The Réalmont Basin is a small, fault-bounded sedimentary depression in the Tarn department, measuring just seven kilometres from north to south and 2.5 kilometres from east to west. Like dozens of similar basins scattered across the remnants of the Variscan orogeny, it formed within the interior zones of the ancient mountain chain that assembled during the late Palaeozoic. Sediments accumulated in alluvial fans, floodplains, and shallow lakes, preserving a snapshot of continental ecosystems at the Carboniferous-Permian transition. Until now, the only hints of fossil life from this basin were mentions of plant fragments and unidentified fish scales recorded in unpublished reports from the 1970s, which later publications repeated without further detail.

That changed when prospections in 2025 and 2026 at the Cantemerle locality, where the Cantemerle Formation outcrops along a roadside, brought to light a rich collection of vertebrate remains now housed at the Muséum d’Histoire Naturelle de Toulouse. The fossils come from finely laminated black shales interbedded with sandstones, interpreted as deposits of a lacustrine environment periodically interrupted by coarse alluvial debris flows. The articulated fish skeletons are concentrated at the base of one of these shale layers, suggesting rapid burial events that entangled whole bodies before decay could scatter the bones.

Among the most striking finds is a partially preserved fin spine belonging to a hybodontiform shark of the so-called Lissodus morphotype. The spine, measuring an estimated 35 to 45 millimetres in total length, displays smooth longitudinal costae, a strong anterior keel, and a single median row of hooked denticles along its posterior face. These features are characteristic of Palaeozoic hybodontiforms traditionally assigned to the genus Lissodus, though the authors caution that a comprehensive revision of Palaeozoic material has never been undertaken. The Réalmont spine shows a unique combination of seven proximal costae and approximately twelve denticles per centimetre, differing from all comparable Palaeozoic species, and is therefore conservatively assigned to Hybodontiformes indeterminate pending additional remains.

Equally significant is a suite of acanthodid remains, including cranial fragments, scapulocoracoids, fin spines, gill rakers, branchiostegal rays, and several partial postcranial skeletons. Acanthodians, often called spiny sharks, were toothless zooplanktivores that persisted into the early Permian in European freshwater basins. The Réalmont specimens share the combination of a bent, toothless mandibular splint, an inverted Y-shaped scapulocoracoid, smooth sabre-shaped fin spines with a longitudinal groove, and small rhombic scales that define the European Stephano-Permian acanthodids Acanthodes and Westrichus. However, the absence of complete specimens and the plesiomorphic nature of most preserved characters prevent a specific identification, and the material is assigned to Acanthodidae indeterminate.

The centrepiece of the discovery, however, is an abundance of material belonging to the ray-finned fish Progyrolepis cf. heyleri. This predatory actinopterygian, which could reach 70 to 80 centimetres in length, is represented by a fragment of maxilla bearing the diagnostic vermicularly curved ridges of the genus, isolated conical teeth with ornamented ganoine collars, five postcranial skeletons showing signs of skeletal immaturity, and hundreds of isolated scales. The scales display a distinctive ornamentation of diagonally oriented ridges that divide each scale into anteroventral and posterodorsal halves, a pattern shared with the macrodont predatory fish of the late Variscan basins. The postcranial skeletons preserve details of the anal, dorsal, and caudal fins, including basal fulcral scales and grooved lepidotrichia segments, matching the anatomy described for Progyrolepis heyleri from the Bourbon-l’Archambault and Brive basins in central France.

Perhaps the most evocative specimens are four black, shiny coprolites, fossilised faeces, recovered from the same layers. Micro X-ray fluorescence analysis conducted at the Museum für Naturkunde Berlin revealed that the coprolitic matrix is composed primarily of calcium and phosphorus, consistent with calcium phosphate minerals, in sharp contrast to the aluminium, silicon, and iron of the surrounding sediment. Embedded within the matrix of one coprolite are three quadrangular scales whose ornamentation is identical to the mid-body flank scales of Progyrolepis cf. heyleri. The absence of a spiralled structure, which would be expected from a chondrichthyan producer, combined with the crushing dentition of the Lissodus-type shark, points to Progyrolepis itself as the most plausible producer. If correct, this would suggest that the Réalmont predator was not only hunting other fish but consuming members of its own kind, an early hint of cannibalism in a Palaeozoic freshwater ecosystem.

The faunal assemblage also carries important biostratigraphic weight. Progyrolepis heyleri is known only from the early Autunian of the French Massif Central, in deposits lying just above the Autunian-Stephanian C boundary. In the Buxières Formation of the Bourbon-l’Archambault Basin, the vertebrate-bearing level has been radiochronologically dated to 298.592 plus or minus 0.075 million years ago, an early Asselian age close to the Carboniferous-Permian boundary. The presence of the same fish in the Cantemerle Formation confirms the early Autunian age previously proposed on the basis of the macroflora, anchoring the Réalmont Basin firmly in the earliest Permian.

Beyond dating, the discovery reshapes our understanding of Progyrolepis itself. A critical review of the genus’s fossil record reveals that several historical attributions are invalid. The species Progyrolepis tricessimalaris from the early Permian of Texas differs fundamentally from the genus in its cranial anatomy and must be excluded. Fragmentary remains from New Mexico and Colorado lack the diagnostic characters of Progyrolepis and should be treated as indeterminate actinopterygians. Even the long-cited occurrence of Progyrolepis speciosus in the Puertollano Basin of Spain, never described or illustrated, requires verification. When these questionable records are removed, Progyrolepis emerges as a genus restricted to the Stephanian B through early Autunian of the Bohemian Massif and the French Massif Central, with P. speciosus in Bohemia and P. heyleri in France.

This restricted but still substantial distribution challenges a broader pattern. Most families of late-Variscan actinopterygians, including the amblypterids, igornichthyids, and elonichthyids, exhibit strong species-level endemism, with individual species confined to single basins or small clusters of neighbouring basins. The pronounced endemism of these small-bodied fish has been linked to their size, which implies high rates of evolution and rapid local differentiation. Progyrolepis, by contrast, spans both the French and Bohemian massifs, suggesting that its larger body size and correspondingly lower evolutionary rate allowed it to maintain wider geographic ranges even as the landscape dynamics of the Variscan range provincialised its species into separate French and Bohemian populations.

The affinities of Progyrolepis have also been clarified. Previous classifications placed it among the Palaeoniscidae, a wastebasket group, or within the Pygopteridae and Acrolepidae, but the new material and a review of skull anatomy show that these placements are untenable. Recent cladistic analyses of late Variscan actinopterygians place Progyrolepis in a clade of macrodont taxa alongside Briveichthys chantepieorum and the Elonichthyidae, sharing characters such as a large flat maxilla, two rows of marginal teeth, numerous branchiostegal rays, and identical scale ornamentation. The authors refrain from formally assigning Progyrolepis to the Elonichthyidae, noting differences in the lacrimal bone, the absence of a presupracleithrum, and the shape of the maxilla, but the possibility of a close evolutionary link between all Variscan macrodont forms is now firmly on the table.

Finally, the palaeobiogeographic implications of the Réalmont assemblage are profound. The ichthyofauna of the Cantemerle locality is identical to that of the Buxières-les-Mines locality in the Bourbon-l’Archambault Basin, demonstrating an active aquatic connection between the northern and southern basins of the French Massif Central during the early Autunian. At a larger scale, the similarity between the Réalmont fauna and that of the Bohemian basins, where Lissodus and Progyrolepis speciosus co-occur in the Stephanian C, indicates a direct dispersal corridor between the French Massif Central and the Bohemian Massif. Because none of these basins were connected by marine routes at this time, the dispersal must have occurred through continental freshwater pathways, painting a picture of interconnected lake systems threading through the Variscan highlands that allowed large predatory fish to move across hundreds of kilometres of ancient mountain terrain.

Subject of Research: Early Permian fossil fish assemblage and coprolites from the Réalmont Basin, France, and their palaeobiogeographic and taxonomic implications

Article Title: New palaeontological remains from the early Permian Réalmont Basin (Tarn, France): implications for late-Variscan palaeobiogeography and Progyrolepis taxonomy ( Actinopterygii )

Article References: New palaeontological remains from the early Permian Réalmont Basin (Tarn, France): implications for late-Variscan palaeobiogeography and Progyrolepis taxonomy ( Actinopterygii ). (n.d.). https://doi.org/10.3897/fr.29.197176

Image Credits: AI Generated

DOI: 10.3897/fr.29.197176

Keywords: Progyrolepis, Réalmont Basin, early Permian, Variscan Belt, palaeobiogeography, actinopterygii, hybodontiform, acanthodid, coprolites, French Massif Central, Carboniferous-Permian boundary, fossil fish

The two specimens that define the new species were collected in 1978 by the German paleontologist Jürgen Boy at Ruthweiler, near the town of Kusel, but their true distinctiveness went unrecognized for decades. Embedded in light brownish-gray, mica-rich mudstones of the Ruthweiler black shale, the fossils preserve light brown bone with traces of black skin and a faint coating of pyrite. Because the bones and sediment share such similar coloration, the animals are difficult to discern in photographs, which may partly explain why they sat unnoticed within a collection dominated by other, more familiar branchiosaurids.

Branchiosaurids are small, wholly aquatic temnospondyl amphibians that rank among the most abundant vertebrate fossils in the Saar-Nahe Basin, where nearly 200 years of fieldwork by geologists and private collectors has produced an exceptional record of Paleozoic life. Their remains sometimes preserve soft tissues such as skin and even larval gills, making them iconic animals in museum displays. For a long time, the basin’s branchiosaurid fauna was thought to consist almost entirely of a single abundant species, Apateon pedestris, a tiny plankton feeder that apparently spent its entire life in eutrophic lakes set in a mountainous tropical landscape.

The two new specimens overturn that simple picture. The holotype, catalogued as NHMMZ-PW 2022-83-LS, is a larval skull exposed in dorsal view and measuring just 11 millimeters long. The paratype, NHMMZ-PW 2022/399-LS, is a larger, 17-millimeter skull preserved in ventral view, accompanied by the anterior thirteen neural arches, ribs, both clavicles, and an impression of the left humerus. Together they display a suite of features seen in no other regional temnospondyl: a deltoid skull outline, unusually small orbits, and remarkably short frontal bones that occupy only about a quarter to a third of the skull’s length, compared with roughly 0.35 to 0.4 in Apateon.

The maxilla is particularly telling. In both specimens, even the more heavily ossified and presumably mature paratype, the maxilla fails to contact the cheek and carries tiny, closely set, pin-shaped teeth. This open-cheeked, weak-jawed configuration indicates that Schoenfelderpeton brevifrons was likely a plankton feeder, filtering minute food particles from the water much like Apateon pedestris. Because the feature persists in the larger individual, it appears to be a genuine adult condition rather than a larval stage, a case of neoteny in which aquatic, larval traits were retained throughout life. Both specimens also preserve rows of branchial denticles, the pharyngeal teeth characteristic of gill-bearing aquatic larvae.

Detailed anatomical comparison reveals further autapomorphies that justify naming a new species. The squamosal embayment, the notch at the back corner of the skull where the jaw muscles once passed, is unusually shallow and sits at an unusually large distance from the orbit, and it bears a small rounded pocket wedging between the squamosal and tabular bones. The posterior margin of the postparietal is strongly convex, and the parasphenoid, a bone forming much of the braincase floor, carries a wide, short basal plate and a straight, unexpanded cultriform process. The palatine is exceptionally narrow, framing a slit-like choana, the internal nostril, an almost unique construction among branchiosaurids.

To determine where the new animal belongs on the amphibian family tree, the researchers added its anatomical characters to an existing matrix of 48 taxa and 115 characters and analyzed it with the software TNT, using New Technology search algorithms including Sectorial Search, Ratchet, and Tree Fusing across 1,000 random addition sequences. All characters were left unordered, since evolutionary transformation sequences are hard to assess in a group so pervasively shaped by neoteny. The analysis recovered a single most parsimonious tree of length 337, with a consistency index of 0.397 and a retention index of 0.731.

The result places Schoenfelderpeton brevifrons as the sister species of Schoenfelderpeton prescheri, a highly paedomorphic form previously known from Thuringia and Saxony, and nests the pair within a larger clade alongside Leptorophus and the species informally known as “Melanerpeton” humbergense. Because the earlier subclade name Leptorophinae turned out to be preoccupied by a reptiliomorph group based on the Russian genus Leptoropha, the study proposes the new name Leptobranchinae, from the Greek words for light and gill, for this branchiosaurid subclade. The broader tree also reshuffles relationships among other Permian lineages, revealing, for the first time, two distinct sister groups of early Permian branchiosaurids: one uniting Melanerpeton sensu stricto, Xerodromeus, and Apateon, and the other comprising the leptobranchines.

The evolutionary implications are rich. The leptobranchines shared slender palatine and ectopterygoid bones with weak dentition, and some, including the Leptorophus subclade, sported large conical teeth indicating they were small predators of insects, crustaceans, and even other branchiosaurid larvae. The two Schoenfelderpeton species, by contrast, evolved or re-evolved the open cheek and pin-like dentition of plankton feeders, paralleling the condition seen in certain populations of Apateon. This repeated convergence, in which distant relatives independently arrived at the same filter-feeding anatomy, illustrates how strongly the ecological pressures of Permian lake ecosystems molded these animals’ bodies.

Schoenfelderpeton brevifrons inhabited the Ruthweiler black shale, a regionally restricted lake deposit laid down during the final phase of the M8 sequence, above the famous Odernheim limestone unit. The Meisenheim Formation is believed to be of Asselian age, roughly 295 to 300 million years ago. The new species lived in a tetrapod community strikingly devoid of fishes, dominated by the abundant “Melanerpeton” humbergense and the large Apateon caducus, a putative apex predator of that setting. In this fishless lake, a modest plankton-eating amphibian with a delta-shaped head found its own niche. Known so far only from this single locality, Schoenfelderpeton brevifrons stands as a reminder that even collections gathered nearly half a century ago can still yield species entirely new to science, and that the diversity of Permian amphibians in Europe remains far from fully charted.

Subject of Research: A new branchiosaurid temnospondyl amphibian from the early Permian of Germany

Article Title: A new delta-headed branchiosaurid temnospondyl from the early Permian Meisenheim Formation of Germany

Article References: Schoch, R. R. (2026). A new delta-headed branchiosaurid temnospondyl from the early Permian Meisenheim Formation of Germany. Fossil Record, 29(2), 765-775. https://doi.org/10.3897/fr.29.209145

Image Credits: AI Generated

DOI: 10.3897/fr.29.209145

Keywords: Schoenfelderpeton brevifrons, branchiosaurid, temnospondyl, early Permian, Saar-Nahe Basin, Meisenheim Formation, paleontology, neoteny, Germany, amphibian evolution, phylogenetics, Ruthweiler

The specimen was recovered from a gray siltstone layer overlying Coal No. 4 at the Antaibao Surface Mine near Shuozhou City, a site within the famed Shanxi Formation. That formation represents a marine-nonmarine transitional environment where swampy peat accumulated on floodplains fed by braided rivers. The regional plant assemblage, characteristic of the Cathaysian Flora, includes index species such as Emplectopteris triangularis and Taeniopteris mucronata, and high-precision uranium-lead dating of nearby strata has pinned the Shanxi Formation to the Asselian stage of the Early Permian. The fossil itself consists of two facing parts of a cast, each preserving three partially preserved fruits arranged in parallel and oriented the same way.

Morphologically, each fruit is strap-shaped, with nearly parallel sides, preserved lengths of 4.2 to 4.8 millimeters and widths of 1.2 to 1.3 millimeters. All three structures display a terminal zone free of any reproductive bodies, a detail the team interprets as a seed-free, styleless fruit tip. Inside the fertile regions are three to six globular bodies of varying size and form. Larger bodies measure 1.15 to 1.4 millimeters across, while smaller ones range from 0.36 to 0.57 millimeters. The larger bodies bear vermiform, worm-like surface sculpture interpreted as a seed coat, and their discoid, biconcave form is striking, since such doubly concave seeds are known in some living angiosperms, such as the moonseed family Menispermaceae, but have never been documented in any extinct or extant gymnosperm.

To examine the internal anatomy without damaging the precious specimen, the researchers subjected it to micro-computed tomography at the State Key Laboratory of Continental Dynamics at Northwest University in Xi’an. Scanning at 170 kilovolts and 200 microamperes, they collected 2,000 projections at a resolution of 26.22 micrometers and reconstructed the fossil in three dimensions using VG Studio 3.2. The virtual sections revealed seeds and ovules attached along two lines following the margins of the fruit walls, with some bodies positioned along the upper, adaxial side and others along the lower, abaxial side. This arrangement of multiple enclosed bodies at different developmental stages is central to the team’s case that the organs represent genuine carpels rather than any known gymnospermous structure.

The key diagnostic trait is what botanists call angiospermy, or angio-ovuly: the enclosure of ovules, and ultimately seeds, within a closed organ before pollination. This feature has defined flowering plants since the term angiosperm was coined in 1690 by the German botanist Paul Hermann, and it was the basis on which the celebrated Jurassic fossil Archaefructus liaoningensis was recognized as an early angiosperm in 1998. The presence of both smaller bodies, interpreted as unfertilized ovules or abortive seeds, and larger, seed-coated bodies within the same fruit satisfies both the original and stricter versions of this criterion. The authors acknowledge caveats: some conifers and the seed fern Caytonia do enclose seeds after pollination, but they argue that seed position and overall organ morphology in those groups differ fundamentally from what Permocarpelloides displays.

Beyond the mere presence of enclosed ovules, the orderly, parallel arrangement of the three fruits carries deep evolutionary significance. Because the three structures lie so close together and share the same orientation, the team infers they belonged to a single gynoecium, the female reproductive apparatus of one flower. Their clear separation from one another implies that this gynoecium was apocarpous, meaning it was composed of multiple free, distinct carpels, exactly the condition seen in magnoliids such as magnolias and in the basal angiosperm Amborella. Traditional theories of angiosperm evolution, stretching back to Arber and Parkin’s influential 1907 hypothesis, have held that such free, apocarpous gynoecia were ancestral, making Permocarpelloides the first Permian fossil to match those theoretical expectations so directly.

Yet the fossil also complicates the old textbooks. Classical theory, reinforced by Canright’s 1960 study of magnoliaceous carpels and by the celebrated Cretaceous fossils Archaeanthus and Archaefructus, expects seeds to be borne along the adaxial, upper side of the carpel, reflecting the idea that a carpel is a folded megasporophyll, a seed-bearing leaf. In Permocarpelloides, seeds attach along both sides of the fruit, and the coexistence of seeds at markedly different developmental stages in a single organ is highly unusual. The authors note that ovule position has proven far more variable than the classical model allows: recent work on living Illicium and Michelia, reinvestigation of the Archaeanthus holotype, and studies of the Jurassic fossil Xenofructus all indicate that the placenta may originally have been an ovule-bearing branch independent of any leaf margin, casting doubt on the venerable megasporophyll concept itself.

Permocarpelloides is not an isolated anomaly. It joins a growing roster of putative Early Permian angiosperms from China, including Taiyuanostachya, reported in 2023; Yuzhoua juvenilis, described in 2025; and Dengfengfructus maxima, announced earlier in 2026. Compared with those fossils, Permocarpelloides more closely resembles the carpels of modern flowering plants, thanks to its free carpels and apocarpous organization. The striking morphological diversity among these four distinct Permian fossils suggests that flowering plants were already diversified, and therefore must have originated even earlier than the Asselian stage. The authors argue that the long-standing no-angiosperms-until-Cretaceous belief can no longer accommodate the accumulating evidence and that the standard scenario of angiosperm evolution urgently needs updating.

Independent lines of evidence appear to converge on the same conclusion. Molecular clock analyses, from early amino acid and chloroplast DNA studies in 1989 through recent phylogenomic work in 2024 and 2025, have repeatedly estimated pre-Cretaceous origins for flowering plants, and a 2026 integrated analysis of fossils and molecular divergence times points to a latest Jurassic origin at minimum. Intriguingly, the earliest pollen-loaded insects are known from the Lower Permian of Russia, hinting that insect pollination relationships may have been established far earlier than assumed. Together, the fossil record, the insect record and molecular dating sketch a coherent picture of a long, hidden history of angiosperms stretching back at least 295 million years. Whether Permocarpelloides ultimately proves to be a bona fide angiosperm or an unprecedented gymnosperm exhibiting angio-ovuly, its discovery blurs a boundary botanists have drawn for centuries and is likely to ignite a new wave of debate and discovery in the study of seed plant evolution.

The naming of the new genus itself carries meaning. The prefix Permo- signals the fossil’s surprising age, while the suffix -oides, meaning resembling, reflects the authors’ caution: the organs are only partially preserved, and no complete fruit was recovered. Because ovules and immature seeds are difficult to distinguish even in living plants, the team uses the terms carpel and fruit, and ovule and seed, interchangeably throughout their description, a pragmatic choice that underscores how much interpretation rests on a single specimen.

The preservation context is also instructive. Coal-bearing sequences of the Shanxi Formation have long been worked for economic coal, with Coal No. 4 averaging roughly thirteen meters thick at the Antaibao mine, but such commercial exposures rarely attract attention from palaeobotanists hunting for delicate reproductive structures. That a carpel-like fossil emerged from a siltstone immediately above a major coal seam suggests that similar material may await discovery in museum drawers and mine sections across North China, where the Cathaysian Flora has been documented for decades yet never before yielded organs of this kind.

The non-destructive imaging approach deserves emphasis. Traditional palaeobotany often relies on dégagement, physically chipping away matrix, or on serial sectioning, both of which consume irreplaceable material. Micro-computed tomography at the 26-micrometer scale achieved here allows seeds to be traced to their attachment points on the fruit wall in three dimensions, providing the positional evidence that two-dimensional photographs alone could not deliver. As such scanning becomes routine, comparable fossils from other Permian localities may be re-examined with fresh eyes.

For now, the specimen rests in the type collection under catalogue number PB206132, available for scrutiny by other researchers. The open-access publication ensures that the images and reconstructions can be independently assessed, an important step given how consequential the claim is for one of botany’s most enduring questions.

Subject of Research: Permian carpel-like fossil organs and the pre-Cretaceous origin of angiosperms

Article Title: Permian carpel-like organs and their implications on origin of angiosperms

Article References: Wang, X., Fu, Q., Huang, W., & Sun, J. (2026). Permian carpel-like organs and their implications on origin of angiosperms. Plant Biosystems, 160(5), Article 257. https://doi.org/10.1007/s44473-026-00267-6

Image Credits: AI Generated

DOI: 10.1007/s44473-026-00267-6

Keywords: angiosperm origin, carpel, Permian, Permocarpelloides, fossil, paleobotany, enclosed ovules, apocarpous gynoecium, China, Shanxi Formation, seed plant evolution, Early Permian