taxonID	type	description	language	source
03C687D1FF9A3010A297FE54FAC8BDA7.taxon	description	IMAGING Fossil specimens were scanned at the University of Chicago X-ray μCT facility, on a GE Phoenix 240 / 180 scanner. Parameters used were voltage 130 kV, current 170 μA, timing 500 s and a 0.5 mm Cu filter for P 10419 a & b, and voltage 130 kV, current 170 μA, timing 500 s and no filter for FMNH UF 462 and FMNH UF 464. Wet specimens were stained in 5 % phosphotungstic acid (PTA) in alcohol for 2 weeks and then scanned with the parameters voltage 75 kV, current 140 uA, timing 1000 s and no filter. Segmentation and anatomical reconstruction were accomplished using MATERIALISE MIMICS v. 19 (Materialise Mimics, 2016). Imaging of digital models was completed using BLENDER v. 2.8 (Community, 2018). Photography of fossils was accomplished with a Leica DFC 490 camera attached to a Zeiss Stemi SV 6 microscope and the images processed in IMAGE-PRO PLUS v. 6.2 using the enhanced depth of field function to align and process multiple z-stacks of images. PHYLOGENETIC METHODS	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF993013A2CBFD4AFE1BBEC5.taxon	materials_examined	Holotype: NHMUK PV P 10419 a & b. Rhadinichthys planti part and counterpart.	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF97301CA11DFD3DFDDBB840.taxon	description	preservation. Circumorbital bones are known only from an unusually broad left nasal and an approximately triangular jugal (posterior infraorbital). The jugal portion of the infraorbital canal produces four short posterior branches. Cheek bones recovered include maxillae, a preopercular and quadratojugal. The maxilla is broad and bears a short anterior process, around half the length of the posterior, expanded portion of the bone. An anteriorly restricted lamina descending from the rear margin of the maxilla overlaps the lower jaw. The preopercular has a deeply notched anterior edge. The posterior limb is around one-third of anterior limb length, and the angle between limbs is barely pronounced. The preopercular sensory canal (po. sc, Fig. 5 B) runs along the dorsal margin and exits dorsally prior to the anterior extremity of the preopercular bone. Ventrally, the canal passes into NEUROCRANIUM The braincase of Ph. mossae is barely distorted: the neurocranial roof is present in NHMUK P 10419 a and the bulk of the neurocranium in NHMUK P 10419 b (Fig. 6). External, perichondral, surfaces are well preserved, but the internal walls and details of the endocranial space are lacking, perhaps unmineralized or eroded. Limited exposure of the cancellate endochondral bone is present posterior to the basipterygoid processes. No structural evidence indicates separate ossification centres; the entire neurocranium is thus reconstructed as one coherent entity. A primary neurocranial roof is present, although it is partially obscured by overlying dermal bones as seen in Figure 4 B. The orbits are large, and the interorbital septum is unossified ventral to the olfactory tracts. The anterior of the neurocranium is well preserved, making Phoebeannaia the third Carboniferous actinopterygian with a detailed anterior ethmoid region, joining Kansasiella eatoni and Kentuckia deani (Rayner, 1951). Phoebeannaia resembles Kansasiella in many aspects including general proportions, extent of fissures and location of articular surfaces and foramina. The following regional descriptions will therefore first detail Phoebeannaia and subsequently list salient differences observed in Kansasiella. Notable general differences in our Kansasiella specimens with reference to Phoebeannaia include the lack of a preserved ethmoid region (here considered incomplete relative to Poplin’s 1974 material) and dissociation of the occiput in FMNH UF 464. Note that separation of the occipital region from the anterior neurocranium has been previously documented in Kansasiella (Poplin, 1974). OCCIPITAL	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF96301EA297FAD2FD41BF98.taxon	description	In posterior view (Fig. 8), the occipital plate is taller than it is wide, with lateral boundaries gently curving out from a narrow base to small craniospinal processes (csp. p; Figs 7 A, 8 A, 9 A, B) level with the top of the foramen magnum (f. m; Figs 7 B, 8 A). Internally the foramen magnum is pierced by two ventral foramina for the occipital nerves (oc. n; Fig. 7 B), which exit the braincase laterally. The notochord canal (nc. c; Figs 7 B, 8 A) is separated from the foramen magnum dorsally and the aortic canal ventrally, and terminates anteriorly before the otico-occipital fissure. An intermuscular septum is marked by a faint ridge that runs dorsoventrally through the craniospinal fossa (csp. f; Fig. 8 A, E): the broad, posterior-facing shallow depression for the attachment of epaxial musculature flanking the foramen magnum. No prominent horizontal ridge or midline crest edging the fossa is present. The left craniospinal process bulges slightly anteriorly around the opening for the vagus nerve, but the right-side equivalent shows no evidence of this protrusion. The occipital plate is narrow relative to the rest of the braincase, approximately 60 % as wide as the distance between the post-orbital processes. The occiput is not as wide as in Kansasiella or other contemporaneous species; the narrower dimensions in Phoebeannaia reveal much of the otic wall in posterior view including the parampullary fossa, jugal canal, and articular surfaces for the pharyngobranchials and hyomandibula. The basioccipital is dominated by the median dorsal aortic canal (ao. c; Figs 7 B, 8 A). There is a midline foramen approximately level with the exit of the vagus nerve, which must have transmitted some efferent branchial arteries (e. br. a; Figs 7 B, 9 A). The enclosed canal ends where the grooves for the lateral dorsal aorta diverge (lat. d. ao. c; Fig. 9 A), level with the glossopharyngeal nerve exit (IX, Fig. 7 A). No associated indentations mark the reception of further efferent branchial arteries, although they might have joined ventral to the neurocranium. A prominent articular surface for the first infrapharyngobranchial (art. 1 st. i; Figs 7 A, 9 A) projects from the ventrolateral extremity of the basioccipital. Anteriorly, this surface is delimited by a deep groove marking the passage of the efferent hyoid artery up into the jugal canal.	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF943019A297FC80FE28BEF8.taxon	description	Unlike Phoebeannaia, in Kansasiella the midline dorsal rim of the occipital plate projects anteriorly into the posterior dorsal fontanelle (pos. d. fon; Fig. 9 D); thus, the fontanelle outline is bean-shaped rather than ovoid. The vestibular fontanelles (v. fon; Figs 7 C, 8 E, 9 C) are smaller than in Phoebeannaia. The interior space, in Kansasiella, is bounded anteriorly by a tall dorsum sellae (d. s; Fig. 7 D) which rises into the cranial cavity from the basisphenoid, continues across the midline tracking the ventral fissure and bears foramina allowing the abducens nerves (VI, Figs 7 D, 10 B, D) to pass through to the rear of the posterior myodome. The notochord canal (nc. c; Figs 7 D, 8 E) is clearly separated from the foramen magnum above and dorsal aortic canal below and terminates at approximately the same position as in Phoebeannaia contra Poplin (1974) (p 63; fig. 20) where it was shown to reach the ventral fissure. The ventral fissure (v. f; Figs 7 D, 9 C) and associated lateral foramina (VII. pal, o. a; Figs 7 D, 9 C, 10 C) are curved as in Phoebeannaia but are more widely separated from the vestibular fontanelles. There is no lateral projection for the first infrapharyngobranchial. The lateral dorsal aortae diverge within a long enclosed bony canal and exit separately (lat. d. ao. c; Fig. 9 A) just posterior to the ventral fissure. Paired foramina for efferent branchial arteries (e. br. a; Figs 7 D, 9 C) are positioned anteriorly relative to the single midline passage in Phoebeannaia. Foramina for the occipital arteries (oc. a; Fig. 7 D) are located posterior to the level of the efferent branchial foramina. OTIC	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF933019A2A5FC0AFC05B949.taxon	description	In lateral aspect, a well-defined triangular parampullary fossa (pa. f; Figs 7 A, 8 A) spans the area between the rear of the hyoid facet (overlying the external semicircular canal ampulla) and the prominence capping the posterior ampulla. This is the likely site of opercular adductor muscle origin. The crest forming the roof of the parampullary fossa descends anteriorly, aligned with the dorsal margin of the kidney-shaped, posteroventrally directed hyomandibular facet (hm. fa; Figs 7 A, 8 A, 9 A). A narrow spiracular bridge (spir. br; Figs 7 A, 9 A) ascends from the anterior rim of the hyoid facet to the posterior of the post-orbital process (po. p; Figs 7 A, 9 A, 10 A), enclosing (canalizing) a shallow spiracular groove. The free posterior surface of the post-orbital process likely provided the attachment surface for the hyoid constrictor muscles in the absence of a clearly defined dilatator fossa. The ventral rim of the post-orbital process tapers smoothly into a broad lateral commissure. The groove for the efferent hyoid artery, anterior to the projection for the first infrapharyngobranchial, ascends the lateral commissure and joins the jugular canal via a foramen (e. hy. a; Figs 7 A, 8 A, 9 A). On the left side of the neurocranium there are three posterior exits from the jugal canal: a large medial opening primarily for the jugular vein (j. c; Figs 7 A, 8 A, 9 A, 10 A), and two small lateral foramina that might have transmitted branches of the facial nerve (VII. hy, VII. op; Fig. 8 A). There is only one lateral foramen on the right side, which could indicate plasticity of this feature or merely post-mortem breakage. The groove for the jugular vein continues posteriorly ventral to the exit of the vagus nerve and is roofed by a broad horizontal shelf with an anterior concavity — the subtemporal fossa (st. f; Figs 7 A, 8 A, 9 A). The glossopharyngeal nerve foramen, ventral to the jugal groove, is closely associated with a broad lateral projection for articulation with the first suprapharyngobranchial. Both the foramen and the articulation are aligned with the rear of the vestibular fontanelle, which is bounded posteriorly by a ventral tongue-like projection from the otic wall (v. t; Figs 7 A, 8 A, 9 A). Similar examples are present in Kansasiella, Kentuckia deani and Pteronisculus.	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF933018A11DFBF1FDC0BB60.taxon	description	ORBITAL	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF92301BA297F9B3FD4EBB63.taxon	description	The supraorbital shelf extends medio-dorsally from the post-orbital processes (po. p; Figs 7 A, 9 A, B, 10 A, B) and descends at the midline beneath the olfactory nerves and forebrain. There is no ossified orbital septum, and the optic nerves consequently exit the intracranial space together through a large midline foramen (II; Fig. 10 A, B). The basisphenoid pillar (bs. p; Fig. 7 B) transmits the hypophysis from the cranial cavity ventrally through the floor of the neurocranium (hyp. f; Figs 7 B, 9 A), and flares out laterally into buttresses (lat. b; Fig. 10 A, B) which connect directly with the base of each large, anteriorly directed basipterygoid process (bpt. p; Figs 7 A, 8 A, 9 A, 10 A, B). A third, slender, medial buttress extends anteriorly a short distance before merging into the midline crest of the basisphenoid (ant. b; Figs 7 B, 10 B). Fenestrae between each lateral strut allow passage between the orbital space and the large unpaired posterior myodome. Anterior openings of the jugal canal (j. c; Figs 7 A, 8 A, 9 A, 10 A), walled by the lateral commissure (lat. com; Fig. 7 A), flank the posterior myodome laterally. The many foramina and fossae in the orbital region can be understood by comparison to extant and extinct taxa. The post-orbital wall, dominated by the jugular vein canal, bears a medial foramen for the exit of the facial nerve (VII; Fig. 10 A, B) and a recess marking the location of the geniculate ganglion. Dorsal to the exit of the jugal canal, another recess likely for the gasserian and lateralis ganglia (V, lat; Fig. 10 A, B) is associated with a broad communion with the intracranial space. Within this recess, a lateral passage seems to have guided the otic branch of the trigeminal nerve (V. ot; Fig. 10 B) back into the otic wall. A shallow depression above and lateral to the trigeminal recess might have served as a site of attachment for jaw musculature, though it lacks clear boundaries and is not as large as the corresponding fossa in Moythomasia. Small foramina for the oculomotor nerves (III; Fig. 10 A, B) flank the optic foramen. The posterior wall of the expansive single posterior myodome is pierced on either side by a foramen for the abducens nerve (VI; Figs 7 D, 10 B, D) and, more ventrally, by a passage to accommodate the descending palatine branch of the facial nerve and / or the ascending orbital artery (VII. pal, o. a; Figs 7 B, 9 A, 10 A, B). The roof of the myodome bears a midline opening permitting the hypophysis to descend from the intracranial space into the groove of the basisphenoid pillar before reaching the hypophysial canal ventrally (hyp. f; Figs 7 B, 9 A). Paired passages (int. car; Figs 9 A, 10 A, B) located ventromedially to the basipterygoid processes enclose the internal carotids (after giving off the orbital arteries). These canals run medially and dorsally into the space between the basisphenoid pillar and anterior buttress. Grooves on the central pillar (gr. a. int. car; Fig. 10 A, B) mark the ascent of the internal carotids to a midline foramen, ventral to the anterior buttress, that opens to the intracranial space. On the ventral surface of the basisphenoid, narrow grooves mark the passage of the efferent pseudobranchial arteries (ep. a; Fig. 9 A) prior to anastomosis with the internal carotids. A narrow midline groove runs anteriorly from the hypophysial canal and is laterally joined by a groove for the palatine artery and nerve (gr. p. a, n; Fig. 9 A).	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF91301BA2A5F9C6FB5ABEDA.taxon	description	ETHMOID	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
03C687D1FF903005A297FB22FBADBB65.taxon	description	PHYLOGENETIC RESULTS Specimens NHMUK PV P 10419 a & b, FMNH UF 464 and FMNH UF 462 were coded into a morphological matrix and analysed using parsimony and Bayesian inference. The unconstrained parsimony analysis retrieved 60 equally parsimonious trees of 163 steps. The strict consensus (Fig. 12 A) recovered Phoebeannaia in a polytomy with Coccocephalus wildi, Luederia kempi, Lawrenciella, a branch leading to Kansasiella eatoni and Pteronisculus magnus, and a clade consisting of the neopterygian crown plus Boreosomus piveteaui Stensiö, 1921 and Australosomus kochi Stensiö, 1932. This fairly unresolved node has a Bremer Index of two, and is supported by seven character state changes, four of which are unambiguous and none of which have a Consistency Index (CI) of one. The strict consensus has a tree length of 173, CI of 0.3526, Homoplasy Index (HI) of 0.6474, Retention Index (RI) of 0.7248 and Rescaled Consistency Index (RC) of 0.2556. The topology of the strict consensus tree contradicts known morphological and molecular evidence regarding the branching order of crown Actinopterygii, thus alternative phylogenetic placements under parsimony were investigated using a constrained backbone search. The backbone-constrained parsimony analysis retrieved 12 equally most parsimonious trees of 165 steps, and the strict consensus (Fig. 12 B) recovered Phoebeannaia in the same position, supported by the same characters, with a Bremer Index of two. The strict consensus has a tree length of 172, CI of 0.3547, HI of 0.6453, RI of 0.7273 and RC of 0.2579. While the recovered topology is no longer contradictory, the insertion for Phoebeannaia is still relatively unresolved and supported by low CI characters. The backbone-constrained analysis was therefore re-run after character reweighting by RC. The reweighted and backbone constrained parsimony analysis found three equally most parsimonious trees, and the strict consensus (Fig. 13 A) is nearly fully resolved; Phoebeannaia is sister to Kansasiella and Pteronisculus magnus (Bremer Index = 2, Bootstrap = 66.52), with the clade comprising the Neopterygian crown plus Coccocephalus wildi, Luederia kempi, Lawrenciella, Boreosomus piveteaui and Australosomus kochi as their sister group and Pteronisculus stensiöi as the nearest outgroup. The strict consensus has a higher CI (0.5347), RI (0.8513) and RC (0.4552), and a lower HI (0.4653) than the previous analyses, as expected, which could be interpreted as a partial correction for high levels of homoplasy in the dataset. The clade of Phoebeannaia, Kansasiella and Pteronisculus magnus is supported by one character, the presence of a ‘ buttressed’ basisphenoid pillar (CI = 1). This feature manifests via the lateral expansion of the basisphenoid pillar with excavation of paired recesses through to the posterior myodome, as seen in Figure 10. It might have allowed for larger ocular muscles while simultaneously bracing the basipterygoid processes to withstand increased stress from palatoquadrate motion. Kansasiella and Pteronisculus magnus are further held together by three unambiguous character state changes, and five characters (four unambiguous) bind Phoebeannaia, Kansasiella and Pteronisculus magnus in place. The short branch lengths calculated in the most parsimonious trees (one example shown in Fig. 13 B) emphasize the similarity between these taxa. The unconstrained Bayesian analysis (Fig. 14 A) retrieved Phoebeannaia as sister to Kansasiella and Pteronisculus magnus (Posterior Probability 0.59). That clade was placed in a large polytomy with Coccocephalus wildi, Luederia kempi, Lawrenciella and the group of Boreosomus piveteaui, Australosomus kochi and crown Neopterygii, with Pteronisculus stensioi as the nearest outgroup. The placement of polypterids and chondrosteans contradicting modern consensus justified re-running the analysis with an imposed backbone topology (Fig. 14 B), resulting in the addition of the Chondrostei plus Birgeria groenlandica Stensiö 1932 and Saurichthys sp. (Argyriou et al., 2018) to the large polytomy.	en	Caron, Abigail, Venkataraman, Vishruth, Tietjen, Kristen, Fls, Michael Coates (2023): A fish for Phoebe: a new actinopterygian from the Upper Carboniferous Coal Measures of Saddleworth, Greater Manchester, UK, and a revision of Kansasiella eatoni. Zoological Journal of the Linnean Society 198: 957-981
