Restionaceae seedlings: Morphology,anatomy and systematic implications

Restionaceae differ from most monocot families in having both epigeal and hypogeal germination. The green cotyledons associated with epigeal germination have a central vascular strand as found in most epigeal monocotyledons. In some genera the cotyledon may have a hairpin‐like structure, also described for Anthericaceae. The cotyledon of the hypogeal seedlings is short, without green pigment and largely remains embedded in the seed coat. Hypogeal germination is correlated with large, woody, indehiscent, frequently myrmecochorous nuts, while epigeal germination is found in species with smaller indehiscent nutlets or seeds, dispersed in a variety of ways. The primitive condition is most likely epigeal germination. In hypogeal seedlings of some African and Australian taxa an epicotyledonary rhizome is found between the primary root and the first leaves. Seedlings of African Restionaceae frequently have elongated culm internodes, whereas in the Australian species studied, internodes are very short, resulting in a cluster of seedling leaves. The leaf blades, which in most species are only found on the seedlings, are very simple anatomically. However, they appear to be unifacial, similar to the leaf blades of Anarthria (Anarthriaceae). The anatomical specialisations in the blades mirror those recorded for the culm anatomy. These observations are consistent with the hypothesis that Centrolepidaceae may be neotonous Restionaceae. They also corroborate the morphology of the African Restionaceae, and the presently accepted phylogeny of the African genera of Restionaceae.     

Contrasting patterns of radiation in African and Australian Restionaceae

The floras of the Mediterranean-climate areas of southern Africa and southwestern Australia are remarkably species rich. Because the two areas are at similar latitudes and in similar positions on their respective continents, they have probably had similar Cenozoic climatic histories. Here we test the prediction that the evolution of the species richness in the two areas followed a similar temporal progression by comparing the rates of lineage accumulation for African and Australian Restionaceae. Restionaceae (Poales) are typical and often dominant elements in the fynbos vegetation of the Cape Floristic Region of southern Africa and the kwongan vegetation of the Southwestern Floristic Province of Western Australia. The phylogeny of the family was estimated from combined datasets for rbcL and trnL-F sequences and a large morphological dataset; these datasets are largely congruent. The monophyly of Restionaceae is supported and a basal division into an African clade (approximately 350 species) and an Australian clade (146 species) corroborated. There is also support for a futher subdivision of these two large sister-clades, but the terminal resolution within the African clade is very weak. Fossil pollen records provided a minimum age of the common ancestor of Australian and African Restionaceae as 64-71 million years ago, and this date was used to calibrate a molecular clock. A molecular clock was rejected by a likelihood ratio test; therefore, rate changes between the lineages were smoothed using nonparametric rate smoothing. The rate-corrected ages were used to construct a plot of lineages through time. During the Palaeogene the Australian lineage diversity increased consistent with the predictions of the constant birthrate model, while the African lineage diversity showed a dramatic increase in diversification rate in the Miocene. Incomplete sampling obscures the patterns in the Neogene, but extending the trends to the modern extant diversity suggests that this acceleration in the speciation rate continued in the African clade, whereas the Australian clade retained a constant diversification rate. The substantial morphological and anatomical similarity between the African and Australian Restionaceae appear to preclude morphological innovations as possible explanations for the intercontinental differences. Most likely these differences are due to the greater geographical extent and ecological variation in temperate Australia than temperate Africa, which might have provided refugia for basal Restionaceae lineages, whereas the more mountainous terrain of southern Africa might have provided the selective regimes for a more rapid, recent speciation.     

The structure and evolution of the female flower of the African Restionaceae

UNDER, H. P., 1992. The structure and evolution of the female flower of the African Restionaceae . The anatomical details of the gynoecium and perianth of the African Restionaceae are described. The same basic pattern of stylar and ovular vascular traces is found throughout the group. Variation in the number of fertile locules is the result of several reduction sequences. The most common sequence is the loss of the fertility of locules, followed by the loss of the infertile locules. These carpels are still represented by the stylar vascular traces. In different lineages, different locules become sterile. A second form of reduction is fusion of carpels. This occurred in only one lineage. The variation in the structure of the ovary walls is discussed, and three different types of ovary wall construction are demonstrated. The anatomical construction of the tepals is briefly mentioned, but it is apparent that the sample is too small for confident interpretation. Variation in ovary structure is shown to be largely consistent with the phylogeny and classification of the group. However, the relationship between Thamnochortus and Staberoha may need re-evaluation, and the status of Calopsis as a monophyletic group distinct from Restio is questioned.     

Correlations between flavonoid chemistry,anatomy and geography in the restionaceae

Comparisons of the flavonoid patterns in stems and inflorescences between Australasian and South African members of the Restionaceae indicate significant differences with geography. Nine of 14 Australasian species contain gossypetin or a related 8-hydroxyflavonoid and proanthocyanidins are uncommon. By contrast, the 33 South African taxa studied contain common flavonols, flavones and glycoflavones, while proanthocyanidins are present in 29. Two anatomically related South African genera, Chondropetalum and Elegia, contain, in addition, myricetin 3-galactoside, together with the 3-galactosides of the myricetin methyl ethers, larycitrin and syringetin. These results confirm the conclusions derived from anatomy that members of Hypolaena, Leptocarpus and Restio, genera represented in both Australia and South Africa, have the distinctive flavonoids characteristic of their geographic origin rather than of their systematic position. The family as a whole is different in flavonoid pattern from other monocotyledonous families with which it is sometimes associated.     

Ontogeny and evolution of the flowers of South African Restionaceae with special emphasis on the gynoecium

 The South African Restionaceae make up a highly diverse group of genera displaying several reductive trends in the configuration of the flower, especially in the gynoecium. In this paper the floral ontogeny of fourteen species representing nine of the 11 genera of the Restio clade is studied with the SEM. Although flowers are basically simple, the variability in both mature and developmental stages is striking. Differences between species are the result of changes in growth rate, coupled with differential pressures of organs. Trends in the elaboration of bracts, perianth, androecium and gynoecium are compared. Together with data that have been presented elsewhere about the other clade of African Restionaceae, viz. the Willdenowia-clade, a scheme with potential developmental pathways is proposed and the most evident routes are selected based on ontogenetic evidence. Nine possible reductions are presented arising through three main routes. Received August 27, 2001 Accepted October 26, 2001     

Billburttia, a new genus of Apiaceae (tribe Apieae) endemic to Madagascar

The genus Billburttia is described to include two new species endemic to Madagascar, B. capensoides and B. vaginoides. Both species were tentatively placed within the problematic genus Peucedanum L. (as Peucedanum sp. A and Peucedanum sp. B) based on their dorsally compressed fruits lacking prominent dorsal ribs and with winged marginal ribs. Recently, however, the African members of Peucedanum have been shown to be only distantly related to the type of the genus and have therefore been segregated into six African endemic genera. While the Malagasy species appear superficially similar to members from one of these segregate genera, viz. Notobubon, they differ markedly in their fruit anatomical characters, notably a narrower commissure, six commissural vittae, vascular tissue in the tip of the ribs and sphaerocrystals distributed in and around the epidermis. The latter two characters apparently represent apomorphies for Billburttia. The non-peucedanoid affinity of the genus, as suggested by the fruit anatomical data, was confirmed using ITS and rps16 intron sequences. Both parsimony and Bayesian analyses of these data place Billburttia within the tribe Apieae of subfamily Apioideae and not closely related to either Peucedanum (Selineae) or the African peucedanoid genera (Lefebvrea clade of Tordylieae).     

国产嵩草属(莎草科)植物秆的解剖学研究

对嵩草属27种(亚种)植物秆的解剖学研究证明,嵩草属植物秆的解剖学性状具有系统学意义.在该属中,秆的横切面外形为三角形、圆三角形、圆形或扁圆形.在横切面上分为2个区域;外部区域包括绿色组织、外韧维管束和气腔,内部区域为薄壁组织或其碎裂形成空腔.秆表皮的横切面观和表面观均与叶的下表皮相似.以上特征与莎草科其它类群植物秆的解剖特征一致,不支持将嵩草属和其近缘属另立为嵩草科.同时,秆的解剖学特征可以做为某些在外部形态上难于区分的种之间的分类依据.     

Morphology and development of the gynoecium in Centrolepidaceae: The most remarkable range of variation in Poales

This paper explores the relative impacts of reduction and polymerization on the evolution of reproductive structures in the small but morphologically diverse family Centrolepidaceae. Centrolepidaceae are closely related to Restionaceae and belong to the large order Poales, which also includes the grasses. In the largest genus of Centrolepidaceae, Centrolepis, the reproductive structures are viewed either as highly unusual aggregations of reduced flowers (the pseudanthial interpretation) or as unique flowers evolved through extreme reduction in the androecium, usually accompanied by a drastic increase in carpel number and elaboration of the entire gynoecium. Comparative data are here presented on gynoecia of all three genera of Centrolepidaceae; these data strongly support the latter (euanthial) interpretation. The combined phenomenon of carpel multiplication and decrease in stamen number is unexpected in a predominantly wind-pollinated lineage. Applying a pseudanthial interpretation would create a considerable morphological gap with reproductive structures of other Poales, whereas accepting a euanthial concept allows an almost continuous morphological series with related taxa.     

Phylogenetic divergence of the Australian allodapine bees (Hymenoptera: Apidae)

Allodapine bees are most diverse in Africa but are distributed throughout the Old World tropical and Austral regions. They are considered useful for studies into the evolution of social behaviour since they exhibit the full range of social organisation from solitary to highly eusocial (sensu; ). Five genera are found in Australia, namely Braunsapis, Exoneurella, Exoneura, Brevineura, and Inquilina. Sociality and life histories are well documented for the exoneurine genera (review in ) and Inquilina is an obligate social parasite of species of Exoneura (). In this paper, maximum parsimony and maximum likelihood methods using molecular sequence data from two mitochondrial gene regions (cyt b and COI) and a single nuclear gene region (EF-1alpha) are used to reconstruct phylogenetic relationships of the Australian allodapine genera. Results suggest that the exoneurine group (Brevineura, Exoneurella, and Exoneura+Inquilina) diverged very rapidly and are monophyletic to the exclusion of other (primarily African) allodapine genera. A clade containing Australian species of Braunsapis is also monophyletic to the exclusion of African congeners. Braunsapis is not phylogenetically close to, and is a more derived group than the exoneurine group and probably came to occupy the Australian plate via a later dispersal through the southern Asian region. It is unclear at this point how the exoneurine group came to occupy the Australian plate and possible scenarios are discussed.     

Evolution of lamina anatomy in the palm family (Arecaceae)

The unique properties of tree building in Arecaceae strongly constrain their architectural lability. Potentially compensating for this limitation, the extensive diversification of leaf anatomical structure within palms involves many characters whose alternate states may confer disparate mechanical or physiological capabilities. In the context of a recent global palm phylogeny, we analyzed the evolution of 10 such lamina anatomical characters and leaf morphology of 161 genera, conducting parsimony and maximum likelihood ancestral state reconstructions, as well as tests of correlated evolution. Lamina morphology evolves independently from anatomy. Although many characters do optimize as synapomorphic for major clades, anatomical evolution is highly homoplasious. Nevertheless, it is not random: analyses indicate the recurrent evolution of different cohorts of correlated character states. Notable are two surface layer (epidermis and hypodermis) types: (1) a parallel-laminated type of rectangular epidermal cells with sinuous anticlinal walls, with fibers present in the hypodermis and (2) a cross-laminated type of hexagonal cells in both layers. Correlated with the cross-laminated type is a remarkable decrease in the volume fraction of fibers, accompanied by changes in the architecture and sheath cell type of the transverse veins. We discuss these and other major patterns of anatomical evolution in relation to their biomechanical and ecophysiological significance.     

What lies beneath: molecular phylogenetics and ancestral state reconstruction of the ancient subterranean Australian Parabathynellidae (Syncarida, Crustacea)

The crustacean family Parabathynellidae is an ancient and significant faunal component of subterranean ecosystems. Molecular data were generated in order to examine phylogenetic relationships amongst Australian genera and assess the species diversity of this group within Australia. We also used the resultant phylogenetic framework, in combination with an ancestral state reconstruction (ASR) analysis, to explore the evolution of two key morphological characters (number of segments of the first and second antennae), previously used to define genera, and assess the oligomerization principle (i.e. serial appendage reduction over time), which is commonly invoked in crustacean systematics. The ASR approach also allowed an assessment of whether there has been convergent evolution of appendage numbers during the evolution of Australian parabathynellids. Sequence data from the mtDNA COI and nDNA 18S rRNA genes were obtained from 32 parabathynellid species (100% of described genera and ~25% of described species) from key groundwater regions across Australia. Phylogenetic analyses revealed that species of each known genus, defined by traditional morphological methods, were monophyletic, suggesting that the commonly used generic characters are robust for defining distinct evolutionary lineages. Additionally, ancestral state reconstruction analysis provided evidence for multiple cases of convergent evolution for the two morphological characters evaluated, suggesting that caution needs to be shown when using these characters for elucidating phylogenetic relationships, particularly when there are few morphological characters available for reconstructing relationships. The ancestral state analysis contradicted the conventional view of parabathynellid evolution, which assumes that more simplified taxa (i.e. those with fewer-segmented appendages and setae) are derived and more complex taxa are primitive.     

The comparative leaf anatomy of Agave, Beschorneria, Doryanthes and Furcraea species (Agavaceae: Agaveae)

The leaf anatomy is compared of 35 ppecies and one variety of Agave , three species of Furcraea , one species of Beschorneria and one species of Doryanrhes , four genera assigned to the tribe Agaveae by Hutchinson in his classification of the family Agavaceae. The genera can be readily differentiated by leaf anatomy. There are close similarities between Agave, Beschorneria and Furcraea , but Doryanthes differs widely from these genera. Sufficient anatomical differences exist to differentiate the Agave species examined.     

Leaf anatomy of Bruniaceae: ecological, systematic and phylogenetic aspects

CARLQUIST, S., 1991. Leaf anatomy of Bruniaceae: ecological, systematic and phylogenetic aspects. Quantitative and qualitative data are given for 60 species of the 12 genera of Bruniaceae; most data are based on liquid-preserved material. Leaves of Bruniaceae are basically linear (broader forms are probably derived) with an apicula that contains phellogen activity. Most bruniaceous leaves have some degree of isolateral construction, with transition to normal bifacial construction in a few species, but more commonly transition to 'inverse' bifacial structure (stomata on adaxial face, palisade on abaxial face). The latter type is correlated with the tendency for leaves to be appressed to stems. Tannins and very likely other dark-staining materials are very characteristic of mesophyll cells. Six genera have a large strand of fibres on the midvein and rhomboidal crystals in bundle sheath cells. The other six genera have few or no fibres on veins and have druses in mesophyll cells (but not in bundle sheath cells). These distinctions may relate to intrafamilial taxonomy, but they also support the primitive position usually accorded to Audouinia, Thamnea and Tittmannia. A key to genera based on leaf antomy is offered. Details of epidermal cell shape, cuticular relief and trichome form and structure based on scanning electron microscopy are given. Leaf anatomy, combined with other features, favours a relationship between Bruniaceae and Grubbiaceae in particular and in broader contexts allies Bruniaceae to rosalean and possibly hamamelidalean families.     

Anatomical adaptations of shrews

The family Soricidae is reviewed with special regard to anatomical structures related to habitat and life-style. Some 266 species in 20 genera are classified into six feeding and foraging categories: terrestrial, semifossorial, scansorial, semiaquatic, psammophilic and anthropophilic. Examples of corresponding anatomical adaptations are given as well as notes on adaptive radiation and convergent evolution.     

国产节柱竹族地下茎的解剖及其在分类中的意义

观察了我国节柱竹族（广义的）５属２７种的地下茎解剖特徵,发现地下茎,尤其是竿柄,在本族的分属中有相当重要的意义。作者根据地下茎以及其它器官的特徵,对国产节柱竹族５个属之间的关系作了探讨,认为这５个属以各自独立为宜。     

Convergent evolution of shell shape in freshwater limpets: the African genus Burnupia

Convergent evolution of shell shape is a well known phenomenon in gastropods that has caused much confusion in taxonomic and systematic studies. A paradigm is the patelliform shell shape in several taxa of the order Basommatophora. Historically, most freshwater limpets were assigned to the family Ancylidae. Based on anatomical data, some taxa were subsequently moved to different families. However, there are still doubts about the monophyly of the remaining ancylids. This is also true for the African limpets and particularly for species of the genus Burnupia Walker, 1912. In the present paper, two independent molecular markers (COI, 18S rRNA) are used to (a) infer the position of Burnupia within the Basommatophora and (b) to test whether the shell shape of Burnupia evolved independently. Bayesian and maximum likelihood analyses of 12 genera of limnic Basommatophora indicate that Burnupia not only appears to be distinct from the Ancylidae, but also from all other representatives of the superfamily Planorboidea studied here. Based on a generally well-supported phylogeny and preliminary anatomical data, it is concluded that patelliform shell shapes evolved at least three times in the taxa studied here and that the shell shape of Burnupia represents another case of convergent evolution. Previously proposed functional and evolutionary scenarios for the evolution of patelliform shell and their relevance for freshwater limpets are discussed.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 577–586.     

MEGAGAMETOPHYTE AND NUCELLUS IN RESTIONACEAE AND FLAGELLARIACEAE

Restionaceae are characterized by Polygonum-type development, a well-developed hypostase, the presence of starch grains in the mature female gametophyte, and a thick nucellus that is uniseriate at the micropylar end. Flagellaria indica differs in these characters. In general, proliferation of antipodals occurs in South African taxa of Restionaceae but not in Australasian taxa. The presence of multiplied antipodals is the usual condition in the allied family Poaceae, but the sporadic occurrence of this character in Restionaceae cannot be used to support the view that the two families are sister groups.     

中国伞形科天胡荽亚科果实解剖特征及其系统学意义

经对中国伞形科天胡荽亚科 (Hydrocotyloideae)中天胡荽属 (HydrocotyleL .)积雪草属 (CentellaL .)和马蹄芹属(DickinsiaFranch .) 10种植物果实解剖结构的观察比较 ,发现 3属的果实横切面以合生面为基准可以分成 2个类型 ,即两侧压扁类型 (天胡荽属和积雪草属 )和背腹压扁类型 (马蹄芹属 )。除此之外 ,中果皮的外侧是否有色素块沉积、内侧是否有木化细胞层或厚壁细胞层以及最内层是否形成晶体细胞层 ,内果皮细胞的层数、排列方向以及有无棱槽油管等性状 ,可以作为区分 3属的重要解剖学特征。在此基础上 ,结合外部形态特征 ,讨论了该亚科及其内部各属的解剖学特征、属间区别及其演化关系。认为马蹄芹属应归属于Mulineae族 ,另外 2个属应归属于天胡荽族(Hydrocotyleae) ;3属中积雪草属原始而马蹄芹属进化 ;天胡荽亚科可能不是自然的类群 ,它的分类系统和演化地位有待深入研究。     

Convergent evolution and paraphyly of the hawk-eagles of the genus Spizaetus (Aves, Accipitridae) – phylogenetic analyses based on mitochondrial markers

The phylogenetic relationships within the New and Old World hawk-eagle assemblage (genus Spizaetus ; Aves: Accipitridae) were studied using mitochondrial DNA sequences ( cytochrome b , control region). Eighty-four specimens representing all Spizaetus species and almost all currently distinguished subspecies as well as 11 other booted and non-booted 'eagle' genera from the Neotropics, Africa, Eurasia, South Asia and Australasia ( Oroaetus , Harpia , Morphnus , Lophaetus , Stephanoaetus , Hieraaetus , Aquila , Ictinaetus , Spilornis , Pithecophaga , Harpyopsis ) were investigated. Although the basal branching could not be resolved, our investigations clearly indicate that hawk-eagles represent a paraphyletic assemblage and thus their external similarities have to be ascribed to convergent evolution. The New World taxa of Spizaetus cluster together, but the South American species Oroaetus isidori appears embedded within this clade. The taxa from Southeast to East Asia form a clearly separated monophyletic group. It is further divided into two subgroups, which are also characterized by distinct juvenile plumage patterns. Spizaetus africanus , the only African representative of the genus, is found in a mixed cluster consisting of members of the genera Aquila and Hieraaetus . These findings are in accordance with previous studies of other authors based on various molecular markers and different sets of taxa, but disagree with current taxonomy. Therefore, we suggest assigning the species of the genus Spizaetus to three different genera: (1) Spizaetus (including Oroaetus isidori ) in Central and South America and (2) Nisaetus for the Southeast to East Asian group. (3) The African taxon ( Spizaetus africanus ) is discussed to be included into the genus Aquila. Furthermore, we propose to use the former genus name Lophotriorchis Sharpe, 1874, for the monotypic species Hieraaetus kienerii , which has an isolated phylogenetic position.