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The Sunken Laboratory: Ancient Penguins and the Lost World of Zealandia

Penguins stand on mossy rocks by a turbulent sea. A large bone rests nearby. A map overlay appears over the ocean, under a cloudy sky.

1. Introduction: The Archipelagic Laboratory



1.1. Zealandia as an Evolutionary Crucible


The submerged continent of Zealandia, Te Riu-a-Māui, represents one of the Earth's most significant yet enigmatic biological provinces. Separated from the supercontinent Gondwana approximately 80 million years ago, this continental fragment drifted into the isolation of the South Pacific, carrying with it a cargo of ancient lineages that would evolve in splendid seclusion.1 While often characterized by its terrestrial oddities—the flightless moa, the nocturnal kiwi, and the tuatara—Zealandia’s marine realm has been equally prolific in generating evolutionary novelties. For the past 60 million years, the archipelago has functioned as a global center for seabird evolution, serving as both a "cradle" for new lineages and a "museum" for ancient forms that persisted long after their continental counterparts vanished.3

Among the marine avifauna, the Sphenisciformes—penguins—stand as the crown jewels of Zealandia’s fossil record. The region preserves a nearly continuous archive of penguin history, from the Paleocene epoch, shortly after the extinction of the non-avian dinosaurs, to the present day.5 It was here that the earliest giant penguins, such as Kumimanu fordycei, evolved to fill the ecological vacuum left by marine reptiles, reaching sizes that rivaled adult humans.7 These archaic giants demonstrated that the penguin body plan was capable of supporting massive bulk early in its evolutionary history. However, for decades, paleontologists believed that the era of the "monster penguins" ended in the Paleogene, replaced by smaller, more agile modern forms as ocean circulation patterns shifted and marine mammals radiated.9


1.2. The Pliocene Anomaly


Recent discoveries from the late Neogene of New Zealand have shattered this linear narrative of declining size. In September 2025, a groundbreaking study published in the Journal of Paleontology by Alan Tennyson, Felix Marx, Daniel Ksepka, and Daniel Thomas unveiled evidence of a colossal penguin surviving well into the Pliocene epoch, just 3 million years ago.10 This fossil, a nearly complete skull assigned to the genus Aptenodytes (the lineage of modern King and Emperor penguins), reveals that giant penguins did not merely survive in the remote past; they thrived in the subtropical waters of New Zealand virtually until the dawn of the Ice Ages.11

The discovery of this specimen, cataloged as NMNZ S.048857, presents a profound paradox. The modern members of the genus Aptenodytes are the epitome of cold-adaptation, restricted to the sub-Antarctic and Antarctic zones. The Emperor Penguin (Aptenodytes forsteri) breeds on sea ice in the depths of the polar winter, while the King Penguin (Aptenodytes patagonicus) patrols the cool waters of the Antarctic Convergence. Yet, their close relative from the Tangahoe Formation lived in a world of warm seas, sharing the coast with ancestral monk seals and diverse tubenose seabirds.12


1.3. Scope of Analysis


This report provides an exhaustive analysis of this discovery and its broader implications. It moves beyond a simple description of the fossil to reconstruct the entire ecosystem of the Mid-Piacenzian Warm Period (mPWP) in New Zealand. By synthesizing geological data from the Tangahoe Formation, morphometric comparisons of the fossil skull, and ecological modeling of the co-occurring fauna, we aim to answer two fundamental questions: How did a lineage now defined by its tolerance for extreme cold thrive in a subtropical greenhouse? And what force was powerful enough to drive these giants to extinction, clearing the way for the modern seabird communities of the Southern Ocean? The answers lead us to a dramatic "turnover hypothesis," where climate is secondary to the rise of a terrifying new guild of aerial predators.11



2. Geological Context: The Tangahoe Formation



2.1. Stratigraphy and Age


The fossil was recovered from the Tangahoe Formation, a spectacular sedimentary sequence exposed along the coastal cliffs of South Taranaki on the North Island of New Zealand.14 These cliffs, constantly eroded by the high-energy waves of the Tasman Sea, provide a continuously refreshed cross-section of Pliocene marine life.

The formation is biostratigraphically assigned to the Waipipian Stage of the New Zealand geological timescale. Through the integration of oxygen isotope stages, magnetostratigraphy, and biostratigraphic index fossils (such as the scallop Phialopecten triphooki), the age of the Tangahoe Formation has been constrained with remarkable precision to between 3.36 and 3.06 million years ago (Ma).13 This places the deposition of the fossil-bearing beds squarely within the late Pliocene (Piacenzian), a period of critical importance to climate science.


2.2. Lithology and Preservation


The Tangahoe Formation consists primarily of massive, bluish-grey siltstones and fine-grained sandstones, representing the accumulation of terrigenous sediment on a continental shelf.16 The depositional environment was a shallow marine setting, likely at mid-shelf depths (50–150 meters), characterized by moderate currents and high biological productivity.18

A key feature of the formation is the presence of concretions—hard, cemented nodules of calcium carbonate that form within the softer mudstone matrix. These concretions often nucleate around organic remains, sealing them in a chemically stable environment that protects them from compaction and dissolution.19 When the surrounding soft cliffs erode, these hard concretions fall onto the beach, where they are often found by collectors. The preservation quality within these nodules is often exceptional, retaining three-dimensional shape and fine surface details of bone that are typically lost in compression fossils.21 The giant penguin skull NMNZ S.048857 was preserved in exactly this manner, found inside a marine mudstone concretion loose on Ohawe Beach.20


2.3. The Mid-Piacenzian Warm Period (mPWP)


The temporal window of the Tangahoe Formation (c. 3.2 Ma) coincides with the Mid-Piacenzian Warm Period, the most recent time in Earth's history when atmospheric CO2 levels were comparable to today's (approx. 400 ppm).10

  • Global Climate: During the mPWP, global mean temperatures were 2–3°C higher than pre-industrial levels. Polar amplification meant that high latitudes were significantly warmer, leading to reduced ice sheet volume in Antarctica and Greenland.23

  • Sea Levels: Eustatic sea levels were estimated to be 10 to 25 meters higher than present, resulting in the inundation of coastal lowlands and the expansion of shallow shelf seas around Zealandia.11

  • Local Oceanography: Crucially for the penguin story, sea surface temperatures (SST) in the Tasman Sea and around Taranaki were 10–20°C warmer than the waters currently inhabited by Aptenodytes species.11 Reconstructions suggest local water temperatures of 14–20°C, comparable to the modern subtropical waters north of New Zealand today.

This paleo-environmental context is vital. It establishes that the Tangahoe Aptenodytes was not living on the fringe of a glaciated world, but in a warm, productive subtropical sea, fundamentally challenging the "ice-obligate" view of the genus.11



3. The Discovery and the Specimen



3.1. A Citizen Science Triumph


The recovery of NMNZ S.048857 highlights the essential role of amateur paleontologists and citizen scientists in New Zealand. The specimen was discovered in 2015 by John Buchanan-Brown, a local collector who regularly monitors the erosion of the Taranaki cliffs.20 Recognizing the shape of bone within a loose concretion on Ohawe Beach, Buchanan-Brown collected the heavy nodule and undertook the initial preparation.

The collaboration between collectors like Buchanan-Brown and professional curators such as Alan Tennyson at Te Papa Tongarewa is a cornerstone of New Zealand paleontology.1 Without the regular patrolling of beaches by locals, many significant fossils—exposed by storms and destroyed by the next tide—would be lost to the ocean. Buchanan-Brown's preparation revealed the unmistakable contours of a massive avian skull, prompting its transfer to the national collection for detailed study.20


3.2. Specimen Description: NMNZ S.048857 - an Ancient Penguin


The fossil consists of a largely complete cranium and rostrum (beak), though it lacks the mandible (lower jaw) and some posterior elements of the braincase.12 Despite these missing pieces, the preservation of the facial region and the beak is exquisite, allowing for precise taxonomic identification.


3.2.1. Taxonomic Diagnosis


The research team assigned the skull to the genus Aptenodytes based on a suite of diagnostic features that distinguish it from all other penguin lineages:

  1. Temporal Fossae: The depressions on the side of the skull that anchor the jaw muscles are widely separated at the midline of the skull roof. In contrast, stem penguins and the genus Spheniscus (e.g., Humboldt penguins) possess temporal fossae that meet at the midline to form a sagittal crest.12

  2. Salt Gland Fossae: Marine birds possess supraorbital glands to excrete excess salt. In the fossil, the fossae for these glands lack a raised lateral border. This feature is present in other crown genera like Eudyptes (crested penguins), Megadyptes (yellow-eyed penguins), and Pygoscelis (brush-tailed penguins), making its absence a key identifier for Aptenodytes.12

  3. Beak Morphology: The rostrum is exceedingly elongate, a trait shared with the King Penguin (A. patagonicus) but distinct from the shorter, stouter bills of most other penguins.11


3.2.2. Dimensions and Scaling


The most immediate impression of the fossil is its size. Comparative morphometric analysis reveals that the skull is approximately 35% larger than that of the Emperor Penguin (A. forsteri).10

  • Emperor Penguin: The largest living penguin, standing up to 1.3 meters tall and weighing 22–45 kg.

  • Tangahoe Giant: While post-cranial bones (femurs, humeri) are needed for a precise mass estimate, scaling isometrically from the skull suggests a bird of immense proportions. It would have undoubtedly exceeded the 45 kg upper limit of modern Emperors, likely occupying a mass range between the modern Emperor and the colossal 150 kg stem-penguins of the Paleocene.7

This finding is significant because it proves that gigantism was not restricted to the archaic, non-crown penguins of the Paleogene (like Kumimanu and Anthropornis). The modern crown-group lineage, specifically Aptenodytes, also evolved giant forms that persisted until geologically recent times.26


3.3. Distinct Species Status


Although the researchers conservatively labeled the fossil as Aptenodytes sp. indet., it is almost certainly a distinct species from the two living forms.

  • Comparison with A. forsteri: The fossil is significantly larger and has a different beak tip morphology.

  • Comparison with A. patagonicus: The fossil is much larger and more robust.

  • Comparison with A. ridgeni: Aptenodytes ridgeni is the only named extinct species of the genus, described by Simpson in 1972 from Pliocene rocks in Canterbury (South Island).12 However, A. ridgeni is known only from leg bones. Since there are no overlapping skeletal elements (skull vs. leg) between the Tangahoe fossil and the ridgeni holotype, they cannot be synonymized. They may represent the same species, or two different giant species inhabiting different parts of Zealandia.11



4. Functional Morphology: The Unhooked Beak



4.1. Rostral Architecture


The beak of the Tangahoe Aptenodytes possesses a unique combination of features that hint at a specialized feeding ecology.

  • Elongation: The upper beak contributes approximately 60% of the total skull length. This extreme elongation is most similar to the King Penguin, a specialist pursuit-diver.11

  • Robustness: The internarial bar (the bone separating the nostrils) is wider and the overall construction of the bill is stouter than in living Aptenodytes.12

  • The Unhooked Tip: Perhaps the most diagnostic feature is the tip of the premaxilla. In modern King and Emperor penguins, the tip of the beak is decurved, forming a slight hook. This hook is functional, aiding in grasping slippery, soft-bodied prey like squid and fish. The Tangahoe fossil, however, has an unhooked tip.11


4.2. Dietary Implications


The "unhooked" morphology suggests that the Tangahoe giant may not have been an obligate teuthophage (squid-eater) like the King Penguin, nor a generalist like the Emperor.

  • Squid vs. Fish: Hooked beaks are adaptations for seizing soft, elusive prey. An unhooked, robust, spear-like bill might indicate a strategy of capturing different prey types.

  • Pliocene Prey Fields: The productive shelf waters of the Tangahoe Formation were rich in large schooling fish and invertebrates. The robust, straight bill could have been adapted for striking larger, more resistant prey, or perhaps for a distinct mode of capture involving impaling or ram-feeding rather than grasping.11

The divergence in beak shape indicates that the Pliocene Aptenodytes occupied a distinct ecological niche from its modern cousins. It was not merely a "scaled-up" Emperor Penguin but a functionally distinct predator adapted to the subtropical prey guilds of Zealandia.12



5. The Tangahoe Ecosystem: A Pliocene Serengeti


The giant Aptenodytes was but one actor in a complex and vibrant marine ecosystem. The Tangahoe Formation has yielded a "Rosetta Stone" of marine vertebrate fossils, allowing us to reconstruct the community structure of the mPWP in remarkable detail.


5.1. The Dawn Monk Seal: Eomonachus belegaerensis


One of the most spectacular co-discoveries in the Tangahoe beds is Eomonachus belegaerensis, the "Dawn Monk Seal from Belegaer" (named after the sea in Tolkien’s Middle Earth).27 Described in 2020, this species overturned the evolutionary history of the Phocidae (true seals).

  • Southern Origins: Previously, monk seals were thought to have evolved in the North Atlantic. Eomonachus proved that the Monachinae (the subfamily including monk, elephant, and Antarctic seals) actually originated in the Southern Hemisphere.19

  • Ecological Role: Eomonachus was a medium-sized pinniped, roughly 2.5 meters long. Its teeth suggest a diet of fish and squid.

  • Interaction: The coexistence of Eomonachus and the giant Aptenodytes proves that mammalian competition was not an absolute barrier to penguin success in the Pliocene. These two groups likely partitioned the resource base, with seals perhaps targeting different prey depths or sizes than the penguins.29


5.2. The Ancestral Little Penguin: Eudyptula wilsonae


At the opposite end of the size spectrum, the formation preserves the delicate skulls of Eudyptula wilsonae, described in 2023.30

  • The Smallest Giant: E. wilsonae is the smallest extinct crown penguin known, slightly smaller and more slender-billed than the modern Little Blue Penguin (Eudyptula minor).

  • Niche Partitioning: The presence of a ~1 kg penguin alongside a ~50+ kg Aptenodytes illustrates extreme niche partitioning. The giant Aptenodytes would have been an offshore forager, capable of deep dives (modern Emperors dive to 500m), while E. wilsonae likely remained in the shallow coastal zone, hunting small shoaling fish.31 This mirrors the modern stratification of seabirds but with a much larger upper size limit.


5.3. The Dawn Crested Penguin: Eudyptes atatu


The Tangahoe Formation also yielded Eudyptes atatu, an ancestor of the modern crested penguins (rockhoppers, macaronis).32

  • Morphological Shift: Unlike modern crested penguins, which have heavy, bulbous bills for feeding on krill and crustaceans, E. atatu possessed a slender, gracile beak.13

  • Evolutionary Signal: This suggests that the heavy "krill-crushing" bill of modern Eudyptes is a recent adaptation, likely evolving in response to the explosion of krill biomass in the Southern Ocean during the Pleistocene cooling. In the Pliocene, crested penguins were likely piscivorous (fish-eating), competing more directly with the smaller Eudyptula than with the giant Aptenodytes.33


5.4. A Sky Full of Tubenoses


The avian community was completed by a diverse assemblage of Procellariiformes (tubenoses).

  • Macronectes tinae: A fossil Giant Petrel. Modern Giant Petrels ("stinkpots") are aggressive scavengers and predators that feed on seal carcasses and penguin chicks. The presence of M. tinae suggests that this scavenging guild was already established, likely feeding on dead Eomonachus and perhaps harassing the Aptenodytes colonies.18

  • Ardenna buchananbrowni: A diving shearwater species, adding to the mid-sized predator guild.24

  • Procellaria altirostris: A deep-billed petrel related to the White-chinned Petrel.34

  • Pelagornithids: The "bony-toothed" birds were also present. These glider-giants, with wingspans up to 6 meters, possessed pseudo-teeth for snatching squid from the surface. Their presence in the Tangahoe Formation represents some of the last records of this group before their global extinction, marking the end of an avian dynasty that had ruled the oceans since the Eocene.35


Table 1: Trophic Structure of the Tangahoe Marine Ecosystem (c. 3 Ma)


Trophic Guild

Representative Species

Estimated Mass

Primary Prey/Role

Apex Predator (Aquatic)

Carcharodon sp. (White Shark)

>1000 kg

Marine mammals, large penguins

Apex Predator (Aerial)

Hieraaetus moorei (Haast's Eagle)*

~15 kg

Large terrestrial/coastal birds

Meso-Predator (Large)

Aptenodytes sp. indet.

>50 kg

Large fish, squid (Offshore)

Meso-Predator (Large)

Eomonachus belegaerensis

~200 kg

Fish, cephalopods (Coastal/Shelf)

Meso-Predator (Medium)

Eudyptes atatu

~4 kg

Fish (Pelagic)

Meso-Predator (Small)

Eudyptula wilsonae

~1 kg

Small fish (Nearshore)

Scavenger/Predator

Macronectes tinae

~4 kg

Carrion, chicks

Surface Feeder

Pelagornis sp.

~30 kg

Surface squid/fish

*Note: Haast's Eagle appears in the fossil record slightly later/contemporaneously in the terrestrial realm, exerting pressure on coastal breeders.



6. The Turnover Hypothesis: Why Did the Giants Fall?


The presence of Aptenodytes in the Pliocene of New Zealand creates a significant biogeographical problem. If these birds could tolerate the warm waters of the mPWP, they were not physiologically restricted to the Antarctic. Why, then, are they extinct in New Zealand today? Why is the genus now restricted to the coldest environments on Earth?

The study by Tennyson et al. proposes a compelling "Turnover Hypothesis" that links the extinction of these giants not to climate change, but to the rise of a new predation pressure that exploited a fundamental vulnerability in their biology.10


6.1. The Vulnerability of the Surface Brooder


The genus Aptenodytes (Kings and Emperors) possesses a unique reproductive strategy among penguins: foot-brooding.

  • Mechanism: Instead of building a nest in a burrow (like Little Penguins) or under dense vegetation (like Yellow-eyed Penguins), Aptenodytes parents incubate a single egg on top of their feet, covering it with a brood pouch of abdominal skin.

  • Behavioral Requirement: This strategy requires the adult to stand upright, relatively immobile, in open areas (beaches or sea ice) for weeks at a time. They cannot hide; they rely on their size and the absence of terrestrial predators for safety.11

In the early Pliocene of Zealandia, this was a viable strategy. New Zealand had no terrestrial mammals. The beaches were safe havens where a 50kg penguin could stand in the sun, incubating its egg without fear of attack.


6.2. The Arrival of the Raptors


The late Pliocene and early Pleistocene saw a dramatic change in New Zealand’s terrestrial predator guild. Large raptors from Australia dispersed across the Tasman Sea and rapidly evolved into apex predators.

  1. Haast’s Eagle (Hieraaetus moorei): Diverging from the small Australian Little Eagle (Hieraaetus morphnoides) around 2.2 million years ago, this bird became the largest eagle in history.11 It evolved to hunt moa (dinornithids) weighing up to 200 kg.

  2. Forbes’ Harrier (Circus teauteensis): Diverging from the Spotted Harrier around 2.4 million years ago, this was a large, generalist raptor.11


6.3. The "Sitting Duck" Scenario


The arrival of these raptors was catastrophic for the surface-brooding Aptenodytes.

  • Mismatch: An Aptenodytes standing on an open beach is a conspicuous, high-energy target. Unlike a moa, which could run or kick, an incubating penguin is encumbered by its egg. It cannot flee without destroying its offspring.

  • Predation Pressure: Haast’s Eagle, capable of killing a 200kg moa with a crushing grip of its talons, would have found a 50kg penguin colony an easily accessible buffet. The open beaches, once safe, became killing fields.

The authors hypothesize that Aptenodytes was driven to extinction in New Zealand by this top-down predation pressure. The penguins could not evolve a new nesting strategy (e.g., burrowing) fast enough to escape the aerial threat. Their rigid adherence to foot-brooding, a trait deeply embedded in their phylogeny, became their death sentence in the presence of large eagles.11


6.4. Comparative Survival


This hypothesis explains why other penguin lineages survived the raptor invasion:

  • Eudyptula (Little Penguins): Nest in deep burrows, completely inaccessible to eagles.

  • Eudyptes (Crested Penguins) & Megadyptes: Nest under the forest canopy, in rock crevices, or in dense scrub, providing visual cover and physical protection from aerial attack.11

The "turnover" was thus a filter: cryptic nesters survived, while the open-nesting giants were culled. The Aptenodytes lineage survived only in the Antarctic and sub-Antarctic islands, where the extreme cold prevented the colonization of large raptors. The "ice" became their refuge, not their preference.



7. Broader Evolutionary Implications



7.1. Fundamental vs. Realized Niche


The Tangahoe discovery provides a classic case study in ecological niche theory.

  • Fundamental Niche: The physiological limits of Aptenodytes allow it to live in subtropical waters (SST ~20°C).

  • Realized Niche: The actual distribution of Aptenodytes is restricted to cold waters (SST < 8°C).

  • Driver: The contraction from the fundamental to the realized niche was driven by biotic interactions (predation), not abiotic constraints (temperature).

This realization has significant implications for conservation. It suggests that King and Emperor penguins might be physiologically capable of surviving in warmer climates than currently assumed, provided they are safe from predation and have adequate food. However, their behavioral rigidity (breeding on ice/open ground) makes them highly vulnerable to habitat loss and new predators.12


7.2. Zealandia: The Museum and the Cradle


The Tangahoe fauna reinforces the dual role of Zealandia in global biodiversity.

  • Cradle: It was the birthplace of the Monachinae seals (Eomonachus) and likely the center of diversification for crown penguins (Eudyptula, Eudyptes).

  • Museum: It acted as a refugium for ancient lineages like Aptenodytes and Pelagornithids well into the Neogene, preserving functional groups (giant penguins) that had disappeared elsewhere.3


7.3. The Modern Parallel


The extinction of the Pliocene giants offers a somber parallel to the present. The arrival of humans in New Zealand (c. 1280 AD) brought a new wave of predators (rats, dogs, stoats) that decimated the surviving ground-nesting birds. Ironically, humans also caused the extinction of Haast’s Eagle (by exterminating its moa prey).

The extinction of the eagle theoretically "re-opened" the niche for Aptenodytes. Indeed, vagrant King Penguins occasionally wash up on New Zealand shores today, and a colony attempted to establish itself on Stewart Island in the past. However, the presence of introduced mammalian predators (cats, ferrets) now acts as the new barrier, preventing the giants from reclaiming their lost subtropical kingdom. The "predation wall" remains, only the predator has changed.11



8. Conclusion


The skull of Aptenodytes sp. indet., retrieved from a Pliocene concretion on a Taranaki beach, is more than a fossil; it is a key to a lost world. It reveals a Zealandia that was warmer, wilder, and more magnificent than the one we know today—a place where Emperor-sized penguins waddled on sun-drenched beaches, sharing the surf with the ancestors of monk seals and the shadows of giant sharks.

This discovery dismantles the assumption that Great Penguins are inherently creatures of the ice. It shows that they were once thermal generalists, lords of the subtropical shelf, whose retreat to the Antarctic was a survival retreat in the face of evolving predation. The "Turnover Hypothesis" reminds us that extinction is rarely mono-causal; it is the result of complex collisions between changing climates, evolving ecosystems, and immutable behaviors.

As we stand on the precipice of a new climatic era, with temperatures threatening to return to Pliocene levels, the Tangahoe giant serves as both a beacon and a warning. It illuminates the incredible plasticity of life in the past, but also its fragility in the face of rapid ecological change. The giants are gone from New Zealand, not because it got too hot, but because the sky turned against them.



Supporting Data Tables



Table 2: Comparative Cranial Metrics of Aptenodytes Species


Feature

Aptenodytes sp. (Tangahoe)

Aptenodytes forsteri (Emperor)

Aptenodytes patagonicus (King)

Skull Size

~35% Larger than Emperor

Reference Standard

Slightly smaller than Emperor

Beak Length

~60% of Skull Length

~50% of Skull Length

~60% of Skull Length

Beak Tip

Unhooked (Straight)

Decurved (Hooked)

Decurved (Hooked)

Internarial Bar

Wide / Robust

Narrower

Narrower

Temporal Fossae

Widely Separated

Widely Separated

Widely Separated

Niche (SST)

Subtropical (14–20°C)

Polar (< 0°C)

Sub-Antarctic (3–8°C)


Table 3: Geological Timeline of New Zealand Penguin Gigantism


Epoch

Key Species

Estimated Mass

Significance

Paleocene (c. 57 Ma)

Kumimanu fordycei

~154 kg

Stem-penguin; largest known penguin.

Eocene (c. 40 Ma)

Anthropornis nordenskjoeldi

~90 kg

Antarctic giant; stem-lineage.

Oligocene (c. 27 Ma)

Kairuku spp.

~60 kg

"Diverse" giant fauna of NZ.

Miocene (c. 15 Ma)

Palaeospheniscus spp.

~40 kg

South American radiation.

Pliocene (c. 3 Ma)

Aptenodytes sp. indet.

>50 kg

Last giant in NZ; crown-group survivor.

Pleistocene (c. 1 Ma)

Aptenodytes forsteri

~40 kg

Restriction to Antarctic refuge.


Report compiled by [Expert Persona: Evolutionary Paleobiologist]


Date: December 1, 2025

Source Material Identifiers: 10 through 37

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