Ovules and Seeds in Subfamily Phyllanthoideae (Euphorbiaceae): Structure and Systematic Implications

Lingelsheimia ) are distinct from the rest of the subfamily in having a thick inner integument (over six cells thick), an exotegmen composed of cuboidal cells (type II), and vascular bundles in the outer integument and, as molecular evidence also suggests, should be transferred to a separate family Putranjivaceae. Hymenocardieae (Didymocistus and Hymenocardia), whose positions have been controversial, are monophyletic in sharing endotestal seeds with a collapsed exotegmen which is unknown elsewhere in Euphorbiaceae. The genera seem to require separation from the Euphorbiaceae. In addition, a morphological heterogeneity of the two large genera Cleistanthus and Phyllanthus, as well as of tribe Antidesmeae subtribe Scepinae were also discussed. Received 20 October 2000/ Accepted in revised form 14 January 2001     

PATTERNS OF STAMEN VASCULATURE IN THE ARACEAE

A survey of the three-dimensional organization of stamen vasculature in 100 genera and over 350 species of Araceae was made using clearings. The Araceae exhibit highly varied stamen vasculature, with three main patterns: 1) vascular bundles unbranched, 1–3 per stamen, 2) forked bundles in some or all stamens, 3) anastomosing vascular systems with several to many bundles entering a single stamen. Three major groups of taxa in the family can be recognized on the basis of their predominant pattern of stamen vasculature. Virtually all genera with bisexual flowers (most Pothoideae, Monsteroideae, Calloideae, Lasieae) have unbranched bundles, one per stamen, except two to three in some species of Holochlamys, Spathiphyllum, and Scindapsus. Forked stamen bundles are virtually restricted to and occur nearly throughout the monoecious Lasioideae, Philodendroideae, Colocasioideae and among certain Aroideae (sensu Engler), including tribes Arophyteae, Spathicarpeae (Asterostigmateae) and Protareae. No forked bundles were found in tribe Areae (Aroideae), except Theriophonum indicum or any Araceae with bisexual flowers, except two species of Cyrtosperma. Anastomosing systems are virtually limited to members of tribe Areae with larger stamens, such as Arum, Helicodiceros, Eminium and Dracunculus species. A similar pattern occurs in some Amorphophallus, but other patterns occur as well. The distributions of forked bundles and anastomosing systems in the family are notable because they are both highly congruent with Philodendroideae-Colocasioideae, and Aroideae, respectively, in Grayum's new system for the family. Virtually all of the genera with forked bundles are grouped together in the Philodendroideae-Colocasioideae. All of the genera with anastomosing systems are in the Areae, including the complex and variable Amorphophallus, which has an uncertain systematic placement.     

VASCULAR PATTERNS IN STEMS OF THE CYCLANTHACEAE

A survey was made of the distribution of stem vascular bundles in representatives of ten genera of the tropical monocotyledonous family Cyclanthaceae. Films of series of serial transverse sections were used to reconstruct the stem vasculature. Each leaf trace, followed in a basipetal direction from its level of insertion at the stem periphery, describes an obliquely downward course, initially contacting from 1 to 4 (or more) existing axial bundles. The associated bundles form a compound vascular bundle in which the original bundles initially remain discrete, most commonly in a tetrapolar arrangement, with four separate strands. Followed further in the basipetal direction, the strands eventually fuse partly or completely, usually to form a collateral or amphivasal axial bundle which participates in a new structural cycle. Quantitative variation between different taxa includes a simple pattern in Ludovia, in which only bipolar bundles are developed. More elaborate forms have multipolar bundles with more than four separate strands. A systematically useful observation is that stem vasculature in Cyclanthus, representing the subfamily Cyclanthoideae, does not differ significantly from that in subfamily Carludovicoideae although there are some distinctive structural features.     

The phylogenetic significance of the carpophore in Apiaceae

Background and aims

Fruit structural characters have traditionally been important in the taxonomy of the family Apiaceae. Previous investigations using a limited number of taxa have shown that the carpophore may be especially useful in helping to circumscribe subfamily Azorelloideae. The present study examines, for the first time, carpophore structure in 92 species from 43 genera, representing all subfamilies of Apiaceae, and including all genera assigned to subfamily Azorelloideae. Phylogenetic interpretations are made for the first time, using all available information, and a standard terminology is proposed to describe the various character states found in carpophores.

Methods

Carpophore structure was studied in detail using light microscopy.

Key Results

Carpophores, when present, may be categorized into two main groups (B and C) based mainly on the arrangement of the vascular bundles in transverse section, and further divided into six sub-types according to the length of the carpophore (short in B1 and C1) and whether they are entire (B1–B3 and C1) or bifurcate (B4 and C2). Free carpophores are absent in subfamily Mackinlayoideae, and in tribes Lichtensteinieae and Phlyctidocarpeae, which have two opposite vascular bundles (Group A). Entire carpophores with one or two vascular bundles, or bifurcate carpophores with lateral vascular bundles (arranged side by side within the commissural plane), are the main types characterizing Azorelloideae. The short, hygroscopic carpophores found in Choritaenia are unique in Apiaceae and provide additional evidence for the exclusion of this genus from Azorelloideae. Carpophore type C2 is typical for most Apioideae sensu lato (exceptions are, for example, Arctopus and Alepidea, which have type B2).

Conclusions

A single carpophore and ventral vascular bundles not forming free carpophores are proposed to be the ancestral conditions in Apiaceae, while bifurcate carpophores with opposite vascular bundles are the derived state, present in most Apioideae. Secondary reductions seem to have occurred in several unrelated lineages in all major groups, e.g. many Azorelloideae, several protoapioids (including nearly all members of the tribe Saniculeae) and 29 euapioid genera (e.g. some Oenantheae).     

Trichome anatomy of the Saxifragaceae S. l.from the southern hemisphere

Trichome anatomy was examined by light and scanning electron microscopy in 25 genera of Engler's Saxifragaceae from the southern hemisphere. Four principal categories of trichome were recognized: (1) multiseriate with a glandular head; (2) uniseriate with a glandular head; (3) uniseriate, eglandular; (4) unicellular, eglandular. The shape of eglandular hairs ranges from erect to sickle-shaped to T-shaped. The main taxonomic conclusions are as follows: (a) Vahlia should be excluded from tribe Saxifrageae and a possible relationship with Montinia investigated; (b) Francoa and Tetilta (tribe Francoeae) are closely related, although the relationship of the tribe to its parent subfamily, Saxifragoideae, is unclear; (c) Eremosyne is probably allied to the Escallonioideae; (d) subfamily Brexioideae is heterogeneous in trichome anatomy, but the relationships of its constituent genera remain problematic; (e) subfamily Escallonioideae is heterogeneous in trichome anatomy, although similarities between and within the constituent tribes do exist. Thus Cuttsia and Abrophyllum form a natural group (tribe Cuttsieae), to which Carpodetus (tribe Argophylleae) may also be related; similarity in trichome anatomy between Argophyttum and Corokia (tribe Argophylleae) is substantiated by an extensive survey of all the species, and the data tentatively suggest a possible hydrangeoid affinity for these two genera.Forgesia (tribe Forgesieae) is shown to possess the same kind of hairs in the flowers as Quintinia (tribe Escallonieae), and Choristylis (tribe Forgesieae) is shown to be remarkably similar to Escallonia, prompting a suggestion that the two tribes be merged. Trichome data support the inclusion of Anopterus and Polyosma in their own tribes, although their wider affinities remain unclear. Doubts about the inclusion of the glabrous Tribeles in Escallonioideae are expressed. Our material of the following additional genera Lepuropetalon, Tetracarpaea and Brexia was glabrous and little comment could be made about them.     

PETIOLE VASCULARIZATION OF LUXEMBURGIEAE (OCHNACEAE)

Sections through the petiole and blade midrib and sometimes the node of 20 genera of Luxemburgieae have revealed variations in vascularization which appear significant in determining the phylogeny of the tribe. The most primitive group are the pentacarpellate genera; they are characterized by multilacunar nodes and numerous leaf traces which mostly remain free to the leaf apex; some of these become medullary bundles in the petiole. The more advanced bicarpellate genera have multilacunar or trilacunar nodes, still with many leaf traces in the case of Wallacea. These traces fuse in the blade, resulting in a siphonostele. The most advanced genera are tricarpellate and have trilacunar nodes with only three traces; these traces fuse, forming a siphonostele. There are no medullary bundles present in this last group of genera.     

COMPARATIVE ANATOMY OF CHAROPHYTA: II. THE AXIAL NODAL COMPLEX—AN APPROACH TO THE TAXONOMY OF LAMPROTHAMNIUM12

The present investigation on the axial nodes of 32 taxa belonging to the genera Chara, Lamprothamnium, Nitellopsis, Nitella and Tolypella confirms previous reports that within a given taxonomic group the structure of the main axial nodal complex is highly consistent. Besides the genera Nitella and Tolypella of the tribe Nitelleae, Lamprothamnium of the tribe Chareae is the only genus in which the central cells of the main axial nodal complex subdivide. This anatomical feature of the main axial nodal complex thus clearly separates the genus Lamprothamnium from the other genera of the tribe chareae. In 2 controversial species of the genus Chara, C. hornemannii and C. buckellii, the present study reveals that the central cells of their main axial nodal complexes do not subdivide. The transfer of these two species to the genus Lamprothamnium by Daily (1) may not be appropriate.     

New insights in Salicornia L. and allied genera (Chenopodiaceae) inferred from nrDNA sequence data

A phylogenetic analysis was performed based on ITS DNA sequences of fourteen samples from different sources of six species of Salicornia, the three allied genera Arthrocnemum, Sarcocornia and Halocnemum of the same tribe Salicornieae, and other genera of the subfamily Salicornioideae used in previous studies. Bassia hirsuta, Camphorosma monspeliaca (subfamily Chenopodioideae) and four species of Suaeda (subf. Suaedoideae) were chosen as outgroups. Results show that the annual genus Salicornia is a sister group to the perennial genera Sarcocornia, Arthrocnemum and Halocnemum. Moreover, the phylogenetic analysis based on ITS results distinguished two groups of Salicornia species which fitted with ploidy level: one group consisted of diploid species, and the second of tetraploid ones. Sarcocornia and Arthrocnemum are shown to be closely related, even though the species investigated here exhibited an evident distance between their ITS sequences. On the basis of our results, these two genera should be united. Bienertia (already separated as Bienertieae) was confirmed as probable outgroup to the subf. Salicornioideae, while Kalidium (subf. Salicornioideae, tribe Halopeplideae) was an outgroup to the rest of the Salicornioideae (tribe Salicornieae). The group Allenrolfea plus Halocnemum was the most basal of the tribe Salicornieae amongst those investigated in this study. The two samples of Halocnemum strobilaceum used in this work displayed numerous changes (transitions and transversions) in their respective sequences, probably related to their morphological and chorological differentiation. On the basis of our analysis, the most probable basal chromosome number for Salicornieae appears to be 2n = 18. The same number would also be the base number for the annual genus Salicornia and the perennial Arthrocnemum ( + Sarcocornia), with polyploidy arising independently in the two groups.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

SYSTEMATIC ANATOMY OF THE FLOWER AND FRUIT OF COROKIA

Corokia, a genus of shrubs of New Zealand, eastern Australia, and certain Pacific islands, was first placed in Rhamnaceae, later in Cornaceae, and most recently next to Argophyllum, subfamily Escallonioideae, in Engler's monograph of the Saxifragaceae. Most manuals still list Corokia under Cornaceae from which it is readily excluded by several characters, including pluricellular T-shaped trichomes, ligulate petals, vascular bundles running longitudinally through the center of the inferior gynoecium, histology and germination of the woody endocarp, and a conspicuous subepidermal layer of tannin-containing cells. Corokia collenettei, endemic to the isolated island Rapa, retains the most primitive floral characters of the genus. Anatomical comparison of Corokia flowers with flowers of Argophyllum shows similarities that probably indicate affinity, but the relation of these two genera to others in Engler's subfamily Escallonioideae is unclear. Engler's inclusion of Berenice and Carpodetus with Corokia and Argophyllum in a tribe Argophylleae seems especially artificial.     

Wood and bark anatomy of Steganotaenia and Polemanniopsis (tribe Steganotaenieae,Apiaceae) with notes on phylogenetic implications

The wood and bark structure of the distinctive southern African genera Polemanniopsis (including the newly described species P. namibensis) and Steganotaenia have been described. To allow for comparisons with the traditional subfamily Saniculoideae, a shrubby species of Eryngium from the Juan Fernández Islands was also studied. Polemanniopsis and Steganotaenia were recently considered as two closely related genera forming a new tribe Steganotaenieae of subfamily Saniculoideae (Apiaceae), whereas Eryngium is commonly recognized as a member of Saniculoideae. Eryngium differs significantly from the other two genera in the smaller size of intervessel pits, sclerification and radial dilatation in collapsed secondary phloem, the absence of crystals in the phelloderm cells and the occurrence of druse crystals in secondary phloem ray cells. Steganotaenia and Polemanniopsis share features, including the presence of marginal axial parenchyma, the occurrence of radial secretory canals in secondary xylem, dilatation of the secondary phloem by axial parenchyma stretching, cortical periderm initiation and the presence of chambered phelloderm cells containing druse crystals. These characters (especially the occurrence of chambered crystalliferous cells in phelloderm, which has not yet been reported for Apiaceae) support both the monophyly and the isolated position of the Steganotaeniae. No reliable synapomorphic features could be found to support a relationship with Saniculoideae. Steganotaenia is remarkable in the presence of axial secretory canals in the phelloderm: these structures have not yet been found in the periderm of any member of Apiales. Our results do not provide any support for the suggestion that the woody habit in the three genera examined was derived from herbaceous ancestors secondarily. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 55–59.     

A systematic histological analysis of palm fruits VII. The Cyrtostachydinae (Arecaceae)

Fruit specimens representing five taxa of the genusCyrtostachys were examined histologically in order to characterize the pericarp anatomy of the monogeneric subtribe Cyrtostachydinae (tribe Areceae, subfamily Arecoideae), as part of an ongoing survey of the family. The pericarp in this genus can be characterized by a combination of papillate epidermis, heavy layer of tanniniferous/pigmented cells below the epidermis, a system of vascular bundles with thick fibrous sheaths with purely fibrous bundles frequently above and below, absence of brachysclereids, and a very thin sclerified locular epidermis. On the basis of pericarp structure alone, the genus might be most closely related to theGronophyllum alliance of the subtribe Arecinae. This diverges somewhat from the hypothesis of relationship with theAreca group of the Arecinae resulting from two DNA-based phylogenetic studies, and even further from the hypothesis of relationship withIguanura suggested by another DNA-based phylogenetic study.     

THE ORGANIZATION OF THE VASCULAR SYSTEM IN THE STEMS OF THE NYMPHAEACEAE. II. NYMPHAEA SUBGENERA ANECPHYA,LOTOS, AND BRACHYCERAS

The anatomy and organization of the stem vascular system was analyzed in representative taxa of Nymphaea (subgenera Anecphya, Lotos, and Brachyceras). The stem vascular system consists of a series of concentric axial stem bundles from which traces to lateral organs depart. At the node each leaf is supplied with a median and two lateral leaf traces. At the same level a root trace supplies vascular tissue to adventitious roots borne on the leaf base. Flowers and vegetative buds occupy leaf sites in the genetic spiral and in the parastichies seen on the stem exterior. Certain leaves have flowers related to them spatially and by vascular association. Flowers (and similarly vegetative buds) are vascularized by a peduncle trace that arises from a peduncle fusion bundle located in the pith. The peduncle fusion bundle is formed by the fusion of vascular tissue derived from axial stem bundles that supply traces to certain leaves. The organization of the vascular system in the investigated taxa of Nymphaea is unique to angiosperms but similar to other subgenera of Nymphaea.     

VASCULAR PATTERNS IN STEMS OF ARACEAE: SUBFAMILY POTHOIDEAE

The stem vasculature of representative species of ten of the 11 genera in the Pothoideae was analyzed with the aid of films of series of transverse sections. In all species a leaf trace typically diverges from a continuing axial bundle before departing to a leaf, with the possible exception of Heteropsis. However, within this common organizational scheme a considerable range of variation exists, e.g., with respect to degree of branching of axial bundles, and distance from a leaf-trace branch to the point of leaf-trace departure to a leaf. In addition, we have found a wide variety of patterns of bud-trace organization in different genera of the Pothoideae. For example, prominent arcs composed of numerous bud traces occur in the central cylinder of Pothos, Pothoidium, and Heteropsis. Comparative anatomy leads to the conclusion that Pothos and Pothoidium more closely resemble Heteropsis than Anadendrum. Anadendrum should be dissociated from the tribe Pothoeae. With respect to other genera in the Pothoideae, our preliminary results suggest that each of the genera Anthurium, Culcasia, Zamioculcas, Acorus, and Gymnostuchys are highly distinct from each other.     

Hyaloscyphaceae in Japan (1): Non-glassy-haired members of the tribe hyaloscypheae

Discomycetes of the family Hyaloscyphaceae were collected from across Japan and identified. In this report, non-glassy-haired members of the tribe Hyaloscypheae, subfamily Hyaloscyphoideae are described and illustrated. The five species involved include one new species (Phialina lacrimiformis) and four species less well known or newly reported in Japan (Dematioscypha dematiicola, Hamatocanthoscypha laricionis var.Iaricionis, Micropodia chrysostigma, M. grisella). cultural studies were carried out, and teleomorph-anamorph relationships are discussed forD. dematiicola, H. laricionis val.laricionis and their anamorphs. A key to the genera of the tribe Hyaloscypheae in Japan is provided.     

Phylogenetic relationships of Combretoideae (Combretaceae) inferred from plastid,nuclear gene and spacer sequences

Phylogenetic relationships of the subfamily Combretoideae (Combretaceae) were studied based on DNA sequences of nuclear ribosomal internal transcribed spacer (ITS) regions, the plastid rbcL gene and the intergenic spacer between the psaA and ycf3 genes (PY-IGS), including 16 species of eight genera within two traditional tribes of Combretoideae, and two species of the subfamily Strephonematoideae of Combretaceae as outgroups. Phylogenetic trees based on the three data sets (ITS, rbcL, and PY-IGS) were generated by using maximum parsimony (MP) and maximum likelihood (ML) analyses. Partition-homogeneity tests indicated that the three data sets and the combined data set are homogeneous. In the combined phylogenetic trees, all ingroup taxa are divided into two main clades, which correspond to the two tribes Laguncularieae and Combreteae. In the Laguncularieae clade, two mangrove genera, Lumnitzera and Laguncularia, are shown to be sister taxa. In the tribe Combreteae, two major clades can be classified: one includes three genera Quisqualis, Combretum and Calycopteris, within which the monophyly of the tribe Combreteae sensu Engler and Diels including Quisqualis and Combretum is strongly supported, and this monophyly is then sister to the monotypic genus Calycopteris; another major clade includes three genera Anogeissus, Terminalia and Conocarpus. There is no support for the monophyly of Terminalia as it forms a polytomy with Anogeissus. This clade is sister to Conocarpus. Electronic Publication     

Changes in the suprageneric classification of Lasiocampidae (Lepidoptera) based on the nucleotide sequence of gene EF-1α

The nucleotide sequences of nuclear gene EF-1α were determined for 49 species of Lasiocampidae from Eurasia and Africa. Based on these data, the phylogeny of the family was reconstructed using the minimum evolution, maximum parsimony, maximum likelihood, and Bayesian inference methods. The molecular genetic research confirms the monophyly of Malacosominae which is treated as a separate subfamily. The genus Euthrix appears to be paraphyletic. The group of genera similar to Arguda, previously united with Odonestis in the tribe Odonestini, proved to be a separate lineage; contrary to the earlier assumption, the genera do not seem to be related. On the other hand, the genera Argonestis and Odonestis were found to be closely related and therefore were placed in the same tribe. The position of the genus Macrothylacia remains obscure. The genus Stoermeriana de Fr. et Witt is also para- or polyphyletic and consists of several independent lineages whose status remains to be determined. The new classification supports synonymization of Pinarinae with Lasiocampinae. The rank of subfamily is not supported for the Neotropical Macromphaliinae, which is downgraded to a tribe, Macromphaliini stat. n., within Poecilocampinae. The genus Hypopacha, previously considered within Macromphaliinae, is transferred to Poecilocampini; the close relation between Poecilocampini and Macromphaliini is additionally supported by the presence of a member of Poecilocampini in the New World. A new tribe, Argudini Zolotuhin trib. n., is established.