Systematic and ecological wood anatomy of Neotropical Schefflera (Araliaceae), with an emphasis on the Didymopanax group

Wood anatomy has been investigated from 35 species belonging to the Neotropical clade of the polyphyletic genus Schefflera (Araliaceae), representing three of the five subgroups (Didymopanax, Crepinella and Sciodaphyllum). The species examined are rather uniform in their wood structure, sharing the presence of scalariform and simple perforation plates, septate fibres and scanty paratracheal axial parenchyma. The observed variation in many wood characters showed statistically significant differences relative to latitude, climate and, especially, vegetation types. In particular, the intervessel pits are larger in species from higher latitudes and in seasonally dry habitats than those from lower latitudes and rainforests. Latitudinal and ecological trends in the variation of vessel element lengths, bar numbers on perforation plates, intervessel pit sizes and ray widths may be at least partially explained as effects of adaptation to drier environments in the course of dispersal outside the Amazonian region and diversification in the Atlantic Forest subclade and the Savannic subclade within the Didymopanax group. The occurrence of a granular annulus on the intervessel pit membranes in S. chimantensis and S. sprucei (both of the Crepinella group) is the first record of this feature in Araliaceae. In comparisons of Neotropical Schefflera with the other major clades of Schefflera sensu lato, wood anatomical diversity is consistent with the polyphyly of this genus based on molecular phylogenetic analyses. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 452–475.     

Higher level relationships of Apiales (Apiaceae and Araliaceae) based on phylogenetic analysis of rbcL sequences

The two families of the order Apiales (Apiaceae and Araliaceae) represent a classic example of the difficulty in understanding evolutionary relationships between tropical-temperate family pairs. In Apiales, this problem is further compounded by phylogenetic confusion at almost every taxonomic level, including ordinal, interfamilial, and infrafamilial, due largely to difficulties in understanding trends in morphological evolution. Phylogenetic analyses of rbcL sequences were employed to resolve relationships at the ordinal and familial levels. The results of the ordinal analysis confirm the placement of Apiales in an expanded subclass Asteridae as the sister group to Pittosporaceae, and refute the traditional alliance of Apiales with Cornales and Rosidae. This study has also resolved relationships of a number of enigmatic genera, suggesting, for example, that Melanophylla, Aralidium, Griselinia, and Toricellia are close relatives of Apiales. Clarification of phylogenetic relationships has concomitantly provided insights into trends of morphological evolution, and suggests that the ancestral apialean taxon was probably bicarpellate, simple-leaved, woody, and paleotropical. Phylogenetic analysis at the family level suggests that apiaceous subfamily Hydrocotyloideae, often envisioned as an intermediate group between Apiaceae and Araliaceae, is polyphyletic, with some hydrocotyloids closely allied with Araliaceae rather than Apiaceae. With the exception of some hydrocotyloids, Apiaceae appear to be monophyletic. The relationship between Apiaceae and Araliaceae remains problematic. Although the shortest rbcL trees suggest that Apiaceae are derived from within a paraphyletic Araliaceae, this result is only weakly supported.     

Wood anatomy of Penaeaceae (Myrtales): comparative, phylogenetic, and ecological implications

Wood samples of stems, lignotubers, and roots of the majority of species of Penaeaceae were analyzed with respect to qualitative and quantitative features. Virtually no data have hitherto been presented on xylem features of this family, restricted to Cape Province, South Africa. Presence of vestured pits in vessels, septate crystalliferous parenchyma in wood, intraxylary phloem, predominantly erect ray cells in the typically narrow, multiseriate rays and in the uniseriate rays, and amorphous deposits in ray cells place Penaeaceae securely in Myrtales and help to define that order. By comparison of ecological preferences of the species, as observed during field work, with quantitative analysis of conductive tissue, close correspondence of the wood structure to habit and habitat is demonstrated.     

Pollen Morphology of Hydrocotyloideae and Saniculoideae (Apiaceae) in China

Pollen grains of 30 species and 2 varieties from China, belonging to 5 genera (out of 36 species in 5 genera) of Hydrocotyloideae and Saniculoideae, were examined with light microscope and scanning electron microscope. 1.Six types of the pollen shape are recognized in Hydrocotyloideae and Saniculoideae from China, and their evolutionary trend is from rhomboidal→subspheroidal→ellipsoidal→subrectangular→superrectangular to equatorially-constricted. 2.Pollen grains of Hydrocotyloideae are mostly ellipsoidal with P / E ratio 1.07-1.6, the polar axis 22.5-46μm long, the equatorial axis 13.75-27.5μm long, and with the size index 21.1-31.8.They are angulapeturate in the great majority, and the exine sculpture is recticulate under SEM. Those of Saniculoideae are mostly superrectangular or rectangular, with P/E ratio 1.3-2.1, the polar axis 35-65μm long, the equatorial axis 17.5-42.5μm long, and with the size index 28.2-49.7. They are planaperturate in the great majority and the exine sculpture is striate-reticulate or cerebro-reticulate under SEM. Therefore, the latter might be more advanced than the former. 3.Pollen grains of Dickinsia Franch., endemic to China, are ellipsoidal,with P/E ratio 1.5,the polar axis 31.25-37.5μm long, the equatorial axis 20-25μm, and with the size index 27.81. They are angulaperturate, and the exine sculpture is cerebroid-reticulate under SEM. The genus is greatly similar to the members of Hydrocotyloideae in pollen morphology, which supports the treatment of the genus asa member of Hydrocotyloideae by Handel-Mazzetti (1933).     

Systematic and phylogenetic value of wood anatomy in Heteromorpheae (Apiaceae,Apioideae)

The wood anatomy of all four woody genera of the tribe Heteromorpheae (Apiaceae, subfamily Apioideae) has been described and compared, based on 40 wood samples (representing nine species of Anginon, one species of Glia, three species of Heteromorpha and two species of Polemannia). The four genera were found to be relatively similar in their wood anatomy. Helical thickenings on the vessel walls occur in all species investigated and appear to represent an ancestral character state and a symplesiomorphy for the tribes Bupleurieae and Heteromorpheae. Each of four genera has a diagnostically different combination of character states relating to the diameter of vessels, size of intervessel pits, length of fibres, presence and arrangement of banded axial parenchyma, size of rays and ray cells, and presence of septate fibres and crystals in the ray cells. The occurrence of marginal axial parenchyma in Anginon and Glia may be an additional synapomorphy for these taxa. Variation in the wood anatomy of 31 samples from nine species of Anginon is not correlated with habitat (Fynbos or Succulent Karoo Biomes), but instead appears to reflect adaptations to seasonal aridity found in both ecosystems. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 569–583.     

Wood and bark anatomy of lianoid Indomalesian and Asiatic species of Gnetum

Quantitative and qualitative data on wood and bark are offered for 11 species of lianoid Indomalesian and Asiatic species of Gnetum section Cylindrostachys. Material of roots was studied for two species, material of an underground stem for one species, and stem material was studied for all species; wood from inside and outside of a large stem of G. montanum was analysed (quantitative data do not change with age in this species). Roots have shorter, narrower vessel elements, more numerous per mm², compared with those of stems; these trends run counter to those in dicotyledons. Roots and underground stems have more abundant parenchyma and less abundant sclerenchyma than do stems. Parenchyma of both roots and stems is rich in starch. All of the species studied here have both fibre-tracheids and tracheids, but tracheids are not distributed vasicentrically as they are in dicotyledons. Tori are reported for tracheary elements of three species studied here. Vasicentric axial parenchyma (which usually is thick-walled) is present in all species; thick or thin-walled diffuse or diffuse-in-aggregates apotracheal parenchyma is present in almost all of the species studied. Rays are mostly dimorphic in size, but show various conditions with respect to wall thickness, sclerenchyma presence, and crystal presence. As in other lianoid species of Gnetum, the species of the present study show origin of lateral meristem activity in parenchyma of the innermost cortex. Cortex and bark of the species studied here are relatively uniform in distribution of gelatinous fibres, nests of sclereids, the cylinder of brachysclereids that extends around the stem, and sclerenchymatous phelloderm. Laticifers were observed in bark of only two species studied. Although a few species characters are evident, the species that comprise Section cylindrostachys differ from each other mostly in degree rather than in presence or absence character state distributions. Secretory cavities are newly reported for the genus.     

Comparative studies in phytochemistry and fruit anatomy of Thapsia garganica and T. transtagana,Apiaceae (Umbelliferae)

Thapsia garganica L. and T. transtagana Brot. are classified as synonymous in Flora Europaea. In the present investigation significant differences between the two taxa with regard to fruit anatomy and phytochemistry are demonstrated and they are considered as two separate species. Microscopic studies revealed a distinct difference in number and position of secretory spaces in the pericarp of fruits from the two species and in addition pronounced differences were found in the presence of thapsigargins, the bioactive constituents of the two species. Quantitative HPLC analyses of thapsigargins were carried out on different plant organs from T. garganica and T. transtagana collected from various locations. Thapsigargin, thapsigargicin, nortrilobolid and thapsivillosin I and J were dominant compounds in all organs of T. garganica whereas none of these compounds could be detected in any organ of T. transtagana. On the contrary, thapsitranstagin and trilobolid were main thapsigargins of T. transtagana. T. garganica may include two chemotypes, as trilobolid was detected in some specimens of T. garganica only. As this is the first time trilobolid has been detected in these two species, it was isolated from both and identified by ¹H NMR-spectroscopy.     

Wood anatomy of the tribe Podalyrieae (Fabaceae,Papilionoideae): Diversity and evolutionary trends

Detailed wood anatomical data for 32 species from all nine genera of the tribe Podalyrieae are presented, together with numerical analyses and the mapping of character states onto the latest available molecular phylogeny. It was found that trees (Cadia, Calpurnia and Virgilia) have vessels in small isolated groups, whilst fynbos shrubs (the remaining genera: Amphithalea, Cyclopia, Liparia, Podalyria, Stirtonanthus and Xiphotheca) commonly show highly grouped narrow vessels (frequently in a dendritic pattern), and helical thickening on the vessel walls. Comparisons of the main character state changes with the molecular phylogeny of the tribe show that the wood structure of trees probably represents the basic condition in the tribe; character states present in shrubs appear to have arisen a few times and very likely represent adaptations to seasonal water stress. In general, the wood anatomy is congruent with current subtribal and generic delimitations. Fire-survival strategy is reflected in the rays, with seeders having mostly procumbent cells whilst sprouters have square and upright cells. The close similarity in wood anatomy between Cadia and Calpurnia is in agreement with the transfer of Cadia to the Podalyrieae. A remarkable diversity of crystals was found, including prismatic, acicular and navicular crystals, the last two of which may occur singly or in sheaf-like aggregates.     

Wood and bark anatomy of Muntingiaceae: A phylogenetic comparison within Malvales s. l.

Quantitative and qualitative data on wood and bark anatomy are given for Muntingia calabura L. and Dicraspidia donnell-smithii Standley. These data are compared with phylogenetic schemes, based on DNA analysis, in which Muntingiaceae belong to the “dipterocarp clade” within Malvales. The data are consistent with this hypothesis, although Muntingiaceae lack pit vestures in vessels, which are seen in the other malvalean families (Cistaceae, Dipterocarpaceae, Neuradaceae, Sarcolaenaceae, Thymeleaceae), and this may represent a loss of pit vestures. All families of the dipterocarp clade agree with both genera of Muntingiaceae in having tracheids as the imperforate tracheary element type (at least ancestrally), although fiber-tracheids also occur in some Dipterocarpaceae and Thymeleaceae. The large size of some malvalean families (with attendant greater diversity in character states) and a paucity of wood studies in those families make for difficulty in comparison of features such as axial parenchyma and ray types with those of Muntingiaceae; character states of these features are consistent with placement of Muntingiaceae in the dipterocarp clade of Malvales. Banded phloem fibers in bark of Muntingiaceae are much like those of other Malvales. Wood of Muntingiaceae is highly mesomorphic according to quantitative vessel features.     

The phylogenetic position of Peucedanum sensu lato and allied genera and their placement in tribe Selineae (Apiaceae, subfamily Apioideae)

ITS sequences of members of Peucedanum and its segregates (Cervaria, Holandrea, Imperatoria, Oreoselinum, Pteroselinum, Thysselinum, Tommasinia, and Xanthoselinum), as well as representatives of other members of the previously identified Angelica and Arracacia clades, were analysed using distance, maximum likelihood, and maximum parsimony methods in order to establish their phylogenetic relationships. The Angelica and Arracacia clades comprise a strongly supported monophyletic group that is formally recognised as tribe Selineae. The genus Holandrea, expanded to include Cervaria caucasica and Peucedanum achaicum, is distantly related to the other peucedanoid genera. The remaining segregates are closely related to one another and to Peucedanum sensu stricto. The genera Seseli and Selinum are polyphyletic. Members of the genus Angelica form a single clade that also includes representatives of Selinum and Sphenosciadium. New combinations in Holandrea (H. caucasica, H. achaica) and Angelica (A. capitellata) are proposed, and Selinum pyrenaeum is recognised in Angelica (A. pyrenaea).     

Variation in native pollinators in the absence of honeybees: implications for reproductive success of an Australian generalist-pollinated herb Trachymene incisa (Apiaceae)

Most plants are generalist in terms of pollination, with the potential for significant spatial and temporal variation in their pollinators. Few studies have investigated how variable pollinators are in a generalist pollination system or how this variation affects plant reproduction. We investigated the degree of variation in pollinators and resulting reproductive success among populations of a widespread generalist-pollinated herb, Trachymene incisa (Apiaceae). The European honeybee was unexpectedly absent from the pollinator assemblages, providing the unique opportunity to study the native Australian pollinators. Insect visitation rates and the taxonomic composition of the pollinator assemblage varied significantly across populations, indicating that populations of T. incisa are not equally serviced and are not equally generalist. This highlights that sampling one population would not characterize the extent of species generalization. There was no positive correlation between insect visitation rate and reproductive success, with the Agnes Banks population receiving the highest visitation rate but producing the lowest reproductive output, and the Myall Lakes population receiving the lowest visitation rate and producing the highest seedling emergence. This study shows that variation in pollinators can have measurable effects on populations of generalist-pollinated plants, therefore there is potential for large-scale change in all plant–pollinator interactions.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 479–490.     

Spermatozoa and spermiogenesis of Liphistius cf. phuketensis (Mesothelae, Araneae, Arachnida) with notes on phylogenetic implications

The present study deals with the spermatozoa and spermiogenesis of Liphistius cf. phuketensis, a representative of the most primitive and enigmatic spider group Mesothelae. The general organization of the spermatozoa is very similar to the condition known from Amblypygi supporting a sister-group relationship between Araneae and Amblypygi. Besides plesiomorphic characters such as, e.g., an elongated and corkscrew shaped nucleus, the sperm cells are characterized by several apomorphic characters, e.g., the giant body and conspicuous membranous areas which are formed at the end of spermiogenesis. As the transfer form, coenospermia are formed at the end of spermiogenesis, which strongly supports the idea that this type of sperm aggregation is the primitive transfer form within spiders. A very remarkable character of the spermatozoa of some groups of arachnids is the coiling of the main cell organelles at the end of spermiogenesis. Previously, the Mesothelae were believed to be the only spider group which does not show a complete coiling of the main cell organelles. With the present study the first evidence of a complete coiling of spermatozoa within this primitive spider group could be documented, indicating that this character is part of the ground pattern of spider spermatozoa. Consequently, the incomplete coiling seems to be a synapomorphy of certain species of Mesothelae, which sheds new light on the discussion of the phylogenetic relationships of this group.     

Juvenile cranial anatomy of Nimravidae (Mammalia, Carnivora): biological and phylogenetic implications

The anatomy of the spectacular sabretoothed carnivorans has been studied many times. Their behavioural and biomechanical implications are well known in adults, but not in juvenile individuals, which are poorly represented in the fossil record. This work reports on a newly discovered Hoplophoneini (Nimravidae Nimravinae) of 5–6 months of age, which represents the earliest and one of the most complete juvenile crania discovered. We discuss and compare the juvenile anatomy of Palaeogene nimravids relative to that of other mammals, especially other carnivorans including the Miocene nimravids (i.e. Barbourofelinae). We also discuss the ontogenetic development of this individual and other sabretoothed predators, which provides important data for reconstructing the biology of these extinct predators. Our study of the juvenile anatomy of the Hoplophoneini reveals an original association of basicranial features: the ectotympanic rotates away from the cranium earlier than in other mammals, which may be an additional autapomorphy distinguishing the Palaeogene Nimravidae from other Carnivora; within Palaeogene Nimravidae, at a similar ontogenetic stage, there is a major difference in development and maximum ventral extension of ossification of the caudal entotympanic; in contrast to the Miocene nimravid Barbourofelis, there is no strong relationship between the eruption of the upper canines and the development of other sabretoothed features (especially the mandibular flange) in the individual described. These results confirm the conclusions of recent studies which suggested that Palaeogene Nimravidae (Nimravinae) and Miocene Nimravidae (Barbourofelinae) are not closely related. Muscular reconstruction and biomechanical models also confirmed that juvenile individuals were probably not able to kill or hunt any prey. They may have fed on carcasses, eating muscles by using the incisors, and associated with a pulling or twisting action of neck. © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 138 , 477–493.     

Species boundaries,phylogenetic relationships,and ecological differentiation inAnthriscus (Apiaceae)

Species boundaries and phylogenetic relationships of 17 taxa ofAnthriscus (Apiaceae), with special emphasis on the critical sect.Cacosciadium, were explored using morphological data with principal component analysis, phenetics, and phylogenetics. The analyses did not provide satisfactory resolution of taxa from sect.Cacosciadium and only four species were retained. The total number of species was reduced to nine. Sect.Cacosciadium is distinguished by only two synapomorphies while sects.Anthriscus andCaroides are better supported. Present geographic and ecological variation suggests that the radiation ofAnthriscus occurred through divergence of peripheral isolated populations adapting to different habitats: high montane meadows and screes, shady climax forests, and seasonally dry habitats at lower altitudes. The adaptive significance of particular morphological traits is discussed.     

Phylogeny and biogeography of Chinese Heracleum (Apiaceae tribe Tordylieae) with comments on their fruit morphology

The Sino-Himalayan Mountain Region is a major center of diversity of Heracleum (Apiaceae, Tordylieae, Tordyliinae), yet the phylogenetic relationships among its rich diversity of endemic taxa are largely unknown. In this study, we (1) investigate the monophyly of Heracleum and ascertain the phylogenetic placements of those species of Heracleum from China that fall outside of the group; (2) estimate phylogenetic relationships within Heracleum sensu stricto, particularly among those species occurring in the Hengduan Mountain Region of southwestern China; and (3) reconstruct the biogeographic history of Heracleum sensu stricto. To achieve these objectives, Bayesian inference and maximum-parsimony analyses of the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) region and five noncoding chloroplast DNA (cpDNA) loci (rps16-trnK, rps16 intron, trnQ-rps16, rpl32-trnL, and psbA-trnH), separately and in combination, were carried out. Fruit morphology of the Chinese Heracleum species was examined. Statistical dispersal-vicariance analysis (S-DIVA) was used to reconstruct the biogeographic history. In total, 114 accessions were considered, which included 88 species representing eight genera of subtribe Tordyliinae, 28 of the 29 species of Heracleum occurring in China, and 4 species of Angelica as outgroups. The genus Heracleum is monophyletic upon the exclusion of nine species of Heracleum from China and the inclusion of the Caucasian monotypic genera Mandenovia and Symphyoloma. These nine excluded species show affinities to Tetrataenium, Semenovia, and Angelica. The species of Heracleum endemic to southwestern China form a moderately to strongly supported clade; however, major discordance between the ITS- and cpDNA-derived phylogenies, lack of resolution in the trees, and polyphyly of several species preclude unequivocal hypotheses of phylogenetic relationship. The distinctly clavate vittae, shorter than the length of the mericarp, is a diagnostic feature for Heracleum sensu stricto. Heracleum originated in the West Palearctic, with three possible migration routes leading to its present-day distribution. Major topological conflicts between the ITS and cpDNA trees may be due to interspecific hybridization and/or incomplete lineage sorting. The taxonomic limits of many Chinese Heracleum species remain unclear, possibly because of recent radiation in the Hengduan Mountains.     

First phylogenetic analysis of the tribe Oligaphorurini (Collembola: Onychiuridae) inferred from morphological data,with implications for generic classification

Oligaphorurini represent tribe of the subfamily Onychiurinae, which currently comprises 5 genera and 53 species. The present study evaluated the monophyly of Oligaphorurini genera. We investigated phylogenetic relationships among 39 species, representing all extant genera of Oligaphorurini. Both equal- and implied-weighting parsimony analyses were used in phylogenetic reconstruction. The cladistic analyses were based on comprehensive survey of adults’ morphological characters because specimens suitable for molecular studies were not available for the majority taxa. The phylogenetic analysis resulted in the recognition of a monophyletic Chribellphorura, and strongly supported non-monophyly of the previously recognized genera Archaphorura, Dimorphaphorura, Micraphorura, and Oligaphorura. The following new synonymy is recognized: Oligaphorura = Dimorphaphorura syn. nov., = Micraphorura syn. nov., = Archaphorura syn. nov. The general classification of Oligaphorurini is followed by the diagnoses of genera and key to the all known species.     

Articulation of the pelvic spine in acanthopterygian fishes,with notes on its phylogenetic implications

The pelvic spine articulation in 88 non-squamipinne acanthopterygian species belonging to 59 families and 8 orders was examined. For most acanthopterygians, the pelvic bone lacks a foramen for the attachment of the pelvic spine flanges and this structural condition is considered a primitive acanthopterygian state. Only in Caproidae, Leiognathidae, and Polynemidae among the examined species do the pelvic spine flanges fuse through a foramen in the pelvic bone and this articulation is considered most derived. In the remaining acanthopterygians examined, the pelvic spine flanges butt against each other through the pelvic foramen. However, these flanges do not fuse. This structure is considered an intermediate state. The possible phylogenetic implication of these articulating states regarding acanthopterygian phylogeny is discussed.     

Ultrastructural observations of spermatozoa and spermiogenesis in Wandella orana Gray, 1994 (Araneae: Filistatidae) with notes on their phylogenetic implications

Spermatozoa and spermiogenesis of the prithine filistatid spider Wandella orana are described. The spider produces coenospermia, i.e. sperm aggregations that include several single sperm cells commonly surrounded by a secretion sheath. One sectioned coenospermium in W. orana contains at least five spermatozoa. During copulation many coenospermia are transferred into the female. Coenospermia are regarded as a peculiar transfer form of sperm which occurs in early derivative spiders such as Liphistiomorphae and Mygalomorphae. The only exception which was found in Araneomorphae until now was the filistatine spider Filistata insidiatrix. Our observation is the second case and supports the view that Filistatidae represent an early derivative taxon. Furthermore, the individual sperm cells show characteristics which also may be regarded as being plesiomorphic. There is a cone-shaped acrosomal vacuole, a very long acrosomal filament, a rather stout nucleus and a small implantation fossa. The axoneme shows the 9x2+3 pattern of microtubules which is synapomorphic in Megoperculata (Uropygi, Amblypygi and Araneae). The finding of coenospermia in two distant taxa of Filistatidae may have consequences for phylogenetic and systematic considerations.