Comparative wood anatomy of Andromedeae s.s., Gaultherieae, Lyonieae and Oxydendreae (Vaccinioideae, Ericaceae s.l.)

The wood anatomical structure of 11 out of 13 genera from four tribes of the Vaccinioideae, namely Andromedeae s.s. , Gaultherieae, Lyonieae and Oxydendreae (Ericaceae s.l. ), is described using light and scanning electron microscopy. Several features of the secondary xylem support the tribal classification based on molecular data: arrangement of vessel-ray pitting, height of multiseriate rays and the shape of the body ray cells. Oxydendreae are clearly defined from the other representatives by various wood anatomical features. Gaultherieae can be distinguished from Lyonieae by differences in vessel perforation plates, vessel-ray pitting, height and structure of multiseriate rays, and occurrence of prismatic crystals, but the wood of Andromedeae s.s. is similar to Gaultherieae. Moreover, Andromedeae s.s. , Oxydendreae and Vaccinieae are characterized by their pith structure, whereas considerable variation in the pith cells is found in Lyonieae and Gaultherieae.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 161–179.     

Oldest shrimp and associated phyllocarid from the Lower Devonian of northern Russia

A mass occurrence of schooling, weakly sclerotized eocaridid shrimps in the Early Devonian pelagic environment of northern Russia suggests that the body-plan of the Eumalacostraca originated as an adaptation to swimming in the open sea. They probably replaced their ecological analogues, the archaeostracan phyllocarids Caryocaris , as a consequence of the rebuilding of the global marine environment during the later Ordovician. This may explain the rarity of the eumalacostracans in the Devonian and the difficulty with documenting their links with the archaeostracans. The new eocaridid shows similarities to the Carboniferous Anthracophausia in the general appearance and weak sclerotization of the carapace and appendages, but lacks its derived characters. The morphology of the pleural lobes, which have minute spines, is the only identified distinguishing character of the proposed new genus and species Archangeliphausia spinosa gen. et sp. nov. Its carapace was probably firmly connected dorsally with the thoracic segments. The only associated fossils of the assemblage represent a bizarre archaeostracan Pechoracaris aculicauda gen. et sp. nov., with a weakly sclerotized carapace lacking any hinge or rostral plates but with reduced furca, an extremely long caudal spine and pleopods transformed into spines.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 83–90.     

Anatomical features, monomer lignin composition and accumulation of phenolics in 1-year-old branches of the Mediterranean Cistus ladanifer L.

Mediterranean shrub species are described as having phenology, habitus , reproductive biology and anatomical alterations in certain tissues, allowing their survival during the dry season and protecting them from herbivory. Anatomical and chemical analyses were conducted in 1-year-old branches of Cistus ladanifer L. in order to investigate the role played by shoot structure in the adaptive strategies of this species in the Mediterranean environment. Results showed that both xylem and pith underwent lignification. Pith parenchyma cells had thickened walls, higher lignin content than xylem and different monomer composition. Xylem presented features aiding safe water transport. A large accumulation of phenolic substances was found in xylem, pith and cortical parenchyma. Observations reported in this paper suggest the occurrence of adaptive strategies in 1-year-old branches of C. ladanifer whose structural features: (1) allow mechanical reinforcement of tissues to withstand drought without suffering permanent damage; (2) favour safety rather than efficiency in water transport; (3) defend the plants from animal predation and pathogens by accumulating phenolics in various tissues, and (4) protect inner tissues against UV-B radiation through deposition of phenolic compounds in cortical layers.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 361–371.     

Rotafolia songziensis gen. et comb. nov., a sphenopsid from the Late Devonian of Hubei, China

A sphenopsid from the Upper Devonian (Famennian) Xiejingsi Formation, south-western Hubei Province, China, previously named as various species in Sphenophyllum , Hamatophyton , Bowmanites and Sphenophyllostachys , is now reinvestigated and assigned to a new taxon, Rotafolia songziensis gen. et comb. nov. Its ribbed axes are anisotomous and possess slightly expanded nodes. Lateral axes are inserted at nodes on main axes. Whorls of much divided vegetative leaves are attached at nearly right angles to nodes of basal axes, and at acute angles to nodes of terminal axes. There are six leaves per whorl. The terminal strobilus includes a central axis and verticils of fertile units. Each fertile unit consists of a bract and numerous sporangia. The margin of the elongate-cuneate bract bears a distal and many lateral elongate segments. Clusters of elongate sporangia are abaxially attached to the base of the bract at the same level. The axis has an actinostele, composed of a three-ribbed, exarch primary xylem and radial secondary xylem. Although Rotafolia songziensis closely resembles Hamatophyton verticillatum in axis character, leaf morphology and primary xylem type, they are quite different in strobilar structure. Taxonomically, Rotafolia is placed in the order Sphenophyllales by three well-defined characters: 1) whorled appendages; 2) ribbed protosteles; 3) exarch primary xylem maturation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 21–37.     

A new species of Lepidostrobus from the Upper Devonian of Xinjiang, China and its bearing on the phylogenetic significance of the order Isoëtales

Lepidostrobus xinjiangensis sp. nov. is described from Upper Devonian rocks of the eastern Junggar Basin, Xinjiang, north-west China. It provides new insights into the reproductive diversification and phylogenetic relationships of lycopsids in the Late Devonian. The strobilus shares more characteristics with arborescent lycopsids than any herbaceous forms, and falls within the diagnosis of Lepidostrobus . Each sporophyll consists of a deltoid pedicel and a triangular lamina. Sporophylls are horizontally inserted into the strobilus axis in low spirals. The pedicel, with lateral alations and an abaxial keel, extends distally into the upturned lamina and downturned heel, producing a peltate appearance. A single sporangium with terminal longitudinal dehiscence is a radially elongate, dorsiventrally flattened ovoid and is attached along its length to the adaxial surface of the pedicel. A column-like subarchesporial pad is found in the sporangium. A possible ligule occurs on the adaxial surface of the pedicel distal to the sporangium. The strobilus is microsporangiate, containing Lycospora -type spores with granulate ornamentation and an equatorial flange. Based on this new species, the reproductive diversification and evolutionary pattern in arborescent lycopsids from the Devonian through the Carboniferous are discussed in a phylogenetic framework. We suggest that the reproductive strategies represented by bisporangiate- and monosporangiate-strobili had proliferated by the Late Devonian, which implies that phylogenetically advanced arborescent lycopsids bearing Lepidostrobus strobili had an earlier origin than previously thought.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 55−67.     

Clarification of the name Nothofagus alpina and a new epithet for a Nothofagus hybrid

Arguments in favour of using Nothofagus alpina (Poepp. & Endl.) Oerst. (Fagaceae) rather than N. procera Oerst. or N. nervosa (Phil.) Krasser as the correct name for rauli are presented. We also refute suggestions that N. alpina is based on hybrid material. The hybrid of N. alpina with N. obliqua (Mirb.) Blume is formally described as N . ×  dodecaphleps Mike L. Grant & E. J. Clement and a key to the deciduous taxa of Nothofagus is provided.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 447–451.     

The origin of snakes (Serpentes) as seen through eye anatomy

Snakes evolved from lizards but have dramatically different eyes. These differences are cited widely as compelling evidence that snakes had fossorial and nocturnal ancestors. Their eyes, however, also exhibit similarities to those of aquatic vertebrates. We used a comparative analysis of ophthalmic data among vertebrate taxa to evaluate alternative hypotheses concerning the ecological origin of the distinctive features of the eyes of snakes. In parsimony and phenetic analyses, eye and orbital characters retrieved groupings more consistent with ecological adaptation rather than accepted phylogenetic relationships. Fossorial lizards and mammals cluster together, whereas snakes are widely separated from these taxa and instead cluster with primitively aquatic vertebrates. This indicates that the eyes of snakes most closely resemble those of aquatic vertebrates, and suggests that the early evolution of snakes occurred in aquatic environments.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 469–482.     

Thermal physiology and the origin of terrestriality in vertebrates

The adaptive reasons for the evolutionary transition between obligatorily aquatic lobe-finned fish and facultatively terrestrial early tetrapods have long been debated. The oldest adequately known amphibians, Acanthostega and Ichthyostega , from the final stage in the Upper Devonian (Famennian), can be clearly distinguished from the most advanced choanate sarcopterygian fish from the next older stage (Frasnian) by the presence of large pectoral and pelvic girdles, limbs generally resembling those of later Palaeozoic land vertebrates, and the absence of bones linking the back of the skull with the shoulder girdle. Upper Devonian and most Lower Carboniferous amphibians, like their aquatic predecessors, differed significantly from modern amphibians in their much larger size, up to a metre or more in length. Animals of this size, resembling modern crocodiles and the marine iguana, could have raised their body temperatures by basking in the sun and sustained them upon re-entry into the water. It is hypothesized that the physiological advantages of thermoregulation were a major selective force that resulted in the increasing capacity for the ancestors of tetrapods to move into shallow water, and later to support their bodies against the force of gravity and increase the size and locomotor capacities of the limbs.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 345–358.     

A new species of Estinnophyton from the Lower Devonian Posongchong Formation, Yunnan, China; its phylogenetic and palaeophytogeographical significance

A new species of early vascular land plant, Estinnophyton yunnanense , is described from the Lower Devonian (upper Pragian) Posongchong Formation at Zhichang village, Gumu Town, Wenshan District, Yunnan Province, China. This plant possesses aerial stems with isotomous branching. The leaves are slender, once- or twice-bifurcated, and arranged in low spirals or pseudowhorls. Fertile leaves are morphologically identical to the vegetative ones, clustered on the axis, forming a loose strobilus-like structure. There are three to six fertile leaves in each gyre. Two ovoid-elongate sporangia are singly attached to the adaxial surface each subtended by a recurved stalk, either at the base of each segment in a once-bifurcate fertile leaf or below the points of successive divisions in a twice-bifurcate fertile leaf. Estinnophyton yunnanense differs from two other species described previously from Europe, E .  gracile and E .  wahnbachense , mainly in possessing two single stalked sporangia on each fertile leaf rather than two pairs of sporangia, as well as in the dimensions of its leaves. In view of the complex taxonomy and nomenclature surrounding the type material of Estinnophyton , we considered possible emendation of the generic diagnosis based on the new Chinese material. Following comparison with typical members of the Protolepidodendrales and other taxa, Estinnophyton is considered to have had a closer affinity with primitive sphenopsids than the protolepidodendralean lycopsids. A comparison between Chinese and European floras containing Estinnophyton indicates that South China and Western Europe had floristic elements in common although they were in different phytogeographical units during the Early Devonian epoch.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 201–216.     

A new spiny, cushion-like Euphorbia (Euphorbiaceae) from south-west Iran with special reference to the phytogeographic importance of local endemic species

Euphorbia acanthodes Akhani is described as a new gypsophilous species from south-west Iran. The affinity, morphological and carpological features, and habitat of the species are discussed. A list of a further 36 interesting endemic species of south-western Iran and adjacent areas is given with notes on their affinities and distribution. These include Centaurea gudrunensis Boiss. & Hausskn., as a new record from Iran. The specific status of Pteropyrum naufelum Al-Khayat is confirmed and it is suggested that P. noëanum Boiss. ex Meisn. is synonymous with P. aucheri Boiss. The biogeographic importance of the so-called 'Persian foothills', and biodiversity conservation status of the area are discussed, together with distribution maps of 12 species. It is concluded that the majority of the endemic species in the southern and south-western parts of Iran are Irano–Turanian or have their origins in the Irano–Turanian region. Therefore, it is questionable to consider the area as part of either the Saharo–Sindian, Sudanian or Sudano–Zambezian regions.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 107–121.     

A new Spanish species of Tragopogon (Asteraceae: Lactuceae)

Tragopogon cazorlanum (Asteraceae: Lactuceae) is proposed as a new Spanish species with distribution restricted to the Baetic mountains (south-eastern Spain). Its morphological, palynological, chromosomal, biogeographical and ecological features are discussed, as well as its main relationships and differences with reference to other Spanish species of the genus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 505–511.     

Anatomy of the vegetative organs and secretory structures of Rhaponticum carthamoides (Asteraceae)

Roots, stems, rhizomes and leaves of Rhaponticum carthamoides (Willd.) Iljin (a Siberian adaptogenic plant, originating from the Altai and Saian Mountains) of different ages were investigated by means of light and electron microscopy. Schizogenous secretory reservoirs occurred in every organ, and were located within the secondary xylem (adventitious roots and rhizome of young plants), at the interface of endodermis/cortical parenchyma (roots and hypocotyl), along phloem and primary xylem (older rhizome), around the vascular bundles (inflorescence stem, petiole and leaf midrib veins) and along phloem (cotyledonary and leaf veins). At the interface of endodermis/inner parenchyma, secretion accumulated in the intercellular spaces prior to the formation of proper epithelial cells. The secretion as observed by transmission electron microscopy comprised three components: soluble (i.e. absent from sections; probably phenolic), insoluble and strongly osmiophilic (probably phenolic) and insoluble, moderately osmiophilic (probably lipidic). Numerous osmiophilic oil droplets, similar to the lipidic secretion inside the reservoirs, local proliferation of rough endoplasmic reticulum and numerous multivesicular bodies characterized epithelial cells in all organs. Leucoplasts (in subterranean organs) with osmiophilic inclusions and peroxisomes with crystalloid inclusions were specific for parenchyma cells. Peltate glandular hairs were formed on leaf blades.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 207–233.     

Intercontinental distribution of Plagiochila corrugata (Plagiochilaceae, Hepaticae) inferred from nrDNA ITS sequences and morphology

Plagiochila sect. Vagae is a large pantropical clade that is characterized morphologically by frequent terminal branching, vegetative distribution by propagules on the ventral surface of the leaves and a capsule wall with thickenings in all layers. Plagiochila corrugata from Brazil is characterized by strongly undulate, toothed leaf margins and represents the only known neotropical species of sect. Vagae with unispiral elaters. Plagiochila cambuena from Madagascar is distinguished by the same features. Maximum likelihood and parsimony analyses of 38 nrDNA ITS sequences of Plagiochila reveal P. corrugata and P. cambuena in a weakly (ML) to well (MP) supported monophyletic lineage within P.  sect.  Vagae . As an outcome of the morphological and molecular investigation, P. cambuena is relegated to the synonymy of P. corrugata. Plagiochila corrugata is placed in a Vagae -subclade with 11 further American species. The range of P. corrugata can be ascribed to long-range dispersal from the Neotropics rather than a Gondwanan distribution. Species from tropical Asia and Africa are placed at the base of the Vagae clade. Branch length within P.  sect.  Vagae points to a sudden radiation.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 469–481.     

On the presence of extracellular calcium oxalate crystals on the inflorescences of Araceae

This study presents a survey of the species of the Araceae where extracellular production of calcium oxalate crystals has been observed and discusses the patterns of production of the crystals in different genera. For all Araceae studied using SEM, the oxalate crystals exuding on the epidermal surface correspond to extended aggregate/druses or crystal sand and the oxalate crystals mixed with pollen correspond to raphides or styloids (prismatic crystals). The type of crystals associated with pollen varies among genera. However, the presence of crystals associated with pollen is a specific rather than a generic characteristic. Our results show that the presence of raphides mixed with pollen seems to be a widespread phenomenon in the aroid family.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 181–190.     

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.     

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.     

Seed morphology and anatomy of Austrotaxus spicata (Taxaceae) and its systematic position

The anatomy and ultrastructure of seed envelopes of a New Caledonian endemic Austrotaxus spicata were examined for the first time. The systematic position and phylogenetic relations of Austrotaxus were analysed in light of these data. The structure of aril and spermoderm were investigated to demonstrate the similarities with Phyllocladus as well as with Taxus and Pseudotaxus . On the basis of all female reproductive organ characters, Austrotaxus appeared to be fairly isolated and its placing in the independent family Austrotaxaceae was confirmed from the standpoint of comparative anatomy of the seed coat. Taking into consideration that the heterobathmy of features can be the most distinctively traced in the structure of reproductive organs, evaluating the extent of evolutionary advancement of Austrotaxus seems to be rather difficult. However, it is evident that the relationship of Austrotaxus either with Taxaceae or with Podocarpaceae s.l . is considerably remote.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 437–443.