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Although the record of Paleozoic drillholes is long and extensive, evidence pertaining to the identity of the drillers is sparse. The most conclusive evidence, a driller “caught in the act”, has been documented only once (Baumiller 1990). In that example, a drillhole in the calyx of a crinoid was found directly beneath an attached platyceratid gastropod. Additional evidence for drilling by platyceratids has been circumstantial, i.e., based on the association of platyceratids with certain blastoids and crinoids, and the presence of drillholes in other crinoid and blastoid taxa. To a skeptic, the lack of congruence between drilled and platyceratidinfested crinoids and blastoids is not sufficient evidence that platyceratids were the drillers. More conclusive evidence requires examples of drillholes in taxa that are known to have been platyceratid−infested, preferably from localities where both infested specimens and drilled specimens co−occur.
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Roman Kozłowski (1889-1977)

63%
The position of the Palaeozoic Anthracoptilidae has been a major problem of insect systematics for over a century. The previous hypotheses suggested affinities of Anthracoptilidae with the Palaeodictyopteroida, Protorthoptera, Hypoperlida, stem-Mantodea, Paraneoptera, Holometabola, or Eoblattida. Herein we put forward a new hypothesis based mainly on a comprehensive re-evaluation of the wing venation characters and re-examination of the type material of Anthracoptilus perrieri and Mesoptilus dolloi. The Anthracoptilidae are considered as belonging to the paoliid lineage, sister group of the Dictyoptera. In particular this result refutes the presence of Mantodea in the Paleozoic fossil record. The two families Strephocladidae and Strephoneuridae, are considered as junior synonyms of the Anthracoptilidae, while the previous synonymy of the Anthracoptilidae with the Ischnoneuridae is rejected. We consider the Permarrhaphidae, previously considered as synonym of the Anthracoptilidae, (and Permarrhaphus) as Insecta incertae sedis. The following new taxa are proposed: Mesoptilus carpenteri sp. nov. from the early Permian of Wellington Formation in Oklahoma (USA) which extends the range and distribution of the genus; Pseudomesoptilus gen. nov. is designated to include Mesoptilus sellardsi; Strephocladus permianus sp. nov. from the middle Permian of South of France; Westphaloptilus gallicus gen. et sp. nov. from the Bashkirian of the North of France.
The fusellar tissue of Palaeozoic rhabdopleurid pterobranchs has been studied using the SEM techniques. The fibrillar material of Ordovician Kystodendron ex gr. longicarpus and Rhabdopleurites primaevus exhibits a distinct dimorphism, comprising: (1) thinner, wavy and anastomosing/branching fusellar fibrils proper, producing a tight three-dimensional meshwork; and (2) long, more or less straight and unbranched cortical fibrils, sometimes beaded, and arranged in parallel. These fibrils are similar to the fusellar and cortical fibrils of graptolites, respectively. Until now, dimorphic fibrils and their arrangement within fusellar tissue were regarded as unique characters of the Graptolithina. In general, the fibrillar material of these fossils is partially preserved in the form of flaky material (new term) composed of flakes (new term). Flakes are interpreted as flattened structures originating from the fusion of several neighbouring tightly packed fibrils. A Permian rhabdopleurid, referred to as Diplohydra sp., reveals a fabric and pattern of fusellar tissue similar to that of both Ordovician rhabdopleurids but devoid (?) of cortical fibrils. The results presented here question views that: (1) substantial differences in fabric and pattern of fusellar tissue exist between fossil pterobranchs and graptolites; and (2) the ultrastructure of pterobranch periderm has remained unchanged at least since the Ordovician. The Palaeozoic rhabdopleurids investigated are closer ultrastructurally to graptolites than to contemporary pterobranchs. The pterobranchs and the graptolites should be treated as members of one class - the Graptolithoidea.
Anomalous development of shell ornamentation and repaired shell injury in the Early Devonian dacryoconarid tentaculites are documented and interpreted as either a repaired injury of the shell (evidence of unsuccessful predation obscured by recrystallization), or as a result of an anomalous function of the mantle, caused by injury of the soft body. The manner of shell repair, which resembles the way that some modern marine animals, such as mollusks, repair their shells, is discussed. The issue of phylogenetic affinities of tentaculites has been also outlined. These findings represent the first documentation of unsuccessful predation on the Middle Paleozoic plankton.
The teeth of a well known late Palaeozoic cladodont chondrichthyan, “Cladodus” occidentalis from Russia, USA, and England are restudied and a new generic name, Glikmanius gen. nov., is proposed for this species. Yet another tooth−based species, formerly described as ?Symmorium myachkovensis, occurring on the Russian Platform and in Nebraska, is considered to belong to the newly erected genus. Although there is no direct evidence that Glikmanius possessed fin spines, the broad similarity between its teeth and those of Ctenacanthus compressus suggests it had a ctenacanthiform affinity. The possible relationships between Glikmanius, Cladodus sensu stricto, “Ctenacanthus” costellatus, and Heslerodus, are suggested. However, the proposition put forward by an earlier author that the teeth of Heslerodus might represent the lower jaw dentition of G. myachkovensis, is rejected. The overall resemblance of Glikmanius teeth and those of Cladoselache and Squatinactis is recognised as convergent.
Listracanthus pectenatus sp. nov. represents the youngest record of the enigmatic chondrichthyan Listracanthus. This new species is the only Mesozoic record of this genus and highlights survival of a rare and enigmatic group of cartilaginous fishes across the Paleozoic–Mesozoic boundary. In the Vega−Phroso Siltstone Member of the Sulphur Mountain Formation (western Canada), two kinds of numerous dermal denticles identified as Listracanthus occur predominantly in strata probably of early Smithian age. The new species differs from all other known species of the genus in the structure of the anterior and posterior borders of the large denticles. The small denticles appear to be less diagnostic than the large ones and are readily distinguished from small denticles generally assigned to the genus Petrodus. Histology reveals that the largest denticles were originally hollow, probably secondarily ossified as acellular bone. The conclusion drawn by previous authors that Listracanthus may be a petalodontid shark, based on ambiguous non−skeletal associations with Deltoptychius, Petrodus, or Calopodusis not supported by this study. The large number of denticles, the size of both types of denticles and their arrangement suggest that Listracanthus pectenatus sp. nov. was a large chondrichthyan of aberrant body shape and yet uncertain systematic position.
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Stylophoran supertrees revisited

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Supertree analysis is a recent exploratory method that involves the simultaneous combination of two or more character−based source trees into a single consensus supertree. This method was recently applied by Ruta to a fossil group of enigmatic Palaeozoic forms, the stylophoran echinoderms. Ruta’s supertree suggested that mitrates are polyphyletic and originated from paraphyletic cornutes. Re−examination of Ruta’s data matrix strongly suggests that most source trees were based on dubious homologies resulting from theory−laden assumptions (calcichordate model) or superficial similarities (ankyroid scenario). A new supertree analysis was performed using a slightly corrected version of Ruta’s original combined matrix; the 70% majority−rule consensus of 24,168 most parsimonious supertrees suggests that mitrates are monophyletic and derived from paraphyletic cornutes. A second new supertree analysis was generated to test the influence of the pruning of three taxa in some calcichordate source trees; the 70% majority−rule consensus of 3,720 shortest supertrees indicates that both cornutes and mitrates are monophyletic and derived from a Ceratocystis−like ancestor. The two new supertree analyses demonstrate the dramatic influence of the relative contributions of each initial assumption of plate homologies (and underlying anatomical interpretations), in original source trees, on the final topology of supertrees.
Although bored invertebrates have been described from every period of the Paleozoic, little information on the frequency and nature of Late Paleozoic drill holes exists. Our examination of the Permian silicified fossils, which were bulk collected by G.A. Cooper from the Glass Mountains of west Texas, revealed numerous drilled brachiopods and bivalve mollusks. Drill holes are perpendicular to the shell, smooth sided, sometimes beveled, and have other characteristics consistent with a predatory/parasitic origin. The frequency of drilling is significantly lower (p ≤ 0.05) for brachiopods (1.07%, n = 7597) than for bivalves (7.43%, n = 619). This study confirms that drilling predators and/or parasites were present in the Late Paleozoic. However, the drilling frequencies reported here—rarely exceeding 5%—are much lower than those reported for the Late Mesozoic and Cenozoic, which typically exceed 20%. The low Late Paleozoic frequencies are consistent with a majority of estimates reported previously for the older periods of the Paleozoic and suggest that the intensity of drilling predation/parasitism in marine benthic ecosystems remained low throughout the Paleozoic and did not increase until some time in the Mesozoic. Our data suggest that prey/host types with a higher nutritional return (bivalve mollusks) may have been preferentially selected for attack by predator(s)/parasites(s) already in the Permian.
The revision of shark teeth from the Pennsylvanian black shales of central USA, ascribed to “Phoebodus heslerorum” Williams, 1985 and Symmorium reniforme Cope, 1893, shows that “Ph. heslerorum” is a junior synonym of “Cladodus” divergens Trautschold, 1879. This species belongs neither to Phoebodus nor to Cladodus, so a new genus Heslerodus is proposed. Very common, robust cladodont teeth with a deep labio−basal depression and two buttons, often referred to as S. reniforme, do not belong to the latter species, but to “Cladodus” occidentalis Leidy, 1859. The generic affinity of “C.” occidentalis is yet undetermined, but it is possible that it represents ctenacanthoids.
We present discoveries of internal bodies in problematic Silurian and Devonian organic−walled microfossils classified traditionally as polygonomorph, acanthomorph, sphaeromorph, and herkomorph acritarchs. These bodies are comparable with reproductive structures (autoand/or aplanospores) of modern unicellular green algae (Chlorococcales). Our findings suggest that many of these microfossils may represent asexually reproducing (sporulating) vegetative cells of chlorococcalean algae. The presence of spore−like bodies in the studied acritarchs supports earlier suggestions, based on ultrastructural and biomarker studies, that some acritarchs can be affined with green algae.
The affinities of the Early Permian synapsid Tetraceratops insignis have been reevaluated several times since the early 20th century, being considered as an eothyridid, a sphenacodontid, or a therapsid. This controversy continues into the 21st century, with recently raised doubts about the interpretation of Tetraceratops as the oldest known therapsid, a hypothesis supported by the only redescription of this fossil in the second half of the 20th century. Our study examines the arguments proposed to refute therapsid affinities, and concludes that Tetraceratops indeed is the sister−group of all other known therapsids. The most recently published phylogenetic data matrix that includes Tetraceratops fails to confirm its therapsid affinities. However, adding seven characters to that matrix leads to the conclusion that Tetraceratops is the basal−most and oldest therapsid. The recent suggestion of a Laurasian origin of therapsids appears poorly supported; too few data are available on the distribution of Permian synapsids to settle this question.
The fossil record of Paleozoic ophiuroids includes a number of forms which share striking similarities with modern relatives in terms of skeletal morphology. These so called modern-type Paleozoic ophiuroids yield an enormous potential for a better understanding of ophiuroid evolution, yet the scarcity of accurate and sufficiently detailed morphological descriptions available to date precludes any further-reaching assessments. Here, we describe an articulated ophiuroid specimen from the Late Tournaisian (early Carboniferous) of Czatkowice quarry, southern Poland, as a new species Aganaster jagiellonicus sp. nov. The good preservation of the specimen allowed for a morphological analysis at a level comparable to recent ophiuroid descriptions. It shows remarkable morphological similarities with extant former ophio-lepidids Ophiomusium and Ophiosphalma. The new find thus contributes to a solid basis for future investigations on the position of the modern-type Paleozoic ophiuroid in the phylogeny of the class.
The epidermal anatomy of Pseudoctenis samchokense is described revealing non−bennettitalean characters of these leaves from the Permo−Carboniferous of China (and Korea). The specimens were originally described as Pterophyllum samchokense suggesting a bennettitalean affinity. They can no longer be considered bennettitalean since their cuticles lack the distinctive brachyparacytic stomata of that clade. Pterophyllum was originally erected as a morphogenus for segmented leaves from the Mesozoic and has subsequently been clearly assigned to the Bennettitales. The segmented leaves from the Permo−Carboniferous of Shanxi and Hebei, China described herein require a new ordinal and generic assignment since the non−bennettitalean cuticular characters documented reinforce the uncertainties in attribution of any foliage older than Late Triassic to the Bennettitales. Based on leaf−architecture and epidermal anatomical characters, the specimens are at best assigned to the cycadalean genus Pseudoctenis. This genus was formerly known only from Mesozoic rocks. Consequently, the specimens are highly significant, as they are among the oldest known vegetative remains of cycads. Re−evaluation of the affinities of all specimens assigned to Pterophyllum from Palaeozoic rocks is thus essential. Based on a review of other Permian–Carboniferous fossil leaves assigned to Pterophyllum, we conclude that none yet reveals definitive bennettitalean characters.
A new interpretation of the wing venation of the fossil insect order Caloneurodea is proposed. Complex fusions of veins are demonstrated, supporting the assignment of the Caloneurodea to the Panorthoptera, i.e. the order is closely related to the Orthoptera. A new diagnosis of the order is proposed. The following taxa are re-described after new interpretation, material or preparation, or are newly described: Caloneura dawsoni Brongniart, 1885 (= Boltonaloneura subtilis (Bolton, 1925) syn. nov.), Gigagramma carpenteri gen. and sp. nov.; Sthenarocera pachytyloides Brongniart, 1885, Pruvostiella lecomtei (Pruvost, 1919), Apsidoneura flexa Carpenter, 1943, Homaloptila similis (Meunier, 1911), Pleisiogramma medialis Carpenter, 1943 (=Pleisiogramma reducta Carpenter, 1943 syn. nov.), Paleuthygramma acuta Carpenter, 1943, Paleuthygramma sharovi sp. nov., Euthygramma parallelum Martynov, 1928 (= Euthygramma curvatum Martynov, 1931 syn. nov.; = Paleuthygrammella aberrans (Martynov, 1938) syn. nov.), Anomalogramma parva Carpenter, 1943, Nanogramma gandi gen. and sp. nov., and Ligogramma sinuosa gen. and sp. nov. The family Gelasopteridae Carpenter, 1976 is revised and its ordinal relationships are discussed. After the proposed phylogenetic frame, mutation(s) occurring in homeotic genes are expected to be responsible of the unusual wing morphology of the Caloneurodea, mixing fore- and hind-wing 'panorthopteriď features.
The connecting ring in orthoceratids is composed of two calcified layers: an outer spherulitic−prismatic and an inner calcified−perforate. The spherulitic−prismatic layer is a direct continuation of that layer in the septal neck, whereas the calcified−perforate layer is a structurally modified continuation of the nacreous layer of the septal neck. The latter layer is traversed by numerous pores which are oriented either transversally to the siphuncular surface, or have a somewhat irregularly anastomosing course. The connecting ring structure is positively correlated to the dorsal position of the scars of the cephalic retractor muscles. A similar type of connecting ring and a dorsal postion of retractor muscle scars also occur in lituitids, previously assigned to tarphyceratids, and in baltoceratids, previously assigned to ellesmeroceratids. These two taxa are therefore included in the suborder Orthoceratina, which, together with the suborder Actinoceratina, are assigned to the order Orthoceratida Kuhn, 1940.
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