including the reassignment of Groebertherium to Meridiolestida based on new specimens from the . Other notable genera include (a small, shrew-like form from the early ), Coloniatherium (with bunodont molars suggesting omnivory), and Necrolestes patagonensis (a survivor extending the clade's temporal range by nearly 45 million years and demonstrating persistence amid radiations). Recent discoveries, such as Orretherium tzen from the Campanian-Maastrichtian Dorotea Formation in and Yeutherium pressor from subantarctic deposits, underscore their biogeographic extent and evolutionary innovations in tooth replacement and occlusal wear for processing hard foods. Phylogenetically, Meridiolestida is positioned as a derived dryolestoid within , sister to other gondwanan dryolestoids, with analyses supporting their divergence in the early following isolation of . Their survival across the Cretaceous-Paleogene boundary into the and illustrates a mosaic of faunal turnover in , where they coexisted with early metatherians before declining due to competition from invading therians. This clade's and morphological adaptations provide critical insights into the diversity of outside the therian-dominated northern record.

Overview

Definition and Etymology

Meridiolestida is an extinct of dryolestoid mammals within the larger group , endemic to and primarily documented from fossil remains in (especially ), with possible Eocene occurrences in ; it ranges temporally from the stage of the to the early . This group represents a significant radiation of non-therian mammals closely related to the broader Dryolestoidea superfamily. Members of Meridiolestida were particularly dominant in the mammalian faunas of , where they comprised a substantial portion of the known diversity as one of the non- clades phylogenetically closest to the therian lineage. Their persistence into the , as evidenced by Miocene records, underscores their evolutionary success in isolation on the southern continent. The name Meridiolestida was established by José F. Bonaparte in 1990.

General Characteristics

Meridiolestida displayed considerable morphological diversity, encompassing small insectivores comparable to modern in size (approximately 10–20 g and 10–15 cm long), such as Cronopio dentiacutus, to larger herbivores or omnivores approaching the mass of a large (exceeding 20 kg), exemplified by Peligrotherium tropicalis. This range reflects adaptations to varied ecological niches within Gondwanan terrestrial habitats, from insectivory in forested settings to more robust forms suited for browsing in open environments. Meridiolestidans share a distinctive reversed triangular occlusal pattern in the cheek teeth, lacking the protocone and basined talonid of tribosphenic therians, with variations including sectorial in basal forms and bunodont molars (low, rounded cusps suited for grinding or crushing food) in derived lineages such as mesungulatoids; this distinguishes them from the more sectorial dentitions of basal dryolestoids. Upper molars lack a parastylar hook, facilitating broader occlusal surfaces for processing diverse diets. Additionally, the lower exhibits an absence of the Meckelian groove in adults, signaling an advanced configuration and divergence from other dryolestoid lineages. These dental and mandibular features underscore the clade's evolutionary specialization within stem therians. The overall body plan of Meridiolestida was quadrupedal, with limb proportions and skeletal robusticity adapted for terrestrial locomotion across the landscapes of , where the group achieved faunal dominance. Fossorial or semi-fossorial tendencies in some smaller taxa, inferred from cranial and dental specializations, complemented their ground-dwelling , enabling exploitation of burrowing niches amid dinosaur-dominated ecosystems.

Taxonomy and Phylogeny

Classification History

The earliest known fossils attributable to Meridiolestida were described by Florentino Ameghino in 1891, based on fragmentary remains of Necrolestes patagonensis from the early of , ; Ameghino classified it as an extinct placental unrelated to modern South American mammals. Subsequent discoveries of fossils in the 1980s, including Mesungulatum houssayi described by F. Bonaparte and María F. Soria in 1984 from the Campanian-Maastrichtian Los Alamitos Formation, expanded the group's temporal range and initially led to interpretations as primitive ungulate-like mammals or basal therians, distinct from Laurasian lineages. Bonaparte formally erected the clade Meridiolestida in 1990 within Dryolestoidea, based on additional material from the same formation, emphasizing shared dental features like reversed triangular molars among taxa such as Mesungulatum, Coloniatherium, and Peligrotherium. Early classifications often placed meridiolestidans near sparassodont metatherians due to superficial resemblances in postcranial or as basal therians outside crown , reflecting limited material and uncertainties in phylogeny. A pivotal shift occurred in studies by Guillermo W. Rougier and colleagues in 2011–2012, which incorporated new specimens like and revised Reigitherium, firmly establishing Meridiolestida as a monophyletic of dryolestoid cladotherians through phylogenetic analyses highlighting synapomorphies such as single-rooted molars and specific angular process . These works resolved prior ambiguities by demonstrating their as the dominant southern Gondwanan dryolestoids, sister to northern forms. Recent analyses from 2021 to 2024 have refined this framework, addressing debates on and broader affinities. A 2013 proposal by Alexander O. Averianov and colleagues suggested meridiolestidans as non-cladotherian trechnotherians based on alternative character scorings, but this was refuted by subsequent datasets. The description of Orretherium tzen in 2021 supported a cladotherian placement within Mesungulatidae, as the earliest diverging member, reinforcing through dental and mandibular traits. Similarly, reassessment of Groebertherium in 2024 integrated it into based on new material, resolving prior exclusion as a dryolestid and stabilizing the against concerns. Necrolestes is now recognized as a survivor of this radiation.

Phylogenetic Position

Meridiolestida is positioned within as a clade of Dryolestoidea, belonging to the broader group , where Dryolestoidea serves as the to (encompassing marsupials and placentals). This placement reflects their status as non-therian mammals that diverged before the origin of crown , with Meridiolestida representing a distinct southern n radiation adapted to South American ecosystems during the . Phylogenetic analyses incorporating dental and cranial characters consistently recover this relationship, emphasizing Meridiolestida's role in illuminating early cladotherian diversification in . Key synapomorphies linking to other dryolestoids include specialized dental features such as the presence of a neomorphic cuspulid (cusp e) on lower molars, oriented mesiodistally, and a reduction in the number of premolars, alongside shared traits like a long rostrum and compressed, triangular-cusped teeth. However, exhibits unique adaptations, such as bunodont crowns and broad cingulids in derived forms like mesungulatoids, reflecting a trophic shift toward omnivory or herbivory distinct from the more insectivorous northern dryolestids. These features underscore their endemic evolution in isolation from Laurasian mammal faunas. The phylogenetic position of Meridiolestida remains debated, with some analyses suggesting it as a stem-therian group outside , potentially sister to spalacotheriids within a novel Alethinotheria. This alternative view, based on analyses of trechnotherian characters, challenges the of Dryolestoidea by rendering it paraphyletic. In contrast, more recent studies affirm the of Meridiolestida as a cohesive, endemic South American lineage within Dryolestoidea, supported by expanded morphological datasets that resolve internal relationships into monophyletic subgroups like Cronopioidea and Mesungulatoidea.

Included Taxa

Meridiolestida encompasses several families and approximately 10–12 valid genera, primarily known from South American deposits spanning the to the early . The clade is divided into major groups such as Cronopioidea and Mesungulatoidea, with taxa exhibiting diverse adaptations from small insectivores to larger herbivores. The family Necrolestidae includes the genus Necrolestes, represented by such as N. patagonensis and N. mirabilis, which are notable as survivors of an otherwise radiation, extending the group's temporal range by nearly 45 million years. These small, mole-like forms are characterized by specialized burrowing adaptations and are the most recent known meridiolestidans. Mesungulatidae comprises larger herbivorous taxa, including Mesungulatum houssayi, Coloniatherium cilinskii, and Orretherium tzen, all from deposits in Patagonia and ; these genera feature robust dental structures suited for grinding . Paraungulatum is also tentatively placed within this family based on shared molariform traits. Orretherium tzen represents the earliest diverging member of Mesungulatidae. Reigitheriidae contains enigmatic carnivorous or omnivorous forms such as Reigitherium bunodontum from the of ; these small to medium-sized taxa exhibit crenulated enamel and complex premolars indicative of a varied . Yeutherium pressor, described in 2025 from subantarctic deposits, is tentatively placed here based on shared crushing , further extending the biogeographic range. Peligrotheriidae is monotypic, consisting of Peligrotherium tropicalis from the of ; this large, dog-sized genus is distinguished by bunodont molars adapted for a herbivorous diet and represents one of the largest known meridiolestidans. Basal or cronopioidean genera include from the early () Candeleros Formation and Leonardus from the late Late Cretaceous (Campanian-Maastrichtian) Los Alamitos Formation of , both small insectivores with acute, sectorial teeth for piercing prey. Groebertherium from the Late Cretaceous Allen Formation has been recently reassigned to Meridiolestida from Dryolestidae based on new specimens showing compatible dental morphology. Several genera remain within Meridiolestida, including Amarillodon meridionalis from the early of , known from a single suggesting advanced dental angulation, and Bondesius ferox from the , with fragmentary remains indicating possible carnivorous habits. Lakotalestes luoi from the of has been proposed as a potential extralimital relative based on shared dryolestoid traits, though its exact affinities remain debated.

Anatomy and Morphology

Dental Features

The dentition of Meridiolestida is characterized by a reversed triangular pattern on the molars, lacking tribosphenic features such as a protocone or basined talonid, which distinguishes them from more derived therians. Upper molars are typically bunodont, featuring low, rounded cusps arranged in a trigon with a prominent paracone and stylocone, but without a metacone or parastylar hook—a key difference from dryolestids, where the parastylar hook connects to the stylocone via the preparacrista. In larger, more derived forms, the upper molars develop expanded cingula that facilitate grinding, though a true hypocone is absent across the . Tooth counts vary, but commonly include 3–5 premolars and 3–5 molars per quadrant; for instance, Reigitherium exhibits 4 premolars and 3 molars, while Peligrotherium has 2 premolars and 5 molars. Lower molars in Meridiolestida feature a triangular trigonid formed by the protoconid, paraconid, and metaconid, adapted for shearing, paired with a low talonid that supports crushing functions through crenulated and accessory cuspulids. The talonid often includes an enclosing (enceinte) with intense enamel crenulation, enhancing in bunodont taxa. Premolars are generally simple and blade-like in basal forms but become molarized in derived species, contributing to overall dental complexity. Dental variations reflect dietary shifts within Meridiolestida, with insectivorous members like displaying sharp, pointed cusps on both upper and lower teeth—such as a tall paracone on uppers and acute protoconid on lowers—for precise cutting and piercing of prey. In contrast, herbivorous forms like Peligrotherium tropicalis possess broader, low-crowned molars with blunt cusps and fused cingula forming transverse ridges, suited for crushing fibrous plant material. Evolutionary trends in meridiolestidan dentition show a progression from carnassial-like, shearing morphologies in insectivores to more bunodont, grinding adaptations in herbivores, bridging plesiomorphic sharp-toothed forms with derived crushing dentitions across the Cretaceous-Paleogene boundary. This shift correlates with increasing body size and ecological diversification in southern continents.

Skeletal Morphology

The skeletal morphology of Meridiolestida reflects a conservative relative to mammals, retaining several primitive non- features while showing adaptations for terrestrial life in Gondwanan environments. Cranial morphology is characterized by a robust structure, including a globular braincase and an upturned rostrum in some taxa, as seen in the survivor Necrolestes patagonensis. The exhibits a reduced Meckelian groove, a diagnostic feature distinguishing Meridiolestida from more basal dryolestoids. Basicranial elements, such as the petrosal bone and large ventral opening of the cavum epiptericum, further align with non- patterns, supporting enhanced low-frequency hearing potentially linked to subterranean habits in derived forms. Postcranial elements indicate adaptations for terrestrial , with forelimbs showing a parasagittal evidenced by a ventrally facing glenoid and well-developed humeral trochlea in mesungulatid taxa. Long bones, including the and , are robust in larger species to support body weight, while smaller forms display relatively elongated proportions suited to agile movement. In Necrolestes, the is stout and distally expanded, the features a modified , and the has a shallow patellar groove, all contributing to capabilities with shortened limbs and inferred strong claws for digging. Body size varies widely across Meridiolestida, estimated from dimensions and cranial measurements, ranging from approximately 15 cm in length for small insectivores like to over 1 m in large mesungulatoids like Mesungulatum ferox, with Peligrotherium tropicalis reaching 50–60 cm.

Distribution and Fossil Record

Temporal Range

The fossil record of Meridiolestida primarily spans the to the early , with the earliest definitive records dating to the stage (approximately 89–86 Ma) from the Los Bastos Formation in Patagonia, , representing the oldest known mesungulatoid member of the clade. Earlier potential dryolestoid relatives appear in the (about 100 Ma), marking the onset of their radiation in , though unambiguous meridiolestidans are confined to the later . The group underwent its main diversification during the stages (83–66 Ma), achieving peak diversity in South American ecosystems, as evidenced by diverse assemblages from formations such as the . Following the , Meridiolestida persisted into the (66–63 Ma), exemplified by the genus Peligrotherium from Patagonian deposits, indicating survival of non-therian lineages amid the rise of tribosphenic mammals. Possible records extend into the Eocene (56–34 Ma), including a tentative brandoniid fragment from the La Meseta Formation on the , though this material is now lost and requires confirmation. The clade's final known representatives occur in the early (23–16 Ma), with Necrolestes patagonensis from the Santa Cruz Formation in Patagonia, extending the group's by nearly 45 million years beyond previous expectations. Meridiolestida likely declined due to competitive pressures from immigrant mammals following the erosion of South America's Gondwanan isolation in the late , with the last confirmed fossils dating to around 18 . This temporal distribution underscores their role as a relict Gondwanan radiation, bridging and faunas in southern continents.

Geographic Distribution

Meridiolestida fossils are primarily known from in , encompassing regions of present-day and . In , key occurrences include the Neuquén Basin in , where specimens have been recovered from the Los Bastos Formation, and Río Negro Province, with finds from the Campanian-Maastrichtian Anacleto and Allen Formations. Extensions into southern Chile are documented by taxa such as Orretherium tzen from the Dorotea Formation in the Río de Las Chinas Valley, . A possible record from exists based on a tentative identification of a lower molar fragment as ?Meridiolestida from the Eocene La Meseta Formation on [Seymour Island](/page/Seymour Island), though the specimen was lost prior to formal description and requires confirmation. Meridiolestida exhibit strong Gondwanan , with all confirmed records restricted to southern continents and no verified occurrences in the , aside from the dubious Lakotalestes from , whose affinities remain uncertain. Their distribution reflects vicariance following the breakup of Pangea and subsequent Gondwanan fragmentation, limiting dispersal to isolated southern landmasses during the .

Major Discoveries

The earliest significant discovery of a meridiolestidan was that of Mesungulatum houssayi in the 1980s from the Los Alamitos Formation in northern , , , where an upper molar was initially described as the first from the region, initially misinterpreted as a condylarth before being recognized as a dryolestoid. This find, based on material collected from outcrops in , marked a pivotal moment in understanding diversity in , as it represented one of the first skeletal elements attributed to the group. A major advance came in 2011 with the unearthing of from the Los Llanos site in , , within the early , yielding two well-preserved skulls and associated postcranial elements that revealed specialized saber-like canines and insectivorous adaptations. This discovery, from the La Buitrera Paleovertebrate Locality, provided the first complete cranial material for a South American meridiolestidan, bridging gaps in early mammalian evolution during the and highlighting nocturnal predatory behaviors. More recent excavations have further enriched the record, including the description of additional dental and dentary fossils of Reigitherium bunodontum in the of , which clarified its bunodont dentition and mesungulatoid affinities, expanding knowledge of dietary diversity within the clade. In 2021, Orretherium tzen was reported from a partial lower with five cheek teeth discovered in the Dorotea Formation near Cerro Guido in southern Chilean , representing the southernmost known meridiolestidan and suggesting wider Gondwanan distribution. An important early specimen is the of Necrolestes patagonensis, described in 1891 from early strata in Patagonia, , which consists of a partial including fragments, vertebrae, and limbs recovered from the Santa Cruz Formation. This near-complete material, long enigmatic and initially classified as a placental , was reassigned to Meridiolestida in 2012, confirming it as the sole known survivor of the lineage and providing insights into post-Cretaceous persistence. The 2024 recovery of new mammalian specimens from the Upper Cretaceous Allen Formation in Patagonia, including material referable to Groebertherium, prompted its reassignment from Dryolestidae to , significantly expanding recognized diversity and refining the phylogeny of Mesungulatoidea through enhanced resolution of dental and mandibular features. These finds from screenwashing efforts at Cerro Tortuga underscore the untapped potential of the for elucidating late therian evolution in southern continents. In 2025, Yeutherium pressor was described from a partial bearing an upper in the Dorotea Formation, Río de las Chinas Valley, subantarctic Chile. This new reigitheriid taxon, closely related to Reigitherium, features specialized crushing and highlights independent evolution of such adaptations in Meridiolestida, further demonstrating their biogeographic extent and dietary innovations.

Paleobiology and Evolutionary Role

Diet and Ecology

Members of Meridiolestida exhibited diverse diets inferred primarily from dental morphology. Small-bodied taxa, such as Cronopio dentiacutus, were likely insectivorous, possessing sharp, triangular molars and elongated snouts adapted for capturing and processing insects and other small invertebrates. In contrast, larger forms like Peligrotherium tropicalis displayed omnivorous to herbivorous adaptations, featuring bunodont molars suited for grinding tough plant material, seeds, and possibly some animal matter, with biomechanical analyses indicating masticatory forces comparable to those of herbivorous ungulates such as the black rhinoceros (Diceros bicornis). Meridiolestids occupied terrestrial habitats in warm temperate to subtropical environments, including forested floodplains and riverine settings across during the and early . Some taxa, notably Necrolestes patagonensis, likely adopted lifestyles, burrowing in soft soils to exploit subterranean resources, as suggested by their robust cranial features and specialized . The group's persistently low diversity in post-Cretaceous implies effective niche partitioning with co-occurring early marsupials, allowing relictual survival in marginal ecological roles amid the isolation of the continent following the K-Pg boundary. During the , small meridiolestids such as Cronopio probably served as prey for larger vertebrates, including dinosaurs, in predator-rich ecosystems. Their endurance into the , exemplified by Peligrotherium and Necrolestes, reflects opportunistic exploitation of vacated niches in isolated after the end-Cretaceous , where geographic barriers limited competition from northern immigrants.

Evolutionary Significance

Meridiolestida represents a key clade in mammalian evolution, highlighting diversity among non-therian lineages. As cladotherians within Dryolestoidea, sister to the therian lineage within Cladotheria, they exhibit morphological features that illuminate parallel developments in anatomy to those in therians, particularly in dentition. Their molars often display a reversed triangular cusp pattern lacking a protocone and basined talonid, though some taxa exhibit bunodont features indicating dietary specialization, while retaining plesiomorphic dryolestoid traits such as reduced premolar count and specialized cingulids. This dental morphology underscores their role in documenting the evolution of complex mastication in early mammals, providing evidence for the acquisition of features that enabled dietary versatility in parallel lineages. The clade's Gondwanan radiation exemplifies the underestimated diversity of mammals during the , prior to the global dominance of s following the Cretaceous-Paleogene extinction. Meridiolestidans were the predominant mammals in , diversifying into a range of body sizes and ecologies that filled niches later occupied by s in northern continents. Their persistence into the , as exemplified by the survival of lineages like Necrolestes, demonstrates remarkable resilience amid faunal turnovers, allowing non- forms to coexist with incoming immigrants for tens of millions of years. This prolonged Gondwanan persistence highlights a parallel evolutionary trajectory in the south, independent of Laurasian developments. Recent discoveries challenge traditional northern-biased narratives of mammal evolution by revealing greater taxonomic and morphological diversity among Meridiolestida than previously recognized, informing broader biogeographic patterns of early mammalian dispersal. Studies from 2024, including new material from the , document additional morphotypes and reassign taxa like Groebertherium to the , indicating a more extensive herbivore-omnivore guild in and suggesting that southern mammalian radiations were more precocious and varied. A 2025 discovery of Yeutherium pressor, a reigitheriid from deposits, further highlights specialized crushing as a functional in the for processing hard foods. These findings imply that Gondwanan non-therians played a crucial role in shaping continental faunas, with their innovations contributing to the ecological foundations later exploited by therians.

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