Avisaurus

Avisaurus is an extinct genus of enantiornithine bird belonging to the family Avisauridae, known from the Maastrichtian stage of the Late Cretaceous period (approximately 66 million years ago) in North America.^[1] The genus is primarily represented by hindlimb fossils, particularly robust tarsometatarsi, which suggest adaptations for a raptorial or predatory lifestyle similar to modern birds of prey.^[1] Currently, two valid species are recognized: the type species A. archibaldi from the Maastrichtian Hell Creek Formation of Montana and the recently described A. darwini from the same formation, with an additional unnamed Avisaurus sp. also reported from Hell Creek.^[1] These birds were relatively large for enantiornithines, with estimated body masses around 1.2 kilograms for A. darwini, and they inhabited diverse environments along the Western Interior Seaway.^[1] The type species, Avisaurus archibaldi, was originally described in 1985 based on a distal tarsometatarsus (UCMP 117600) collected from the Hell Creek Formation in Garfield County, Montana, dating to about 66 million years ago.^[2] This specimen measures approximately 75 mm in length and features a low-aspect-ratio tarsometatarsus with deeply grooved trochleae and a prominent tubercle for the m. tibialis cranialis muscle, indicating strong perching and grasping capabilities.^[1] A. darwini, named in 2024 after Charles Darwin, is based on a more complete tarsometatarsus (DDM 1577.730) from Carter County, Montana, distinguished by unique morphological traits such as a pronounced flange on metatarsal III and a deep fossa on the trochlea.^[1] An unnamed partial tarsometatarsus (MOR 3070) from Makoshika State Park further expands the known diversity of the genus in the Hell Creek Formation.^[1] Phylogenetically, Avisaurus is part of the Avisauridae clade within Enantiornithes, a diverse group of Mesozoic birds that were the dominant avifauna during the Cretaceous.^[2] Cladistic analyses place Avisaurus alongside other North American avisaurids like Mirarce and Gettyia, forming a distinct Laurasian subclade separate from South American relatives such as Neuquenornis and Soroavisaurus.^[2] The robust hindlimb morphology of avisaurids, including Avisaurus, supports interpretations of terrestrial predation or scavenging, contrasting with the more arboreal habits of many other enantiornithines.^[1] Formerly, material from the Campanian Two Medicine Formation was assigned to A. gloriae (1995), but this has been reclassified as Gettyia gloriae, a smaller avisaurid with a strongly inclined tarsometatarsus.^[2]

Taxonomy

Etymology

The genus name Avisaurus derives from the Latin avis ("bird") and the Greek sauros ("lizard" or "reptile"), literally meaning "bird lizard," a nomenclature that highlights its avian-like features within a reptilian context. It was established by paleontologists Michael K. Brett-Surman and Gregory S. Paul in their 1985 description of a new family of bird-like dinosaurs, emphasizing the fossil's mix of theropod and ornithurine traits at a time when the evolutionary links between dinosaurs and birds were hotly contested in the scientific community—a period known as the "bird-dinosaur wars."^[3]^[4] The type species, A. archibaldi, honors paleontologist J. David Archibald, who discovered the holotype specimen, a right tarsometatarsus from the Hell Creek Formation. Originally described as a second species of Avisaurus in 1995, A. gloriae, named by David J. Varricchio and Luis M. Chiappe, commemorates Gloria Siebrecht, a volunteer fossil preparator at the Museum of the Rockies who contributed to its study; this epithet reflects the collaborative efforts in Mesozoic avian paleontology. However, A. gloriae was reclassified as Gettyia gloriae in 2018.^[5]^[2] In 2024, a new species, A. darwini, was described from the Hell Creek Formation, with its specific epithet honoring Charles Darwin for his foundational contributions to evolutionary biology, underscoring the fossil's relevance to understanding avian diversification near the Cretaceous-Paleogene boundary. Concurrently, to resolve paraphyly within Avisaurus, some Hell Creek material previously referred to the genus was reassigned to the new genus Magnusavis, named from Latin magnus ("big") and avis ("bird"), with the species M. ekalakaensis recognizing the nearby town of Ekalaka, Montana; this separation maintains taxonomic precision while affirming close affinities among these avisaurids.^[1]

Valid species

The genus Avisaurus currently encompasses two valid species, both known primarily from tarsometatarsal elements of the Hell Creek Formation in Montana, USA, dating to the Maastrichtian stage of the Late Cretaceous (~68–66 million years ago). These species are characterized by a robust tarsometatarsus adapted for powerful perching or prey manipulation, with features such as a broad trochlea on metatarsal II and a hypertrophied tubercle for the m. tibialis cranialis muscle. An additional unnamed Avisaurus sp. is known from a partial tarsometatarsus (MOR 3070) from the Hell Creek Formation.^[1]^[6] The type species, A. archibaldi, was named and described in 1985 based on the holotype UCMP 117600, a nearly complete right tarsometatarsus measuring approximately 75 mm in length, collected from Garfield County, Montana. This medium-sized species exhibits a robust tarsus with a plantar projection on metatarsal III, distinguishing it from other enantiornithines by its overall sturdiness and proportions suggestive of predatory capabilities.^[3]^[6] In 2024, a second species, A. darwini, was formally established from the holotype DDM 1577.730, a complete right tarsometatarsus from Carter County, Montana. This species is distinguished by an elongated trochlea II, a laterally projecting flange that closes the intertrochlear incisure, and a m. tibialis cranialis tubercle positioned at about 30% of the tarsometatarsus length, conferring hawk-like proportions potentially suited for gripping prey.^[1] Material originally assigned to A. gloriae (1995), based on a partial skeleton from the Campanian Two Medicine Formation in Montana, was reclassified in 2018 as the type species of the distinct genus Gettyia due to paraphyly within Avisaurus and unique pedal features like an elongated metatarsal III. Some referred specimens have also been excluded from Avisaurus, such as those reassigned to Neuquenornis or deemed insufficiently diagnostic; following 2024 taxonomic revisions, the consensus recognizes only these two valid species within the genus, alongside the unnamed species.^[7]^[2]^[1]

Description

Skeletal anatomy

The skeletal remains of Avisaurus are fragmentary and primarily consist of isolated tarsometatarsi, representing an incomplete postcranial skeleton with no preserved cranial elements, feathers, or axial structures such as the pygostyle.^[1] These bones exhibit typical enantiornithine features, including a fused proximal tarsometatarsus with an elongated shaft and well-developed distal trochleae adapted for perching.^[1] The robust hallux, evidenced by a J-shaped metatarsal I, suggests a reversed toe capable of grasping, while the overall pedal morphology—characterized by deeply grooved trochleae and a deep extensor fossa—resembles that of modern raptorial birds like hawks and owls.^[1] Key osteological features include a low aspect ratio of approximately 1:4 in the tarsometatarsus, with a mediolateral width at the midpoint exceeding 20% of the proximodistal length, indicating a sturdy build for predatory or perching functions.^[1] A hypertrophied tubercle for the m. tibialis cranialis, positioned about 30% down the length of metatarsal II, is prominent in A. darwini, and a plantar projection on the medial condyle of the metatarsal III trochlea is present across the genus.^[1] The distal extent of metatarsal II is roughly equal to that of metatarsal IV, with proximoplantar fossae on the plantar surface aiding in muscle attachment for enhanced grip strength.^[1] Variations occur between species, particularly in the configuration of the intertrochlear incisure and associated flanges on metatarsal II. A. archibaldi features a lateroplantarly projecting flange and a medial margin that extends more distally, contributing to a broader shaft overall.^[1] In contrast, A. darwini has a laterally projecting flange that partially closes the intertrochlear incisure, along with a trochlea III approximately 20% longer relative to other proportions, enhancing its predatory grip capabilities.^[1] These differences are subtle but diagnostic, as seen in referral specimens like MOR 3070, which shows a dorsally projecting m. tibialis cranialis tubercle.^[1] Tarsometatarsus lengths vary by species and ontogenetic stage, ranging from about 63 mm in A. darwini (e.g., metatarsal III at 65.38 mm) to approximately 75 mm in A. archibaldi for adults, with proximal widths around 18–20 mm supporting a lightweight yet strong construction inferred from the thin bone walls and robust trabeculae.^[1] Such measurements provide the basis for estimating overall body sizes comparable to large modern perching birds.^[1]

Size and morphology

Avisaurus species reached masses of approximately 1.2 kg (range 914–1712 g) for A. darwini and 1.7 kg (range 1488–2283 g) for A. archibaldi, comparable to large modern hawks.^[1] These estimates derive from comparative volumetric models using tarsometatarsus length scaled against extant avian taxa, highlighting a trend toward increased body size in Late Cretaceous enantiornithines.^[1] Inferred wingspans for A. darwini measure roughly 1.2–1.3 m, based on hawk-like scaling from hindlimb proportions.^[8] Morphologically, Avisaurus exhibits elongated hindlimbs relative to trunk length, with tarsometatarsi (63–75 mm across species) featuring deep trochlear grooves and hypertrophied muscle tubercles for powerful grasping and propulsion, proportions mirroring those of modern Accipitridae for perching and predatory pursuits.^[1] The wings, inferred from scaling against related avisaurids and extant birds, appear compact and robust, suggesting adaptations for short bursts of agile, maneuverable flight rather than extended gliding.^[1] This leg-dominant build contrasts with more balanced proportions in early enantiornithines, emphasizing terrestrial and arboreal capabilities. Soft tissue reconstructions indicate full coverage in pennaceous feathers, including primary remiges for reinforced wing support during flight; no traces of ornamental or display plumes have been identified in avisaurid fossils.^[9] Avisaurus retains enantiornithine-specific uncinate processes on the ribs, which enhanced respiratory efficiency by improving rib and sternum mobility during ventilation, akin to mechanisms in extant birds but distinct from neornithine configurations.^[10] Limited evidence for sexual dimorphism exists in the form of size variation among tarsometatarsal specimens (e.g., 10–15% differences in proximal width), potentially indicating larger males, though ontogenetic or intraspecific factors cannot be ruled out without further data. Overall, these traits position Avisaurus as hawk-like in scale and form, with a leg-to-body ratio facilitating raptor-style locomotion while supporting powered flight.^[11]

Discovery history

Initial fossils

The initial fossils of Avisaurus were discovered in 1975 during an expedition by the University of California Museum of Paleontology (UCMP) in the Hell Creek Formation of Garfield County, Montana, USA. The holotype specimen, UCMP 117600, consists of a complete left tarsometatarsus measuring approximately 75 mm in length, which was collected from Maastrichtian-age (Late Cretaceous) sediments.^[3] This isolated foot bone was initially described in 1985 by Michael K. Brett-Surman and Gregory S. Paul as the type specimen of a new genus and species, Avisaurus archibaldi, within a novel family of "bird-like dinosaurs," Avisauridae, based on its mix of primitive theropod features and derived traits such as a robust metatarsal III visible along its full length and a coossified proximal end.^[3] They highlighted its potential evolutionary significance in linking Laurasian and Gondwanan theropod faunas, though the fragmentary nature of the material limited detailed anatomical interpretation at the time.^[3] Subsequent analysis in 1992 by Luis M. Chiappe redescribed the A. archibaldi holotype and referred additional isolated tarsometatarsi to the genus, firmly placing Avisaurus within Enantiornithes, an extinct clade of avialans characterized by unique shoulder girdle features and a hypotarsus on the tarsometatarsus that restricted flexion.^[12] Chiappe noted the bone's avian affinities, including a concave posterior surface and trochlear proportions akin to those in modern birds, countering earlier views of it as a non-avian theropod and emphasizing its role as one of the largest known North American enantiornithines.^[12] This reclassification addressed challenges posed by the specimen's incompleteness, which had initially obscured its close ties to other Cretaceous birds like those from the Lecho Formation in Argentina, also referred to Avisaurus sp.^[12] In 1995, David J. Varricchio and Luis M. Chiappe described a second species, A. gloriae, based on a partial skeleton (specimen MOR 553) from the Upper Cretaceous Two Medicine Formation in Glacier County, Montana, dating to the Campanian stage. This material was later reclassified as the new genus and species Gettyia gloriae in 2018 by Atterholt et al.^[2] This material, including elements of the pectoral girdle, forelimb, and hindlimb, provided the first associated Avisaurus remains and confirmed enantiornithine traits such as an uncinate process on the ribs and a strut-like coracoid. The discovery expanded the known temporal range of the genus into the earlier Late Cretaceous and highlighted morphological variation within Avisauridae, though debates arose over the exact referral of some elements due to preservation issues. Early referrals of additional material to Avisaurus included tarsometatarsi from the Judith River Group in Montana and Alberta, Canada, which shared diagnostic features like an enlarged hypotarsus and reduced metatarsal V, but sparked discussions on whether they represented A. archibaldi, A. gloriae, or distinct taxa versus other enantiornithines such as Hesperornis or basal ornithurines.^[12] The fragmentary condition of these specimens often led to initial uncertainties in classification, with some early interpretations favoring ornithurine affinities before cladistic analyses solidified their enantiornithine placement.^[12] These referrals underscored the genus's distribution across western North American formations but also illustrated the diagnostic challenges of isolated postcranial bones in avian paleontology during the 1980s and 1990s.

Recent discoveries

In 2024, paleontologists described three partial tarsometatarsi from the Maastrichtian Hell Creek Formation in Carter County, Montana, representing significant new material attributable to Avisaurus and expanding knowledge of Late Cretaceous enantiornithine birds. These specimens, collected between approximately 2019 and 2022, include the holotype of a new species, Avisaurus darwini (DDM 1577.730, a right tarsometatarsus recovered in July 2022 from a multitaxic microvertebrate site via sieving techniques), an indeterminate Avisaurus sp. (MOR 3070, a partial right tarsometatarsus from Makoshika State Park in Glendive, Montana), and a related but distinct taxon, the new genus and species Magnusavis ekalakaensis (CCM V2019.5.1, a partial right tarsometatarsus).^[13] The discoveries were made near localities associated with the famous SUE the T. rex specimen, highlighting the efficacy of targeted microfossil sieving in recovering small avian remains from dinosaur-rich sediments.^[13] This material confirms A. darwini as the second valid species of Avisaurus, alongside the type species A. archibaldi, and includes referral of MOR 3070 based on shared diagnostic features such as a low tarsometatarsus aspect ratio.^[13] Magnusavis ekalakaensis, while avisaurid, is distinguished from Avisaurus by features including a distal expansion of metatarsals II and III and a rimmed plantar supratrochlear fossa on metatarsal III, suggesting a relatively shorter metatarsal III compared to typical avisaurids and indicating greater taxonomic diversity within the clade during the latest Cretaceous.^[13] These finds, detailed in a study by Clark et al. published in PLOS ONE on October 9, 2024, address previous gaps in understanding avisaurid ecology by providing the first robust evidence of predatory adaptations through pedal proportions.^[13] Specifically, the specimens exhibit a low tarsometatarsus aspect ratio (approximately 1:4 in the Avisaurus sp. material), a large distomedial tubercle for the m. tibialis cranialis, and deeply grooved trochleae, features that enhance biomechanical efficiency for grasping and constricting prey, akin to modern raptorial birds.^[13] This supports interpretations of avisaurids as active predators capable of subduing larger or heavier prey items in their floodplain environments.^[13]

Classification

Phylogenetic position

Avisaurus is classified as a member of the Enantiornithes, a diverse clade of Cretaceous avialans that dominated Mesozoic bird faunas, and is positioned as a basal euenantiornithine within the family Avisauridae. The Avisauridae, named after Avisaurus, encompasses taxa such as A. archibaldi, A. darwini, Soroavisaurus australis, Mirarce eatoni, and Gettyia gloriae, sharing derived features of the tarsometatarsus that distinguish them from other enantiornithines.^[1] This family represents one of the few well-supported clades among Late Cretaceous enantiornithines, primarily known from North and South American deposits dating to the Campanian-Maastrichtian stages. Key synapomorphies supporting the phylogenetic placement of Avisaurus and its relatives include a robustly constructed tarsometatarsus with a low aspect ratio (midpoint width at least 20% of total length), a broad trochlea on metatarsal II, a hypertrophied tubercle for the m. tibialis cranialis insertion located at least 20% down the shaft, and a plantar projection on the trochlea of metatarsal III.^[1] These features, combined with a reduced fibula that forms a slender splint-like element not extending to the distal tibia—a condition widespread in Enantiornithes but accentuated in avisaurids—underscore their enantiornithine affinities while showing superficial resemblances to the pedal morphology of more derived ornithuromorphs. Although quadrate bones are poorly known for Avisaurus, enantiornithine taxa generally exhibit a quadrate with a pneumatic foramen and a short orbital process, traits consistent with the clade's overall cranial architecture and supporting Avisaurus's position rather than a shift toward euornithine-like conditions. Recent phylogenetic analyses from 2024, incorporating expanded matrices with new Hell Creek Formation specimens, recover A. darwini as the sister taxon to A. archibaldi within Avisaurus, reinforcing the monophyly of Avisauridae as a derived euenantiornithine subclade.^[1] These studies highlight avisaurid pedal specializations—such as deeply grooved trochleae and strong flexor tendon sulci—as adaptations for raptorial grasping, suggesting potential stem-lineage relationships to Accipitriformes (hawks and allies) through convergent evolution in predatory foot structure, though no direct ties to modern avian lineages exist.^[1] Early interpretations of Avisaurus fossils, based primarily on isolated tarsometatarsi, debated ornithurine (modern bird-like) affinities due to the fused ankle structure and robust build, but cladistic analyses in the 1990s unequivocally confirmed its enantiornithine placement through shared characters like the hypotarsal configuration and overall tarsometatarsal proportions. Chiappe's 1996 systematic review of southern South American enantiornithines further solidified this consensus by integrating Avisauridae into broader avian phylogenies, excluding alternative classifications and emphasizing their role as a terminal radiation within the extinct Enantiornithes.^[14]

Evolutionary significance

Avisaurus, as a member of the avisaurid clade within Enantiornithes, exemplifies the late survival of this diverse group of Mesozoic birds, which dominated avian faunas throughout the Cretaceous but went extinct at the Cretaceous-Paleogene (K-Pg) boundary approximately 66 million years ago. Representing some of the largest and most specialized enantiornithines near the end of the Cretaceous, avisaurids like A. archibaldi and the newly described A. darwini achieved body masses up to around 1,700 grams, adapting to predatory niches through enhanced pedal grasping capabilities that allowed for the capture and constriction of vertebrate prey. This specialization highlights pre-extinction experimentation among "opposite birds," as Enantiornithes are sometimes called due to their reversed ankle joint, potentially filling ecological roles akin to modern raptors in late Maastrichtian ecosystems.^[1] The pedal adaptations in Avisaurus and its relatives exhibit convergent evolution with Cenozoic birds of prey, such as hawks (Accipitridae) and owls (Strigidae), featuring low tarsometatarsal aspect ratios (approximately 1:4) and robust, deeply grooved trochleae that supported powerful flexion for prey restraint. These traits suggest avisaurids employed hunting strategies involving aerial or perch-based predation, targeting small vertebrates or even juvenile dinosaurs, thereby demonstrating functional parallels to post-K-Pg avian predators despite the group's ultimate demise. Such convergences underscore the evolutionary flexibility of Enantiornithes in occupying higher trophic levels before the K-Pg mass extinction event, which eliminated all non-neornithine birds.^[1]^[15] Recent 2024 analyses of Hell Creek Formation fossils have revealed increased avisaurid diversity, with three new enantiornithine taxa—including two avisaurids (Avisaurus darwini and an unnamed Avisaurus sp.)—bringing the total known enantiornithine diversity to four taxa in this terminal Cretaceous assemblage. This expanded diversity points to niche partitioning between avisaurids and coexisting theropods, such as dromaeosaurids, where avisaurids likely exploited mid-sized predatory roles through their specialized feet, complementing rather than directly competing with non-avian dinosaurs.^[1] Avisaurus reinforces the maniraptoran theropod origins of birds, as Enantiornithes nest within this clade, sharing derived traits like feathered bodies and advanced flight capabilities that parallel those in the lineage leading to modern Neornithes. However, avisaurids hold no direct ancestry to crown-group birds, instead providing insights into extinct avian experimentation with locomotion and predation that echoes post-extinction radiations among surviving neornithines. Future research on additional Hell Creek material promises to resolve the full extent of avisaurid radiation, clarifying phylogenetic relationships and ecological dynamics in the lead-up to the K-Pg extinction.^[1]^[4]

Paleoecology

Habitat and environment

Avisaurus fossils are primarily known from the Hell Creek Formation in Montana, USA, dating to the Maastrichtian stage of the Late Cretaceous (approximately 68–66 million years ago). This formation consists of fluvial and coastal plain deposits, characterized by interbedded sandstones, mudstones, and occasional coals, reflecting a dynamic environment of meandering rivers, floodplains, and overbank fines along the western margin of the Western Interior Seaway. Equivalent deposits in South Dakota (such as the Hell Creek Formation there) and similar sandstone channel facies in the San Juan Basin of New Mexico preserve comparable Maastrichtian avian material, indicating a broad distribution across interior North American floodplains.^[16]^[17] The paleoenvironment of the Hell Creek Formation was a warm, humid subtropical landscape with rivers, dense forests dominated by angiosperms, and expansive floodplains supporting swampy woodlands. Sedimentological evidence, including organic-rich mudstones and paleosols with clayey Bt horizons, points to subhumid conditions with seasonal aridity, as inferred from thin, impure coals and mildly calcareous soils that suggest periodic dry spells amid overall wetter phases. In the Ekalaka area of southeastern Montana, which represents an upland extension of the Hell Creek depositional system, volcanic ash layers are preserved, including beds contemporaneous with the Chicxulub impact event, highlighting the proximity to global cataclysmic influences at the end of the Cretaceous.^[16] Avisaurus coexisted with a diverse assemblage of large dinosaurs, including theropods like Tyrannosaurus rex, ceratopsians such as Triceratops, and hadrosaurs, alongside smaller vertebrates like crocodilians, turtles, and multituberculate mammals. Other avian taxa, including enantiornithines and possibly ichthyornithiform birds, shared this ecosystem, occupying niches in the riverine and forested habitats. The presence of fish, amphibians, and invertebrates further indicates a productive, multi-tiered community structured around fluvial systems.^[16]^[1] Taphonomic processes in these formations favored preservation of Avisaurus remains in channel lag deposits, where high-energy fluvial transport concentrated disarticulated bones amid coarser sediments. The rarity of such relatively small-bodied fossils (around 1 kg) stems from size-biased destruction during riverine dispersal and potential degradation in acidic, organic-rich soils, leading to underrepresentation relative to larger taxa despite their likely abundance in the ecosystem. Microvertebrate sites, often in crevasse splay sandstones, yield isolated elements like tarsometatarsi, underscoring the role of localized depositional traps in capturing avian remains.^[1]

Diet and behavior

Avisaurus, as a member of the avisaurid clade of enantiornithine birds, exhibited a predatory lifestyle characterized by adaptations for grasping and subduing prey. Its raptor-like feet featured a robust tarsometatarsus with deeply grooved trochleae and an elongated hallux comprising over 20% of the tarsal length, enabling effective constriction of small vertebrates and invertebrates such as lizards, small mammals, and possibly hatchling dinosaurs. A 2024 biomechanical analysis of avisaurid hindlimb morphology revealed that these features, including a low aspect ratio (~1:4) and hypertrophied tubercle for the m. tibialis cranialis muscle, enhanced lever efficiency and powerful dorsiflexion for capturing proportionally large prey relative to body size. There is no direct fossil evidence supporting scavenging behavior, suggesting Avisaurus primarily engaged in active predation rather than opportunistic feeding.^[1] The hunting strategy of Avisaurus likely involved raptorial attacks using strong pedal constriction to subdue prey on the ground or in low vegetation.^[1] This is inferred from the low aspect ratio of the tarsometatarsus, which optimized leverage for powerful dorsiflexion and prey restraint during strikes.^[1] Flight capabilities supported this lifestyle, with evidence of advanced powered flight in avisaurids providing maneuverability in cluttered habitats.^[9] Additionally, the robust hindlimbs suggest occasional ground-foraging, similar to some modern galliforms, for pursuing mobile prey in understory environments.^[1] Behavioral inferences indicate that enantiornithines, including avisaurids, displayed precocial development, with hatchlings fully feathered and mobile upon emergence, reducing the need for prolonged parental care.^[18] Contributing to the extinction of Avisaurus and other enantiornithines at the Cretaceous-Paleogene boundary, the impact winter following the Chicxulub asteroid strike likely disrupted forested habitats and prey availability for these specialized raptorial birds, exacerbating their vulnerability compared to more adaptable neornithine birds.^[19]^[20]