identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038D5604FFDC0447FF70034CF075FCB1.text	038D5604FFDC0447FF70034CF075FCB1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Acanthodii Owen 1846	<div><p>Class ACANTHODII Owen, 1846</p> <p>REMARKS</p> <p>The order Climatiiformes is thought to contain the most primitive acanthodian species, including Lupopsyrus pygmaeus. The diagnosis of the Climatiiformes provided by Denison (1979) included acanthodians with enlarged cranial tesserae and scales, a dermal shoulder girdle including pinnal and lorical plates, and in some cases, prepectoral spines, plus all climatiiforms possessed two dorsal fins. Not surprisingly, the diagnosis of the order has changed with each new publication as new taxa are shoe-horned into the classification scheme. Gagnier &amp; Wilson (1996a) revised this diagnosis and limited the climatiiform character list to include: scales with, or derived from Nostolepis Pander, 1856, type histological structure, the presence of two dorsal fins, and the presence of fairly large head scales to accommodate Kathemacanthus rosulentus Gagnier &amp; Wilson, 1996a, and Brochoadmones milesi Bernacsek &amp; Dineley, 1977; note that K. rosulentus has been recently reclassified as a putative chondrichthyan based primarily on scale growth (Hanke &amp; Wilson 2010). Gagnier &amp; Wilson (1996a) excluded pinnal and lorical plate armour and prepelvic spine presence from their climatiiform character list, to incorporate their new taxa and because mesacanthids also possess prepelvic spines (see: Egerton 1861; Miles 1966, 1973; Denison 1979; Gagnier 1996; Upenice 1996; Cumbaa &amp; Schultze 2002; Hanke 2008). Some mesacanthids also have fairly large head scales, so we think that this feature too is not unique to climatiiforms. Support for Gagnier &amp; Wilson’s decision to eliminate prepectoral and prepelvic spines as a climatiiform characteristic follows the discovery of several new taxa from MOTH which possess these spines but lack characteristic scales and perichondral bone of acanthodians (Hanke &amp; Wilson 1998, 2004, 2010; Wilson&amp; Hanke 1998). This leaves us with: 1) scales derived from a Nostolepis - type of histology; and 2) two dorsal fins, as potential features defining the order Climatiiformes.Two dorsal fins are present in non climatiid acanthodians such as: diplacanthids, ischnacanthids and also gyracanthids plus early chondrichthyans, osteichthyans and sarcopterygians (Janvier 1996). Furthermore, most “ Nostolepis ” species are known only from isolated microremains, with the exception of a few taxa (Valiukevičius 2003a; Burrow &amp; Turner 2010), and acritolepid ischnacanthiforms have Nostolepis - type scale histology (Valiukevičius &amp; Burrow 2005). As a result we cannot support the definition of climatiiform acanthodians using histological features now known to exist outside the group. This historical perspective shows there are no synapomorphies to unite the climatiiforms as historically defined (Janvier 1996; Hanke 2001; Davis 2002; Hanke &amp; Wilson 2004; Burrow &amp; Turner 2010), and as a result, the higher classification of L. pygmaeus is left open pending detailed reexamination of climatiiform fishes.</p> </div>	https://treatment.plazi.org/id/038D5604FFDC0447FF70034CF075FCB1	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	Hanke, Gavin F.;Davis, Samuel P.	Hanke, Gavin F., Davis, Samuel P. (2012): A re-examination of Lupopsyrus pygmaeus Bernacsek & Dineley, 1977 (Pisces, Acanthodii). Geodiversitas 34 (3): 469-487, DOI: 10.5252/g2012n3a1, URL: http://dx.doi.org/10.5252/g2012n3a1
038D5604FFDC0447FC85018BF06DFA92.text	038D5604FFDC0447FC85018BF06DFA92.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lupopsyrus Bernacsek & Dineley 1977	<div><p>Genus Lupopsyrus Bernacsek &amp; Dineley, 1977</p> <p>REVISED DIAGNOSIS. — As for the type and only species. Note that Bernacsek &amp; Dineley (1977) provided a diagnosis of the genus Lupopsyrus, and suggested that characteristics of the single included species were “as for the genus”. Here the species is rediagnosed and the characteristics of the genus are dependent on features of the included species, not the reverse.</p> <p>AGE. — Early Devonian (Lochkovian).</p></div> 	https://treatment.plazi.org/id/038D5604FFDC0447FC85018BF06DFA92	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	Hanke, Gavin F.;Davis, Samuel P.	Hanke, Gavin F., Davis, Samuel P. (2012): A re-examination of Lupopsyrus pygmaeus Bernacsek & Dineley, 1977 (Pisces, Acanthodii). Geodiversitas 34 (3): 469-487, DOI: 10.5252/g2012n3a1, URL: http://dx.doi.org/10.5252/g2012n3a1
038D5604FFDC044DFCF10737F7B8FDD3.text	038D5604FFDC044DFCF10737F7B8FDD3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lupopsyrus pygmaeus Bernacsek & Dineley 1977	<div><p>Lupopsyrus pygmaeus Bernacsek &amp; Dineley, 1977 (Figs 1-6)</p> <p>HOLOTYPE. — NMC 22715 (Bernacsek &amp; Dineley 1977: text-fig. 3A, pl. 1).</p> <p>REFERRED MATERIAL. — NMC 22700B, C, 22718, 22719, 22700D-F, 22701C, D, 22716, 22717, 22720, 22745. — UALVP 19260, 32420, 32442, 32456, 32458, 32474, 32476, 32480, 32482, 39065, 39067, 39079-</p> <p>39082, 39121, 41493, 41629, 41632, 41665, 41931, 41939, 41945, 42000, 42002, 42008, 42012, 42013, 42027, 42046, 42061, 42113, 42142, 42150, 42173, 42208, 42274, 42518, 42524, 42529, 42530, 42533, 42538, 42453-42455, 42544, 42597, 42605, 43064, 43091, 43092, 43094, 43095, 43256, 43409, 43456, 45154, 45155.</p> <p>LOCALITY AND AGE. — Lupopsyrus specimens are recovered from talus below a Lower Devonian (Lochkovian) horizon between 430-435 m in the MOTH locality section (as measured in 1996), central Mackenzie Mountains, N.W. T., Canada (see Hanke et al. 2001b: figs 1, 2); approximately 411 m in the section measured by the Geological Survey of Canada (Gabrielse et al. 1973). The Devonian fish layer in the MOTH locality section is equivalent to GSC locality 69014 in section 43 of Gabrielse et al. (1973) and locality 129 in the UALVP catalogue system. Although previous authors have suggested habitats ranging from intertidal lagoons to deep-water shelf settings, recent sedimentological, ichnological and taphonomic study suggests an oxygen-poor, intra-shelf topographic low below storm wave base (Zorn et al. 2005) on a shelf that fringed western Laurussia (combined Laurentia and Baltica; Li et al. 1993).</p> <p>REVISED DIAGNOSIS. — Acanthodians with two longitudinal rows of enlarged keeled scutes situated along the posterior half of each side of the body and caudal fin axis; largest keeled scutes located below the second dorsal fin base; three dermal opercular gill plates per side; pectoral, pelvic, anal and dorsal spines with widely separated ribs with fine nodular ornamentation; single prepectoral spine positioned over lateral end of each procoracoid; prepelvic fin spines with unornamented blade-like posterior lamina; circumorbital scales identical to head scales; head and body scales have crown with prominent median keel and one lateral flange per side; central keel and lateral flanges of each scale terminating posteriorly in a single point; scales are monodontode with mesodentine, Stranggewebe-like crown histology.</p> <p>DESCRIPTION</p> <p>Much of the dermal covering over the head of Lupopsyrus pygmaeus is preserved on UALVP 41493 (Fig. 1), and other new Lupopsyrus specimens in the University of Alberta collections. The head of L. pygmaeus as preserved is short at 10 % of body length, but the rostrum and jaws either have lost scale cover in all specimens, or lacked scale cover in life and therefore, the full snout length cannot be estimated (Figs 1; 2B); the jaws may also have fallen clear of the carcass in all specimens during decay. The dorsal surface of the head is covered with small scales which are similar in structure to body scales. These head scales differ slightly from those on the body in that the posterior apex of the crown of each scale is short, blunt and does not extend posterior to the basal rim surrounding the pulp canal. The gradual transition between head and body scales occurs over the branchial chamber; enlarged head scales and/or tesserae are absent (Figs 1; 2).</p> <p>The orbits of L. pygmaeus lack ossified sclerotic plates and are highlighted by an area of silveryblack material within the ring of micromeric circumorbital scales (note that terminology follows Burrow et al. [2011]). The dorsal and posterior margin of the orbit has scales which are identical to normal head scales, and there are no enlarged circumorbital scales (Figs1; 2). There are no scales anterior and ventral to the orbit in any of the available specimens, therefore the lachrymal and labial portions of the head are unknown and may have lacked scales in life.</p> <p>The jaws, branchial arches and endocranium are not mineralized. Teeth are absent and there is no indication of jaw extent or shape. The position of the otic capsule, and therefore, the position near the posterior end of the braincase is indicated by small masses of sandy, otic material, which Sahney&amp; Wilson (2001) interpret as otic statoconia. When scale cover is complete, these masses appear only as paired bulges (Figs 2B; 4B), but where scales are lost, the light coloured sandy material is exposed. Many L.pygmaeus specimens have heads preserved as dorsoventral compressions, and the masses of otic material are well-separated (Figs 2B; 4B). This preservation suggests that the braincase and head of L. pygmaeus was fairly broad rather than laterally compressed; a broad, depressed braincase is characteristic in several clades of gnathostomes (Janvier 1996) and likely is a primitive feature for gnathostomes.</p> <p>Sensory lines preserved on very few L. pygmaeus specimens run between scales (Figs 1; 2). The supramaxillary sensory canal and preopercular sensory canals converge posteriorly across the cheek anterior to the hyoidean gill plates (Fig. 1); the latter canal meets the main sensory line dorsally. The supraorbital sensory canal traces converge posteriorly, but its entire course cannot be determined in the available specimens (Fig. 2A, B). One specimen shows the trace of the infraorbital canal posteroventral to the orbit (Fig. 2B). There also are traces of short, paired, converging, middle pit lines, and posterior pit lines preserved near the level of the otic region of the braincase (Fig. 2A). Behind the posterior pit lines are a pair of gaps in the scale cover which may indicate the position of the external endolymphatic duct openings (Fig. 2A). The occipital cross-commissure is seen as a short canal trace leading dorsally from the main lateral canal level with the origin of the pectoral fin spine (Fig. 1B).</p> <p>The branchial chamber of Lupopsyrus pygmaeus is not well-preserved in any specimens, but appears compact relative to the orbito-otic region; the extent of the branchial chamber is estimated from the position of the otic material and the position of the pectoral girdle (Fig. 2B). There is no evidence of gill openings, single or multiple, but the presence of three dagger-like dermal plates located in an arc over the opercular region indicates that a single opercular flap was present rather than a series of narrow separate opercular flaps (Figs 1; 2A, B). The middle plate of the operculum is larger than the dorsal and ventral plates, and each plate has a single longitudinal keel, surrounded by small tubercles. This keel is serrated, and its summit is near the mid-length of the plate. The underside of each opercular plate possesses a shallow trough which continues along the entire length of the plate; the basal rim is tear-drop shaped, widest anteriorly, and tapers posteriorly.</p> <p>The dorsal fins and spines were described in detail by Bernacsek &amp; Dineley (1977), although they did not mention that the anterior dorsal fin spine had a shallow insertion area (Fig. 1). This basal portion presumably anchored the spine into the epaxial musculature. Lupopsyrus pygmaeus lacks enlarged scales around the base of each fin spine.</p> <p>The anterior and posterior dorsal fin spines support fin webs which possess a convex trailing margin extending posterior to the apex of the fin spine, and also have irregularly-arranged scales (Figs 1; 3A). The distal half of each dorsal fin web appears to be detatched from the dorsal fin spine (Fig. 3A), but this may be an artefact of preservation or preparation. Fin-web scales are minute, and apart from size, are identical to typical body scales. There is a gradual size transition between typical body scales and small fin web scales (Fig. 3A, B).</p> <p>The pectoral fin spine was described in detail by Bernacsek &amp; Dineley (1977) and no new information can be added in our account. However, details of the pectoral dermal plate armour require further clarification. Bernacsek&amp; Dineley (1977) mentioned that a single lorical plate, with a single, median spine was present on the holotype of Lupopsyrus pygmaeus. They also indicated that the lorical plate was poorly preserved, as is evident in the photographs published with the species description. A median lorical plate is not known in any University of Alberta Lupopsyrus specimens (Figs 2A, B; 4), although in one, a small pile of displaced scales positioned posterior to the pectoral girdle creates the impression of a small, elevated node (Fig. 4B). It may be possible that the “lorical plate” described by Bernacsek &amp; Dineley (1977) was a similar mass of scales.Specimens examined at the Canadian Museum of Nature lack both lorical plates and median “lorical” spines.</p> <p>The margins of the prepectoral spines cannot be distinguished in the figures presented by Bernacsek&amp; Dineley (1977), and they suggest that the prepectoral spines are completely fused to what they described as compound pinnal plates. The University of Alberta specimens show that the prepectoral spines have a distinct basal rim which denotes the perimeter of each spine (Figs 2A; 4), and these spines sit over the lateral end of procoracoid bones (see below). The prepectoral spines are curved, have a broad basal cavity, and possess longitudinal ribs which are ornamented with fine nodes (Figs 2A; 4).</p> <p>Bernacsek &amp; Dineley (1977) wrote that L. pygmaeus had compound pinnal plates, and went so far as to suggest a homology between the first and second pinnal plates of Climatius reticulatus. In fact, the bones that subtend both prepectoral spines and contact the anterior-most base of the pectoral spines are smooth, unornamented, and have a surface texture similar to that of the perichondrally-ossified scapulocoracoids, not like the dermal tuberculated plates which are characteristic of climatiid pectoral dermal armour (Miles 1973; Watson 1937; Denison 1979). In L.pygmaeus, these bones extend anteromedial to the scapulocoracoid and meet at the ventral midline (Figs 2A, B; 4). We interpret these smoothly ossified structures to be procoracoids, based on structure and similarity of position to procoracoids of other acanthodians. The procoracoids have a small fossa on the lateral margin (Fig.4) which may have served as the point of articulation for the proximal end of the pectoral fin spine. A revised reconstruction of the pectoral girdle of Lupopsyrus pygmaeus, based on UALVP 32476, 39079 and 39080 (Figs 2; 3) is presented in Figure 5. Any apparent ornamentation on the procoracoids (Fig. 4A) consists of overlying scales which had settled on the bone during decay and preservation of the carcass.</p> <p>Two specimens (UALVP 39080 and 41493) show that a pectoral fin web was present posterior to the pectoral fin spine.The fin web is covered with minute scales, and the trailing edge of the fin probably was convex (Fig. 1). The fin web is not attached to the fin spine in UALVP 41493; additional specimens will be needed to determine whether the detatchment of the pectoral fin web in UALVP 41493 is a taphonomic artefact.</p> <p>Little can be added to the descriptions of the prepelvic and pelvic fin spines. However, since the material available to Bernacsek &amp; Dineley (1977) was poorly preserved, they could not determine which prepelvic spine had a posterior, flat lamina. Based on the material at the University of Alberta, all prepelvic spines of Lupopsyrus pygmaeus possess this flat, unornamented, trailing lamina, which is most prominent on the posterior-most prepelvic spine pair (Fig. 1).</p> <p>The pelvic fin web has a convex distal margin which extends beyond the apex of the fin spine, and overlaps the origin of the anal fin spine (Fig. 1). The anal fin web also has a convex posterior margin, is sub-triangular, and extends posterior to the distal tip of the anal fin spine. Both the pelvic fins and anal fin are covered with scales which are identical to those of the dorsal fins, and the transition between typical body scales and those of the pelvic and anal fins is gradual (Fig. 3B).</p> <p>The epicercal heterocercal caudal fin is elongate with only a slight dorsal deflection relative to the body axis (Figs 1; 3C). The hypochordal lobe is elongate and the entire tail superficially resembles those of several extant slow swimming sharks such as the frilled shark, hexanchids and scyliorhinids (see examples in Castro 1983; Compagno et al. 2005). The posterior end of the caudal fin axis extends beyond the hypochordal lobe of the fin (Fig.3C). The leading edge of the hypochordal lobe is covered with scales which are similar to those on the rest of the fin (Fig.3D); this condition is in contrast to that of other acanthodians where slightly enlarged scales reinforce the leading edge of the caudal fin.Body scales on the caudal fin axis grade into typical caudal fin scales with no abrupt change in scale size, and these irregularly-arranged scales decrease in size along the caudal axis toward the posterior tip (Figs 1A; 3C).</p> <p>The scales of L. pygmaeus change little in size over the body (Fig. 1), although as mentioned above, smaller scales are found on the fin webs, the posterior portions of the caudal fin axis, and on the head (Figs 1A; 2; 3). Bernacsek &amp; Dineley (1977) described the external ornamentation of the scales of L. pygmaeus, but their photographs did not show the fine details of scale structure and pattern. All scales are ornamented with a central crest and a pair of lateral flanges which possess smooth edges (Fig.6 A-D) and merge posteriorly to form the trailing tip of each scale. The pulp cavity extends into the posterior portions of the central crest (Fig. 6C). Lupopsyrus scales have a developed neck, but no neck canals were identified in the scales which were sectioned. Internally, the scales of L. pygmaeus are simple monodontode structures (Fig. 6G). The scale crowns appear to be mesodentinous with parallel cell spaces resembling that of Stranggewebbe; the tissue around the basal rim of the neck also may be cellular (Fig. 6G), but the poor preservation of histological detail prevents detailed comparison. The underside of the scale neck forms a concave, rhombic rim which lacks traces of Sharpey’s fibres or cellular basal tissue and the pulp cavity remains open (Fig. 6A, B). The apparent lack of Sharpey’s fibres is problematic given that MOTH fishes have poor histological preservation. It is also impossible to determine whether the scales of L. pygmaeus were aligned in life, although the rhombic shaped neck and basal rim (Fig. 6B) suggests that scales could have aligned in oblique rows.</p> <p>Similar monodontode scales lacking basal tissue are known in thelodonts and certain chondrichthyans as well as modern elasmobranchs (Karatajute-Talimaa 1973, 1992, 1998; Turner 1991; Hanke &amp; Wilson 2004; Märss et al. 2002). Bernacsek&amp; Dineley(1977) suggested that the scales of L. pygmaeus were simple as a secondary specialization rather than a primitive feature relative to the complex, layered scales of other acanthodians (and compound scales of some putative chondrichthyans).</p> <p>Little can be added to the description of the flank scutes of Lupopsyrus pygmaeus (Fig.6E, F). The scutes are largest at the level of the second dorsal fin and on the caudal peduncle (Figs 1; 3C). This region corresponds to the region where scales are first added during ontogeny in acanthodid species (Zidek 1985, 1988), and may indicate a similar origin for scale development in L. pygmaeus. There is no evidence that the scutes conducted the main sensory canal as discussed by Bernacsek &amp; Dineley (1977), but the scutes may have functioned to direct water along the caudal peduncle and reduce turbulence as do scales of extant sharks (Reif 1978; Reif &amp; Dinkelacker 1982; Dean &amp; Bhushan 2010).</p> </div>	https://treatment.plazi.org/id/038D5604FFDC044DFCF10737F7B8FDD3	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	Hanke, Gavin F.;Davis, Samuel P.	Hanke, Gavin F., Davis, Samuel P. (2012): A re-examination of Lupopsyrus pygmaeus Bernacsek & Dineley, 1977 (Pisces, Acanthodii). Geodiversitas 34 (3): 469-487, DOI: 10.5252/g2012n3a1, URL: http://dx.doi.org/10.5252/g2012n3a1
