New Silurian cooksonias from dolostones of north-eastern North America

New specimens of Cooksonia and Hostinella are described from the Bertie Group of Ontario and New York State, which is dated by faunas as latest Silurian (Přídolí). The rare plant fossils are unusual in that they are preserved in fine-grained, slightly argillaceous dolostones ('waterlime') rather than clastic rocks. At least two species of Cooksonia are present, one with ± globular sporangial morphology close to C. hemisphaerica Lang. Those with ellipsoidal/discoidal sporangia are compared with C. pertoni Lang, C .  paranensis Gerrienne et al . and C. bohemica Schweitzer, the latter represented by a single specimen from the Přídolí of the Czech Republic. However, the paucity of specimens, which prevents assessment of taphonomic influences on shape, combined with the absence of any anatomical features and the gross morphological simplicity of the fossils, precludes specific assignment. Specimens of Hostinella include one in which apices and a lateral basal structure resembling a root are preserved. It is concluded that the Laurentian assemblage of Ontario and New York State is less diverse and disparate than coeval assemblages, which are also preserved in marine rocks. Its preservation in limestones may have been facilitated by the hypersalinity inferred from various sedimentary features, which would restrict the activity of many decomposers.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 399–413.     

A new species of the marattialean fern Scolecopteris (Zenker) Millay from the uppermost Permian of Guizhou Province, south-western China

Several isolated marattialean synangia and sporangia are reported from coal balls collected from Coal Seam No.1 (C605) in the uppermost Permian Wangjiazhai Formation in Guizhou Province, south-western China. The synangia are radially symmetrical with diameters between 0.8 and 1.2 mm and are 1.7 mm long, consisting of 3–4 elongate sporangia that are fused basally, free distally and possess a pointed apex. The outer-facing sporangial wall is 4–5 cells thick and conspicuously differentiated. Spores are trilete, have a granular ornamentation and are nearly round equatorially with a diameter of 55–60 µm. Comparisons with other anatomically preserved Palaeozoic marattialean synangia from the Euramerican and Cathaysian floras permit their assignment to the genus of Scolecopteris (Zenker) Millay. In this species the thick, outer-facing sporangial walls and large trilete spores are features consistent with those of the Oliveri Group within Scolecopteris , a group that has previously been considered primitive within this genus. Distinctions from all other previously recognized species within the Oliveri Group lead to the creation of a new species, S. guizhouensis sp. nov. This species is the youngest of the reported species of Scolecopteris recognized from the Euramerican and Cathaysian floras, and provides important evidence on the organization of marattialean ferns from the Upper Permian strata of south China.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 279–288.     

A diverse assemblage of early land plants from the Lower Devonian of the Welsh Borderland

FANNING, U., EDWARDS, D. & RICHARDSON, J. B., 1992. A diverse assemblage of early land plants from the Lower Devonian of the Welsh Borderland. Nine rhyniophytoid taxa are described from an early Gedinnian locality ( middle micromatus-newportensis spore Biozone) near Ludlow, England. They include Cooksonia pertoni, C. hemisphaerica, C. cambrensis, Tortilicaulis transwalliensis and three new taxa, Salopella marcensis sp. nov., Uskiella reticulata sp. nov. and Tarrantia salopensis gen. et sp. nov. Isolated sporangia of reniform shape, and those subtended by short lengths of axis, contain spores of Apiculiretusispora type and may belong to C. caledonica or Renalia . Morphologically distinctive forking, terminal sporangia lacking identifiable spores are not placed in a new taxon, because evidence based on in situ spores from elsewhere suggests they may belong to Salopella . Structures previously interpreted as clusters of sporangia of Yarravia- type are shown to be ± globular sporangia longitudinally split into valves. Sterile axes are dominated by smooth forms; although rare examples possess small enations. Tracheids have not been seen in axes of fertile specimens nor in sterile coalified compressions. A single pyrite permineralization contains tracheids of zosterophyll type. The assemblage demonstrates diversity among rhyniophytoids in the early Devonian and the existence of low vegetation 'alongside' the much larger zosterophyll dominated type.     

Hsüa deflexa sp. nov. from the Xujiachong Formation (Lower Devonian) of eastern Yunnan, China

A new vascular plant, Hsüa deflexa sp. nov., is documented from the Lower Devonian ((upper) Pragian-lower Emsian) Xujiachong Formation, near Xujiachong village, Qujing District, eastern Yunnan, South China. In three dimensions, the branching system comprises a robust creeping main axis and comparatively slender erect lateral axes inserted oppositely or alternately. The lateral axes depart at right angles from the main axis. Towards the apex, the main axis is perhaps erect with the lateral axes attached at smaller angles. The lateral axes dichotomize equally one to three times in different planes and decrease in diameter and length acropetally. They bear sparse and irregular tiny spines. Apices of a pair of distal lateral axes curve in opposite directions and each terminates in a rounded to reniform sporangium. The sporangium dehisces into two equal valves along an indistinct convex marginal thickening. The xylem is possibly centrarch. This plant fits Hsüa in branching pattern, sporangial characters and xylem maturation. Hsüa deflexa sp. nov. differs mainly from the type species H.  robusta in the presence of axial spines, perpendicular extension of lateral axes from the main axis and curving of distal lateral axes. Based on the possibly centrarch xylem and terminal sporangium, this plant relates to the rhyniophytes ( sensu Banks, 1968). In view of the equal sporangial valves with marginal thickenings it resembles the zosterophyllophytes ( sensu Banks, 1968). Hsüa is now treated as incertae sedis .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 255−271.     

Autochory in ferns,not all spores are blown with the wind

Dispersal is a key process in plant population dynamics. In ferns, two successive vectors are needed: the sporangium catapulting mechanism, and wind or gravity. However, some rock ferns have a growth habit that suggests a kind of autochory by placing spores on the rock surface. Moreover, some ferns show modifications of the sporangial dehiscence. To determine the role of growth habit in spore dispersal, we checked the sporangial opening mechanism and explored the spatial distribution of plants on the walls. The presence of spores of Asplenium celtibericum, a rupicolous fern, in the rock surface was checked. In addition, its sporangial dehiscence, plant size and position in the wall were analysed. Spores and indehiscent sporangia were present on walls at each sampling moment. Their highest number was found close to the plants. There was a positive correlation between crack width and plant size. However, most plants occupy the upper half of the cliffs. The growth habit of A. celtibericum is instrumental to deposit the spores over the neighbouring rock surface, thus enhancing the probability of spores to find suitable crevices for germination. Furthermore, dispersal of indehiscent sporangia might promote intergametophytic mating, and the modified sporangial opening mechanism extends the dispersive period.     

Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia

We report on and name two new taxa of basal crocodylomorph archosaurs from the Lower Jurassic, Litargosuchus leptorhynchus gen. et sp. nov. , from the upper Elliot Formation (Stormberg Group) of South Africa, and Kayentasuchus walkeri gen. et sp. nov. , from the Kayenta Formation (Glen Canyon Group) of Arizona, USA. Examination of this material led to a reconsideration of basal crocodylomorph interrelationships. A phylogenetic analysis found no support for the monophyly of Sphenosuchia.  © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 136 , 77–95.     

A new genus for isolated bivalved sporangia with thickened margins from the Lower Devonian of the Welsh Borderland

A new taxon Sporathylacium salopense gen. et sp. nov . is based on small isolated coalified sporangia from the Lower Devonian (Lochkovian: micrornatus—newportensis spore zone) from the Welsh Borderland. The sporangia have two equal valves with multi-layered walls and thickened borders, and contain trilete crassitate, non-curvaturate isospores that are completely covered by a microgranular ornament with possible distal verrucate/murornate structures. They differ from zosterophyll sporangia in details of the presumed dehiscence zone extending around the entire free convex margin, particularly in the presence of a wedge of amorphous material between the valves, and in spore characters. Absence of any information relating to water-conducting cells prevents further assignment within the embryophytes. Spherical bodies associated with spores and a resilient sporangial lining are compared with similar structures in extant free-sporing plants and with Ubisch bodies. The mode of sporangial dehiscence involving anatomical modifications of the valve margins and the novel wedge of tissue which connects them remains speculative.     

A new rhyniopsid with novel sporangium organization from the Lower Devonian of South Wales

SHUTE, C. H. & EDWARDS, D., 1989. A new rhyniopsid with novel sporangium organization from the Lower Devonian of South Wales. Re-investigation of permineralized plants originally called Cooksonia sp. from the Lower Old Red Sandstone (Siegenian) of Gwent, S. Wales shows them to be rhyniopsids with simple isotomous branching in smooth axes and ellipsoidal terminal sporangia that are longer than wide and that possess a complex wall organization. They are thus placed in a new genus. The sporangial wall is several cells thick, the outermost comprising a layer with pronounced thickening of the anticlinal and outer periclinal walls, which is interrupted by a zone of thinner-walled cells parallel to the longest dimension of the ellipsoidal organ and considered to be involved in its dehiscence into two equal halves. The alete isospores have a bilayered wall, the outer interpreted as an ornamented perispore. Similar granular ornament seen on sheets and globules in the vicinity of the spores and on the innermost surface of the sporangium wall possibly demonstrates the activity of a periplasmodial tapetum. The permineralized sporangia are considered conspecific with those in compression fossils with elliptical outlines and pronounced borders. Comparison of presumed dehiscence mechanisms in a number of Silurian and early Devonian fossils suggests that splitting into two equal valves along the longest dimension, so that a maximum area of spores was exposed to the atmosphere, arose independently in a number of unrelated plants.     

A review of the platanaceous woods from the Eocene paratropical rainforest of south-east England

Small diameter pyritized axes, commonly referred to as 'twigs', of fossil platanaceous wood are described from the Lower Eocene London Clay Formation of south-east England. These twigs are characterized by solitary vessels with scalariform perforation plates, opposite intervessel pits, and tall, multiseriate rays that dilate in the phloem region. The wood anatomy supports close relationship to members of extant Platanaceae and the material is placed in the organ genus Plataninium Unger erected for fossil woods with close anatomical similarity to Platanus L. This material supplements the fossil record of platanaceous type wood from the Eocene London Clay and documents the first record of Plataninium decipiens Brett in the twig flora.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 181–191.     

Crepidula beklemishevi gen. et sp. n. and Dimeiospora palustris gen. et sp. n. (Microspora: Amblyosporidae)--new microsporidian genera and species from blood-sucking mosquitoes (Diptera: Culicidae) from the south of the western Siberia

Microsporidia parasitizing the adipose body of mosquito larvae of Anopheles beklemishevi and Aedes punctor has been studied. Two new genera of microsporidia are described based on lightmicroscopic and ultrastructural characteristics of spores and sporogony stages. The spore wall of Crepidula beklemishevi gen. n. et sp. n. is formed by two-membrane exospore, thick exospore, bilayer endospore and thin plasmolemma. Spores with single nucleus, polar filament anisofilar, with 6-7 coils (2+ 4-5), polaroplast consisting of three parts: macrochelicoidal, microhelicoidal and lamellar. Fixed spores 4.2 +/- 0.22 x 2 +/- 0.01 microns. The sporogony of Dimeiospora palustris gen. et. n. results in spore formation of two different types. Spores of the first type are oviform, with thick wall, single-nuclear, 6.1 x 4.9 microns. Spore wall with three layers, about 370 nm. Exospore electron-dense, subexospore moderately electrondense. Exospore and subexospore irregularly pleated on the almost spore surface and slightly thinner on anterior end only. Endospore electron-translucent. Polar filament anisofilar, with 9 coils (3 + 6). Polaroplas consists of three parts: lamellar, fine bubbled, and coarse bubbled. Spores of the second type broad-ovate, with apical pole narrower, distal pole concave, 4.6 x 3.7 microns. Spore wall with three layer, 355 nm. Exospore on the apical end irregularly pleated, consists of thin electrondense exospore, subexospore of variable electron density, endospore electron-translucent. Polar filament anisofilar, with 13 coils (3 + 10). Polaroplast has two parts: lamellar and vesicular.     

A giant rhinocerotoid (Mammalia, Perissodactyla) from the Late Oligocene of north-central Anatolia (Turkey)

A giant rhinocerotoid is described for the first time south of the Black Sea, in Turkey. The single specimen, a fragmentary radius referred to Paraceratherium sp., originates from conglomerates nearby at Gözükizilli, in the Çankiri–Çorum Tertiary basin. These layers correspond to the Lower member of the Kizilirmak Formation. The same locality (Gözükizilli-2) yields also the small rhinocerotid Protaceratherium sp., cf. P. albigense (Roman, 1912). Three other mammal localities (Gözükizilli-1, in the Lower Member of the formation, with several rodent species; Tepe 641 and Kizilirmak, in the Upper Member, with a diversified micro- and macro-mammal fauna) allow us to refer the Kizilirmak Formation as a whole to the Late Oligocene. All the observed taxa have strong Asian and/or European affinities, which precludes any geographical insulation for this part of Anatolia during the Late Oligocene.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152 , 581–592.     

First record of fossil Mesozoic Ctenopoda (Crustacea, Cladocera)

Pre-Pleistocene representatives of the crustacean order Ctenopoda Sars, 1865 are unknown. Here we describe Mesozoic fossil specimens of ctenopods from two localities in Mongolia: Khotont (Jurassic/Cretaceous boundary, about 145 Mya) and Khutel Khara (Lower Cretaceous, about 129 Mya). All specimens apparently belonged to the tribe Latonini Korovchinsky, 1986 of the subfamily Sidinae Baird, 1850. At the Khotont site, ctenopods were the most numerous microscopic animal fossils. We assigned these cladocerans to a new genus and species Archelatona zherikhini gen. nov., sp. nov. Our findings support a pre-Cretaceous origin for cladocerans.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 146 , 269–274.     

A new late Westphalian fossil marattialean fern from Nova Scotia

Sydneia manleyi gen. et sp. nov. is based on part of a fertile frond from the upper Westphalian D of the Sydney Coalfield, Nova Scotia, Canada. It has small synangia composed of laterally fused sporangia that are elongate and with a circular cross-section. The sporangia yielded variably sized monolete and trilete spores with laevigate and microspinate ornamentation; intermediate forms were also observed. The spores can be correlated with the sporae dispersae species Latosporites minutus , Punctatosporites oculus and Laevigatosporites minimus . Size distribution of the spores is variable and highly skewed, suggesting heterogeneity of the spores within the sporangium. Spore ultrastructure indicates that the fossil is part of a fern, and the morphology of the spores and synangia indicate marattialean affinities.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 199–212.     

Fossil palm flowers in Dominican and Baltic amber

Five palm flowers in Dominican amber and one in Baltic amber are described or characterized. Palaeoraphe dominicana gen. et sp. nov. in the subtribe Livistoninae, is described from one perfect flower in Dominican amber. Roystonea palaea sp. nov. is described from one staminate and one pistillate flower in Dominican amber. Three other palm flowers, two perfect flowers from Dominican amber and one staminate flower from Baltic amber, are briefly characterized and figured.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 361–367.     

The histology and affinities of sinacanthid fishes: primitive gnathostomes from the Silurian of China

On the basis of well preserved specimens from the Lower Silurian of the Tarim Basin, Xinjiang Uygur Autonomous Region and Shiqian County, Guizhou Province, People's Republic of China we describe in detail the histological structure of sinacanthid spines, the only known remains of a group of fishes common in Silurian strata from China. The sinacanthids have previously been assigned either to the acanthodians or to the chondrichthyans. The spine structure is composed of an outer layer of atubular dentine and an inner layer of globular calcified cartilage, and the nature and distribution of these tissues indicates that the spines were formed as a result of interaction between the endoskeleton and dermoskeleton. The tissue distribution and style of growth described herein places the sinacanthids crownwards of the placoderms, and possibly within the total group Chondrichthyes. However, before they can be firmly placed within a phylogenetic scheme, further evidence is required both on the general anatomy of sinacanthids and on the nature of chondrichthyan apomorphies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 379–386.     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age ( micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.     

On the osteology and phylogenetic affinities of the Pseudasturidae – Lower Eocene stem-group representatives of parrots (Aves, Psittaciformes)

The osteology of the early Eocene (about 50 mya) avian taxon Pseudasturidae Mayr, 1998 is revised and its phylogenetic affinities are analysed. Members of the Pseudasturidae are known from abundant and excellently preserved skeletal material, both complete skeletons on slabs as well as isolated, three-dimensional bones. Although this taxon is thus among the best represented of all small early Tertiary birds, its systematic affinities were unknown so far. Derived osteological characters which are visible in newly recognized specimens from the Lower Eocene London Clay of England most convincingly support classification of the Pseudasturidae into the Psittaciformes (parrots). Both, in overall morphology and in terms of derived characters, the tarsometatarsus of the Pseudasturidae closely resembles that of the Eocene Quercypsittidae, which were assigned to the Psittaciformes by Mourer-Chauviré (1992 ). The Pseudasturidae are considered to be stem-group representatives of the Psittaciformes and the sister taxon of all other known psittaciform birds. The Eocene taxon lacks the specialized bill morphology of crown-group Psittaciformes of the Psittacidae. Several other osteological differences between the Pseudasturidae and the Psittacidae probably are also functionally correlated with the specialized feeding technique of the latter.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 715–729.     

Gynoecium, fruit and seed structure of Paullinieae (Sapindaceae)

Despite an emphasis on fruit characters in Paullinieae taxonomy, few detailed morphological and anatomical studies of the gynoecia, fruits and seeds exist. The aims of the present study were (1) to provide a detailed documentation of gynoecium, fruit and seed structure and ontogeny in selected Paullinieae taxa; (2) to determine whether the gynoecium, seed and seedling provide additional characters of systematic value within the tribe; and (3) to relate the structural findings to mechanisms of fruit dehiscence and dispersal within these taxa. Newly described characters of systematic value within Paullinieae are shape and surface of the obturator, type of pollen tube transmitting tract, indumentum of the inner and outer surface of the carpels, ovary wall anatomy, aril anatomy, pseudo-hilum form, seedling germination mode and structure of first leaves. The fruits of Paullinia are septifragal, and conspicuous colour contrasts between the pericarp, aril and seed in most species of this genus are suggestive of a bird dispersal syndrome. Interestingly, it appears that relatively minor structural changes are associated with switches to rodent dispersal in Paullinia sphaerocarpa and water dispersal in P. clathrata and P. hystrix. Anemochorous fruits are septifragal ( Cardiospermum and Urvillea ) or schizocarpic ( Houssayanthus , Lophostigma , Serjania ). They are structurally similar and Cardiospermum with septifragal capsules may also show septicidal dehiscence.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 159–189.     

Patagial complex evolution in hummingbirds and swifts (Apodiformes): a molecular phylogenetic perspective

Studies of the role of flight in vertebrate evolution often have focused on the propatagial muscle complex because this structure forms the wing's leading edge. However, historical narratives about the evolution of flight anatomy are compromised because traditional higher-level taxonomies typically are based in part on the propatagium itself. To avoid this circularity, I used a consensus molecular phylogeny to examine propatagial evolution in the highly aerial sister groups, hummingbirds and swifts (Apodiformes). Mapping of anatomy on molecular-based phylogeny indicates that structural variation in M .  tensor propatagialis pars brevis (TPB) is congruent with the major subclades of both hummingbirds and swifts. However, the humeral tendon and broad attachment of the fleshy belly of TPB with M .  extensor metacarpi radialis (EMR) most likely underwent parallel change in hummingbirds and swifts, while the distal tendon present only in hummingbirds changed from a thin sheet to a strong tendon. The combination of divergent (within hummingbirds or swifts) and parallel (between hummingbirds and swifts) evolutionary patterns implies that the taxonomic value of the propatagial complex in apodiformes depends on anatomical component and level of divergence. The congruence of anatomy with molecular phylogeny provides independent criteria for designating relatively ancestral versus derived clades of apodiformes. Based on these polarities, living hummingbirds and swifts express additional parallel trends from arboreal to more aerial foraging styles, and from depauperate to species-rich clades. Within apodiformes, the link of flight anatomy with taxonomic and ecologic diversity suggests that elaboration of locomotor modes was important for apodiform diversification, echoing a similar pattern for birds relative to their reptilian ancestors.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 211–219.     

STUDIES OF PALEOZOIC MARATTIALEANS: AN EARLY PENNSYLVANIAN SPECIES OF THE FERTILE FERN SCOLECOPTERIS

Remains of the fossil Marattiales are very rare in Lower Pennsylvanian sediments. The present report describes a new species of the fertile fern foliage Scolecopteris from the Lewis Creek, Kentucky locality (Lower or lower Middle Pennsylvanian). Scolecopteris conicaulis n. sp. has radial synangia composed of a ring of 4–7 elongate, exannulate sporangia. Most features of the synangia of S. conicaulis were previously hypothesized to be primitive in Scolecopteris based on geologically younger species. Supposed primitive characters include the large synangium pedicel with fiber core, an outer-facing sporangial wall lacking differentiation or zonation, and large spores. The anatomy of the sporangium walls, pinnule morphology, and general spore type support an association with the Minor group of Scolecopteris. The new species is similar in several important features to Scolecopteris (Cyathotrachus) altus, the only other anatomically preserved fertile marattialean known from this early time, and indicates a considerably earlier origin for fertile foliage of this type.