Araucariaceae (Pinopsida): Aspects in palaeobiogeography and palaeobiodiversity in the Mesozoic

The paper examines recent information on the history of the ‘southern hemisphere’ conifer family Araucariaceae and gives supplementary notes to previously published monographs. Important data from the Mesozoic fossil record are presented and summarized to describe the ancient diversity and distribution of Araucariaceae. Information on the origin of the family and oldest fossil records of the present-day genera are assembled. Existing models on phylogenetic relationships within the family inferred from molecular data are compared with relationships between the genera determined from morphological data. Reasons for disappearance of representatives of this conifer family in the latest Cretaceous in North America and Europe are briefly discussed.     

Yezonia,a new section ofAraucaria (Araucariaceae) based on permineralized vegetative and reproductive organs ofA. vulgaris comb. nov. from the upper cretaceous of Hokkaido,Japan

A new specimen of an araucarian cone,Araucaria nihongii, was found attached to the vegetative organs ofYezonia vulgaris, and is described asAraucaria vulgaris comb. nov. Thick branches show characteristic bark structure with lenticular patches. Secondary wood is usually araucarioid. Leaves are arranged helically on shoots, which are imbricate, appressed and fused to surface of the stem. External and anatomical features of leaves closely resembleBrachyphyllum. The seed cone is spherical with winged bracts and thin ovuliferous scales. One seed is borne per cone-scale complex. The seed coat and nucellus wall show typical araucarian structure. An araucarian plant that boreBrachyphyllum-like foliage and aEutacta-like seed cone was predicted by Harris in 1979. This reconstructed plant,Araucaria vulgaris, supports this theory and proves the presence of an extinct characteristic-form of the genus. A new section of the genus was proposed forAraucaria vulgaris. Structure and Affinities of the Petrified Plants from the Cretaceous of Northern Japan and Saghalien XV, Consecutive number from previous paper (Nishidaet al. 1993).     

Phylogenetic relationships within Araucariaceae based on rbcL gene sequences

Phylogenetic relationships were determined in the Araucariaceae, which are now found mainly in the Southern Hemisphere. This conifer family was well diversified and widely distributed in both hemispheres during the Mesozoic era. The sequence of 1322 bases of the rbcL gene of cpDNA was determined from 29 species of Araucariaceae, representing almost all the species of the family. Phylogenetic trees determined by the parsimony method indicate that Araucariaceae are well defined by rbcL sequences and also that the monophyly of Agathis or Araucaria is well supported by high bootstrap values. The topology of these trees revealed that Wollemia had derived prior to Agathis and Araucaria. The rbcL phylogeny agrees well with the present recognition of four sections within Araucaria: Araucaria, Bunya, Eutacta, and Intermedia. Morphological characteristics of the number of cotyledons, position of male cone, and cuticular micromorphologies were evaluated as being phylogenetically informative. Section Bunya was found to be derived rather than to be the oldest taxon. Infrageneric relationships of Agathis could not be well elucidated because there are few informative site changes in the rbcL gene, suggesting the more recent differentiation of the species as their fossil records indicate. The New Caledonian Araucaria and Agathis species each formed a monophyletic group with very low differentiation in rbcL sequences among them, indicating rapid adaptive radiation to new edaphic conditions, i.e., ultramafic soils, in the post-Eocene era.     

Morphology,biology, and systematic relationships ofDesmaria (Loranthaceae)

Desmaria mutabilis is unique inLoranthaceae in having dimorphic shoots, the short shoots producing a terminal inflorescence. Other unusual features in the family are well differentiated bud scales and deciduousness. The normal position of mature plants on the trunks of large trees is shown to be a consequence of profuse vegetative reproduction from the epicortical roots, the predominant growth direction of the latter towards the trunk from the original site of establishment on a lateral branch, and the ability of epicortical roots to generate haustorial contacts through heavy host bark. The seedling is heterocotylar, one cotyledon being phanerocotylar, the other cryptocotylar and functioning as a haustorial organ in the endosperm. It is suggested thatDesmaria is a member of the primitive complex of loranthaceous genera which includesGaiadendron.     

Development of mucilage cells of Araucaria angustifolia (Araucariaceae)

The roles of mucilage cells were investigated through morphological and cytological analysis during leaf development in young Araucaria angustifolia plants. Differentiation began in leaf primordia in the shoot apex, when the young cells underwent a greater increase in volume in comparison with other mesophyll cells. The mucilage polysaccharides were synthesized by dictyosomes, from where they were taken by large vesicles and released into a cavity formed by detachment of the tonoplast, which was separated from the cytoplasm. At the end of differentiation, the cell was completely filled with mucilage, a gel consisting of a denser reticular structure surrounding less dense regions. The nucleus and cytoplasm were degenerated in mature cells. The A. angustifolia mucilage cells presented some cytological resemblances to the mucilage cells of members of some dicotyledonous families; however, differences in the dictyosomes and the secretion route were observed. Translocation and water storage of solutes was suggested by the use of the hydroxy pyrenetrisulfonic acid tri-sodium salt apoplastic tracer. The tonoplast detachment, dechromatinization, nuclear condensation, and general degeneration of the membrane systems observed during maturity indicated a programmed cell death process, one not yet described for angiosperm mucilage cells.     

Genetic diversity,differentiation and conservation in <Emphasis Type="Italic">Araucaria bidwillii</Emphasis>(Araucariaceae), Australia's Bunya pine

Habitat fragmentation has resulted in many species becoming geographically restricted, as dispersal among subsequent isolates becomes compromised. This study investigated the effects of historical fragmentation on the genetic diversity and differentiation of Australia's Bunya Pine (Araucaria bidwillii) using random amplified DNA (RAPD) markers. High diversity characterises all Bunya populations sampled, regardless of population size or degree of isolation, which may be either due to similar effective population size among populations, or the lagging effects of the detection of contemporary genetic signal in long-lived conifer species. Large genetic differentiation characterises the northern and southern populations, although all sampled populations are significantly differentiated from each other. The northern population, at Mt Lewis, is responsible for the majority of variation detected among populations, and as such represents a significant genetic reservoir. The conservation of the genotypes of the Mt Lewis population may be imperative for the future of the species, given predicted models of accelerated climate change over the coming decades.     

Diversity variation and tempo-spatial distribution of Otozamites (Bennettitales) in the Mesozoic of China

Otozamites is a representative fossil leaf morphogenus of the extinct Bennettitales, with an extensive distribution during the Mesozoic, especially in China. Understanding the fossil diversity variation and distribution pattern of Otozamites in China will provide information on biodiversity of bennettitalean plants as well as for reconstruction of palaeogeography and palaeoclimate conditions during the Mesozoic. So far, 46 species of this genus have been described in China, excluding unspecified species. The results show that the fossils of Otozamites are extensively recorded in the Late Triassic, and then reach their maximum development in the Early Jurassic, followed by a reduction in diversity in the Middle and Late Jurassic, and finally become extinct at the end of Early Cretaceous. Geographically, they occur in both Northern and Southern Floristic Provinces in the Mesozoic of China, with a relatively higher abundance in the Southern Floristic Province. It implies that the diversity variation and distribution of Otozamites are closely related to the change of the palaeoclimatic conditions. The warm and humid climate prevailed in the Late Triassic and Early Jurassic in South China, propitious to the development of Otozamites. After the Middle Jurassic, dry and hot climate may have caused the lower diversity level and blocked the development of Otozamites; finally at the end of the Early Cretaceous, the frequent arid climate may be a major cause for the extinction of Otozamites.     

The systematic relationships ofAegialitis (Plumbaginaceae) as revealed by pollen morphology

Light and first SEM observations on the pollen ofAegialitis indicate that the two species of the genus are clearly distinguishable from all other genera of the tribeStaticeae to whichAegialitis presently is relegated. Intraspecific exine or aperture dimorphism which appears frequently in this tribe, is not recorded in this genus.A. annulata andA. rotundifolia have distinct monomorphic pollen and show a great resemblance in exine features with the tribePlumbagineae, particularly species ofPlumbago. Considering these and other features, separate subtribal status forAegialitis within thePlumbagineae is proposed.     

Evolution and systematic relationships in theTriticeae (Poaceae)

The evolution and taxonomic relationships in theTriticeae are discussed with the view to highlight aspects of this agronomically important group of plants, which may be of interest to molecular biology. Some of these aspects are addressed in more detail in adjoining papers in which specific genomic loci have been examined at the DNA sequence or isozyme level. Aspects discussed include the systematics and geographic distribution of theTriticeae species, isozyme and chromosome pairing studies on some of the species as well as more recent developments in DNA analyses. A survey of the systematics of theTriticeae indicated that the genomic system ofLöve is probably the most useful starting point for interpreting molecular data even though the system has many problems from a taxonomic point of view. The geographical distribution ofTriticeae species, using both published and unpublished data, suggested that information of this type taken together with the theory of continental drift provides a broad time-span for considering data from DNA sequence studies. The significance, and modes of analyses, of isozyme studies were assessed because they often provide valuable characters in determining relationships between species. The main character underlyingLöve's andDewey's analyses of theTriticeae, namely chromosome pairing, is discussed with particular reference to isozyme studies to show that in some cases, such as species ofHordeum sensu lato, consistent relationships are obtained. Finally, new developments in understanding chromosome structure are considered in relation to the above variables in the taxonomy and evolution of theTriticeae.     

Craniometrical variations among eastern Brazilian marmosets and their systematic relationships

The craniometrical variations among marmosets on the eastern coast of Brazil or theCallithrix jacchus group (C. jacchus, C. penicillata, C. kuhli, C. geoffroyi, C. flaviceps, andC. aurita) were analyzed to test various hypotheses for the arrangement of the species and phylogenetic relationships ofC. kuhli andC. flaviceps. Q-mode correlation coefficients and principal component analysis were employed to extract important shape factors from the craniometrical data. On this basis, the shape distances among the eastern Brazilian marmosets were found to be larger than those between the marmosets which are recognized as distinct species. This evidence supports the hypothesis that the members of theC. jacchus group are distinct species.C. kuhli shows the strongest resemblance toC. geoffroyi within theC. jacchus group.C. flaviceps is the most similar toC. aurita in its cranial shape. These findings indicate thatC. kuhli is most closely related toC. geoffroyi, whileC. flaviceps is toC. aurita.     

Leaf micromorphology inAgathis and its taxonomic implications

This study surveys the micromorphological surface structure of the leaves of the conifer genusAgathis (Araucariaceae) from throughout the range of the genus (Malaysia to New Zealand and Fiji) as seen with the scanning electron microscope. These data confirm that the vegetative parts ofAgathis are taxonomically highly conservative, but suggest the Melanesian and New Zealand elements to be distinctive from those of the rest of the genus, and from one another. Conclusions are compared with those that have been derived from studies based on other characters.     

Partial shoot reiteration in Wollemia nobilis (Araucariaceae) does not arise from 'axillary meristems'

Background and Aims

Conifers are characterized by the paucity of axillary buds which in dicotyledonous trees usually occur at every node. To compensate, conifers also produce ‘axillary meristems’, which may be stimulated to late development. In juvenile material of Wollemia nobilis (Araucariaceae: Massart''s model) first-order (plagiotropic) branches lack both axillary buds and, seemingly, axillary meristems. This contrasts with orthotropic (trunk) axes, which produce branches, either within the terminal bud or as reiterated orthotropic axes originating from axillary meristems. However, plagiotropic axes do produce branches if they are decapitated. This study investigated how this can occur if axillary meristems are not the source.

Methods

The terminal buds of a series of plagiotropic branches on juvenile trees were decapitated in order to generate axillary shoots. Shoots were culled at about weekly intervals to obtain stages in lateral shoot development. Serial sections were cut with a sliding microtome from the distal end of each sample and scanned sequentially for evidence of axillary meristems and early bud development.

Key Results

Anatomical search produced no clear evidence of pre-existing axillary meristems but did reveal stages of bud initiation. Buds were initiated in a group of small starch-rich cortical cells. Further development involved de-differentiation of these small cells and the development of contrasting outer and inner regions. The outer part becomes meristematic and organizes the apex of the new branch. The inner part develops a callus-like tissue of vacuolated cells within which vascular cambia are developed. This kind of insertion of a branch on the parent axis seems not to have been described before.

Conclusions

Axillary meristems in Wollemia characterize the leaf axils of trunk axes so that the origin of reiterated shoots is clear. Plagiotropic axes seemingly lack axillary meristems but still produce axillary branches by distinctive developmental processes. These observations demonstrate limited understanding of branch initiation in trees generally.     

Sporulation and diversity of arbuscular mycorrhizal fungi in Brazil Pine in the field and in the greenhouse

The aim of this work was to assess the sporulation and diversity of arbuscular mycorrhizal fungi (AMF) at different forest sites with Araucaria angustifolia (Bert.) O. Ktze. (Brazil Pine). In addition, a greenhouse experiment was carried out to test the use of traditional trap plants (maize + peanut) or A. angustifolia to estimate the diversity of AMF at each site. Soil samples were taken in two State Parks at southwestern Brazil: Campos do Jordão (Parque Estadual de Campos do Jordão [PECJ]) and Apiaí (Parque Estadual Turístico do Alto Ribeira [PETAR]), São Paulo State, in sites of either native or replanted forest. In PECJ, an extra site of replanted forest that was impacted by accidental fire and is now in a state of recuperation was also sampled. The spore densities and their morphological identification were compiled at each site. In the greenhouse, soil samples from each site were used as inoculum to promote spore multiplication on maize + peanut or A. angustifolia grown on a sandy, low-fertility substrate. Plants were harvested, respectively, after 4 months or 1 year of growth and assessed for mycorrhizal root colonization. Spore counts and identification were also performed in the substrate, after the harvest of plants. Twenty-five taxa were identified considering all sites. Species richness and diversity were greater in native forest areas, being Acaulospora, the genus with the most species. Differences in number of spores, diversity, and richness were found at the different sites of each State Park. Differences were also found when maize + peanut or A. angustifolia were used as trap plants. The traditional methodology using trap plants seems to underestimate the diversity of the AMF. The use of A. angustifolia as trap plant showed similar species richness to the field in PECJ, but the identified species were not necessarily the same. Nevertheless, for PETAR, both A. angustifolia and maize + peanut underestimated the species richness. Because the AMF sporulation can be affected by many conditions, it is impossible to draw detailed conclusions from this kind of survey. More precise experiments have to be set up to isolate the different factors that modulate the ecophysiological interactions between host plant and endophyte.     

Morphology,biology and pesticide tolerance ofChelotogenes ornatus [Acari: Cheyletidae]

BioControl - This paper reports on the morphology, biology and pesticide tolerance ofCheletogenes ornatus (Canestrini &;amp; Fanzago), the most common predator ofPinnaspis aspidistrae (Signoret)...     

Phylogenetic relationships among the ciliate arthrodontous mosses: Evidence from chloroplast and nuclear DNA sequences

Parsimony and maximum likelihood analyses of combinedtrnL (UAA) 5 exon —trnF (GAA) andrps4 exon cpDNA, and 18S nrDNA sequences of 60 arthrodontous moss taxa indicate strong support for the monophyly of a clade containing theSplachnineae, Orthotrichineae, and diplolepideous alternate sub-orders. A clade including theSplachnineae, Meesiaceae andLeptobryum (Bryaceae) is similarly well supported and forms the sister group to a clade comprising theOrthotrichineae and the other diplolepideous alternate mosses. Within this latter clade a number of well supported lineages are identified, but relationships among these remain poorly resolved. These analyses indicate that the Splachnaceous and Orthotrichaceous peristomes have been independently derived from an ancestral perfect bryoid peristome.     

Sectile pollinia and relationships in theOrchidaceae

The sectile state, in which pollinia are subdivided into units larger than tetrads, is one of several systematically important orchid pollinium characters. We recognize two types of sectile pollinia, based upon form and arrangement of massulae. One type is found primarily in members ofOrchidoideae-Spiranthoideae, while the other characterizes epidendroid genera. Additional characters that have systematic potential are massula dimorphism, caudicle composition, and degree of tetrad packing. Hollow sectile pollinia uniteArethusa andCalopogon; Stereosandra andEpipogium have dissimilar pollina, but other characters support a close relationship between them. The sectile condition is interpreted as resulting from a shift in the timing of cohesion of developing pollen cells.     

The embryology and relationships ofOliniaceae

Two of the five species ofOliniaceae (Olinia emarginata andO. ventosa), a monotypic and problematic family of theMyrtales, were investigated embryologically.Oliniaceae clearly agree with otherMyrtales in their basic embryological characteristics, and are characterized further by having an ephemeral endothecium, a campylotropous ovule, and a thick, three-five-layered, outer integument. A combination of these three characteristic features is unknown elsewhere inMyrtales, so that embryological features do not support a close relationship with any other member of the order. Shared distinctive anther characteristics (i.e. ephemeral endothecium) suggest thatOliniaceae are derived from the common ancestor ofCrypteroniaceae s. str.,Rhynchocalycaceae, Alzateaceae, andPenaeaceae.     

Phylogenetic relationships in theCampanulales based onrbcL sequences

Phylogenetic relationships within the angiosperm orderCampanulales were investigated by comparative sequencing of the chloroplast generbcL. CompleterbcL sequences were obtained for ten species in six families within the order. These data were analyzed along with previously publishedrbcL sequences from other taxa (for a total of 117 species) within the subclassAsteridae and outgroups, producing 32 equally parsimonious trees. A subset consisting of 44 of these taxa was then chosen and more rigorous analyses performed, resulting in four equally parsimonious trees. Results indicate that two major clades roughly corresponding to traditionally circumscribedAsterales andCampanulales exist as sister taxa. In particular, therbcL trees indicate thatSphenoclea is not a member ofCampanulales orAsterales, thatPentaphragma is more closely allied toAsterales thanCampanulales, that theCyphiaceae are not monophyletic, thatCampanulaceae andLobeliaceae are not sister taxa, and thatStylidiaceae are correctly placed withinCampanulales.     

Structure-function relationships amongLathyrus lectins

The structure-function relationship of eight structurally very similarLathyrus lectins, which are two-chain lectins from theVicieae tribe, has been studied. Hemagglutination activity, inhibition of hemagglutination, glycoprotein reactivity and mitogenic activity were determined for each of the lectins. Despite the close structural resemblance, marked differences in the carbohydrate binding activity of theLathyrus lectins were recorded. These functional differences were related to small conformational changes due to minor differences in amino acid sequence of eachLathyrus lectin.     

Phylogenetic relationships of theJuglandaceae

A cladistic analysis of molecular data from the chloroplast generbcL was used to examine the taxonomic relationships of the walnut family (Juglandaceae). In addition, chemical and morphological data from a previous study byHufford (1992) were incorporated, expanded, and analyzed independently and in combination with the molecular data. The results of these analyses suggest that theJuglandaceae are more closely related to theFagaceae, Betulaceae, Casuarinaceae, andUrticaceae and their relatives (sensuCronquist 1981) than they are to theAnacardiaceae (sensuThorne 1983). However, sequence data fromrbcL also suggest a relationship between the higherHamamelidae and certain families in theRosidae sensuCronquist 1981 (such asRosaceae andRhamnaceae), an outcome which would add credence to the widely accepted view of the polyphyletic nature of theHamamelidae.