taxonID	type	description	language	source
03B72C1C3020902CFEFBFC483893FAE9.taxon	description	urn: lsid: zoobank. org: act: 7407645 D- 1643 - 4 B 4 E-ADCE-FF 29 C 837 CC 56	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020902CFEFBFC483893FAE9.taxon	etymology	ETYMOLOGY. — Aemilia from the Latin aemulus meaning emulate, referencing the completeness of its hyobranchial skeleton, rivalling with that of the extant Latimeria.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020902CFEFBFC483893FAE9.taxon	type_taxon	TYPE SPECIES. — Aemilia stellata n. sp.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020902CFEFBFC483893FAE9.taxon	diagnosis	DIAGNOSIS. — Same as for the type species, by monotypy.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	description	(Figs 2 - 13) urn: lsid: zoobank. org: act: 15405 ABA- 87 C 4 - 4 D 88 - 9209 - 2 A 6 C 8 DB 118 C 4	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	materials_examined	TYPE MATERIAL. — Holotype. United States • Texas, Jack County, northeast of Jacksboro, TXV 200 Mapes locality; AMNH FF 20686, nodule comprising a partially preserved braincase and associated elements of the skull roof, cheek, hyoid and branchial arches, and shoulder girdle in close articulation (Fig. 2 A-D).	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	etymology	ETYMOLOGY. — stellata from the Latin stella meaning star, referencing the Lone Star State of Texas (United States) where the specimen was found.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	diagnosis	DIAGNOSIS. — Coelacanth characterized by the following unique association of characters: descending process in the form of a longitudinal ridge on the ventral surface of the postparietal; presence of pores on the posterior margin of the postparietal corresponding to anterior branches of the occipital commissure from the median extrascapular; squamosal and postorbital ornamented with pointed tubercles; dentition of the ectopterygoid composed of a single row of small teeth and a large fang; braincase composed of separate ossifications (parasphenoid, basisphenoid, prootics, and basioccipital); buccohypophysial canal open on the parasphenoid; prominent prefacial eminence on the prootic; presence of a postero-dorsal process on the prootic for the contact with the tabular; complex suture between the prootic and the basioccipital; cleithrum and extracleithrum ornamented with tubercles; fully ossified boomerang-shaped first pharyngobranchial; fully ossified fourth epibranchial.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	distribution	TYPE LOCALITY. — United States, Texas, Jack County, northeast of Jacksboro, TXV 200 Mapes locality.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	biology_ecology	TYPE HORIZON AND AGE. — Finis Shale member, Graham Formation, Cisco Group, Lower Gzhelian (Virgilian), Upper Carboniferous (Pennsylvanian).	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	description	DESCRIPTION Dermal skull The dermocranium is only poorly represented and comprises parts of the skull roof (postparietals, tabulars, and possible supraorbitals) and the cheek (postorbital, squamosal). Skull roof Postparietal. The two postparietals (Pp; Figs 2 E; 3 E, H; 5 A-D) are broad and overall flattened bones but are poorly preserved, lacking most of their anterior and lateral margins; only the medial margin displaying a straight median suture (m. s; Fig. 5 B, D) and the posterior margin of the right element can be identified confidently. The inner surface is smooth and otherwise featureless, but a reduced descending process (d. p. Pp; Fig. 5 D) in the form of a longitudinal ridge is present on the left postparietal, running diagonally across the anterior portion of the bone. The course of the otic canal on the antero-lateral margin cannot be reconstructed. The presence of pores in the posterior portion of both postparietals (po. a. occ; Fig. 5 B) are associated with anterior branches of the occipital commissure (corresponding to the supratemporal commissure). Tabular. The two disarticulated tabulars (Ta; Figs 2 E; 3 E, H; 5 A-D; corresponding to the supratemporal of Forey 1998) are preserved; the right one (Ta. r; Figs 2 E; 3 E, H; 5 F, H) is fairly complete, whereas the left one (Ta. l; Figs 3 E, H) is only represented by a small portion of a canal-bearing process. The exposed surface of the right tabular is densely pitted and pierced by large pores associated with the trajectory of the otic sensory canal (po. oc; Fig. 5 F). The medial and lateral margins display articulation surfaces for the adjoining bones in the form of depressed smooth flanges separated from the elevated exposed and ornamented surface of the bone. The lateral margin shows ventrally a moderate process, probably for the attachment of the opercular ligament (op. lig; Fig. 5 F, H) as in Latimeria chalumnae and other fossil coelacanths (Forey 1998). The posterior margin carries a small shelf (po. sf; Fig. 5 F) projecting postero-ventrally with a distinctive concave, oval depression, certainly for the articulation of a convex process of the unpreserved lateral extrascapular. The ventral surface is smooth, and no distinctive ventral process has been identified. The otic canal (oc; Fig. 5 F, H) penetrates the tabular through an antero-medial process and exits posteriorly through a large single opening. There is no evidence of a triple junction of the otic, occipital commissure and main lateral line canals in the tabular, which most probably occurred in the lateral extrascapular. Supraorbitals. A small bony element, ornamented with sparse rounded tubercles, lies dorsal to the postorbital on the right side of the specimen. Given its small size and ornamentation, it is similar to a spiracular (corresponding to the postspiracular of other sarcopterygians). However, it carries a well-developed canal visible in internal view and thus it may correspond to the posteriormost supraorbital bone (So. r; Figs 2 E, G; 3 E, H). A similar element lies on the left side of the specimen (So. l; Figs 2 H; 3 E, H), it is a small ossification with an enlarged middle section, probably for the course of the supraorbital sensory canal. However, as opposed to the one on the right side, the left element is not ornamented, possibly due to a greater erosion of the left side of the fossil nodule. Several small, ossified elements displaced from their original position are located between the branchial arches, but are not visible in the figures (see Appendix 2). They are roughly square elements, usually pierced by pores related to a possible internal canal and display dorsal and ventral gutters. We tentatively assign them to the supraorbital series due to their similarity with the supraorbitals of Graulia branchiodonta (Manuelli et al. 2024). Cheek Postorbital. The right postorbital (Po. r; Figs 2 E, G; 3 E, G) is incompletely preserved, slightly displaced from its original position and overlapping the antero-ventral portion of the squamosal. The preserved fragment is roughly triangular and does not show any trace of a lateral sensory canal, usually found in the anterior most region, suggesting that the bone lacks the anterior portion and was quite large posteriorly. The external surface is profusely ornamented by rounded to pointed tubercles. Squamosal. The right squamosal (Sq. r; Figs 2 E, G, 3 E-G) is the largest bone retrieved from the cheek; however, it is incompletely preserved. The external surface is richly ornamented with tubercles, more elongate and antero-posteriorly oriented dorsally, and more rounded ventrally. The internal surface displays a prominent canal for the horizontal course of the jugal canal. The ventral margin shows an overlapping facet for a putative preopercular. Palate Several elements of the palate have been identified: both palatoquadrates, a left ectopterygoid, and the parasphenoid. Palatoquadrate. The palatoquadrate complex (Pq; Figs 2 G, H; 3 E-H) is a single ossification, roughly triangular in shape, comprising both dermal (entopterygoid) and endoskeletal (quadrate and metapterygoid) elements. Only the entopterygoids appear to be preserved and no distinctive autopalatines have been found. Entopterygoid. Both entopterygoids (Entp; Fig. 6 D, E) are almost entirely preserved, only the anterior most portion is missing. The lingual side is slightly concave, covered in a shagreen of small denticles arranged in concentric lines following the postero-dorsal margins of the bone (Fig. 2 C, D). In posterior view, a groove on the postero-dorsal margin corresponds to the spiraculo-hyomandibular recess (re. sh; Fig. 6 E, F). The ventral margin of the anterior projection is straight and carries a narrow antero-posterior depression, likely for the contact with the lingual lamina of the ectopterygoid. The postero-ventral corner displays a ventral projection probably fitting inside a groove for the articulation with the unpreserved quadrate. Ectopterygoid. The right ectopterygoid (Ectp; Fig. 6 L, N, P, Q) is elongate, triangular in dorsal view, carrying teeth on a horizontal lamina close to the labial side. Its small size may result from a lack of preservation of its most anterior portion. The dentition is organised into a single row of small teeth (t; Fig. 6 L, N, R) associated with a large, pointed, posteriorly recurved fang (f; Fig. 6 L, N, R), flanked by two replacement sockets (r. s; Fig. 6 L, N, R). The dorsal surface carries a deep groove for the articulation with the ventral margin of the anterior projection of the entopterygoid (gr. Entp; Fig. 6 L, P). Parasphenoid. The parasphenoid (Psph; Figs 3 E, F, H; 6 H, J; 9 A-E) is only partially preserved posteriorly; the anterior denticulated section is missing. The ventral surface is convex and displays an hour-glass-shaped median section that broadens posteriorly. The dorsal surface is open, composed of two laterally depressed flanges delimiting a median furrow between their parallel ridges. The posterior margin is broad and comprises a deep pit to accommodate the anterior expansion of the basisphenoid, dorsally articulating with the latter. A foramen in the anterior extremity of the preserved section of the parasphenoid indicates that the buccohypophysial canal (bh. c; Figs 3 E, F, H; 6 H, J) was open. Braincase Numerous endoskeletal elements of the neurocranium have been preserved and include: the basisphenoid, basioccipital, and the paired prootics. The zygal series comprises the anazygal, and the anterior and posterior catazygals. The separate ossifications of the parasphenoid, basisphenoid, prootics, and basioccipital confirm that the braincase was not ossified as a single unit. No supraoccipital or exoccipitals have been found. Basisphenoid. The basisphenoid (Bsph; Figs 2 E, H; 3 E, H, 7 B, D; 9 A-E) is incompletely preserved, with its dorsal surface exposed in the nodule. The paired processus connectens (pr. con; Fig. 7 B) are mildly developed as blunt, elongate, and anteriorly tapering lateral margins; they did not contact the parasphenoid and were certainly capped by large cartilage pads to articulate with the otic shelves of the prootics. The antero-lateral margins of the basisphenoid are broken with only the broad proximal portion of the antotic processes being preserved. A pair of sphenoid condyles (sph. co; Fig. 7 B, D) are well developed posteriorly and separated by a gap to articulate with the anazygal. The dorsum sellae (ds; Fig. 7 B) forms the elevated merging surface between two triangular slight depressions dorsal to the processus connectens laterally and the sphenoid condyles posteriorly. Anterior to the dorsum sellae, two thin parallel ridges frame a narrow groove likely for the course of the hypophysial canal (gr. hyp; Fig. 7 B). In ventral view, a large notochordal pit (ncp; Fig. 7 D) is anteriorly bordered by a small ridge and laterally by the ventral edge of the processus connectens. Anterior to it, two lateral deep notches, seemingly converging anteriorly, mark the trajectory of the internal carotid arteries (gr. car; Fig. 7 D). No distinct foramina can be reconstructed suggesting that the oculomotor (III) and profundus (V 1) nerves left the cranial cavity anterior to the basisphenoid. There is no evidence of a basipterygoid nor of a suprapterygoid process. Prootic. The two prootics (Pro; Figs 3 E, H; 8; 9 A-E) are fully preserved as separate elements of the otico-occipital portion of the neurocranium. The otic shelf (ot. sh; Fig. 8 D, F, H) is short, with a concave dorsal margin, and carries a groove to accommodate the processus connectens of the basisphenoid. The dorsal and postero-lateral margins appear jagged owing to the lack of preservation of an important cartilaginous connection with the rest of the braincase. The hyomandibular facet (hyo. fa; Fig. 8 B) is postero-laterally directed but cannot be confidently reconstructed as the entire lateral commissure (lat. co; Fig. 8 B, F) was entirely made of cartilage. Similarly, any putative articulatory surfaces for the pharyngo- and epibranchial elements of the branchial series have not been preserved. Dorsal to the otic shelf, a prominent antero-dorsal projection (the prefacial eminence [pref. e]; Fig. 8 B, D, F, H) carries an overlapping surface, probably for the contact with the ventral ridge of the postparietal. Medial to it, a postero-dorsally oriented process displays a small overlapping surface on its dorsal edge, probably for a contact with the tabular (ov. Ta?; Fig. 8 D, F, J), but this is difficult to ascertain. The medial surface is smooth and slightly concave, framing the notochord. The ventral surface carries a groove for the course of the basicranial muscle (gr. m. bc; Fig. 8 B), which probably inserted on the posterior wing of the prootic, medially framed by a sharp ridge forming the medial wall of the notochordal canal (n. c; Fig. 8 D, H) and laterally by the lateral commissure. The canal for the otic ramus of the facial nerve (c. n. VII; Fig. 8 D, F) opens dorsally through a foramen at the medial base of the prefacial eminence, dorsal to the posterior edge of the otic shelf, and exits laterally through a large common foramen for the jugular canal (c. ju; Fig. 8 B, F, H). The canal for the jugular vein (c. v. ju; Fig. 8 B, J) crosses the prootic antero-posteriorly through a mildly-developed groove, entering the bone between the lateral commissure and the prefacial eminence, and exiting lateral to the saccular chamber through a large foramen associated with the passage of the hyomandibular ramus of the facial nerve. The large saccular chamber (or otic capsule [ot. ca]; Fig. 8 F, J) is ovoid, postero-dorsally open, and laterally framed by the irregularly delimited lateral commissure. The medial wall (m. w; Fig. 8 D, J) separates the saccular chamber from the notochordal canal and displays a serrated outline, with a convex postero-dorsal expansion. The floor of the otic capsule projects posteriorly into the posterior wing of the prootic (p. w. Pro; Fig. 8 B, D, F, J), which is medially inclined and terminated by a complex interdigitated margin for the suture with the basioccipital. Basioccipital. The basioccipital (Boc; Figs 7 R, T; 9 A-E) is entirely preserved, detached from the rest of the braincase. It has a semilunar shape, dorsally concave and ventrally convex. The dorsal surface is smooth and perfectly semicircular, accommodating the notochord. The ventral surface displays a median ridge that bifurcates anteriorly towards two antero-lateral projections. These antero-lateral projections have an interdigitating anterior margin to match the posterior wings of the prootics. The lateral margins are incompletely ossified, suggesting the occurrence of a cartilaginous connection with the prootics. Zygals. The complete zygal series is preserved, represented by the anazygal and the anterior and posterior catazygals. The anazygal lies dorsal to the notochord, articulating with the basisphenoid. The catazygals ventrally frame the notochord, occupying the basicranial fenestra, and they are posteriorly followed by the basioccipital. Anazygal. The anazygal (Az; Figs 7 F, H; 9 A-E) has a quadrangular saddle-shape, ventrally concave and dorsally convex with a faint midline groove. It articulates with the basisphenoid condyles via two well-developed anterior depressions separated by a narrow gap. The bone is well ossified, but the lateral margins are unfinished suggesting the occurrence of cartilaginous lateral extensions. Catazygals. The catazygals are crescent-shaped bones, dorsally concave and ventrally convex, without articulation facets as they do not contact any bone of the neurocranium. The anterior catazygal (a. Cz; Figs 7 J, L; 9 A-E) is slightly larger than the posterior catazygal (p. Cz; Figs 7 N, P; 9 A-E), which is less ossified along its median portion than the anterior catazygal. Hyoid and branchial arches The hyoid and branchial arches are overall complete on each side of the skull. The hyoid arch comprises the interhyal, symplectic, and fragments of the ceratohyal. There are five branchial arches composed of five ceratobranchials, four epibranchials and three pharyngobranchials. No distinctive hypohyals or hypobranchials have been identified. A single basibranchial is entirely preserved while the anterior portion of the urohyal is missing. Hyoid arch Interhyal. The interhyals (Ih; Figs 3 E-H; 4 E-H; 10 D, H; 12 A-E) consist of small, elongate, weakly-ossified tubular bones made of thin periosteal walls. They lie near to the dorsal fragment of the ceratohyals and the symplectics and would have articulated ventrally with the latter. Symplectic. The symplectics (Sy; Figs 3 E-H; 4 E-H; 10 F, J; 12 A-E) are also elongate with thin periosteal walls, and they display a characteristic inverted tripodal shape, slightly curved inwards in dorsal view. The ventral portion is cylindrical and hollow with a circular ventral extremity. The enlarged triangular dorsal portion shows a concave inner surface, probably for the course of the posterior mandibulohyoid ligament (gr. lig. pmh; Fig. 10 J), running on the medial side of the symplectic, connecting the retroarticular to the interhyal (Forey 1998). Ceratohyal. The two ceratohyals (Ch; Figs 2 F-H; 3 E-H; 4 E-H; 10 B; 12 A-E) are incompletely preserved. Most of the bones’ contour has been preserved as imprints in the outer surface of the specimen and only fragments of the posterior (dorsal) portion were modelled. The ceratohyals are curved and elongate, with a well-developed and characteristic ventro-lateral flange (v. fl. Ch; Figs 2 H; 10 A), visible as an imprint on the left side of the nodule. The preserved dorsal portion of the left ceratohyal is triangular in shape with a concave lateral surface and a large extremity to accommodate a probably well-developed cartilage pad for the articulation with the symplectic and interhyal, as in Latimeria chalumnae (Millot & Anthony 1958). Branchial arches Pharyngobranchials. Three pairs of pharyngobranchials (Pb; Figs 3 E-H; 4 E-H; 11 G, I; 12 A-E) are preserved on each side of the skull. The first pharyngobranchial (Pb 1; Figs 3 E-H; 4 E-H; 11 G; 12 A-E) has a distinctive boomerang shape and displays three surfaces of articulation: a large ventral one in the shape of a bean for the first epibranchial (art. Eb 1; Fig. 11 G) and two reduced dorsal surfaces at the extremities of long processes, which can be recognised as the infra- and suprapharyngobranchial portions. The infra- and suprapharyngobranchial processes differ in width, with the suprapharyngobranchial portion (s. Pb 1; Fig. 11 G) being larger and cylindrical in outline, whereas the infrapharyngobranchial portion (i. Pb 1; Fig. 11 G) is slender and gently tapers anteriorly. The articulation surface of the suprapharyngobranchial portion might have contacted the so-called parampullary process of the prootic (art. pa. pr; Fig. 11 G) as known in other coelacanths like Diplocercides, Rhabdoderma or Latimeria (Jarvik 1954; Forey 1998). The infrapharyngobranchial portion carries a small articulation surface anteriorly, probably contacting an unpreserved articulatory facet on the postero-lateral margin of the prootic (art. Pro; Fig. 11 G). The rest of the pharyngobranchial series is difficult to identify and may correspond to separate supra- and infrapharyngobranchials from the second arch (? Pb 2; Figs 3 E-H; 4 E-H; 11 I; 12 A-E;? Pb 2 l, r. 14 A, C-D), represented by small rod-like elements decreasing in size medially of which only a small portion of the middle section was lightly ossified. Epibranchials. The epibranchials (Eb; Figs 3 E-H; 4 E-H; 11 K, M, O, Q, S; 12 A-E) are robust bones, four in number on each side of the specimen and variable in size and shape. The first two epibranchials (Eb 1, Eb 2; Figs 3 E-H; 4 E-H; 11 K, M; 12 A-E) are somewhat pear-shaped with a well-developed longitudinal ridge and associated dorsal groove (gr. a. br; Fig. 11 M). Their anterior (dorsal) articulation head is expanded, roughly triangular in outline, and larger than the more circular proximal (ventral) extremity, articulating with the ceratobranchial (art. Cb; Fig. 11 K, M). The third epibranchial (Eb 3; Figs 3 E-H; 4 E-H; 11 O, Q; 12 A-E) is the largest of the series and carries a large canal for the passage of the efferent branchial artery (ca. a. br; Fig. 11 Q) that pierces the bone antero-posteriorly, with the posterior opening prolonging itself into a postero-dorsal groove (gr. a. br; Fig. 11 O). The most medial epibranchial (Eb 4; Figs 3 E-H; 4 E-H; 11 S; 12 A-E) is again more elongate with a slightly-developed dorsal process displaying an antero-dorsal foramen for the efferent branchial artery (f. a. br; Fig. 11 S), as in Latimeria chalumnae (Millot & Anthony 1958). There is no fifth epibranchial. Ceratobranchials. The ceratobranchials (Cb; Figs 3 E-H; 4 E-H, 11 B, D, F; 12 A-E) are the largest elements of the branchial arches. They are five in number on each side, decreasing in size postero-medially, and all display a similar curved shape with an elongate, cylindrical anterior (ventral) section and a shorter, cylindrical, oval or trapezoidal (depending of the element) posterior (dorsal) section. The cross sections reveal a weakly developed ossification, decreasing medially along the series, with a hollow internal cavity and thin periosteal walls. The anterior (ventral) tips are circular in outline and appear open since they were probably capped with large cartilaginous heads. The first pair of ceratobranchials (Cb 1; Figs 3 E-H; 4 E-H; 11 B, D; 12 A-E) are the longest ones but both are anteriorly broken with the missing section preserved as an imprint in the fossil nodule. The posterior (dorsal) tips of the ceratobranchials 3 - 5 are also missing due a posterior breakage of the nodule. Ceratobranchials 1 and 2 are completely preserved posteriorly (dorsally) but the lack of cartilage heads hampers a confident reconstruction of their relationship with the epibranchials (art. Eb 2; Fig. 11 B). The ventral surface of the ceratobranchials 1 - 4 (Cb 1 - 4; Figs 3 E-H; 4 E-H; 11 B, D; 12 A-E) carries a large groove (gr. Cb; Fig. 11 D) for the passage of the afferent and efferent branchial arteries. The first four ceratobranchials (Cb 1 - 4) were connected to the basibranchial (art. Bb; Fig. 11 B). The fifth ceratobranchial (Cb 5; Figs 3 E-H; 4 E-H; 11 F; 12 A-E) is the smallest of the series, lacks a groove and is inwardly curved, articulating anteriorly and posteriorly with the ceratobranchial 4 (art. Cb 4; Fig. 11 F) as in Latimeria chalumnae (Millot & Anthony 1958; Datovo & Johnson 2025) and other sarcopterygians (Janvier 1996). Basibranchial series Basibranchial. A single basibranchial (Bb; Figs 3 E-H; 4 E, F; 10 P, R; 12 A-E) is entirely preserved. It is mainly perichondrally ossified and roughly hexagonal in outline, without traces of partial division. In lateral view, the anterior portion is thickened, becoming more flattened towards the rear. The anterior margin presents a large, inverted trapezoidal surface towards the symphysis of the lower jaw that carries two dorsal depressions for the articulation with the ceratohyals (art. Ch; Fig. 10 R). The dorsal surface is slightly convex with two lateral depressions bordering a more elevated median section. The ventral surface displays a ventral process in the shape of a shelf, anteriorly bordering the circular facet for the articulation with the urohyal (art. Uh; Fig. 10 R). The lateral margins display two slightly concave articulatory surfaces for the ceratobranchials 1 - 4 (art. Cb 1 - 4; Fig. 10 R) that were probably covered by large cartilage pads, as in Latimeria chalumnae (Millot & Anthony 1958). Urohyal. The well-ossified urohyal (Uh; Figs 2 F; 3 F-H; 4 E-H; 10 L, N; 12 A-E) is only partially preserved on its posterior third while the middle section is visible as an imprint in the fossil specimen. It is dorso-ventrally flattened and is composed of a wide bifid posterior portion narrowing towards an elongate middle and anterior portion (unpreserved). The posterior expansion displays two lateral flanges with small ridges on the ventral surface framing the median gap and a prominent median ridge on the dorsal surface (m. r. Uh; Fig. 10 N). The imprint in the nodule reveals that the ridge does not reach the anterior end, which is partially missing. Branchial dentition. Numerous scattered branchial platelets have been modelled (but are not shown in the figures, see Appendix 2). These dental plates were certainly associated with the ceratobranchials and the large basibranchial. They display a similar square to rectangular shape. Unfortunately, their precise number, arrangement, and distribution on the branchial elements cannot be reconstructed. Pectoral girdle The ventral part of both pectoral girdles is well-preserved comprising the clavicle, the extracleithrum, the ventral blade of the cleithrum, and the scapulocoracoid. There is no evidence of an interclavicle. The dorsal series of post-temporal, supracleithrum and anocleithrum has not been retrieved. Clavicle. The clavicles (Cla; Figs 2 F; 3 F-H; 13 C, D, G, H) are overall well-preserved, but parts of the antero-ventral blade are missing from both sides of the skull. The clavicle can be divided into two distinct portions: a large dorso-lateral section overlapping the ventral edge of the cleithrum and the extracleithrum, and a dorso-ventrally flattened antero-ventral flange. The internal surface of the dorso-lateral section is strongly concave and is funnel-shaped. The anteriormost portion of the dorso-lateral section displays a wing-like pointed process embracing a groove present in the antero-lateral portion of the cleithrum and overlapping it. Cleithrum. Both cleithra (Cl; Figs 3 E-H; 13 C, D, G, H) are incompletely preserved, lacking the dorsal blade. The cleithrum is narrow throughout its length and its external surface displays a smooth anterior margin and a more rugose and ornamented postero-dorsal corner. The external surface is slightly convex, except for the anterior narrow groove accommodating the clavicle. The internal surface is concave and displays a prominent ridge towards the medial margin certainly housing a large cartilaginous blade supporting the scapulocoracoid, as in Latimeria chalumnae (Millot & Anthony 1958). Extracleithrum. The extracleithra (Ecl; Figs 3 F-H; 13 C, D, G-H) are narrow and elongate bones. The external surface is ornamented with tubercles as on the cleithrum, and similar to those present on the postorbital and squamosal. The dorsal end is levelled with the articulatory head of the scapulocoracoid. The ventral margin is serrated and is overlapped by the clavicle abutting a ventro-lateral buttress, while the major part of the bone fits in a postero-lateral depression of the cleithrum. The postero-dorsal edge displays a small depression concomitant with a similar depression in the posterior margin of the cleithrum. Scapulocoracoid. The ossified articulatory portion of both scapulocoracoids (Scc; Figs 3 F-H; 13 C, D, G-H) is completely preserved. It is a remarkably large, shoe-shaped element, internally hollow with extremely thin periosteal walls. The base is quadrangular, slightly narrowing ventrally. The medial columnar section is dorso-posteriorly inclined and the proximal edge would have likely housed a convex cartilaginous articulatory head for the humerus. There are no traces of perforations in the outer surface of the bone.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
03B72C1C3020903DFE91FACE3924FEFD.taxon	discussion	Phylogenetic results The phylogenetic position of Aemilia stellata n. gen., n. sp. was evaluated through maximum parsimony and Bayesian inference using the revised and enhanced matrix of Clement et al. (2024). The differences between the parsimony (Appendix 3) and Bayesian (Fig. 15) analyses will only be briefly described here. The parsimony analyses yielded unreliable results (Appendix 3). The strict consensus produced a poorly resolved tree from 34 equally parsimonious trees (length = 2033; consistency index = 0.18; retention index = 0.23) as well as the 50 % majority rule consensus tree (tree length = 1475; consistency index = 0.25; retention index = 0.49). The resulting large polytomy is certainly due to the fact that Aemilia stellata n. gen., n. sp. lacks numerous important phylogenetic characters owing to the limited available material. The bootstrap replications yield an average support value of 7.9 for the resulting tree, which falls below the threshold required to consider the tree as robust. For a fully resolved parsimony analysis and the characters supporting each node, seeFerrante & Cavin (2025). The Bayesian analyses provided a clearer outcome (Fig. 15). Aemilia stellata n. gen., n. sp. is reconstructed as the sister group and earliest representative of a Permian-Triassic-Jurassic clade of coelacanths (clade 3; Fig. 15), comprising wellknown taxa like Diplurus newarki Newberry, 1878 (Jurassic), Piveteauia madagascarensis Lehman, 1952, Rieppelia heinzfurreri Ferrante & Cavin, 2023, and Foreyia maxkuhni Cavin, Mennecart, Obrist, Costeur & Furrer, 2017 (Triassic), among others. This clade is in turn reconstructed as the sister group of an exclusively Triassic clade (clade 4; Fig. 15) mainly comprising the different species of Whiteia Moy-Thomas, 1935, as well as Rebellatrix divaricerca Wendruff & Wilson, 2012, Sassenia tuberculata Stensiö, 1921, and Heptanema paradoxum Belloti, 1857. Several clades can be associated with previously identified families of coelacanths, although their composition may vary with respect to earlier studies (e. g., Schultze 1993, 2004; Forey 1998; Manuelli et al. 2024; Ferrante & Cavin 2025). Among the most clearly recognizable families, we note the Laugiidae Berg, 1940 (clade 1; Fig. 15), Axeliidae Ferrante & Cavin, 2025 (clade 2; Fig. 15), Mawsoniidae Schultze, 1993 (clade 5; Fig. 15), and Latimeriidae Berg, 1940 (clade 6; Fig. 15). Other traditional coelacanth families like Miguashaiidae Schultze, 1993, Diplocerciidae Stensiö, 1921, Hadronectoridae Lund & Lund, 1984, Rhabdodermatidae Berg, 1958, Whiteiidae Schultze, 1993, and Sasseniidae Forey, 1998 were not retrieved as monophyletic and will not be further discussed here. The family Laugiidae Berg, 1940 (clade 1; Fig. 15) maintains its traditional composition (e. g., Forey 1998; Schultze 2004) and corresponds to the clade including Laugia groenlandica Stensiö, 1932 and Coccoderma suevicum Quenstedt, 1858, with the exlusion here of Piveteauia madagascarensis Lehman, 1952, usually retrieved among the laugiids (e. g., Toriño et al. 2021 a; Ferrante & Cavin 2023, 2025; Manuelli et al. 2024). The family Axeliidae Ferrante & Cavin, 2025 (clade 2; Fig. 15; reconstructed as a subset of Whiteiidae (node E 1) by Arratia & Schultze (2015) and defined by Ferrante & Cavin (2025 )) is composed of Axelia robusta Stensiö, 1921, Wimania sinuosa Stensiö, 1921, and Atacamaia solitaria Arratia & Schultze, 2015; its sister group is here represented by Coelacanthus granulatus Agassiz, 1839. Ferrante & Cavin (2025) reconstructed the family Whiteiidae Schultze, 1993 including Whiteia Moy-Thomas, 1935, Guizhoucoelacanthus Liu, Yin, Luo, Wang & Wang, 2006, and Garnbergia Martin & Wenz, 1984. In our analysis, all surveyed species of the genus Whiteia (W. durabilis Wendruff, 2011, W. lepta Wendruff, 2011, W. woodwardi Moy-Thomas, 1935, W. uyenoteruya Yabumoto, Brito, Iwata & Abe, 2019, W. nielseni Forey, 1998, and W. oishoii Yabumoto & Brito, 2016) cluster close together in a larger clade (clade 4; Fig. 15) that also includes Rebellatrix divaricerca Wendruff & Wilson, 2012, Sassenia tuberculata Stensiö, 1921, and Heptanema paradoxum Belloti, 1857. However, Guizhoucoelacanthus is found within clade 3, whereas Garnbergia is reconstructed among the Mawsoniidae. The families Mawsoniidae Schultze, 1993 (clade 5; Fig. 15) and Latimeriidae Berg, 1940 (clade 6; Fig. 15) are also clearly recognizable as sister clades whithin the suborder Latimerioidei Schultze, 1993 (for a different composition see Dutel et al. 2012; Cavin & Grădinaru 2014; Arratia & Schultze 2015; Toriño et al. 2021 a; Ferrante & Cavin 2023, 2025; Manuelli et al. 2024). One puzzling result is that Graulia branchiodonta Manuelli, Mondéjar Fernández, Dollman, Jakata & Cavin, 2024 is no longer found as a basal Mawsoniidae (as opposed to Manuelli et al. 2024) but appears well-nested within the Latimeriidae, whereas Dobrogeria aegyssensis Cavin & Grădinaru, 2014 also switches position from the Latimeriidae (Ferrante & Cavin 2023, 2025; Manuelli et al. 2024) to the Mawsoniidae. Within the Latimeriidae, the subfamily Ticinepomiinae Ferrante & Cavin, 2023, previously containing the species Ticinepomis peyeri Rieppel, 1980, Foreyia maxkuhni Cavin, Mennecart, Obrist, Costeur & Furrer, 2017, and Rieppelia heinzfurreri Ferrante & Cavin, 2023 known exclusively from the Triassic of Switzerland (Ferrante & Cavin 2023, 2025) has surprinsingly exploded, with Ticinepomis peyeri retrieved as an early latimeriid in a clade containing Holophagus gulo Egerton, 1861, Undina (U. penicillata Münster, 1834 and U. cirinensis Saint-Seine, 1949), and Macropoma willemoesii Vetter, 1881. On the other hand, Rieppelia heinzfurreri and Foreyia maxkuhni were reconstructed as crownward members of the clade including Aemilia stellata n. gen., n. sp. (clade 3; Fig. 15). The analysis also revealed probable polyphyletic taxa, notably Rhabdoderma Newberry, 1856, Sassenia Stensiö, 1921, Changxingia Wang & Liu, 1981, Whiteia Moy-Thomas, 1935, and Macropoma Agassiz, 1835, as well as paraphyletic genera like Diplocercides Stensiö, 1922, Euporosteus Jaeckel, 1927, Undina Münster, 1834, and Axelrodichthys Maisey, 1986. Among the multispecific genera of coelacanths surveyed, only Miguashaia (M. bureaui Schultze, 1973 and M. grossi Forey, Ahlberg, Lukševičs & Zupiņš, 2000) and Mawsonia (M. gigas Woodward, 1907, M. soba Brito, Cupello, Yabumoto, Hell, Brunet & Otero, 2018, M. brasiliensis Yabumoto, 2002, and M. tegamensis Wenz, 1973) were retrieved as monophyletic. It is not in the scope of this study to discuss the probable causes for this odd distribution of taxa, but we encourage future surveys to thoroughly review these polyspecific genera and shed light on these problematic phylogenetic reconstructions.	en	Fernández, Jorge Mondéjar, Mansuit, Rohan, Flament, Richard, Cloutier, Richard, Mapes, Royal, Clément, Gaël, Pradel, Alan (2026): A new Carboniferous coelacanth illuminates the evolution of the actinistian hyobranchial skeleton. Geodiversitas 48 (11): 185-226, DOI: 10.5252/geodiversitas2026v48a11
