identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03C687D1FF9A3010A297FE54FAC8BDA7.text	03C687D1FF9A3010A297FE54FAC8BDA7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kansasiella eatoni	<div><p>Kansasiella eatoni</p><p>The two specimens of  Ka. eatoni here examined are FMNH UF464 and FMNH UF462, both phosphatic nodules loaned from the Field Museum in Chicago. They were collected from a thin band of yellow-grey shale above the marine Haskell Limestone and below the brackish Robbins Shale of the Douglas Group near Lawrence, Kansas, along with over 400 other nodules ascribed to  Kansasiella (Poplin, 1974) . These nodules were likely formed in a near-shore shallow marine subtropical environment, and have been dated to the Virgilian, Upper Pennsylvanian (305–299 ± 0.8 Mya) (Miller &amp; Swineford, 1957; Poplin, 1974; Hamel &amp; Poplin, 2008). Neither specimen is complete, though they complement one another near perfectly. The exterior surface of FMNH UF462 is well preserved including the occipital, otic, sphenoid and ventral portion of the ethmoid, but the large crack running through the centre of the nodule obscures the interior and back of the orbit. In contrast, FMNH UF464 has an exquisitely detailed otic and sphenoid region, including fine internal and postorbital features, but is entirely lacking the occipital, ethmoid and ventral parts of the neurocranium. Digital superposition of the two specimens revealed an exact morphological match (after accounting for a 2% global upscaling of FMNH UF162), and the resulting combined reconstruction is shown in all subsequent figures.</p><p>IMAGING</p><p>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 P10419a &amp; b, and voltage 130 kV, current 170 μA, timing 500 s and no filter for FMNH UF462 and FMNH UF464. 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.</p><p>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 DFC490 camera attached to a Zeiss Stemi SV6 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.</p><p>PHYLOGENETIC METHODS</p></div>	https://treatment.plazi.org/id/03C687D1FF9A3010A297FE54FAC8BDA7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF993013A2CBFD4AFE1BBEC5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phoebeannaia mossae Caron & Venkataraman & Tietjen & Fls 2023	<div><p>PHOEBEANNAIA MOSSAE SP. NOV.</p><p>Zoobank registration: urn:lsid:zoobank. org:act: 25295052-B714-4DE7-854A- C451F9CA9085</p><p>Holotype: NHMUK PV P 10419a &amp; b.  Rhadinichthys planti part and counterpart.</p></div>	https://treatment.plazi.org/id/03C687D1FF993013A2CBFD4AFE1BBEC5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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
03C687D1FF98301DA10FFE85FAAABF16.text	03C687D1FF98301DA10FFE85FAAABF16.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phoebeannaia mossae Caron & Venkataraman & Tietjen & Fls 2023	<div><p>Phoebeannaia mossae</p><p>Specimen P10419a &amp; b includes several disarticulated dermal bones spread across the plane of the break in the nodule (Fig. 4). Ornamentation of all dermal bones consists of fine, close-packed, sinuous ridges. Of the skull table, left and right frontals and parietals can be easily identified. The suture between the frontal and parietal is W-shaped with the deep anterior projection of the parietal enclosing the passage of the supraorbital sensory canal (spo.sc; Fig. 5A). The canal itself bears a distinct medial kink on the frontal, mirroring the concave lateral edge of the bone. The parietal pit lines are unknown due to poor a distinctively large subrectangular quadratojugal. Of the mandible, clearly preserved bones include the dentary, angular, articular and coronoids (Fig. 4). The presence or absence of the surangular is unknown. Several branchiostegal rays are preserved (Fig. 4, reconstruction Fig. 5C), although scattered in such a way it is impossible to count them. The articular is well ossified and bears a double glenoid facet for articulation with the quadrate. The suture between the angular and dentary is visible on the right mandible (Fig. 4B, D), and a line of coronoids is visible on both mandibles. Fragments of the shoulder girdle are also evident, but detailed reconstruction is difficult due to compaction, breakage and dissociation of the individual bones.</p></div>	https://treatment.plazi.org/id/03C687D1FF98301DA10FFE85FAAABF16	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF97301CA11DFD3DFDDBB840.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kansasiella eatoni	<div><p>Kansasiella eatoni</p><p>Part of the parasphenoid (psp; Fig. 9C) is attached to the neurocranium of FMNH UF462 and imaged in place due to difficulty distinguishing the exact boundaries between dermal and endoskeletal components. The parasphenoid extends from the ventral fissure to the ethmoid palatal articulation and bears high ascending processes (a.p.psp; Figs 7C, 9C) that nearly reach the spiracular canal. The anterior process is more extensive and straight sided than estimated in Poplin’s (1974: fig. 8) reconstruction. No other dermal bones are evident.</p><p>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. 5B) runs along the dorsal margin and exits dorsally prior to the anterior extremity of the preopercular bone. Ventrally, the canal passes into</p><p>NEUROCRANIUM</p><p>The braincase of  Ph. mossae is barely distorted: the neurocranial roof is present in NHMUK P10419a and the bulk of the neurocranium in NHMUK P10419b (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.</p><p>A primary neurocranial roof is present, although it is partially obscured by overlying dermal bones as seen in Figure 4B. 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 UF464. Note that separation of the occipital region from the anterior neurocranium has been previously documented in  Kansasiella (Poplin, 1974) .</p><p>OCCIPITAL</p></div>	https://treatment.plazi.org/id/03C687D1FF97301CA11DFD3DFDDBB840	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF96301EA297FAD2FD41BF98.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phoebeannaia mossae Caron & Venkataraman & Tietjen & Fls 2023	<div><p>Phoebeannaia mossae</p><p>The occipital region is well preserved barring minor damage on the posterior right wall of the aortic canal. The otico-occipital fissure (ot.oc.f; Figs 7A, 9A, B) delimits the dorsal portion of the occipital unit from the otic region. The fissure spans an ovoid posterior dorsal fontanelle (pos.d.fon; Fig. 9B) at the dorsal midline and extends ventrolaterally, expanding to accommodate the exit of the vagus nerve (X; Figs 7A, B, 9A) and terminates in the large vestibular fontanelles ( v.fon; Figs 7A, 8A, 9A). A narrow bridge of bone separates the vestibular fontanelles from the ventral fissure (v.f; Figs 7B, 9A). The ventral fissure bounds the anterior extent of the basioccipital and is curved with an anterior-facing concavity, allowing it to skirt around but not intersect with the posterior myodome. The perichondrally-lined ventral fissure is laterally fringed by, but not contiguous with, foramina through which the palatine branch of the facial nerve and/or branches of the orbital artery (VII.pal,o.a; Figs 9A, 10A, B) might have communicated with the posterior myodome.</p><p>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 7A, 8A, 9A, B) level with the top of the foramen magnum (f.m; Figs 7B, 8A). Internally the foramen magnum is pierced by two ventral foramina for the occipital nerves (oc.n; Fig. 7B), which exit the braincase laterally. The notochord canal (nc.c; Figs 7B, 8A) 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. 8A, 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.</p><p>The basioccipital is dominated by the median dorsal aortic canal (ao.c; Figs 7B, 8A). 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 7B, 9A). The enclosed canal ends where the grooves for the lateral dorsal aorta diverge (lat.d.ao.c; Fig. 9A), level with the glossopharyngeal nerve exit (IX, Fig. 7A). 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. 1st.i; Figs 7A, 9A) 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.</p></div>	https://treatment.plazi.org/id/03C687D1FF96301EA297FAD2FD41BF98	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF943019A297FC80FE28BEF8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kansasiella eatoni	<div><p>Kansasiella eatoni</p><p>The occipital unit is complete with only the interior dorsolateral margins of the foramen magnum incompletely recovered. All data presented in this section are from FMNH UF462, as the entire region is absent in FMNH UF464. The occipital plate of  Kansasiella has more pronounced craniospinal processes (csp.p; Figs 7C, 8E, 9C, D) than  Phoebeannaia and a consequently sharper concave taper. The basioccipital is dorsoventrally thicker and encloses the vestibular fontanelle to a greater extent than in  Phoebeannaia . The entire occiput is smaller relative to the rest of the braincase than is depicted in Poplin (1974); the craniospinal processes are approximately 90% as wide as the post-orbital processes, and the basioccipital does not expand ventrally to the same degree as previously described.</p><p>Unlike  Phoebeannaia, in  Kansasiella the midline dorsal rim of the occipital plate projects anteriorly into the posterior dorsal fontanelle (pos.d.fon; Fig. 9D); thus, the fontanelle outline is bean-shaped rather than ovoid. The vestibular fontanelles ( v.fon; Figs 7C, 8E, 9C) are smaller than in  Phoebeannaia . The interior space, in  Kansasiella, is bounded anteriorly by a tall dorsum sellae (d.s; Fig. 7D) 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 7D, 10B, D) to pass through to the rear of the posterior myodome. The notochord canal (nc.c; Figs 7D, 8E) 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 7D, 9C) and associated lateral foramina (VII.pal,o.a; Figs 7D, 9C, 10C) 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. 9A) just posterior to the ventral fissure. Paired foramina for efferent branchial arteries (e.br.a; Figs 7D, 9C) are positioned anteriorly relative to the single midline passage in  Phoebeannaia . Foramina for the occipital arteries (oc.a; Fig. 7D) are located posterior to the level of the efferent branchial foramina.</p><p>OTIC</p></div>	https://treatment.plazi.org/id/03C687D1FF943019A297FC80FE28BEF8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF933019A2A5FC0AFC05B949.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phoebeannaia mossae Caron & Venkataraman & Tietjen & Fls 2023	<div><p>Phoebeannaia mossae</p><p>The otic wall extends from the otico-occipital fissure to the post-orbital processes. The roof of this region is partially obscured by displaced frontal and parietal bones, although the semicircular canals and fossa bridgei are evident on the right side of the specimen (ant.ssc, pos.ssc, f.br; Fig. 9B). The ridge overlying the anterior semicircular canal is longer than the posterior, as is common in Carboniferous actinopterygians, and extends forward to the level of the post-orbital process. Consequently, the circumscribed fossa bridgei is large and contiguous with the dorsal exit of the spiracular canal (spir.f; Fig. 9B). There does not appear to be any connection between the intracranial space and fossa bridgei, nor could the fossa bridgei have housed epaxial musculature due to the height of the neurocranial roof overlying the posterior semicircular canal.</p><p>In lateral aspect, a well-defined triangular parampullary fossa (pa.f; Figs 7A, 8A) 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 7A, 8A, 9A). A narrow spiracular bridge (spir.br; Figs 7A, 9A) ascends from the anterior rim of the hyoid facet to the posterior of the post-orbital process (po.p; Figs 7A, 9A, 10A), 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.</p><p>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 7A, 8A, 9A). 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 7A, 8A, 9A, 10A), and two small lateral foramina that might have transmitted branches of the facial nerve (VII.hy, VII.op; Fig. 8A). 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 7A, 8A, 9A). 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 7A, 8A, 9A). Similar examples are present in  Kansasiella,  Kentuckia deani and  Pteronisculus .</p></div>	https://treatment.plazi.org/id/03C687D1FF933019A2A5FC0AFC05B949	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF933018A11DFBF1FDC0BB60.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kansasiella eatoni	<div><p>Kansasiella eatoni</p><p>Otic inner and dorsal surfaces are more completely preserved than in  Phoebeannaia, revealing the semicircular canals, ampullae and details of the neurocranial roof. There is a small midline tectosynotic fossa (t.syn.f; Fig. 9D) between the posterior semicircular canals, and a long anterior dorsal fontanelle (ant.d.fon; Fig. 9D) that stretches forward to the level of the basipterygoid processes. An expansive fossa bridgei (f.br; Figs 7C, 9D) houses an anterior spiracular fossa (spir.f; Fig. 9D) and a deep posterior depression containing poorly ossified intermural diverticula that do not appear to enter the intracranial space. The spiracular bridge (spir.br; Figs 7C, 9C, D) is deeper and more obliquely angled than that of  Phoebeannaia, extending anterodorsally from the base of the hyoid facet. The enclosed spiracular canal is narrower. The lateral commissure curves sharply in to meet the post-orbital processes, forming a more hooked shape in lateral view than the smooth taper of  Phoebeannaia . The vestibular fontanelle and the posterior descending ventral tongue are proportionately smaller than in  Phoebeannaia, and the projection for articulation with the first suprapharyngobranchial (art.1 st.s; Fig. 7C) is located more anteriorly. Our reconstruction is more detailed than prior depictions of the area surrounding the hyomandibular articulation, revealing a facet (hm.fa; Fig. 7C) like that of  Phoebeannaia and clearly delimited from the adjacent spiracular bridge and parampullary fossa.</p><p>ORBITAL</p></div>	https://treatment.plazi.org/id/03C687D1FF933018A11DFBF1FDC0BB60	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF92301BA297F9B3FD4EBB63.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phoebeannaia mossae Caron & Venkataraman & Tietjen & Fls 2023	<div><p>Phoebeannaia mossae</p><p>The orbital region is loosely defined here as anterior to the ventral fissure and lateral commissure, and posterior to the olfactory capsules and ethmoid processes. Functionally, this region of the braincase transmits cranial nerves and vasculature, houses the extraocular muscles and forebrain, frames the orbits and supports the palatoquadrate.</p><p>The supraorbital shelf extends medio-dorsally from the post-orbital processes (po.p; Figs 7A, 9A, B, 10A, 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. 10A, B). The basisphenoid pillar (bs.p; Fig. 7B) transmits the hypophysis from the cranial cavity ventrally through the floor of the neurocranium (hyp.f; Figs 7B, 9A), and flares out laterally into buttresses (lat.b; Fig. 10A, B) which connect directly with the base of each large, anteriorly directed basipterygoid process (bpt.p; Figs 7A, 8A, 9A, 10A, B). A third, slender, medial buttress extends anteriorly a short distance before merging into the midline crest of the basisphenoid (ant.b; Figs 7B, 10B). 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 7A, 8A, 9A, 10A), walled by the lateral commissure (lat.com; Fig. 7A), flank the posterior myodome laterally.</p><p>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. 10A, 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. 10A, 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. 10B) 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. 10A, 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 7D, 10B, 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 7B, 9A, 10A, 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 7B, 9A).</p><p>Paired passages (int.car; Figs 9A, 10A, 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. 10A, 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. 9A) 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. 9A).</p></div>	https://treatment.plazi.org/id/03C687D1FF92301BA297F9B3FD4EBB63	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF91301BA2A5F9C6FB5ABEDA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kansasiella eatoni	<div><p>Kansasiella eatoni</p><p>The general proportions of the orbital region of  Kansasiella eatoni mirror  Phoebeannaia excepting the supraorbital shelf, which bears a deep groove and laterally encloses the forebrain to a greater extent, and smaller basipterygoid processes (bpt.p; Figs 7C, 8E, 9C, 10C, D), consistent with Poplin’s (1974) description. The posterior myodome and buttressing of the basisphenoid pillar are remarkably similar to  Phoebeannaia, though details surrounding the passage of the hypophysis are lacking due to cracks in the specimen. Ventrally, the surface of the basisphenoid is mostly obscured by the parasphenoid. Foramina for the oculomotor (III; Fig. 10C, D) and trochlear (IV; Fig. 10C, D) nerves are clearly preserved. The channel through which the trigeminal nerve and gasserian ganglia must have passed (V.so,lat; Fig. 10C, D) is somewhat recessed into the jugal canal and enclosed by a bridge of bone similar to the ‘alisphenoid pedicel’ (a.ped; Fig. 10C) described in  Moythomasia durgaringa (Gardiner, 1984) . While this recess primarily opens ventrally into the jugal canal, there is another passage that opens dorsally into the deep groove on the supraorbital shelf that probably marks the path of the supraopthalmic branch of the trigeminal and lateralis nerves (groove for the superopthalmic branch of the trigeminal nerve [gr.V.so]; Fig. 10D). The groove runs the length of the orbital region, though there is a dorsal foramen (lat.f; Fig. 10C) directly above the passage from the trigeminal chamber that might have transmitted one or more nerves into the intracranial space.</p><p>ETHMOID</p></div>	https://treatment.plazi.org/id/03C687D1FF91301BA2A5F9C6FB5ABEDA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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
03C687D1FF91301AA11DFC29FD1DB9F8.text	03C687D1FF91301AA11DFC29FD1DB9F8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phoebeannaia mossae Caron & Venkataraman & Tietjen & Fls 2023	<div><p>Phoebeannaia mossae</p><p>The ethmoid region preserves details of the olfactory system, optic musculature and anterior palatal articulation. Given that most known Carboniferous actinopterygian fossils lack a preserved ethmoid region, it is notable that the morphology displayed by  Phoebeannaia mossae does not closely resemble the complete ethmoid known from  Kentuckia deani . Instead, it is more similar to incompletely recovered specimens of  Lawrenciella schaefferi Poplin, 1984 (Hamel &amp; Poplin, 2008) and  Kansasiella eatoni (Poplin, 1974) . The olfactory tracts descend slightly as they progress anteriorly before passing separately though foramina in the postnasal wall (I; Fig. 11A, B). Above and lateral to the canal for the olfactory nerves, depressions mark the anterior dorsal myodome (a.d.myo; Fig. 11A, B); these depressions extend forward through a notch in the postnasal wall and might have allowed the musculature for the superior obliques to meet across the midline as suggested in  Kansasiella (Poplin, 1974) . The ventral anterior myodome (a.  v.myo; Figs 7A, B, 11A, B), located just below the olfactory tracts, appears as an indentation in the postnasal wall with two distinct depressions—likely origins for the inferior oblique muscles—and a dorsal midline passage that has been hypothesized to transmit the profundus nerve (V.pr; Fig. 11A, B) in the contemporaneous  Lawrenciella schaefferi (Hamel &amp; Poplin, 2008) . Ventral to the ventral anterior myodome is a thin midline ridge (l.r; Fig. 7A, B) rising from the basisphenoid. The postnasal wall broadens ventrally and forms rectangular, posterolateral-facing processes for articulation with the palatoquadrate (eth.p.a; Figs 9A, 11B). The anterior surface of the postnasal wall supports two lateral depressions that form the posterior wall of the nasal capsules (p.n.w; Fig. 11A).</p></div>	https://treatment.plazi.org/id/03C687D1FF91301AA11DFC29FD1DB9F8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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.text	03C687D1FF903005A297FB22FBADBB65.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kansasiella eatoni (Poplin 1974)	<div><p>Kansasiella eatoni</p><p>The ethmoid region is incompletely preserved in the present specimens; only the ventralmost part could be recovered. There is a small midline longitudinal ridge (l.r; Fig. 7C, D) which terminates anteriorly, presumably allowing for an unpaired anterior ventral myodome (a.  v.myo; Fig. 7C, D). The articulation for the palatoquadrate (eth.p.a; Figs 7C, 9C) is less pronounced than that of  Phoebeannaia, though positioned similarly with respect to the midline longitudinal ridge and postnasal wall. The probable lack of endochondral bone in the central ethmoid region is consistent with observations in other early actinopterygians (e.g.  Moythomasia,  Mimipiscis Choo, 2011,  Lawrenciella) (Gardiner, 1984; Hamel &amp; Poplin, 2008; Pradel et al., 2016).</p><p>PHYLOGENETIC RESULTS</p><p>Specimens NHMUK PV P 10419a &amp; b, FMNH UF464 and FMNH UF462 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. 12A) 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.</p><p>The backbone-constrained parsimony analysis retrieved 12 equally most parsimonious trees of 165 steps, and the strict consensus (Fig. 12B) 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.</p><p>The reweighted and backbone constrained parsimony analysis found three equally most parsimonious trees, and the strict consensus (Fig. 13A) 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. 13B) emphasize the similarity between these taxa.</p><p>The unconstrained Bayesian analysis (Fig. 14A) 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. 14B), resulting in the addition of the Chondrostei plus  Birgeria groenlandica Stensiö 1932 and  Saurichthys sp. (Argyriou et al., 2018) to the large polytomy.</p></div>	https://treatment.plazi.org/id/03C687D1FF903005A297FB22FBADBB65	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Caron, Abigail;Venkataraman, Vishruth;Tietjen, Kristen;Fls, Michael Coates	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
