New perspectives on the Mesozoic seed fern order Corystospermales based on attached organs from the Triassic of Antarctica

A new Triassic corystosperm is described from the Shackleton Glacier region of Antarctica. The compression fossils include cupulate organs (Umkomasia uniramia) and leaves (Dicroidium odontopteroides) attached to short shoot-bearing branches. The cupulate organs occur in groups near the apices of the short shoots, and each consists of a single axis with a pair of bracts and a subapical whorl of five to eight ovoid cupules. This unique architecture indicates that the cupules are individual megasporophylls rather than leaflets of a compound megasporophyll. A branch bearing an attached D. odontopteroides leaf provides the first unequivocal evidence that Umkomasia cupulate organs and Dicroidium leaves were produced by the same plants. Although this had previously been assumed based on organ associations, the new specimens are important in demonstrating that a single species of corystosperm produced the unique cupulate organs described here and the geographically and stratigraphically widespread and common D. odontopteroides leaf. Therefore, biostratigraphic, paleoecological, and phylogenetic studies that treat Dicroidium leaf morphospecies as proxies for biological species of entire plants should be reconsidered. Phylogenetic analysis suggests that the corystosperm cupule is an unlikely homologue for the angiosperm carpel or outer integument.     

Arachnids from the Carboniferous of Russia and Ukraine,and the Permian of Kazakhstan

New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carboniferous of Russia and Ukraine and one complete, albeit poorly preserved, specimen from the Permian of Kazakhstan, are described. The spider genus Arthrolycosa is reported from the Late Carboniferous (Late Pennsylvanian: Kasimovian–Gzhelian) of Chunya in the Tunguska Basin of Siberia; it is the first find of a spider outside the Carboniferous tropics. Another fossil assigned to the same genus comes from the Late Carboniferous (Early Pennsylvanian: Bashkirian) of Kamensk–Shakhtinsky in the Donets Basin of Russia; it is probably the oldest fossil spider known. A thelyphonid (whip scorpion) carapace is described from the Late Carboniferous (Late Pennsylvanian: Kasimovian) of the adjacent Lugansk Province of the Donets Basin of Ukraine.     

<Emphasis Type="Italic">Dicroidium bandelii</Emphasis> sp. nov. (corystospermalean foliage) from the Permian of Jordan

Dicroidium bandelii sp. nov. (Corystospermales) is described from the Um Irna Formation (Permian) of the Dead Sea Region, Jordan. The type and only specimen is a compressed frond fragment with cuticle. The new species is unique in showing pinnate frond architecture with elongate, lanceolate pinnae with a dense, near-parallel venation. In situ epifluorescence-microscopic analysis of the cuticle revealed evenly distributed, superficial stomata with differentiated lateral subsidiary cells typical of the genus. Epidermal and cuticular features readily distinguish D. bandelii from some superficially similar species of Ptilozamites or Thinnfeldia. Our report demonstrates that the diversity of Corystospermales in the unusual ‘mixed flora’ of the Permian palaeotropics is still greater than previously thought, and that the Um Irna Formation in particular holds a great potential for future palaeontological research.     

Trend of changes in species diversity of Permian radiolarians of the genus <Emphasis Type="Italic">Entactinia</Emphasis> Foreman

The trend of species diversity and spatiotemporal distribution of spherical radiolarians of the genus Entactinia Foreman, 1963 in the Permian Period is analyzed. The center of origin of Permian entactinians in the Cis-Ural Paleobasin is established in the area of the modern interfluve of the Ural River and latitudinal current of the Belaya River on the Southern Urals. It is shown that the species composition of this genus almost completely changed in certain Permian epochs. Two forefather species of each new pleiad of Early, Middle, and Late Permian are revealed: Entactinia parapycnoclada and E. tyrrelli. Two new species of the Lower Permian radiolarians from the Southern Urals and Northern Mugodzhary, E. chernykhi sp. nov. and E. subquadrata sp. nov., are described.     

A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa

The basal clade Burnetiamorpha is known from only two specimens representing two genera, Proburnetia from the Severodvinskian horizon of the Vyatka River Basin in the Kotelnich district of Russia, and Burnetia from the Dicynodon Assemblage Zone of the Beaufort Group of South Africa. Both genera are of Late Tatarian (Late Permian) age. This paper describes the cranial morphology of a new genus of burnetiamorph, Bullacephalus , from the Late Permian Tapinocephalus Assemblage Zone of the Beaufort Group of South Africa. It is known from a relatively complete skull and lower jaw and is the best preserved burnetiamorph yet discovered. Apart from being the oldest member of the clade, Bullacephalus is also morphologically the least derived and provides new evidence on the phylogeny of this poorly understood group of basal therapsids.     

Dicynodontids of Eastern Europe

Dicynodontids are the most widespread and diverse group of higher anomodonts in the Late Permian of Eastern Europe. Dicynodon trautscholdi Amalitzky, 1922 is the first dicynodont described in Russia (Amalitzky, 1922). The taxonomic position of this and several closely related forms described by V. P. Amalitzky and P. P. Sushkin (Sushkin, 1926) remains an open question. A reexamination of cranial morphological features in East European members of the family Dicynodontidae allows the assignment of Dicynodon trautscholdi to a new genus, Fortunodon gen. nov., and corroborates the isolated position of Dicynodon amalitzkii established by Sushkin (1926), which is also assigned to the genus Fortunodon gen. nov. Beginning from the Sokolki Assemblage, Late Permian dicynodontids adapted to various ecological niches and had a noticeable effect on the general appearance of the tetrapod fauna.     

河南固始下石炭统杨山组华夏木一新种

描述河南固始杨山煤矿下石炭统杨山组的一个华夏木新种Cathaysiodendronyangshanensesp.nov.,这是近年来继在甘肃靖远早石炭世臭牛沟组中发现华夏木之后该属在早石炭世地层中的又一发现。华夏木属是华夏植物群的特有分子,它在早石炭世的发现对探讨华夏植物群的起源有重要意义。     

Family Linoproductidae Stehli, (Brachiopoda) from the Upper Moscovian of the Middle Carboniferous of the Moscow Region

Species from the family Linoproductidae sensu str. from the Podolskian-Myachkovian deposits of the Moscow Region are revised and redescribed. The peculiarities of the arrangement of spines on the ventral valve are used as a basis for a new taxonomic structure. A new genus Linipalus, which appeared in the Podolskian time and was widespread in the Late Carboniferous and Early Permian, is established. Three new species of the genus Linispinus Lazarev, 2006, and one new species of the genus Linipalus gen. nov. are described.     

The first tunicate with a calcareous exoskeleton (Upper Triassic,northern Italy)

The first tunicates with a calcareous exoskeleton are reported from Late Triassic buildup‐slope deposits of the Dolomites. Although examples of this group have been known since the early 1900s from the middle–upper Permian of eastern Asia and Sicily as Khmeria, they were erroneously attributed to rugose corals. These early representatives are small, double‐valved, conical skeletons, which evolved into multi‐plated capsules with up to 35 opercula. The latter are joined along zigzag margins, which in life could probably be opened for the atrial and branchial siphons. The construction and shape of these skeletons distinguish them from plants or other invertebrate phyla, while they share several similarities with living tunicates, specifically to sessile ascidians. Apart from a soft‐bodied genus from the lower Cambrian of China, ascidians are known only from isolated spicules, which occur sporadically from the Lower Jurassic onwards. The calcareous skeleton of these Late Triassic tunicates consists of aragonitic fibres, which form spherulitic or clinogonal microstructures. It seems that the stellate aragonitic spicules of Jurassic to Recent ascidians are a vestige of Permian–Triassic ancestors, which after the Carnian lost the ability to construct compound solid skeletons but partly still retain a soft double‐valved or multi‐operculate cellulose‐like tunic. The following taxa are described as new: Order Khmeriamorpha with the genera Khmeria Mansuy and Zardinisoma gen. nov., and the following species: Khmeria stolonifera (late Permian), Khmeria minima (Late Triassic), Zardinisoma japonicum (late Permian), Z. cassianum, Z. pyriforme, Z. polyplacophorum and Z. pauciplacophorum (all Late Triassic).     

新建迭部蛤(新名) (Tewomorpha)替代晚出名)(Tewoella Ding and Li, 1987 (非)(Tewoella Sun, 1979)

1993年, 于菁珊和董国义为辽宁上三叠统的非海相双壳类化石建立了一个新属辽宁蚌Liaoningia Yu and Dong, 1993, 以Liaoningia opima Yu and Dong, 1993作为模式种。然而, 早在1979年, 邢裕盛和刘桂芝就已为辽宁晚前寒武纪南关岭组所产的化石标本创建了新属名辽宁水母属Liaoningia Xing and Liu, 1979, 模式种是Liaoningia fuxianensis Xing and Liu, 1979。然而, 多数学者反对将它视为水母类化石, 有人怀疑辽宁水母属是假化石, 迄今学术界并无统一意见, 故本文暂将它归为疑问化石(Problematica)。根据“国际动物命名法规”, 我们提出一个新的属名Liaoningoconcha nom. nov., 用以替代Liaoningia Xing and Liu, 1979的次同义名Liaoningia Yu and Dong, 1993, 仍使用于菁珊和董国义在1993年指定的模式种, 中文译名不变。     

A Late Permian flora with Dicroidium from the Dead Sea region, Jordan

Three new species of Dicroidium, D. irnensis, D. jordanensis and D. robustum, are described from the Um Irna Formation (Upper Permian) of the Dead Sea region, Jordan. The plant remains are preserved as compressions with excellent cuticles. These are the earliest unequivocal records of Dicroidium, a genus that is typical for the Triassic of Gondwana. It is also the northernmost occurrence of this genus that apparently originated in the Permian in the palaeotropics. Middle and Late Permian floras from the Arabian Peninsula and adjacent regions show a remarkable mixture of elements from different floral provinces, i.e. Euramerica, Cathaysia and Gondwana. The climatic amelioration in the Early Triassic apparently enabled Dicroidium to migrate southward and eventually colonise the entire Gondwana region. Dicroidium is one of the very few megaplant genera not affected by the end-Permian biotic crisis, the largest Phanerozoic extinction event.     

First Jurassic occurence of Enoploclytia M’Coy, 1849 (Crustacea: Decapoda: Erymidae)

Among the Erymidae Van Straelen, 1925, known as early as the Late Permian (Changhsingian) and widespread in the Jurassic, the genus Enoploclytia M’Coy, 1849 seems to have a late appearance because of its lack before the Cretaceous. Until now, the oldest representative was E. augustobonae Devillez, Charbonnier, Hy?ný and Leroy, 2016 from the Early Cretaceous (Barremian) of the eastern Paris basin (France). However, a new fossil collected in Normandy (France), showing the typical carapace groove pattern of Enoploclytia, attests the presence of the genus in the Late Jurassic (Oxfordian).     

Linguaserra spandeli sp. nov. (Echinodermata: Ophiocistioidea) from the Late Permian (Zechstein) of Thuringia, Germany

From the Late Permian Zechstein reefs (Lopingian: Wuchiapingian) of eastern Thuringia, Germany, the new ophiocistioid goniodont Linguaserra spandeli sp. nov., is described, figured, and discussed, within the Linguaserridae Reich and Haude, 2004. The (para)genus Linguaserra is redefined; the stratigraphical and regional distribution as well as the phylogenetic position are briefly discussed. Linguaserra spandeli sp. nov. is the stratigraphically youngest record of this echinoderm group (Echinozoa: Ophiocistioidea) worldwide.     

Evolution and systematics of the Late Paleozoic brachiopod family Linoproductidae with descriptions of species from the Lower Permian of the Northern Timan

The analysis of the morphological features of the genera of the family Linoproductidae in the Late Paleozoic substantiates its three subfamilies as three evolutionary trends beginning with the initial subfamily Coopericinae Lazarev, 2004, which is known from the beginning of the Early Carboniferous and two its derivatives: subfamily Linipalinae subfam. nov., which appeared in the Podolskian Time (Upper Moscovian Age), and subfamily Linoproductinae Stehli, 1954, which appeared in the Kasimovian Age. The problems and prospects of the further detailing of the system of these subfamilies are discussed. Three new species of the genus Sublinoproductus are described from the Lower Permian of Northern Timan.     

The status of Jambadostrobus Chandra and Surange (Glossopteridales)

Jambadostrobus was established to accommodate fossils consisting of several multiovulate elliptical–ovate fructifications attached to the midline of glossopterid leaves. Two species have been assigned to the genus: J. pretiosus (the type species) and J. hillii from the Late Permian of India and Early Permian of Australia, respectively. Re-investigation of the original specimens reveals that only solitary fructifications are attached to the subtending leaf in J. pretiosus, hence the species is transferred to Plumsteadia, which was founded on equivalent characters. The putative reproductive organs constituting the single Australian specimen of J. hillii are re-interpreted to constitute a broad area of preservational damage on the leaf, hence Jambadostrobus is considered to be a nomen illegitimum.     

记原始恐头兽类一新属种──甘肃玉门晚二叠世脊椎动物群系列报道之三

记述了恐头兽类一新属种利齿狭头兽（Stenocybusacidentatusgenetspnov）。它以高而窄的头骨,轻巧的结构,大的眼孔,小的颞孔,三角形的上颌骨有别于所有已知的恐头兽类。它的门齿结构,及出现在头骨顶面容纳下颌收肌的凹陷显示了与安泰龙科（Anteosauridae）之间的密切关系。但其狭窄的头骨,大而侧扁的犬齿,及较为长大、侧扁、具边缘锯齿的犬齿后齿表明它代表一个新科,而且是恐头兽类中最为原始的科。     

山东南部太原组一种新的具解剖构造的鳞木目茎--Diaphorodendron rhombicum sp.nov.

报道了产于山东枣庄矿区太原组16号煤层煤核(早二叠世早期或晚石炭世晚期)中一种具解剖构造的鳞木目茎.茎呈压扁状,长轴约6 cm,短轴约2.5 cm.中柱、外部皮层、周皮和叶座均保存.茎具原生中柱,无次生木质部发育,外部皮层由较发育的径向的厚壁的细胞条带和已经毁坏的薄壁的细胞区域交替排列构成.周皮较发育,厚约4～5 mm,由木栓层和栓内层构成.其中栓内层较厚,细胞构造均一,或在有些地方分化为弦向的薄壁的细胞条带和l厚壁的细胞条带,二者交替排列.其中薄壁的细胞条带常保存不好而形成弦向的裂隙.叶座的高和宽近相等,呈近正菱形.叶痕盔形或双凸镜形,位于叶座上部,微微突出于叶座表面之上.叶座表面平坦,未见明显的中脊和横皱纹.叶迹在叶座内近水平状延伸,但在近叶座表面处略向上然后向外弯曲成不太明显的"S"形进入叶痕内.径切面上叶舌穴略向上斜伸,具一宽大的基部,其开口直接位于叶痕顶角处.叶舌大,宽卵球形,长轴达1.2 mm,短轴达1 mm.与目前已知的鳞木目几个属的茎比较,当前标本与Diaphorodendron属的茎最为接近,仅在叶座上有些不同,本文将其归入该属内.Diaphorodendron属目前共有3个种,全分布于欧美植物区晚石炭世中期(威斯法期),它们与当前标本区别明显,因此,将当前标本归入新种D.rhombicum内.本文是国内首次报道Diaphorodendron属在华夏植物区的存在.其高和宽近相等的叶座形态可能表明了一种进化的特征,这从其所在地层的地质时代(早二叠世早期或晚石炭世晚期)较欧美植物区的3个种(晚石炭世中期)较新这一事实得到支持.     

Phylogenetic analysis of Russian Permian dicynodonts (Therapsida: Anomodontia): implications for Permian biostratigraphy and Pangaean biogeography

Dicynodont therapsids have been known from the Upper Permian of Eastern Europe since the beginning of the twentieth century, but the phylogenetic relationships of these taxa have not been examined cladistically. Here we present the results of a phylogenetic analysis that includes eight Permian dicynodonts from Russia as well as 18 taxa best known from southern Africa. Our results do not conflict with much of the established picture of Permian dicynodont phylogeny, but are consistent with several novel hypotheses. Most importantly, our analysis suggests that the genus Dicynodon is paraphyletic, and we question its use in correlating widely separated basins. However, we cannot strongly reject a monophyletic Dicynodon . Our results also indicate that the closest Permian relatives of Kannemeyeria lived in Russia, suggesting a Laurasian origin for the lineage that includes this important Triassic taxon. The phylogeny presented here also suggests a Laurasian origin for several other dicynodont clades, but a Gondwanan origin is equally likely given the data at hand. Regardless of where these groups originated, there appears to be some endemism among Late Permian dicynodont faunas. Although our understanding of dicynodont phylogeny is improving, this study emphasizes the disparity in sampling of the dicynodont record between Gondwana and Laurasia and the need for a large scale phylogenetic analysis of Permian and Triassic dicynodonts.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 157−212.     

Tetrapod Footprint Biostratigraphy and Biochronology

Tetrapod footprints have a fossil record in rocks of Devonian-Neogene age. Three principal factors limit their use in biostratigraphy and biochronology (palichnostratigraphy): invalid ichnotaxa based on extramorphological variants, slow apparent evolutionary turnover rates and facies restrictions. The ichnotaxonomy of tetrapod footprints has generally been oversplit, largely due to a failure to appreciate extramorphological variation. Thus, many tetrapod footprint ichnogenera and most ichnospecies are useless phantom taxa that confound biostratigraphic correlation and biochronological subdivision. Tracks rarely allow identification of a genus or species known from the body fossil record. Indeed, almost all tetrapod footprint ichnogenera are equivalent to a family or a higher taxon (order, superorder, etc.) based on body fossils. This means that ichnogenera necessarily have much longer temporal ranges and therefore slower apparent evolutionary turnover rates than do body fossil genera. Because of this, footprints cannot provide as refined a subdivision of geological time as do body fossils. The tetrapod footprint record is much more facies controlled than the tetrapod body fossil record. The relatively narrow facies window for track preservation, and the fact that tracks are almost never transported, redeposited or reworked, limits the facies that can be correlated with any track-based biostratigraphy.

A Devonian-Neogene global biochronology based on tetrapod footprints generally resolves geologic time about 20 to 50 percent as well as does the tetrapod body fossil record. The following globally recognizable time intervals can be based on the track record: (1) Late Devonian; (2) Mississippian; (3) Early-Middle Pennsylvanian; (4) Late Pennsylvanian; (5) Early Permian; (6) Late Permian; (7) Early-Middle Triassic; (8) late Middle Triassic; (9) Late Triassic; (10) Early Jurassic; (11) Middle-Late Jurassic; (12) Early Cretaceous; (13) Late Cretaceous; (14) Paleogene; (15) Neogene. Tetrapod footprints are most valuable in establishing biostratigraphic datum points, and this is their primary value to understanding the stratigraphic (temporal) dimension of tetrapod evolution.