Placement of Kuhlmanniodendron Fiaschi & Groppo in Lindackerieae (Achariaceae, Malpighiales) confirmed by analyses of rbcL sequences, with notes on pollen morphology and wood anatomy

The phylogenetic placement of Kuhlmanniodendron Fiaschi & Groppo (Achariaceae) within Malpighiales was investigated with rbcL sequence data. This genus was recently created to accommodate Carpotroche apterocarpa Kuhlm., a poorly known species from the rainforests of Espírito Santo, Brazil. One rbcL sequence was obtained from Kuhlmanniodendron and analyzed with 73 additional sequences from Malpighiales, and 8 from two closer orders, Oxalidales and Celastrales, all of which were available at Genbank. Phylogenetic analyses were carried out with maximum parsimony and Bayesian inference; bootstrap analyses were used in maximum parsimony to evaluate branch support. The results confirmed the placement of Kuhlmanniodendron together with Camptostylus, Lindackeria, Xylotheca, and Caloncoba in a strongly supported clade (posterior probability = 0.99) that corresponds with the tribe Lindackerieae of Achariaceae (Malpighiales). Kuhlmanniodendron also does not appear to be closely related to Oncoba (Salicaceae), an African genus with similar floral and fruit morphology that has been traditionally placed among cyanogenic Flacourtiaceae (now Achariaceae). A picrosodic paper test was performed in herbarium dry leaves, and the presence of cyanogenic glycosides, a class of compounds usually found in Achariaceae, was detected. Pollen morphology and wood anatomy of Kuhlmanniodendron were also investigated, but both pollen (3-colporate and microreticulate) and wood, with solitary to multiple vessels, scalariform perforation plates and other features, do not seem to be useful to distinguish this genus from other members of the Achariaceae and are rather common among the eudicotyledons as a whole. However, perforated ray cells with scalariform plates, an uncommon wood character, present in Kuhlmanniodendron are similar to those found in Kiggelaria africana (Pangieae, Achariaceae), but the occurrence of such cells is not mapped among the angiosperms, and it is not clear how homoplastic this character could be.     

蜈蚣草导管分子的观察

对蜈蚣草(Pteris vittata)叶轴中导管分子进行了观察。结果表明,其导管分子十分细长,均具长、且十分倾斜的梯状穿孔板,穿孔没有纹孔膜的残余,与侧壁上的梯纹纹孔有着明显的差异。     

Fossil bananas (Musaceae): Ensete oregonense sp. nov. from the Eocene of western North America and its phytogeographic significance

Fossil seeds of Ensete, a genus presently native to Asia and Africa, have been recovered from the middle Eocene of Oregon, confirming the presence of Musaceae in the North American Tertiary. The seed of Ensete oregonense sp. nov. is operculate, with a well-defined micropylar collar, a pronounced chalazal chamber, and a wide hilar cavity. A survey of seed morphology in extant Zingiberales provides characters for distinguishing Musaceae from other families of the order, furnishes criteria for distinguishing the three extant genera of Musaceae (Musa, Ensete and Musella), and facilitates critical assessment of fossil seed remains. “Musa” cardiosperma Jain from the Cretaceous/Tertiary boundary Deccan Series of India is excluded from Musaceae (although retained in Zingiberales) on the basis of fruit and seed characters, including the lack of laticifers and absence of a chalazal chamber. We reexamined the musaceous seeds from Colombia that previously were described as Tertiary fossils (Musa enseteformis Berry, 1925) and now believe that they are recent, nonfossil remains, evidently from Ensete ventricosum, which is grown in the region where the specimens were originally obtained. In addition, a reputed fossil banana fruit from the Cretaceous of Colombia was reexamined and determined to be a concretion of nonbiological origin. Ensete oregonense is significant therefore, as the first unequivocal fossil record of Ensete and of Musaceae. Although the Musaceae are currently native only to the Old World tropics, this discovery establishes that the family was present in North America about 43 million years ago.     

Multilayered structure of tension wood cell walls in Salicaceae sensu lato and its taxonomic significance

Salicaceae have been enlarged to include a majority of the species formerly placed in the polyphyletic tropical Flacourtiaceae. Several studies have reported a peculiar and infrequently formed multilayered structure of tension wood in four of the tropical genera. Tension wood is a tissue produced by trees to restore their vertical orientation and most studies have focused on trees developing tension wood by means of cellulose‐rich, gelatinous fibres, as in Populus and Salix (Salicaceae s.s.). This study aims to determine if the multilayered structure of tension wood is an anatomical characteristic common in other Salicaceae and, if so, how its distribution correlates to phylogenetic relationships. Therefore, we studied the tension wood of 14 genera of Salicaceae and two genera of Achariaceae, one genus of Goupiaceae and one genus of Lacistemataceae, families closely related to Salicaceae or formerly placed in Flacourtiaceae. Opposite wood and tension wood were compared with light microscopy and three‐dimensional laser scanning confocal microscopy. The results indicate that a multilayered structure of tension wood is common in the family except in Salix, Populus and one of their closest relatives, Idesia polycarpa. We suggest that tension wood may be a useful anatomical character in understanding phylogenetic relationships in Salicaceae. Further investigation is still needed on the tension wood of several other putatively close relatives of Salix and Populus, in particular Bennettiodendron, Macrohasseltia and Itoa.     

蜈蚣草导管分子的观察

对蜈蚣草(Pteris vittata)叶轴中导管分子进行了观察。结果表明,其导管分子十分细长,均具长、且十分倾斜的梯状穿孔板,穿孔没有纹孔膜的残余,与侧壁上的梯纹纹孔有着明显的差异。     

First record of Cercidiphylloxylon(Cercidiphyllaceae)from the Palaeocene of Fushun,NE China

Cercidiphylloxylon spenceri(Brett)Pearson is described from the Lizigou Formation,Palaeocene in China.The growth rings are distinct; pores are diffuse,solitary,with somewhat angular outlines in cross section;vessel elements long with long scalariform perforation plates; intervessel pitting is opposite to scalariform; fibertracheids are present; axial parenchyma is scarce; rays are mostly biseriate and heterogeneous.All wood characters of the fossil specimen fall into the range of those of extant Cercidiphyllum(Cercidiphyllaceae).The finding is one of the earliest fossil wood records of Cercidiphyllaceae.     

The rise of bilaterians: a few closing comments

Freeman (2009, 2010) argue that Ediacara “faunas” should be analysed using “promorphologies” of extant animals, that rangeomorphs and erniettomorphs are colonial animals, and that Vernanimalcula is a bilaterian animal. Recent studies of Ediacara fossils have provided multiple lines of evidence that these fossils represent a sample of phylogenetically diverse marine organisms that included crown-group animals as well as stem-group animals and non-animals. Thus, it is inappropriate to analyse Ediacara fossils using “promorphologies” of extant animals. The interpretation of rangeomorphs and erniettomorphs as colonial animals is inconsistent with the functional morphologies of these Ediacara fossils. Although Vernanimalcula is a fossil, the purported germ layers of Vernanimalcula are of diagenetic origin and there is no morphological evidence in support of its bilaterian affinity.     

First record of Cercidiphylloxylon (Cercidiphyllaceae) from the Palaeocene of Fushun,NE China

Abstract Cercidiphylloxylon spenceri (Brett) Pearson is described from the Lizigou Formation, Palaeocene in China. The growth rings are distinct; pores are diffuse, solitary, with somewhat angular outlines in cross section; vessel elements long with long scalariform perforation plates; intervessel pitting is opposite to scalariform; fiber‐tracheids are present; axial parenchyma is scarce; rays are mostly biseriate and heterogeneous. All wood characters of the fossil specimen fall into the range of those of extant Cercidiphyllum (Cercidiphyllaceae). The finding is one of the earliest fossil wood records of Cercidiphyllaceae.     

Woods of the Vitaceae—fossil and modern

Vitaceoxylon tiffneyi gen. et sp. nov. and Vitaceoxylon carlquistii sp. nov. from the Middle Eocene Clarno formation are the oldest known woods with characteristics of the Vitaceae. They are characterized by a tendency to two diameter classes of vessels, wide and tall rays, and a high proportion of their area is vessel. Other characteristics include septate imperforate elements, scanty paratracheal to vasicentric parenchyma, idioblasts in the rays, crowded alternate intervessel pitting, and vessel-parenchyma pits with reduced borders. Wood anatomy of the major extant genera of Vitaceae was examined and compared to the fossils. Features of secondary xylem useful for distinguishing between genera in the Vitaceae include vessel size and arrangement (two distinct diameter classes or not, radial multiples or tendency to tangential multiples), vessel pitting (scalariform or alternate), crystal type (prismatic, druses, and/or raphides) and location (in chambered parenchyma and/or ray parenchyma), cambial variants (present or absent). Wood anatomy supports the proposed close relationship of Cissus to Cayratia. Pronounced vessel dimorphism occurs in temperate Vitaceae; cambial variant structure occurs in tropical Vitaceae. Despite their conformity with Vitaceae at the family level, the two fossil woods are not comparable to any one extant genus. This contrasts with the Vitaceae seeds from Clarno, all of which are referable to extant genera. Two new combinations for fossil woods of the Vitaceae are proposed: Vitaceoxylon ampelopsoides (Schönfeld) comb. nov., and Vitaceoxylon megyazoense (Greguss) comb. nov.     

Foliar anatomy of neotropical Salicaceae: potentially useful characters for taxonomy

The taxonomy of neotropical Salicaceae, a family that now includes the majority of the former Flacourtiaceae, has been problematic, especially because they display very diverse morphology and have several characteristics in common with many other families. Recent phylogenetic studies have proposed substantial changes at both family and generic levels. Considering the importance of anatomy as an aid for taxonomy, the gathering of anatomical data for the family is fundamental to help clarify the taxonomic problems. Leaves belonging to Abatia americana (four samples), Banara brasiliensis (2), Casearia arborea (4), C. decandra (5), C. gossypiosperma (2), C. obliqua (1), C. sylvestris (3), C. ulmifolia (3), Prockia crucis (3), and Xylosma prockia (4) and the closely related Carpotroche brasiliensis (3) from Achariaceae, were studied by standard microscopy techniques. The leaves were anatomically described, emphasizing their differences and similarities. Similar characters for the neotropical Salicaceae (former Flacourtiaceae) and Salicaceae strictu sensu were recognized, such as the presence of salicoid leaf teeth, brachyparacytic stomata, secondary growth of the petiole, abundance of crystals, collateral and arch-shaped vascular system at the midrib, and sclerenchyma accompanying the bundles. These data demonstrate that leaf anatomy can provide evidence to assist with the taxonomy of Salicaceae, at family, generic, and specific levels.     

Fossils and seed plant phylogeny reanalyzed

In a cladistic analysis of Recent seed plants, Loconte and Stevenson (1990) obtained results that conflict with our 1986 analysis of both extant and fossil groups and argued that fossil data had led us to incorrect conclusions. To explore this result and the general influence of fossils on phylogeny reconstruction, we assembled new “Recent” and “Complete” (extant plus fossil) data sets incorporating new data, advances in treatment of characters, and those changes of Loconte and Stevenson that we consider valid. Our Recent analysis yields only one most parsimonious tree, that of Loconte and Stevenson, in which conifers are linked with Gnetales and angiosperms (anthophytes), rather than with Ginkgo, as in our earlier Recent and Complete analyses. However, the shortest trees derived from our Complete analysis show five arrangements of extant groups, including that of Loconte and Stevenson and our previous arrangements, suggesting that the result obtained from extant taxa alone may be misleading. This increased ambiguity occurs because features that appear to unite extant conifers and anthophytes are seen as convergences when fossil taxa are interpolated between them. All trees found in the Complete analysis lead to inferences on character evolution that conflict with those that would be drawn from Recent taxa alone (e.g., origin of anthophytes from plants with a “seed fern” morphology). These results imply that conclusions on many aspects of seed plant phylogeny are premature; new evidence, which is most likely to come from the fossil record, is needed to resolve the uncertainties.     

猫儿屎导管分子穿孔板新类型的发现

运用扫描电镜(SEM)对木通科(Lardizabalaceae)猫儿屎属(Decaisnea Hook.f. & Thoms.)植物猫儿屎[Decaisnea insignis (Griff.) Hook.f.et Thoms.]茎的次生木质部导管分子进行观察,以期为该属的系统演化提供依据.结果表明,猫儿屎的导管分子具有多个穿孔板,端壁穿孔板除了梯状以外,还有梯-网、梯-网-单、梯-单混合穿孔板等类型;侧壁穿孔板包括梯状、网状、梯-网混合状穿孔板;穿孔板上纹孔膜的残余有丝状、网状和片状.同时对导管分子的长度、宽度及端壁倾斜度等特征进行统计,并讨论了木通科各类群的穿孔板特征.     

Phylogeny of the eudicot order Malpighiales: analysis of a recalcitrant clade with sequences of the petD group II intron

Malpighiales are one of the most diverse orders of angiosperms. Molecular phylogenetic studies based on combined sequences of coding genes allowed to identify major lineages but hitherto were unable to resolve relationships among most families. Spacers and introns of the chloroplast genome have recently been shown to provide strong signal for inferring relationships among major angiosperm lineages and within difficult clades. In this study, we employed sequence data of the petD group II intron and the petB-petD spacer for a set of 64 Malpighiales taxa, representing all major lineages. Celastrales and Oxalidales served as outgroups. Sequence alignment was straightforward due to frequent microstructural changes with easily recognizable motifs (e.g., simple sequence repeats), and well defined mutational hotspots. The secondary structure of the complete petD intron was calculated for Idesia polycarpa as an example. Domains I and IV are the most length variable parts of the intron. They contain terminal A/T-rich stem-loop elements that are suggested to elongate independently in different lineages with a slippage mechanism earlier reported from the P8 stem-loop of the trnL intron. Parsimony and Bayesian analyses of the petD dataset yielded trees largely congruent with results from earlier multigene studies but statistical support of nodes was generally higher. For the first time a deep node of the Malpighiales backbone, a clade comprising Achariaceae, Violaceae, Malesherbiaceae, Turneraceae, Passifloraceae, and a Lacistemataceae–Salicaceae lineage received significant statistical support (83% JK, 1.00 PP) from plastid DNA sequences.     

Phylogenetic Distribution and Identification of Fin-winged Fruits

Fin-winged fruits have two or more wings aligned with the longitudinal axis like the feathers of an arrow, as exemplified by Combretum, Halesia, and Ptelea. Such fruits vary in dispersal mode from those in which the fruit itself is the ultimate disseminule, to schizocarps dispersing two or more mericarps, to capsules releasing multiple seeds. At least 45 families and more than 140 genera are known to possess fin-winged fruits. We present an inventory of these taxa and describe their morphological characters as an aid for the identification and phylogenetic assessment of fossil and extant genera. Such fruits are most prevalent among Eudicots, but occur occasionally in Magnoliids (Hernandiaceae: Illigera) and Monocots (Burmannia, Dioscorea, Herreria). Although convergent in general form, fin-winged fruits of different genera can be distinguished by details of the wing number, texture, shape and venation, along with characters of persistent floral parts and dehiscence mode. Families having genera with fin-winged fruits and epigynous perianth include Aizoaceae, Apiaceae, Araliaceae, Asteraceae, Begoniaceae, Burmanniaceae, Combretaceae, Cucurbitaceae, Dioscoreaceae, Haloragaceae, Lecythidiaceae, Lophopyxidaceae, Loranthaceae, and Styracaceae. Families with genera having fin-winged fruits and hypogynous perianth include Achariaceae, Brassicaceae, Burseraceae, Celastraceae, Cunoniaceae, Cyrillaceae, Fabaceae, Malvaceae, Melianthaceae, Nyctaginaceae, Pedaliaceae, Polygalaceae, Phyllanthaceae, Polygonaceae, Rhamnaceae, Salicaceae sl, Sapindaceae, Simaroubaceae, Trigoniaceae, and Zygophyllaceae. This survey has facilitated the identification of fossil winged fruits such as Combretaceae and Araliaceae in the late Cretaceous of western North America and provides additional evidence toward the identification of various Cenozoic fossils including Brassicaceae, Fabaceae, Polygonaceae, Rutaceae, and Sapindaceae.     

八属木兰科植物木材导管分子的比较解剖

本文比较描述了我国木兰科木莲属、华盖木属,长蕊木兰属,拟单性木兰属,合果木属,观光木属,香木兰属和鹅掌楸属等8属22种植物次生质部的导管分子,这些导管分子的长度和宽度都有差异,木莲属的所有种中都有具梯状穿孔板的导管分子,但在少数种中偶然可见到单穿孔,在其它属中,也都有具梯状穿孔板的导管分子,仅在观光木属中偶见的单穿孔,木莲属大多数种均无螺纹加厚,仅桂南木莲一种除外,另外,除了合果木属和鹅掌楸无螺纹加厚外,其余均有此种加厚,各属的导管分子还存在其它一些差异。     

Spiders in Upper Cretaceous Amber from New Jersey (Arthropoda: Araneae)

The oldest described fossils of the extant spider families Segestriidae, Oonopidae, Oecobiidae, Dictynidae and Linyphiidae, previously known from the Tertiary, are presented from Upper Cretaceous amber of New Jersey. The third and oldest known specimen of the fossil spider family Lagonomegopidae is also described and provides further palaeontological evidence of a common Laurasian fauna. The extant genera Segestria and Oecobius are taken back a further 52 and 69–74 myr respectively in the fossil record. These fossils predict the presence of the Caponiidae, Tetrablemmidae, Orsolobidae, Dysderidae, Hersiliidae, Eresidae, Pimoidae, Scytodoidea s.l. , cyatholipoids, theridioids and symphytognathoids in the Cretaceous. They also extend the known geological range of extant spider families through and beyond the end–Cretaceous extinction. This event, which affected numerous marine and some terrestrial organisms, probably had little effect on the Araneae.     

KLEINODENDRON AND XYLEM ANATOMY OF CLUYTIEAE (EUPHORBIACEAE)

Kleinodendron, a new genus of Euphorbiaceae, was assigned by Smith and Downs to the tribe Cluytieae. A xylem anatomical survey indicates that there are no objections to this placement. Woods of Cluytieae are diverse but may be characterized generally by having pores which average less than 80 μ in diameter and which are well divided between solitary and radial multiple distributions in the same species; simple vessel perforations; alternate intervascular pitting; fiber-tracheids and libriform wood fibers; exclusively uniseriate, or uniseriate and biseriate heterocellular vascular rays in the same species; uniseriate “bridges” linking superposed biseriate ray segments; diffuse, diffuse-in-aggregates, and scanty vasicentric axial parenchyma, sometimes in the same species; and crystal rhomboids. That Microdesmis and Pogonophora diverge sharply from these generalizations in having scalariform vessel perforations and broad vascular rays, is an indication that they may not be closely related to other genera in Cluytieae.     

Fossil forms of Amentiferae

Review of the procedures used in determining fossil plant organs indicates that the many Cretaceous records of extant genera of “Amentiferae” based on leaves should be rejected as theoretically unreliable. Palynological data, in combination with some valid megafossil data, indicate that most recognizable members of “Amentiferae” are no older than the later part of the Late Cretaceous. Juglandales appear to be derivatives of the ancient Normapolles complex and unrelated to other “Amentiferae.” A preliminary account of some of the comparative foliar morphology of extant “Amentiferae” indicates that some—particularly Betulaceae and Fagaceae—are closely related to Hamamelidales but that other families—notably Rhoipteleaceae, Juglandaceae, Didymelaceae, and Leitneriaceae—are unrelated to this order.