identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
E95253443C22D54EFF4C95F5FB3CFC97.text	E95253443C22D54EFF4C95F5FB3CFC97.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Titanokorys gainesi Caron & Moysiuk 2021	<div><p>Titanokorys gainesi gen. et sp. nov.</p> <p>LSID urn:lsid:zoobank.org:act: F07D43CF-9422-4148-B8E9-AF26E0BC895D</p> <p>LSID urn:lsid:zoobank.org:act: 0EE750AC-FB83-4748-ADD9-013EB53459F1</p> <p>3.1.1. Etymology</p> <p>Genus name from Titans, a group of powerful Greek deities of great sizes, in reference to the large size of the central carapace element and from the Greek word Korys meaning helmet; gainesi, after Robert R. Gaines, Professor of Geology at Pomona College, who first joined the ROM-led field expeditions in 2008 as a research collaborator. Robert Gaines was instrumental in the co-discovery of the Marble Canyon fossil deposit in 2012 [26] and several related Burgess Shale outcrops along Tokumm Creek [25], including many fossils in this study.</p> <p>3.1.2. Type material</p> <p>Holotype — ROMIP 65415, a probable moult assemblage consisting of an H-(central) element, a pair of lateral (P) elements, a pair of frontal appendages, oral cone and probable gill blades (figure 3). Paratypes — ROMIP 65168, a complete H-element (figure 1); ROMIP 65741, an individual H-element with partial endites (figure 2); ROMIP 65748 and ROMIP 65749, two fragmentary H-elements (figure 4); other materials: seven additional H-elements, one associated with a P-element and poorly preserved appendages, in various states of preservation and completeness (electronic supplementary material, table 2.1).</p> <p>3.1.3. Locality and stratigraphy</p> <p>The upper part of the ‘Thick’ Stephen (Burgess Shale) Formation, Cambrian (Miaolingian Series, Wuliuan Stage), Ehmaniella biozone, from Marble Canyon [26,41] and Tokumm Creek [2,25] localities.</p> <p>3.1.4. Diagnosis for genus and species</p> <p>H (central)-element ovoid, bearing a broad anterior sagittal spine (ca 7% the length of the entire carapace), flanked by a pair of rounded processes pointing forward. Ocular notches moderately incised and positioned at the posterior of the H-element, flanking a short (ca 10–15% the length of the entire carapace), subtrapezoidal and slightly bilobate axial projection separated by a shallow medial notch. Spinous posterolateral processes are very short, about half the length of the bilobate axial projection. Elongate (sub-equal in length to H-element) and ovoid P (lateral)-element with a very short neck bearing a stout downward-pointing spine. Frontal appendages with elongate spiniform distal endites on podomeres 8–10 and elongate secondary spines on more proximal endites on podomeres 2–6; distal endites and secondary spines at least four times longer than the width of endite 6.</p> <p>3.1.5. Description: paratype</p> <p>ROMIP 65168 preserves a large (L: 27 cm; W: 18.5 cm) complete H-element in dorsal view and is the best example of this element for descriptive purposes (figure 1). The fact that it appears perfectly bilaterally symmetrical suggests that the carapace was buried and compacted with the dorsoventral axis perpendicular to bedding. The H-element is ovoid with the widest point about 60% from the front and width slightly decreasing along the posterolateral processes. Crescentic to linear compression artefacts, starting from the anterior along the midline and extending towards the lateral areas, suggests that the carapace was convex dorsally, curving down gently from the sagittal plane towards the margins. The front of the carapace yields a sagittal spine with a blunt tip (Sa in figure 1 a–c). This spine is about 7% the total length of the carapace and is roughly the shape of an equilateral triangle. Apair of semicircular to somewhat angular bulges, or anterolateral processes, flank the sagittal spine (Ap in figure 1 a–c). The base of each process is comparable in width to the base of the sagittal spine, but they protrude only a few millimetres forward.</p> <p>The posterior margin of the H-element projects into a pair of posterolateral processes and a trapezoidal axial projection, separated by ocular notches (On in figures 1 a,b and 3 a–c). The axial projection is subtly bilobate with smooth rounded posterolateral margins separated by a shallow medial notch (Mn in figures 1 a,b and 3 a–c). It represents ca 10–15% the length of the carapace and 60% of its maximal width anteriorly, becoming narrower towards the rear. This implies that the two ocular notches located on either side, are angled ca 20° anterolaterally from the midline. The posterolateral processes in Titanokorys are very short, extending only half as far posteriorly as the axial projection (Lp in figures 1 a,b and 3 a–c). Each process has a smooth ventral margin and terminates in a large spine (Sl 1 in figure 1 d,e). Asecond shorter spine, about 25% the dimensions of the large spine, is located along the laterodistal margin of the ocular notch, near the base of the large spine (Sl 2 in figure 1 d,e). Both spines are slightly curved such that they point posterolaterally.</p> <p>The entire H-element is ornamented by a series of longitudinal ridges running more-or-less parallel to each other (Ri in figures 1 a,c, 2 b and 4 a–c). The ridges occasionally converge, resulting in a reticulated pattern, similar to Cambroraster [2] but with particularly elongate and angular cells. The number of ridges probably ranges between 30 and 40 in total across the entire width of the H-element as seen from the top. Each ridge consists of a reflective band dotted with chains of repeated structures, each about 1 mm wide and 2 mm long (figures 2 d and 4 a–c). Tubercles are present along the margins of the carapace, in particular anteriorly and along the outside margin of the lateral processes (Tu in figures 1 d, 2 d, 3 a and 4 a–c). By their comparable sizes and arrangements, these seem to represent the lateral expression of the dorsal repetitive spots suggesting that the carapace is covered by rows of small elongated tubercles. These tubercles increase in size towards the terminal spine of the posterolateral processes (figures 1 d and 3 a).</p> <p>The pair of ventrolateral (P-) elements are best preserved in the holotype ROMIP 65745 (figure 3 a). The P-elements share the same overall dimensions (ca L = 20 cm and H = 7 cm) are almost perfect mirror images of each other and show only a few marginal compression artefacts suggesting they were buried more-or-less parallel to bedding. This is unlike the associated H-element, which by comparison with the paratype ROMIP 65168 (figure 1) is not bilaterally symmetrical, suggesting oblique burial. Retrodeformed longitudinally, the H-element, probably would have reached ca 21 cm in length and ca 15 cm in width (assuming a similar L/W ratio as ROMIP 65168), which is subequal to the length of the P-elements (figure 3 c). Each P-element is drop-shaped with a more linear dorsal and truncated posterior margins. The posterodorsal corners of the P-elements are slightly indented, probably representing ocular notches (On in figure 3 a). The paired P-elements are connected to each other by a very short anterior neck section (Pn in figure 3 a) which bears a single broad-based spine on its curved ventromedial surface (Ps in figure 3 a, d). This spine is about a quarter the height of the neck above it. The carapace surface, as in the H-element, bears longitudinal rows of tubercles. In the holotype (figure 3 e), the rows are more visible along the ventromedial surface and appear thinner and with smaller tubercles compared with the H-element in ROMIP 65168 (figure 1). Based on previous studies [1,2,4], the P-elements would wrap tightly laterally and ventrally along the body, with their anterior attachment point below the base of the H-element sagittal spine (figure 5).</p> <p>The holotype also includes a pair of frontal appendages and an oral cone (figure 3 b) as well as a number of indeterminate elements. Smooth plates of the oral cone are faintly visible below or on top of other fossil material (Oc in figure 3 g,h,k) and no fine details can be discerned. One appendage, in lateral view, can be distinguished more clearly using different low-angle light orientations (figure 3 g, h, k–n). It preserves the peduncle (Pd) and all the podomeres (Po), with five long endites (numbered according to their respective podomeres, En2 to En6) bearing distally pointing, gradually tapering secondary (or auxiliary) spines (Se). About 35 secondary spines are visible beyond endite 6, but this number includes overlapping spines from more proximal endites as well, so the true number of spines per endite may be closer to 20–25. The secondary spines are also well preserved in ROMIP 65741 (figure 2 c). Each of these spines on that specimen or the holotype is nearly four times longer than the width of endite 6, a third longer compared with equivalent spines in C. falcatus (figures 2 d and 3 i,j). It is unclear if the distal ends of the secondary spines were hooked like in C. falcatus, although they appear to be curved (figures 2 c and 3 l–n). The holotype shows evidence of unpaired spines and spiniform endites on more distal podomeres (probably 8, 9 and 10) (Sp in figure 3 g,h,k–n). In addition to sclerotized elements, the holotype also preserves bands of lamellae, showing individual gill elements (Gb figure 3 a; Ig figure 3 f). These are similar to those of other hurdiids (e.g. [1,5]) and were probably associated with the trunk and lateral flaps.</p> <p>The reconstruction (figure 5) is based on all of the fossil material available. Details of the trunk, eyes and flaps are not preserved but have been hypothetically reconstructed based on C. falcatus [2].</p></div> 	https://treatment.plazi.org/id/E95253443C22D54EFF4C95F5FB3CFC97	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, J. - B.;Moysiuk, J.	Caron, J. - B., Moysiuk, J. (2021): A giant nektobenthic radiodont from the Burgess Shale and the significance of hurdiid carapace diversity. Royal Society Open Science 8 (9): 210664, DOI: 10.1098/rsos.210664, URL: http://dx.doi.org/10.1098/rsos.210664
E95253443C28D54CFF4C961EFE79FCF7.text	E95253443C28D54CFF4C961EFE79FCF7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pahvantia hastata	<div><p>3.2. Morphological reinterpretation of Pahvantia hastata</p> <p>Lerosey-Aubril and Pates [22] recently described a fossil assemblage of Pahvantia (KUMIP 314089) showing a well-preserved tripartite carapace complex, identifying it as a radiodont, as well as what they interpreted as an unusual type of appendage with extremely long setose endites (their figure 3 a–c). This appendage was interpreted as having seven endites divided into two broadly different types and sizes. The two proximal endites they identified were short with about seven strong auxiliary spines each, with the second endite being about three times wider compared with the first endite. The five distal endites were long—up to four times the size of the proximal endites—and had up to 50–60 setae.</p> <p>There are a number of issues with the interpretation of the putative appendage in this specimen [2]. In particular, there is no evidence of podomeres associated with the elongate setose ‘endites’, and these ‘endites’ are irregularly bent, curving to varying degrees along their lengths, suggesting they were highly flexible. Endites are well sclerotized in other hurdiids and are therefore not typically deformed in this way [2,5]. Hurdiid endites generally exhibit a smooth mesial curvature [2], contrary to the sharp bends seen in the Pahvantia specimen. In addition, no hurdiid shows endites with clearly differentiated secondary spines and secondary setae in the same appendage. One possible exception is Aegirocassis which might have secondary spines on the peduncular endite, but these are not clearly preserved in the figured material (figure 2 a, b in [5]). Finally, the presence of two rather than a single peduncular endite as well as their small relative size is unprecedented [42].</p> <p>As is typical with Burgess Shale-type preservation, KUMIP 314089 is preserved as a part and a counterpart, with the split going through various superimposed layers of the fossilized tissues in such a way that different structures may be visible on each. The part is the best preserved; however, the counterpart shows some remnants of structures missing on the part (figure 6 b, c). When images of both part and counterpart are superposed, the stacked image shows clearly the presence of a distinct appendage, ca 10 mm in height, partly overlapping a larger array of ribbon-like structures (figure 6 b). The appendage bears five endites of similar lengths, each probably bearing seven to eight robust, hooked secondary spines and terminating distally in a similar hooked spine. These structures were partially described by Lerosey-Aubril and Pates [22] who interpreted them as two proximal endites, but we consider their ‘second endite’ to actually consist of three partially stacked endites with visible posterior margins. The most distal elongate endite (number six) was overlooked and is mostly preserved on the counterpart (only its tip is visible on the part) together with the dorsal sections of the podomeres, which are not preserved on the part. Asemicircular structure proximal to the enditebearing podomeres can be identified as the peduncle. Triangular projections dorsal and ventral to the peduncle might represent a dorsal spine and the remnants of a peduncular or shaft endite [42], respectively. Additional possible dorsal spines are present more distally. Overall, the newly recognized appendage is about half the maximum dimension of the structure previously described by Lerosey-Aubril and Pates [22]. The appendage, from the distal tips of its endites to the outer margin of the podomeres, also represents less than one-third of the length of the associated H-element on the same slab, which would make it more consistent with the carapace/appendage size ratio observed in other hurdiid assemblages such as Hurdia [4] and Cambroraster [2].</p> <p>The putative setose endites described by Lerosey-Aubril and Pates [22] are more comparable to structures interpreted as gill blades, associated with flaps or body segments in anomalocaridids and hurdiids [4,12,15] including Hurdia (figure 6 e–h). These bands of lamellate structures are not part of an appendage, nor are they likely to have been used for feeding (contra [22]), probably functioning instead for respiration [15]. In Hurdia, the gill blades are prominent ventrolaterally and can be very elongate as demonstrated in disarticulated or isolated material [4]. Stacks of Hurdia gill blades can be preserved in similar ways to those observed in the Pahvantia material, especially in disarticulated carcasses or moult assemblages (figure 6). In addition, the number of individual elements in the bands in Pahvantia is roughly similar to what is known in the gills of Hurdia [4] suggesting they are equivalent structures. The amorphous strand of material overlapping and extending beyond the margin of the appendage peduncle, originally interpreted as a part of the appendage, is more likely associated with the mass of gills and trunk cuticle.</p> <p>In conclusion, a re-evaluation of KUMIP 314089 based on the photographic material provided by the authors, leads us to demonstrate the presence of a nearly complete and partially unnoticed appendage. This appendage has the greatest similarity in morphology, particularly in terms of the number and form of the secondary spines, to Hurdia [2,4], suggesting an adaptation for capturing larger prey living along or in the sediment and thus probably a nektobenthic lifestyle, contra [22]. However, the subequal length of the five elongate endites as well as the shape of the H- and P-elements sets Pahvantia apart from Hurdia. Together with our phylogenetic results (see below), these differences justify the retention of Pahvantia as a distinct genus.</p> </div>	https://treatment.plazi.org/id/E95253443C28D54CFF4C961EFE79FCF7	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, J. - B.;Moysiuk, J.	Caron, J. - B., Moysiuk, J. (2021): A giant nektobenthic radiodont from the Burgess Shale and the significance of hurdiid carapace diversity. Royal Society Open Science 8 (9): 210664, DOI: 10.1098/rsos.210664, URL: http://dx.doi.org/10.1098/rsos.210664
