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1.
The order Malvales remains poorly circumscribed, despite its seemingly indisputable core constituents: Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae. We conducted a two-step parsimony analysis on 125 rbcL sequences to clarify the composition of Malvales, to determine the relationships of some controversial families, and to identify the placement of the Malvales within Rosidae. We sampled taxa that have been previously suggested to be within, or close to, Malvales (83 sequences), plus additional rosids (26 sequences) and nonrosid eudicots (16 sequences) to provide a broader framework for the analysis. The resulting trees strongly support the monophyly of the core malvalean families, listed above. In addition, these data serve to identify a broader group of taxa that are closely associated with the core families. This expanded malvalean clade is composed of four major subclades: (1) the core families (Bombacaceae, Malvaceae, Sterculiaceae, Tiliaceae); (2) Bixaceae, Cochlospermaceae, and Sphaerosepalaceae (Rhopalocarpaceae); (3) Thymelaeaceae sensu lato (s.l.); and (4) Cistaceae, Dipterocarpaceae s.l., Sarcolaenaceae (Chlaenaceae), and Muntingia. In addition, Neurada (Neuradaceae or Rosaceae) falls in the expanded malvalean clade but not clearly within any of the four major subclades. This expanded malvalean clade is sister to either the expanded capparalean clade of Rodman et al. or the sapindalean clade of Gadek et al. Members of Elaeocarpaceae, hypothesized by most authors as a sister group to the four core malvalean families, are shown to not fall close to these taxa. Also excluded as members of, or sister groups to, the expanded malvalean clade were the families Aextoxicaceae, Barbeyaceae, Cannabinaceae, Cecropiaceae, Dichapetalaceae, Elaeagnaceae, Euphorbiaceae s.l., Huaceae, Lecythidaceae, Moraceae s.l., Pandaceae, Plagiopteraceae, Rhamnaceae, Scytopetalaceae, Ulmaceae, and Urticaceae.  相似文献   

2.
Sequence analyses of the plastid genes atpB and rbcL support an expanded order Malvales. Within this alliance, core Malvales are clearly supported and comprise most genera that have previously been included in Sterculiaceae, Tiliaceae, Bombacaceae, and Malvaceae. Additional well supported malvalean alliances include the bixalean clade (Bixaceae, Diego-dendraceae, and Cochlospermaceae), the cistalean clade (Cistaceae, Dipterocarpaceae, and Sarcolaenaceae) and Thymelaeaceae (including Gonystyloideae and Aquilarioideae). Our results indicate sister-group relationships between (1) Neuradaceae and the cistalean clade; (2) Sphaerosepalaceae and Thymelaeaceae; (3) these two clades (1 and 2); and (4) all these and an alliance comprising the bixalean clade and core Malvales, but this pattern is weakly supported by the bootstrap. The affinities of Muntingiaceae and Petenaea are especially ambiguous, although almost certainly they are Malvales s.l. The traditional delimitation of families within core Malvales is untenable. Instead, we propose to merge Sterculiaceae, Tiliaceae and Bombacaceae with Malvaceae and subdivide this enlarged family Malvaceae into nine subfamilies based on molecular, morphological, and biogeographical data: (1) Byttnerioideae, including tribes Byttnerieae, Lasiopetaleae and Theobromeae (all of which have cucullate petals) and Hermannieae; (2) Grewioideae, including most genera of former Tiliaceae; (3) Tilioideae, monogeneric in our analysis; (4) Helicteroideae, comprising most of the taxa previously included in Helictereae, plus Mansonia, Triplochiton (indicating that apocarpy evolved at least twice within Malvaceae) and possibly Durioneae; (5) Sterculioideae, defined by apetalous, apocarpous, usually unisexual flowers with androgynophores; (6) Brownlowioideae, circumscribed as in previous classifications; (7) Dombeyoideae, expanded to include Burretiodendron, Eriolaena, Pterospermum, and Schoutmia; (8) Bombacoideae, corresponding to former Bombacaceae (without Durioneae) but including Fremontodendreae  相似文献   

3.
Molecular studies indicate that Penaeaceae, Oliniaceae, and the monospecific families Alzateaceae and Rhynchocalycaceae form a clade of Myrtales. Of these four families, Penaeaceae have tracheids with vestured pits, whereas the others have septate fibers lacking vestures; all have vestured pits in vessels. Tracheid presence in Penaeaceae may be related to the arid South African habitats of the family. Presence of vestures on tracheids in families with vestured vessel pits is one indication that imperforate elements are tracheids and are conductive cells, whereas fiber-tracheids and libriform fibers are non-conductive. Tracheids occur widely in angiosperms and may be plesiomorphies or apomorphies. Combretaceae, the first branch of the Myrtales clade, has a great diversity of vesture features in vessels compared to the Penaeaceae alliance families. Alzatea has vestures that spread over the inside of the vessels, whereas in most taxa of the alliance, vestures are confined to the pit cavities and pit apertures. Vestures in the alliance tend to be globular in shape, and are bridged together by strands of wall material. Lignotubers and roots in Penaeaceae have vestures much like those in stems. Only a few species and genera (notably Alzatea) of the alliance have vesture features the pattern of which correlates with the current taxonomic system. Vestured pits should be viewed from the inside surface of vessels as well as the outer surface, and although sectional views of vestured pits are infrequent, they are very informative. Studies that explore diversity from one order or family to another are needed and offer opportunities for understanding the evolutionary significance of this feature.  相似文献   

4.
5.
NAIR, M. N. B. AND MOHAN RAM, H. Y., 1989. Vestured pits and vestured vessel member walls in some Indian dicotyledonous woods. The woods of 144 taxa belonging to 38 families of angiosperms were examined for vestured pits and vestured vessel member walls using scanning electron microscopy. Vestured pits are present in 66 taxa (belonging to ten families) and vestured vessel member walls occur in only six taxa (belonging to three families). In Ehretiaceae and Euphorbiaceae vestures are present only in certain vessel members. In Wrighlia tinctoria , perforation plates containing vestures have been observed in addition to the presence of vestured pits. A classification of vestured pits based on their morphology and distribution is proposed by us. In all the types of vestured pits, vestures are present on the margin of the outer pit aperture or on the pit chamber wall. Occasionally, they are present in the pit canal, on the margin and in the vicinity of the inner pit aperture and rarely over the inner walls of the vessel members. The functions of vestured pits are not clear, although several suggestions are made. Whether or not these structures affect wood processing is not presently understood. It appears that vestured pits and vestured vessel member walls have diagnostic rather than phylogenetic value.  相似文献   

6.
The monophyly of the group comprising the core malvalean families, Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae, was recently confirmed by molecular studies, but the internal structure of this clade is poorly understood. In this study, we examined sequences of the chloroplast ndhF gene (aligned length 2226 bp) from 70 exemplars representing 35 of the 39 putative tribes of core Malvales. The monophyly of one traditional family, the Malvaceae, was supported in the trees resulting from these data, but the other three families, as traditionally circumscribed, are nonmonophyletic. In addition, the following relationships were well supported: (1) a clade, /Malvatheca, consisting of traditional Malvaceae and Bombacaceae (except some members of tribe Durioneae), plus Fremontodendron and Chiranthodendron, which are usually treated as Sterculiaceae; (2) a clade, /Malvadendrina, supported by a unique 21-bp (base pair) deletion and consisting of /Malvatheca, plus five additional subclades, including representatives of Sterculiaceae and Tiliaceae, and Durionieae; (3) a clade, /Byttneriina, with genera traditionally assigned to several tribes of Tiliaceae, plus exemplars of tribes Byttnerieae, Hermannieae, and Lasiopetaleae of Sterculiaceae. The most striking departures from traditional classifications are the following: Durio and relatives appear to be more closely related to Helicteres and Reevesia (Sterculiaceae) than to Bombacaceae; several genera traditionally considered as Bombacaceae (Camptostemon, Matisia, Phragmotheca, and Quararibea) or Sterculiaceae (Chiranthodendron and Fremontodendron) appear as sister lineages to the traditional Malvaceae; the traditional tribe Helictereae (Sterculiaceae) is polyphyletic; and Sterculiaceae and Tiliaceae, as traditionally circumscribed, represent polyphyletic groups that cannot sensibly be maintained with their traditional limits for purposes of classification. We discuss morphological characters and conclude that there has been extensive homoplasy in characters previously used to delineate major taxonomic groups in core Malvales. The topologies here also suggest that /Malvatheca do not have as a synapormophy monothecate anthers, as has been previously supposed but, instead, may be united by dithecate, transversely septate (polysporangiate) anthers, as found in basal members of both /Bombacoideae and /Malvoideae. Thus, “monothecate” anthers may have been derived at least twice, independently, within the /Bombacoideae (core Bombacaceae) and /Malvoideae (traditional Malvaceae).  相似文献   

7.
The placement of a recently discovered South American monotypic genus,Pseudomonotes tropenbosii, in subfam.Monotoideae (Dipterocarpaceae) extends the geographical range of the subfamily from Africa to the Neotropics. Although morphological and anatomical evidence suggest similarities betweenPseudomonotes andMonotes, the close alliance of these two genera was questionable due to their disjunct distribution and a lack of phylogenetic analysis. In the present study, we reconstructed the phylogeny ofPseudomonotes and other putatively related taxa usingrbcL sequence data. The analysis ofrbcL sequences of 20 taxa belonging to 15 genera and eight families recovered a single most parsimonious tree. The genusSarcolaena (Sarcolaenaceae) formed a clade sister to the monophyleticDipterocarpaceae clade.Monotes andPseudomonotes formed a strongly supported group, sister to the monophyletic clade withPakaraimaea and the remaining Asiatic dipterocarp species studied. The study strongly supports the placement ofPseudomonotes within subfam.Monotoideae of theDipterocarpaceae.  相似文献   

8.
An examination of the external morphology of a recent collection from Araracuara, Colombia, has suggested that the plant belongs to the primarily Old World family Dipterocarpaceae. A study of the wood, bark, and pith anatomy of this new taxon,Pseudomonotes tropenbosii Londoño, Alvarez & Forero, was undertaken to help confirm its systematic affinities. Comparisons ofPseudomonotes with data from the literature and reference wood slides, coupled with the use of computer-aided identification keys, support the view that its closest relationships probably are within the family Dipterocarpaceae. Detailed anatomical comparisons have revealed thatPseudomonotes' relationships are most likely to be with the subfamily Monotoideae, comprised of the African generaMonotes A. DC. andMarquesia Gilg.  相似文献   

9.
A broad comparative analysis reveals that the inflorescences of coreMalvales, familiesSterculiaceae, Tiliaceae, Bombacaceae andMalvaceae, include characteristic repeating units. The basic repeating unit is called bicolor unit (afterTheobroma bicolor, where it was first observed). It is determinate and bears three bracts, one of which is invariably sterile, whereas the others subtend lateral cymes or single flowers. Through the demonstration of intermediate steps in closely related taxa the triad of bracts within a bicolor unit and the trimerous malvalean epicalyx are shown to be homologous. Various possibilities for an origin of the bicolor unit are discussed. Bicolor units are variously arranged to form complete inflorescences. In many taxa they are terminal on modules that comprise two (or fewer) prophylls. These modules may be arranged in elongated anthocladia or condensed sympodia, which in turn may constitute components of higher order inflorescence structures. The presence of the bicolor unit or its derivatives linksSterculiaceae, Tiliaceae, Bombacaceae andMalvaceae. It is absent from all other families included in a broader defined orderMalvales and represents one of the rare morphological synapomorphies of coreMalvales. Furthermore, inflorescence morphology provides characters of systematic significance for various taxa within coreMalvales.  相似文献   

10.
11.
Plant habitat associations are well documented in Bornean lowland tropical forests, but few studies contrast the prevalence of associations across sites. We examined habitat associations and community composition of Dipterocarpaceae trees in two contrasting Bornean lowland mixed dipterocarp forests separated by approximately 100 km: Andulau (uniform topography, lower altitudinal range, sandy soils) and Belalong (highly dissected topography, higher altitudinal range, clay‐rich soils). Dipterocarpaceae trees ≥ 1 cm diameter at breast height (dbh) were censused in 20‐m wide belt transects established along topographic gradients at each site. Dipterocarp density, evenness, species richness, and diversity were significantly higher at Andulau than Belalong. Significant site associations (with either Andulau or Belalong) were detected for 19 (52%) of the 37 dipterocarp species tested. Dipterocarpaceae community composition at Belalong correlated with soil nutrient concentrations as well as measures of vegetation and topographic structure, but community composition at Andulau correlated with fewer habitat variables. Within each site, dipterocarp density, species richness, and diversity were consistently higher on ridges than in slopes and valleys. Significant within‐site associations to topographic habitats were less common at Andulau (10% of species tested) than at Belalong (15%). We conclude that edaphic and other environmental factors influence dipterocarp community composition at a local scale, and are more important drivers of community structure in the more variable environment at Belalong. Species richness and diversity of dipterocarps on small plots, however, were higher at Andulau, suggesting that factors other than environmental heterogeneity contribute to contrasts in dipterocarp tree species richness at small scales.  相似文献   

12.
An analysis of rbcL sequence data for representatives of families of putative sapindalean/rutalean affinity identified a robust clade of core “sapindalean” taxa that is sister to representatives of Malvales. The constitution of this clade approximates the broad concept of Sapindales (sensu Cronquist). Five lineages within the order are recognized: a “rutaceae” clade (Rutaceae, Cneoraceae, Ptaeroxylaceae, Simaroubaceae sensu stricto, and Meliaceae); a “sapindaceae” clade (Sapindaceae, Aceraceae, and Hippocastenaceae); Anacardiaceae plus Burseraceae; Kirkiaceae; and Zygophyllaceae pro parte. Relationships among these groups were only weakly resolved, but there was no support for the recognition of the two more narrowly defined orders, Rutales and Sapindales sensu stricto. Several families that have previously been allied to Sapindales or Rutales show no affinity to the core sapindalean taxa identified with the molecular data, and are excluded from the order: viz. Akaniaceae, Bretschneideraceae, Conneraceae, Coriariaceae, Melianthaceae, Meliosmaceae, Physenaceae, Rhabdodrendraceae, Sabiaceae, Staphyleaceae, Stylobasiaceae, Surianaceae, and Zygophyllaceae sensu stricto.  相似文献   

13.
Previous analyses ofAsteropeia andPhysena have not compared the wood anatomy of these genera to those of Caryophyllales s.l. Molecular evidence shows that the two genera from a clade that is a sister group of the core Caryophyllales. Synapomorphies of theAsteropeia-Physena clade include small circular alternate pits on vessels, presence of vasicentric tracheids plus fiber-tracheids, presence of abaxial-confluent plus diffuse axial parenchyma, and presence of predominantly uniseriate rays. These features are analyzed with respect to habit and ecology of the two genera. Solitary vessels, present in both genera, are related to the presence of vasicentric tracheids. Autapomorphies in the two genera seem related to adaptations byPhysena as a shrub of moderately dry habitats (e.g., narrower vessel elements, abundant vasicentric tracheids, square to erect cells in rays) as compared to alternate character expressions that seem related to the arboreal habit and humid forest ecology ofAsteropeia. The functional significance of vasicentric tracheids and fiber-tracheids in dicotyledons is briefly reviewed in the light of wood anatomy of the two genera.  相似文献   

14.
The seeds of dipterocarp trees are the main food resources for many species of weevils, bark beetles and small moths; however, for most seed‐eating insects on dipterocarp tropical trees, seed utilization patterns remain poorly investigated. This study aimed to determine the fruit maturation stages at which eggs are laid by different insect seed predators feeding on the seeds or fruits of the following five dipterocarp species: Dipterocarpus globosus, Dryobalanops aromatica, Shorea beccariana, S. acuta and S. curtisii, which reproduced during the same period. We investigated the occurrence frequencies of the insect seed predators at various growth stages by collecting both unfallen and fallen fruit on several occasions during the period of seed/fruit maturation in a tropical rainforest in Borneo from September to December 2013. Weevils and bark beetles were the dominant insect seed predators of the five tree species. One or two weevil species of Alcidodes, Damnux and/or Nanophyes preyed on the seeds of each of the five tree species, and one bark beetle species, Coccotrypes gedeanus, preyed on the seeds of all five tree species. Many larvae, pupae and adults of each weevil species were found in pre‐dispersal (unfallen) fruit, whereas bark beetles at various growth stages were found in post‐dispersal (fallen) fruit. These results suggested that, among the dominant insect seed predators of the five dipterocarp species, weevil species oviposit on pre‐dispersal fruit and begin their larval growth before seed dispersal, whereas the oviposition and larval development of bark beetle species occurs in post‐dispersal fruit.  相似文献   

15.
Dipterocarpaceae is the dominant family of Southeast Asia's climax tropical rain forest region, and it contains the region's most important commercial timber species. A molecular phylogeny of the Dipterocarpaceae subfamily Dipterocapoideae was constructed using restriction fragment length polymorphisms of polymerase chain reaction-amplified specific genes in chloroplast DNA. A total of 141 site changes were detected among ten genera and 30 species in 11 different genes: rbcL, psbA, psbD, rpoB, rpoC, petB, atpH, 16S, psaA, petA and trnK. Phylogenetic trees constructed by Wanger parsimony and neighbor-joining methods, using Upuna as the outgroup, displayed five monophytelic groups that included Upuna: HopeaShorea-Parashorea-Neobalanocarpus; Dryobalanops; Dipterocarpus; Anisoptera-Vatica-Cotylelobium; and Upuna. The phylogenetic trees clearly separate species with two different base chromosome numbers: the first group is x=7, and the other is x=11. The x=7 group is thought to be in a synapomorphic character state. Parashorea lucida is a sister to most Shorea species. Neobalanocarpus heimii and Hopea from a clade of a sister to two Shorea species, and Cotylelobium and Vatica are closely related species. Our conclusions agree with a phylogeny derived from wood anatomy data analysis, and with Symington's and Ashton's taxonomic classifications.The raw data of the PCR-RFLP analysis can be obtained from the authors  相似文献   

16.
At least four hypotheses regarding the phylogenetic relationships within the Galaginae have been proposed, based on features of karyology, morphology, and behaviour, but these hypotheses share few common elements. Here we investigate erythrocytic allozymes as potential phylogenetic markers, and subject our results to a cladistic analysis. Our study offers little support for the previous models, but suggests instead that the greater galagos andG. alleni form a clade since they share character states for eight of the ten systems examined. The two lesser galago species could not be distinguished using these enzymes, and character states common to them and theallenicrassicaudatus-garnettii clade were found in only four systems. Our resultant cladogram accords well with the rather scant galagine fossil record. The data also show strong concordance with results obtained using highly repeated DNA sequences, which indicate that the galagos form a close-knit genetic group, while the Malagasy lemurids show considerably more inter-taxic variation.  相似文献   

17.
A floristic analysis of the lowland dipterocarp forests of Borneo   总被引:4,自引:0,他引:4  
Aim To (1) identify floristic regions in the lowland (below 500 m a.s.l.) tropical dipterocarp rain forest of Borneo based on tree genera, (2) determine the characteristic taxa of these regions, (3) study tree diversity patterns within Borneo, and (4) relate the floristic and diversity patterns to abiotic factors such as mean annual rainfall and geographical distance between plots. Location Lowland tropical dipterocarp rain forest of Borneo. Methods We used tree (diameter at breast height ≥ 9.8 cm) inventory data from 28 lowland dipterocarp rain forest locations throughout Borneo. From each location six samples of 640 individuals were drawn randomly. With these data we calculated a Sørensen and Steinhaus similarity matrix for the locations. These matrices were then used in an UPGMA clustering algorithm to determine the floristic relations between the locations (dendrogram). Principal coordinate analysis was used to ordinate the locations. Characteristic taxa for the identified floristic clusters were determined with the use of the INDVAL method of Dufrene & Legendre (1997) . Finally, Mantel analysis was applied to determine the influence of mean annual rainfall and geographical distance between plots on floristic composition. Results A total of 77 families and 363 genera were included in the analysis. On average a random sample of 640 trees from a lowland dipterocarp forest in Borneo contains 41.6 ± 3.8 families and 103.0 ± 12.7 genera. Diversity varied strongly on local scales. On a regional scale, diversity was found to be highest in south‐east Borneo and central Sarawak. The most common families were Dipterocarpaceae (21.9% of trees) and Euphorbiaceae (12.2% of trees). The most common genera were Shorea (12.3% of trees) and Syzygium (5.0% of trees). The 28 locations were clustered in geographically distinct floristic regions. This was related to the fact that floristic similarity depended strongly on the geographical distance between plots and similarity in mean annual rainfall. Conclusions We identified five main floristic regions within the lowland dipterocarp rain forests of Borneo, each of which had its own set of characteristic genera. Mean annual rainfall is an important factor in explaining differences in floristic composition between locations. The influence of geographical distance on floristic similarity between locations is probably related to the fact that abiotic factors change with distance between plots. Borneo's central mountain range generally forms an effective dispersal barrier for the lowland tree flora. Diversity patterns in Borneo are influenced by the mid‐domain effect, habitat size and the influence of past climatic changes (ice ages during the Pleistocene).  相似文献   

18.
Wood properties were measured for trees in lowland dipterocarp forests in West Kalimantan. In 1993 and 1994, 353 samples of 286 species were collected from trunk base of trees of approximately 5 cm in diameter, and the specific gravities (SG: oven dry weight/fresh volume) and water contents of wood including bark were measured. The SG of each species ranged from 0.21 to 0.84, and the mean ± SD was 0.53 ± 0.13. The wide range of SG suggests that the forest had a high diversity in wood properties. The most dominant and diversified genus in this area was Shorea, and the SG of 15 species varied from 0.21 to 0.71. The range covered SG of pioneer (six Macaranga, 0.29–0.43) and small trees in primary forests (nine Eugenia and 10 Xanthophyllum, 0.55–0.77). The SG average for tree species of secondary forests of 2–6 years old was 0.31. It was significantly smaller than that of primary forests (0.58). In a primary dipterocarp forest plot, light-wood species grew faster in diameter than heavy-wood species. Water content ranged from 0.26 to 0.76. Heavy wood had low water content. Among light-wood species, some (Shorea, Artocarpus) had low water contents and others (Ficus) had high water contents. Some riverine trees also had high water contents. These wood properties appear strongly related to the life history of trees and successional stage.  相似文献   

19.
To search for cytogenetic signatures that can help to clarify evolutionary affinities among the five orders within the Euarchontoglires clade, we focused on associations of conserved syntenic blocks that have been accumulated in the karyotypes of Primates (Strepsirhini and Haplorhini), five families of Rodentia, Scandentia (Tupaia belangeri), Dermoptera (Galeopterus variegatus) and Lagomorpha (Oryctolagus cuniculus). We examined available chromosome painting data to identify conserved chromosomes and chromosomal segments, and syntenic associations likely to have characterized the ancestral eutherian karyotype. The data set includes 161 characters that have been subjected to a concatenated analysis using maximum parsimony (MP) and Bayesian inference (BI). The phylogenetic pattern recovered is generally consistent with reconstructions based on molecular and morphological data (particularly with respect to higher systematic groupings), but there are several anomalies (e.g., in the position of the lagomorphs). Both MP and BI topologies have weak statistical support, as a consequence of the high number of autapomorphic and homoplastic character states that have evolved during the history of the clade. The vast majority of derived associations are located on the terminal portions of the branches, and very few can be identified to support deeper divergences in the tree, indicating that chromosomal structures are far more fluid that was previously recognized. The high levels of homoplasy reflected in our data suggest that the number of possible syntenic character states is limited by chromosomal structures, and the same associations occur repeatedly.  相似文献   

20.
We measured the terminal velocity of helicopter‐like fruit from the Dipterocarpaceae family and present a model predicting the terminal velocities for all dipterocarp species in the Malesiana region. A ballistic model of seed dispersal using the observed terminal velocities predicted dispersal distances of 17–77 m under normal atmospheric conditions. These data are of applied use in parametizing models of species coexistence, forest regeneration and habitat connectivity in Southeast Asian tropical forests.  相似文献   

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