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Comments on the Squamation of Polish Lower Devonian Porolepiforms
Journal of Verterbrate Paleontology
Abstract
Porolepiform scales from the Lower Devonian of the Holy Cross Mountains, Poland, described from the ‘placoderm sandstone’ of the Daleszyce area, are revised. The aim of the present article is to organize the recently collected, but not formally described, porolepiform material from the Holy Cross Mountains, as well as specimens from erratic boulders collected near Gdynia (northernmost Poland), which can be referred to Porolepis. Previously collected and new material was found to contain at least two sarcopterygian taxa: Porolepis and Heimenia. Based on morphological and histological features, the described material has been divided into two scale assemblages. The first, which is assigned to Porolepis, possesses cosmine cover on the entire exposed area; in the second, assigned to Heimenia, the cosmine cover is reduced or absent. These features prove to be stable regardless of the position of the scale on the body and thus are taxonomically informative.
... Un for tu nately, the sec tion and its sedimentological de tails are not known to have been doc u mented at that time. Dur ing the past two de cades one of us (PS) has made ex ca va tions at the site, pri mar ily for ob tain ing new fos sils, and sev eral hun dred spec i mens have been collected and are un der study (Bur row and Szrek, 2018;Dec, 2019;Wilk et al., 2020;Szrek et al., 2021); it forms the larg est col lec tion of Early De vo nian ver te brates from the re gion as sembled so far. ...
... Mondéjar-Fernández and Clément (2012) noted that the mi gra tion must have ended be fore the clo sure of open ma rine links. The pres ence of Heimenia in the Holy Cross Moun tains was first sug gested by rvig (1969) based on the ma te rial de scribed by Kulczycki (1960) and cor rob o rated by Wilk et al. (2020). The rest of the fauna does not re flect such di rect sys tem atic af fin i ties (see Szrek and Dupret, 2017), and is char ac ter ised as tran si tional be tween Baltica and west ern Eu rope. ...
The Lower Devonian of the Holy Cross Mountains is well-known in the field of vertebrate palaeontology but remains unrecognized as regards palaeoenvironmental reconstructions. We therefore analysed the spatial distribution and relative abundance of fossil vertebrates in this area within one time interval. The fauna from an Early Devonian (Emsian) siliciclastic bone-bearing breccia (the “Placoderm Sandstone”) exposed in four sections of the Łysogóry region and five sections of the Kielce region was analysed with respect to the frequency of the remains and their taxonomic affinity. The relative abundances of agnathans, acanthodians, placoderms, osteichthyans and chondrichthyans suggest more open marine conditions in the Łysogóry region and more terrestrial-influenced in the Kielce region during the Emsian. The results show that the average agnathan and acanthodian content of the Łysogóry region is significantly larger than that in the Kielce region. On the other hand, there are relatively fewer osteichthyans in the Łysogóry region and a significantly higher proportion of bony fishes was recorded in the fauna of the Kielce region. Placoderms are characterized by their generally similar frequency in both regions and from site to site in each of them, though a greater abundance was noted from the Kielce region. Likewise differences in the proportions of particular groups in the Kielce region suggest a large variety of marginal-marine environments under the influence of factors that might have included marine currents and variable conditions around a river mouth.
Helwing (1666-1748)-forgotten pioneer of geology and palaeontology on Polish territory. Prz. Geol., 70: 547-556. A b s t r a c t. Georg Andreas Helwing (1666-1748) spent most of his adult life as a Lutheran pastor in his native town of Wêgorzewo (Angerburg), currently in NW Poland. He was well educated, studied at several universities, focusing on theology and botany. Helwing has been widely acknowledged for his work in botany; however, his achievements in the fields of geology and palaeontology are much less known. Helwing published two books containing illustrations of minerals and fossils from the territory of Poland: Lithographia Angerburgica (1717) and Lithographiae Angerburgicae pars II (1720). His first book contained the first-ever palaeontological illustrations to be published in Poland. It is divided into 7 chapters, with the first two chapters containing general information about Wêgorzewo and its surroundings, and about various types of rocks and soils (Terris, "earths"); these are followed by chapters devoted to various "stones" (Lapidibus). Helwing collected his specimens in the Warmia-Mazury region, usually in close vicinity of Wêgo-rzewo. Specimens described in his books included both autochthonous fossils found in situ, as well as allochthnonous fossils transported from Scandinavia by glaciers. The illustrations in Helwing's book were drawn with high attention to detail and could be used for reliable palaeontological analysis. Helwing maintained close contact with some of the brightest minds in science in Europe, including famous naturalist Johann Jakob Scheuchzer from Zurich, often nicknamed "father of palaeobotany". Four letters from Helwing to Scheuchzer are held in Zentralbibliothek in Zürich. The letters document that both scientists remained in close touch and that Helwing helped Scheuchzer with the preparation of an index for the second edition of one of the Scheuchzer's most famous book Herbarium Diluvianum (1723) that included also Polish names of various species described in it.
Cosmine is a combination of tissues (enamel, dentine, and bone) and a structure (pore-canal system) not found in living vertebrates. Long considered a synapomorphy of sarcopterygians (lobe-finned fishes), new discoveries and phylogenetic interpretations have shown that the assembly of cosmine is a deeper evolutionary event than previously thought and that cosmine constitutes a primitive feature of osteichthyans (bony fishes). A precursor of both cosmine and ganoine might have been present in the last common ancestor of sarcopterygians and actinopterygians (ray-finned fishes). However, despite numerous and thorough studies on cosmine since the late 19th century, its role and biological properties are still a mystery. Currently, the most consensual hypothesis considers cosmine as a metabolic regulator of mineral ions through intricated processes of deposition and resorption of dermal tissues. A new ontogenetic and phylogenetic scenario is emerging from microtomographic data revealing that a pore-canal system initially associated with multiple superimposed odontodes in stem osteichthyans and early sarcopterygians was reorganized and simplified in rhipidistians into a uniform cosmine sheet with a single layer of enamel overlying the dentine (eucosmine). In this review, I will summarize the current knowledge of cosmine origins, structure, development, and possible function based on previous research. I will also present new histological data on poorly known early sarcopterygians to highlight the importance of histological studies for upcoming analyses of sarcopterygian interrelationships. Future efforts to unravel the evolution of cosmine will need new virtual and three-dimensional approaches to paleohistology that will change our current framework of studying early vertebrate evolution and certainly revolutionize our understanding of this distinctive feature.
The site at Orłowo Cliff was used to analyse the stratigraphic position and palaeogeographic interpretations of the properties and depositional conditions of two basal tills from the Late Pleistocene. A multi-proxy approach involved lithofacies, petrographic analysis of the fine gravel fraction, analyses of indicator erratics and till fabric.
TL dating of intra-moraine deposits was used to determine depositional time frames of tills. The sediment profile at Orłowo Cliff
shows a distinct reduction in number of Pleistocene units. Obtained dating results suggest the presence of Middle and Late Pleistocene fluvial units. The main issue discussed is the stratigraphic position of the older till (Unit O-4). It can be assumed that this till was deposited probably during the Middle Weichselian (MIS 4). At Orłowo Horn the till of Unit O-4 reveals incorporation of the erratic material derived from an older till in the surrounded area (according to petrographic composition – probably from MIS 8). The younger till (Unit O-6) was deposited in the Late Weichselian (MIS 2). Moreover, the till of Unit O-6 is characterised by a significant shift towards the south-west in terms of the erratic origin in Unit O-4.
The siliciclastic sequence of the Lower Devonian of the southern part of the Holy Cross Mountains in Poland is renown for abundant vertebrate fossils, including ostracoderm, sarcopterygian, acanthodian, chondrichthyan, and placoderm remains. Study of the vertebrate assemblage from the “placoderm sandstone” from Podłazie Hill in the Holy Cross Mountains reveals that the remains belong to Kujdanowiaspis sp. among other unspecified actinolepids and brachythoracids. The Polish actinolepid material is characterised by sizes bigger than those of the Podolian specimens; this may be related to geographic variation. Owing to the proximity between Podolia and Holy Cross Mountains we suggest that Kujdanowiaspis sp. from the Holy Cross Mountains may be a refugee of some species of Kujdanowiaspis from the Lochkovian–Pragian of Podolia and/or from Spain. Some anterolateral plates provisionally assigned to Arthrodira indet. probably represent a new genus due to the high overlapping surface for median dorsal and anteroventrolateral plates.
Ganoine is a different enamel from “true” enamel. The crystallites end in the middle of the projection of epidermal cell onto the ganoine surface. In contrast, “true” enamel shows the borders of the hexagonal epidermal cells. Cosmine is a combination of tissues (“true” enamel and dentine) and a structure (the pore–canal system). The pore–canal system opens in regular arranged pores on the scale surface and has a network of horizontal canals (mesh canals). Cosmine is limited to sarcopterygians possibly above actinistians and onychodonts, ganoine to actinopterygians. – Actinopterygian scales possess a narrow peg, an anterodorsal extended corner and ganoine either as overlapping or as multilayered enamel, whereas sarcopterygian scales have a broad based peg, no extended anterodorsal corner with two exceptions, but sometimes an anteroventral extension and cosmine. – The genera Andreolepis, Dialipina, Naxilepis, Terenolepis, Ligulalepis and Orvikuina are actinopterygians based on their scale characters. Additionally, Guiyu, Meemannia, Achoania and Psarolepis possess actinopterygian features. According to its scale characters Lophosteus is a stem osteichthyan.
The monograph aims to revise the taxonomy of the East Baltic Silurian-lowermost Devonian vertebrates and determine their stratigraphic and facial distribution. The material on which this monograph is based has been collected during 1969-1982, from 27 Estonian and West Latvian bore cores, and from 57 Estonian outcrops. About 1300 samples from various facies of the East Baltic Silurian Basin have been studied: lagoonal, shoal, open shelf, slope and deep basin zones. The Baltic fauna has been compared with material from North Timan, the Central Urals, Severnaya Zemlya, Gotland and the Welsh Borderland (extracted from the Summary, p. 84).
The placoderm sandstone (Emsian, Holy Cross Mountains) exposed in the abandoned quarry at
Podłazie Hill was revisited and excavated during fieldwork conducted in 2011-2013. Bone-bearing
breccias were identified for the first time at this site and subjected to taphonomic analysis. Vertebrate
remains are dominated by heterostracans, while true placoderms compose less than 20% of the total
vertebrate assemblage. The high degree of fragmentation of the bones and low degree of abrasion
indicate that the remains were reworked and transported before final burial. This is consistent with the
mixed character of the bone accumulations, which comprise both open-shelf forms (acanthodians,
chondrichthyans) as well as those related to marginal-marine environments (placoderms and
sarcopterygians). The bone-bearing succession has been subdivided into five depositional facies
attributed to a coastal lagoon influenced by stormy, possibly tidal conditions. The occurrence of the
invertebrate trace fossil Ilmenichnus sp. accompanied by Lockeia and Monomorphichnus supports this
interpretation.
New material of the “porolepidid” Heimenia ensis (Porolepiformes, Dipnomorpha, Sarcopterygii), sheds light on the evolution of scale morphology in sarcopterygian fishes. This material consists of an articulated, nearly complete body and numerous isolated scales from the Emsian-Eifelian (upper Lower Devonian) of Spitsbergen. Heimenia squamation can be described as ‘transitional’ between that of Porolepis and the more derived Holoptychiidae. The cosmoid scales of Heimenia are divided into three morphotypes: (1) rhombic, (2) intermediate, and (3) rounded. The rounded scales are present in the anterior third of the trunk, intermediate scales occupy its middle third, and rhombic scales extend in the rear and all over the ventral region. The transition between scale morphotypes is progressive along the body. All scales bear traces of a cosmine covering, regardless of their overall shape. Paleohistological study shows that the relative thickness of the cosmine layer decreases whereas the relative thickness of the bony basal layer increases from the rhombic to rounded scales. This unique squamation provides new information about the evolutionary transition from thick/rhombic scales covered with cosmine to thin/rounded scales lacking cosmine in the Porolepiformes. Such morphological and histological changes also occurred by convergence among different groups of Middle–Late Devonian sarcopterygians (e.g., lungfishes and “osteolepiforms”). In Heimenia, the presence of rounded scales in the anterior part of the body is here regarded as an adaptation to a dynamic and agile life style and constitutes a new example to support the hypothesis of the anteroposterior spread of derived characters in fishes.
Recent work in the Late Devonian (Frasnian) Fram Formation of Ellesmere Island, Nunavut, Canada, has produced the first North American record of Laccognathus (Sarcopterygii, Porolepiformes), a taxon previously known only from Latvia and Russia. Represented by a large sample that includes cranial material from 22 individuals, Laccognathus embryi, sp. nov., is uniquely diagnosed by features including the absence of marginal teeth on coronoids 1 and 2, and the lack of a fifth posthyal branchial arch. The large sample allows detailed morphological description of many parts of the cranial skeleton, including the dermal skull, braincase, palatoquadrate, and branchial skeleton. Comparison of the faunas from the Fram Formation in Nunavut, the Escuminac Formation in Quebec, and the Lode and Gauja formations of Latvia confirms a Laurussian (Euramerican) biogeographic province in the late Middle/early Late Devonian and may indicate tolerance of some forms to varied salinities. A cranial reconstruction of the new taxon reveals a wide head with a width-to-length ratio of approximately 2:1, a large spiracular opening, a small orbit, and a lower jaw in which much of the labial surface faced ventrally. The cranial morphology suggests a benthic animal reliant on sit-and-wait predation.
Richard Owen (1804–1892) was a contemporary of Darwin, and like him, attended the University of Edinburgh medical school but left without completing his training. His career as an outstanding palaeontologist began when he was cataloguing the Hunterian Collection of human and animal anatomical specimens which had passed to the Royal College of Surgeons in London. His public lectures on anatomy were attended by Darwin, and he was entrusted with the classification and description of the fossil vertebrates sent back by Darwin from the Beagle voyage. He was responsible for coining many of the terms now used in anatomy and evolutionary biology, including the word 'dinosaur'. Palaeontology (published in 1860) defines, describes and classifies all the fossil animal forms then known, and discusses the origin of species, commenting on the theories of Buffon, Lamarck, the then anonymous author of Vestiges of Creation, Wallace and Darwin.
Newly collected and restudied earlier materials on an enigmatic fish Ventalepis ketleriensis Schultze, 1980 from the upper Famennian (postera - ? Lower expansa conodont zones) of Latvia and central and northwestern Russia support its porolepiform affinities. A new family Ventalepididae fam. nova is established for this genus upon a peculiar combination of characters, including scale structure and dermal bones ornamentation. New records extend the distribution of this genus and the Ventalepis vertebrate assemblage on the whole to a vast geographical zone along the south-eastern coast of the Old Red Sandstone continent. The habitat area of the Devonian vertebrate assemblage over such a large territory within the zoogeographical province of Baltica is established for the first time. Palaeozoogeographical analysis suggests Laurentian affinities of the Ventalepis assemblage demonstrating the major congruency to the Belgian and East Greenland ones. These and Russian localities are separated by a vast ORS continent. Presence of the dipnoan Jarvikia in all three locations, as well as an Ichthyostega-like tetrapod in the Belgian one reveals palaeozoogeographical connections, which might reflect possible dwelling not only in the near-shore continent periphery but also in the river systems of the continent itself.
Recent advances in synchrotron imaging allow us to study the three-dimensional (3D) histology of vertebrate fossils, including microfossils (e.g. teeth and scales) of early jawed vertebrates. These microfossils can often be scanned at submicron resolution (<1 µm) because of their small size. The resulting voxel (3D pixel) stacks can be processed into virtual thin sections revealing almost every internal detail of the samples, comparable to traditional thin sections. In addition, 3D models of the internal microanatomical structures, such as embedded odontodes and vasculature, can be assembled and examined in situ. Scales of two early osteichthyans, Psarolepis romeri from the Early Devonian of China and Andreolepis hedei from the Late Silurian of Sweden, were scanned using propagation phase-contrast synchrotron X-ray microtomography (PPC-SRµCT), and 3D models of internal canal systems and buried odontodes were created from the scans. Based on these new data, we review the evolutionary origin of cosmine and its associated pore-canal system, which has been long recognized as a synapomorphy of sarcopterygians. The first odontode that appeared during growth shows almost identical morphology in the two scales, but the second odontode of the Psarolepis scale shows a distinctive morphology with several pores on the surface. It is suggested that a shift from ridge-like odontode to pore-bearing odontode was the key step in the origin of cosmine, which was then elaborated further in more-derived sarcopterygians. We perform a detailed comparison between the two scales and propose a primary homology framework to generate microanatomical characters, which can be used in the phylogenetic analysis of early osteichthyans when more 3D data become available. Our results highlight the importance of 3D data for the study of histology and ontogeny of the dermal skeleton of early jawed vertebrates, especially scales of the polyodontode type. The traditional microvertebrate collection is not only useful for biostratigraphic studies, but also preserves invaluable biological information about the growth of vertebrate hard tissues. Today, we are only beginning to understand the biological meaning of the new 3D data. The increasing availability of such data will enable, and indeed require, a complete revision of traditional palaeohistological studies on early vertebrates.
An exceptionally well-preserved assemblage of numerous invertebrate and vertebrate trace fossils is described from the Lower Devonian of the Holy Cross Mountains, southern Poland. Two trace-bearing horizons occur in the shallow-marine sequence that is exposed in a small outcrop near Ujazd village. One of the trace fossils is preserved as a bilobate, generally elliptical, epichnial pit is described as Osculichnus tarnowskae isp. nov. and interpreted as a unique example of praedichnia. Neoichnologic experiments and observations indicate that the ichnogenus Osculichnus was produced by feeding fish. The fish producing O. tarnowskae probably hunted bivalves, polychaetes and arthropods, which are represented by invertebrate trace fossils in the same horizons. The overall shape and morphological details of O. tarnowskae suggest that it was made by a lungfish broadly similar to Dipnorhynchus. The trace provides the first direct evidence for Devonian lungfish feeding behaviour, as well as the first record of three-dimensional soft-tissue morphology of the snout area of an Emsian representative of this group. The trace fossils from Ujazd provide new insight into the palaeoecology and taphonomy of the Lower Devonian of the Holy Cross Mountains.
Synopsis
Well preserved material of the crossopterygian fish Eusthenopteron enables fresh reconstructions and interpretations of its postcranial skeleton to be given. Comparisons throughout with other bony fishes show that it may be primitive in many features. Similarities with early amphibians such as the screw-shaped glenoid, the form of the humerus (on which an attempt to restore the pectoral musculature is based), the dorsal bicipital ribs and the possibility of a sacral attachment, throw much light on the origin of the tetrapod postcranial skeleton, particularly of the cheiropterygium. A functional analysis of the skeleton of Ensthenopteron is attempted, suggesting that it resembled the pike ( Esox ) in its mode of life and that it may have been capable of short journeys “walking” overland. The possible selective factors stimulating the evolution of such a fish, and further evolution to the tetrapod stage are discussed.
In the Baltic region, Devonian sedimentary bedrock is exposed in a wide area extending from north-eastern Estonia through Latvia to the south-eastern Baltic Sea. Westwards in the Baltic Sea, the outcrop terminates against a presumably Permian fault, which extends southwards from Gotland. In the southernmost part of the Baltic Sea, the Devonian sedimentary bedrock was eroded during the Permian-Carboniferous uplift event. The Devonian sedimentary bedrock of the investigated area southeast of Gotland, consists of near-shore marine deposits, namely sandstone, siltstone, clay, dolomite, dolomitic marl, and limestone. Gypsum and anhydrite deposits occur frequently in the lower parts. The Silurian-Devonian boundary is marked by a stratigraphic break across the presently investigated offshore area. All regional stages of the Lower and Middle Devonian are present in exposures in the area. In the north-eastern part, offshore Latvia, the basal Devonian Gargždai deposits follow ontop of the Silurian sedimentary bedrock. Further to the south-west, the Gargždai beds are missing, and deposits of the next younger Kemeri Stage rest directly on the Silurian. In the Middle Devonian, the Pärnu Stage contains widespread sandbars, which are locally deeply truncated. This may indicate a shallow nearshore environment. Within the Old Red Sandstone facies of the late Middle Devonian Aruküla-Burtnieki complex, several discontinuity surfaces are present. The reflectors exhibit a mound-like pattern, which again may indicate a shallow nearshore environment with rapid deposition, erosion, and redeposition.
This paper deals with various vertebrates from the Verdalen Member of the Stjørdalen Formation of the Wood Bay Group, Vestspitsbergen, or layers presumably belonging to this member, including the thelodont scales Amaltheolepis winsnesi n.g., n.sp. (earlier known from a locality in Sørkapp Land), the heterostiid brachythoracid Herasmius granulatus n.g., n.sp. and the porolepiform crossopterygian Heimenia ensis n.g., n.sp., the latter with scales intermediate in some ways between those of Porolepis on the one hand, and the Holoptychiidae on the other. Amaltheolepis and Herasmius are not yet known to occur outside Vestspitsbergen; Heimenia is represented in a Devonian material from Anderson River, Northwestern Canada and perhaps also in the Placo-derm Sandstone of Poland. A critical appraisal of the arthrodire faunas of the Wood Bay Group shows that most of the earlier recorded occurrences of Homostius and Heterostius in these layers pertain to euleptas-pid brachythoracids, that the Heterostiidae and Homostiidae are also represented, but only in the Stjordalen Formation, and that the youngest member of the Euleptaspididae, occurring in the Stjordalen Formation, may be closely related to another euleptaspid in the Upper Emsian Heisdorf Beds of the Rhinish sequence (which occurs in association with Ohiolepis scales). The palaeontological evidence may indicate that the Wood Bay Group does not belong to the Lower Devonian in its entirety but includes Middle Devonian strata in its uppermost part.
Subscribe now ». Research-paper. from the with special reference to Powichthys Thorsteinssoni Jessen. Jessen, HL. Palaeontographica Abteilung A Band A167, Lieferung 4-6 (1980), p. 180 - 214 published: 1980-01-01. Keywords
Consequently, a detailed knowledge of these early fossil forms must be of fundamental importance for questions concerning the
The anatomy of the axial skeleton, median fins and skull of the fossil group Porolepiformes (Osteichthyes: Sarcopterygii) is reviewed, and descriptions by previous authors are modified in the light of new evidence. The first dorsal fin support lacks radials, whereas the posterior radials in the second dorsal fin support form a branching structure. In the snout, porolepiforms primitively have two lateral rostral bones on each side, the posterior one corresponds to the “nariodal” of Jarvik, and the infraorbital canal runs along the suture between this bone and the premaxilla. A cladistic analysis based on 54 characters places the Porolepiformes as sister‐group of a clade Powichlhys + Youngolepis + Diabolepis + Dipnoi. The postcranial skeletons of porolepiforms and early lungfishes are very similar, and the skulls of Powichthys, Toungolepis and Diabolepis can be interpreted as forming a transformation series between porolepiform and dipnoan cranial morphologies. Lungfishes thus appear to be derived from ‘rhipidistian’ ancestors. The porolepiform‐lungfish clade is the sister group of another clade containing tetrapods, panderichthyids, osteolepiforms and rhizodonts. Onychodonts and coelacanths are joing unresolved sister groups of the other sarcopterygians. This cladogram is compared with those of other authors, and possible explanations for the differences are discussed. An outline classification of the Sarcopterygii is proposed.
Page 1. <oological Journal of the Linnean Sociely (1989), 96: 119-166. With 16 figures and of the fossil group Porolepiformes (Osteichthyes: Sarcopterygii) PER ERIK AHLBERG, FLS Department
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