VESSELS IN JUNCALES: II. CENTROLEPIDACEAE AND RESTIONACEAE

The occurrence and specialization of vessels in 6 genera and 14 species of Centrolepidaceae and 25 and 72, respectively, of Restionaceae are described. Those in Anarthriaceae (1 and 3) and Ecdeiocoleaceae (1 and 1), newly separated from Restionaceae, were also examined. Vessels in Ecdeiocolea are highly specialized; those in Centrolepidaceae are relatively primitive. Vessels in Restionaceae and Anarthriaceae are intermediate. Vessels in Cyperaceae and Gramineae are compared with those in Juncales in a review of vessels in Glumiflorae. Thurniaceae has most primitive vessels on the average, Juncales has more primitive vessels in leaves and inflorescence axes than does Cyperales; the opposite is true for roots, rhizomes and stems. Vessels in Graminales on the average are much more specialized than those in Cyperales and Juncales.     

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.     

Systematic significance of pollen arrangement in MICROSPORANGIA OF POACEAE AND Cyperaceaei REVIEW AND OBSERVATIONS ON REPRESENTATIVE TAX A

An anatomical survey of anthers of the Poaceae and Cyperaceae and two related families, the Restionaceae and Flagellariaceae, was conducted to determine the taxonomic distribution and possible phylogenetic significance of pollen arrangement. An unusual pollen arrangement was known from a small number of taxa in the Poaceae and Cyperaceae, in which a single, uniseriate cylinder of pollen grains is arranged in the anther locule such that each grain is in contact with the tapetum (termed here “peripheral” pollen). This contrasts with the prevailing arrangement in other angiosperms in which the locule contains a relatively large number of pollen grains in no special configuration, with many interior grains that never touch the tapetum (termed here “central” pollen). A total of 48 species in these four families was examined in this study, and observations on pollen arrangement for numerous additional species in these families were gleaned from the literature. We confirm that the peripheral arrangement is predominant in Poaceae and Cyperaceae, although both families also include species with central pollen, whereas only central pollen is found in Restionaceae and Flagellariaceae. In the peripheral arrangement, the pore of the pollen grain or pseudomonad, when observed, is in contact with the tapetum, but it has not been definitively demonstrated that this is always the case. Peripheral pollen in the Poaceae and Cyperaceae is nonhomologous because of the presence of pseudomonads in the latter family. It remains unexplained why peripherally arranged pollen or pseudomonads, with the attendant reduction in the number of pollen grains, should be associated with anemophily in these two families.     

Monocotyledon fruits and seeds, and an associated palynoflora from Eocene-Oligocene sediments of coastal central Queensland, Australia

Fruits of Restionaceae and seeds of Typhaceae are described from a latest Eocene–Oligocene mudstone underlying oil shales in a subsurface sequence near Rockhampton, coastal central Queensland. The Restionaceae fruits are unilocular and encase a single pendulous orthotropous seed with a structured micropylar cap, which may be tannin filled. These are allocated to Restiocarpum gen. nov., which has as its type Restiocarpum latericum sp. nov., and four other taxa described herein; Restiocarpum tesselatum sp. nov., Restiocarpum verrucatum sp. nov., Restiocarpum laeve sp. nov., and Restiocarpum fusiforme sp. nov. Typhaspermum cooksoniae gen. et sp. nov., which accommodates asymmetric claviform seeds, is interpreted as a member of the Typhaceae based on the presence of a lid-like operculum, bitegmic wall, and chalazal cavity.

Reconstruction of the source plant community emphasizes similarities to restiad swamps of present day Wallum (swamp heathland) vegetation which is extensively developed along the Queensland coast. Biogeographic implications for the Restionaceae and Typhaceae are discussed.     

DIOECISM AND ITS CORRELATES IN THE CAPE FLORA OF SOUTH AFRICA

Floral sexuality is characterized for the flora of the Cape region of South Africa. Among angiosperms (N = 8,497), monostylous hermaphroditism constitutes the largest proportion of species with 77.7%, followed by gynomonoecism (7.1%), dioecism (6.6%), andromonoecism (4.0%), monoecism (2.6%), heterostylous hermaphroditism (1.9%), polygamomonoecism (0.01%) and polygamodioecism (0.01%). The incidence of dioecism is significantly higher (P < 0.001) at the species level for two smaller floras within the Cape flora that consist mostly of species-rich fynbos vegetation (Cape of Good Hope Nature Reserve: 11.6% [N = 1,349], Cape Hangklip: 9.7% [N = 1,046]). The relatively high incidence of dioecism in all three floras compared to other temperate floras is due to the large number of Restionaceae species. Excluding these rush-like plants, dioecism at the species level is only 3.0% for the Cape flora, 4.2% for the Cape Hangklip flora and 3.6% for the Cape of Good Hope flora. At the generic level there are no significant differences in the incidence of dioecism between the three floras. Among dioecious species and genera in the Cape flora, there is a higher than expected incidence of wind pollination, fleshy fruits and nonwoodiness when each variable is analyzed independently. The association between nonwoodiness and dioecy has not been reported for other floras and is due to the large proportion of nonwoody wind-pollinated Restionaceae. If this family is excluded from the analysis, dioecy becomes associated with biotic pollination, fleshy fruits, and woodiness. Interactions among the variables themselves as well as with dioecism indicate the need for a joint analysis of the variables. At the genus level, analyses reveal the following: 1) There is a higher than expected incidence of wind pollination among dioecious plants with dry fruits. If the Restionaceae are excluded from the analysis, wind pollination is more common than expected only for woody plants with dry fruits. 2) Dioecious plants that are biotically pollinated have a significantly higher incidence of fleshy fruits than expected. 3) Dioecious plants with dry fruits that are biotically pollinated have a higher than expected incidence of woodiness. If the Restionaceae are excluded from the analysis, woodiness is more common than expected among both nonwind and wind-pollinated dioecious genera.     

The megagametophyte in Anarthria (Anarthriaceae,Poales) and its implications for the phylogeny of the Poales

The development and structure of the megagametophyte of Anarthria (Anarthriaceae), Aphclia, and Centrolepis (Centrolepidaceae) are described. Anarthriaceae has tenuinucellate ovules and the Polygonum type of megagametophyte development, both characters typical of the Poales. However, it lacks the anticlinally elongated nucellar epidermis and numerous large starch bodies observed in the megagametophyte of Centrolepidaceae, both characters also present in Restionaceae. This relatively generalized megagametophyte structure is consistent with data from the chloroplast genome, which suggest that Anarthriaceae are not as closely related to Restionaceae as previously assumed. New data from the megagametophyte are analyzed cladistically together with other available information on the poalean families. The results show that there are two possible positions for Anarthriaceae: either as sister to Poaceae. Joinvilleaceae, Restionaceae, Ecdeiocoleaceae, and Restionaceae, or as sister to only the latter three families. The new data also allow a critical reevaluation of the phylogenetic position of Centrolepidaceae, which is either basal to the poalean clade (based on microgametophyte data), or embedded in the Restionaceae (based on anther structure and megagametophyte data).     

DEVELOPMENT OF THE OVULE AND FEMALE GAMETOPHYTE IN EUSTACHYS PETRAEA AND E. GLAUCA (POACEAE)

The unilocular pistil in Eustachys contains a single ovule with lateral placentation. In E. petraea and E. glauca, the mature ovule is bitegmic, tenuinucellate, and amphitropous with the endostomic micropyle oriented toward the base of the locule. A single hypodermal archesporial cell enlarges to form the megasporocyte. The chalazal dyad member is larger than the micropylar one, and meiosis II is nonsynchronized. Two-thirds of the tetrads are linear and one-third T-shaped. The chalazal megaspore is functional. Initially ovoid, the two-nucleate female gametophyte becomes curved as it enlarges. The four-nucleate stage becomes wider at its extremities and constricted in the center. Synchronous mitotic divisions establish the eight-nucleate stage with four nuclei at each pole separated by a large central vacuole. In E. petraea, the maturation sequence begins with antipodal differentiation, followed by differentiation of the egg apparatus, migration of the polar nuclei to the center, and division of the antipodals to produce twelve cells. The sequence in E. glauca begins with migration of the polar nuclei followed by differentiation of the antipodals, egg apparatus, and antipodal replication to six cells. The polar nuclei fuse to form a secondary nucleus appressed to the egg cell in E. glauca and separated from it by a vacuole in E. petraea. T-tests for length measurements for various stages of development indicate that the functional megaspore and two-nucleate female gametophyte are significantly larger in E. glauca than in E. petraea. There is no significant difference in gametophyte length at the four-nucleate stage, and at the eight-nucleate stage, length in E. petraea surpasses that in E. glauca. This gap widens significantly at the mature stage. Nuclear volumes are significantly greater in E. glauca than in E. petraea in the functional megaspore and two-nucleate stage, but the volumes are similar at the four-nucleate stage. Consideration of the differences in structural complexity between the sporophyte and gametophyte generations leads to the conclusion that the female gametophytes of these species are more distinctive than are the sporophytes.     

Vicariance, climate change, anatomy and phylogeny of Restionaceae

Cutler suggested almost 30 years ago that there was convergent evolution between African and Australian Restionaceae in the distinctive culm anatomical features of Restionaceae. This was based on his interpretation of the homologies of the anatomical features, and these are here tested against a 'supertrec' phylogeny, based on three separate phvlogenies. The first is based on morphology and includes all genera; the other two are based on molecular sequences from the chloroplast genome-, one covers the African genera, and the other tin-Australian genera. This analysis corroborates Cutler's interpretation of convergent evolution between African and Australian Restionaceae. However, it indicates that for the Australian genera, the evolutionary pathway of the culm anatomy is much more complex than originally thought. In the most likely scenario, the ancestral Restionaceae have protective cells derived from the chlorenchyma. These persist in African Restionaceae, but are soon lost in Australian Restionaceae. Pillar cells and sclerenchyma ribs evolve early in the diversification of Australian Restionaceae, but are secondarily lost numerous times. In some of the reduction cases, the result is a very simple culm anatomy, which Cutler had interpreted as a primitively simple culm type, while in other cases it appears as if the functions of the ribs and pillars may have been taken over by a new structure, protective cells developed from epidermal, rather than chlorenchyma, cells. Cutler suggested that this convergent evolution might have been in response to Tertiary climatic deterioration, but this study finds no strong corroborating evidence for this.     

Ultrastructure of endopolyploid antipodals inAconitum vulparia Rchb.

Summary The ultrastructure of the antipodals ofAconitum vulparia Rchb. was studied in mature embryo sacs. Antipodal cell wall thickness varies in different parts of the cells. The antipodals resemble transfer cells with distinctly marked wall ingrowths which are particularly well developed in the chalazal part and between the antipodals. A few plasmodesmata occur in the cell wall between the antipodals and the central cell. The cytoplasm is rich in ribosomes which occur free or bound to the membranes of the well developed endoplasmic reticulum. Only in the micropylar region of the cells are some larger vacuoles found. The antipodals contain numerous mitochondria, plastids and apparently active dictyosomes. Vesicles with electron dense contents, microbodies, multivesicular bodies as well as lipid droplets and small multiple concentric cisternae are also present in the cytoplasm. The giant endopolyploid nuclei have lobed outlines, especially at the chalazal side of the nuclei.Ultrastructural features, especially the occurrence of numerous free ribosomes and the development of extensive rough endoplasmic reticulum, suggest high metabolic activity in the growing and differentiating antipodals of this species.     

CLADISTIC ANALYSIS OF PATTERNS OF ENDOTHECIAL THICKENINGS IN THE POALES/RESTIONALES

The three-dimensional structure of the endothecial thickenings in the anthers was investigated in 87 species from 70 genera, chosen to provide representative coverage of the families Cyperaceae, Restionaceae, Anarthriaceae, Ecdeiocoleaceae, Centrolepidaceae, Joinvilleaceae, Flagellariaceae, Poaceae, Xyridaceae, and Eriocaulaceae. There is complex variation in the patterns of endothecial thickening: the Eriocaulaceae, Flagellariaceae, and most Poaceae have thickenings with a complete baseplate; the Cyperaceae and most of the Restionaceae are characterized by helical thickenings; some genera in the Bambusoideae have annular thickenings; and U-shaped thickenings occur in the Xyridaceae and Eriocaulaceae and in some Poaceae and Restionaceae. Joinvillea and Ecdeiocolea have unique thickening types. Endothecial characters were subjected to cladistic analysis. Including endothecial characters in an existing cladogram of the group indicates that there is no single, well-corroborated cladogram available for the Poales/Restionales.     

EMBRYO SAC LACKING ANTIPODAL CELLS IN ARABIDOPSIS THALIANA (BRASSICACEAE)

Ultrastructure of the embryo sac lacking antipodals in prefertilization stages in Arabidopsis thaliana has been examined 2 hr before and 5 hr after manual cross pollination. The cytoplasm of both synergids before fertilization is rich in ribosomes, mitochondria, and rough endoplasmic reticulum, and also contains several microbodies and spherosomes. The filiform apparatus includes electron-dense material and a fibrous part. Many cortical microtubules appear in the filiform apparatus area. One of the two synergids degenerates before fertilization. The synergids, the egg cell, and central cell have a rich cytoskeleton of microtubules; only the synergids appear to contain microfilaments. At the chalazal end, the antipodals are initially present but degenerate by the time of pollination in most embryo sacs in the starchless line studied. The embryo sac is completely surrounded by a wall containing an electron-dense layer, separating it from the nucellus, including the chalazal end. When the antipodals have degenerated, the electron-dense layer disappears at the chalazal end only, and the wall between the central cell and the nucellus is homogeneous. Between the central cell and nucellar cells no plasmodesmata are found. The membranes of both antipodal cells at the chalazal end of the embryo sac appear sinuous, like those of transfer cells. The central cell has plastids preferentially distributed around the nucleus, but the other organelles are randomly distributed. The central cell in the embryo sac and the adjacent chalazal nucellar cells show a transfer-cell function in the embryo sac after the antipodals degenerate.     

Variation in δ13C among species and sexes in the family Restionaceae along a fine‐scale hydrological gradient

Consistent, repeatable segregation of plant species along hydrological gradients is an established phenomenon that must in some way reflect a trade‐off between plants' abilities to tolerate the opposing constraints of drought and waterlogging. In C₃ species tissue carbon isotope discrimination (δ¹³C) is known to vary sensitively in response to stomatal behaviour, reflecting stomatal limitation of photosynthesis during the period of active growth. However, this has not been studied at fine‐spatial scale in natural communities. We tested how δ¹³C varied between species and sexes of individuals in the family Restionaceae growing along a monitored hydrological gradient. Twenty Restionaceae species were investigated using species‐level phylogeny at two sites in the Cape Floristic Region, a biodiversity hotspot. A spatial overlap analysis showed the Restionaceae species segregated significantly (P < 0.001) at both sites. Moreover, there were significant differences in δ¹³C values among the Restionaceae species (P < 0.001) and between male and female individuals of each species (P < 0.01). However, after accounting for phylogeny, species δ¹³C values did not show any significant correlation with the hydrological gradient. We suggest that some other variable (e.g. plant phenology) could be responsible for masking a simple response to water availability.     

Light microscopic detection of PAS-positive substances with thiosemicarbazide in freeze-substituted ovaries of Paspalum longifolium before pollination

Summary Localization of polysaccharides in the freeze-substituted, Eponembedded ovaries of Paspalum longifolium prior to pollination was carried out by periodic acid-Schiff's (PAS), periodic acid-thiosemicarbazide-silver proteinate (PA-TSC-AgPr) and periodic acid-thiosemicarbazide-osmium (PA-TSC-OsO₄) reactions. The specificities of these three reactions were also studied. These three reactions are all effective for light microscopic demonstration of polysaccharides in the filiform apparatus, starch grains in the cells and PAS substance in the micropylar region. Nonspecific staining of the nucleoli of the egg and polar nuclei was observed in the PAS reaction. The PA-TSC-AgPr reaction is very specific for polysaccharides but its overall reaction takes a much longer period of time than the PAS reaction. The PA-TSC-OsO₄ reaction colors the cytoplasm and nuclei of most cells and therefore stains of the cell walls, especially those of the egg cell and synergids, do not stand out clearly. The synergid cytoplasm contains some amorphous polysaccharides and thus it colors even in PAS and PA-TSC-AgPr preparations. In the mature embryo sac, the egg and central cell as well as antipodals are vacuolated but the two synergids have no visible vacuoles under light microscope. Each synergid has a prominent filiform apparatus at the micropylar end, which stains intensely in all three preparations. The walls of the central cell and antipodals adjacent to the nucellar cells have many inward papillae which are also intensely stained in all three preparations. Starch grains are abundant in the ovary wall and usually absent in the nucellus and integuments. They are present in the egg, central cell and antipodals, but not in the two synergids.     

CHARACTERIZATION OF INVASIVE CONYZA SPECIES (ASTERACEAE) IN EUROPE: QUANTITATIVE TRAIT AND ISOZYME ANALYSIS

The genus Conyza (Asteraceae) represents one of the foremost examples of intercontinental plant invasions from the New World to the Old World that has resulted in a number of taxonomic problems owing to the dearth of knowledge concerning the biosystematics of the genus. In this study, quantitative trait and enzyme electrophoresis analysis was used to assess the pattern of variation within and among five short-lived taxa of Conyza (C. blakei, C. bonariensis, C. canadenis, C. floribunda, C. sumatrensis), introduced into Europe, and to test the hypothesis of a recent hybrid origin of C. floribunda. The five taxa exhibit marked differences in morphological and life-history characteristics that are concordant with divergence in genes encoding isozymes. In addition, a recent hybrid origin of C. floribunda is clearly not supported by either quantitative or electrophoretic data. These findings suggest that the five taxa differentiated genetically in their native range prior to their introduction in Europe. As formerly suggested by Arthur Cronquist (Bulletin of the Torrey Botanical Club 70: 629–632, 1943), C. canadensis appears to be more closely related to the genus Erigeron than the other taxa. The electrophoretic results also confirm that this particular species is diploid, while the presence of multiple bands and fixed heterozygosity demonstrates that the other Conyza taxa are all genetic allopolyploids. Both quantitative and electrophoretic data suggest the presence of two groups among the polyploid taxa: one comprising C. bonariensis and C. sumatrensis, the other C. blakei and C. floribunda. The latter grouping is supported by the observation that C. blakei and C. floribunda, both native to Argentina, exhibit a striking ability to shift from semelparous to iteroparous reproduction, while the other taxa are strictly semelparous.     

MEGASPOROGENESIS AND MEGAGAMETOGENESIS IN SOYBEAN,GLYCINE MAX

Megasporogenesis and megagametogenesis were examined in Glycine max with light, fluorescence, and electron microscopy. Megasporogenesis results in a linear tetrad of four megaspores. Megagametophyte development is of the Polygonum type, with the functional chalazal megaspore undergoing three successive mitotic divisions to produce an eight-nucleate, seven-celled mature megagametophyte. The central cell becomes packed with starch. At fertilization, the antipodals are degenerate, the polar nuclei have fused, starch is diminished, and the egg occupies most of the micropylar portion of the megagametophyte. Several pollen tubes were occasionally observed at each micropyle, yet only one was involved in fertilization. Pollen tube entry occurs through a slightly reduced, viable synergid cell. Endosperm development precedes embryo growth. These results describing normal development allow important comparison with genetic mutants of soybean that affect female fertility.     

Floral Ontogenetic Evidence in Support of the Willdenowia Clade of South African Restionaceae

Willdenowia clade of Restionaceae was studied to understand patterns of reduction of floral elements and sample evidence for discussing the relationships of the group. All species studied are characterized by a concordant reductive trend involving the retardation/reduction of the perianth, the loss of the anterior carpel and the displacement of the remaining carpels, linked with a strongly compressed spikelet. Different modes of carpel reduction, such as a progressive or immediate loss, or fusion of two neighboring carpels, are presented and discussed. The most parsimonious event of gynoecium evolution for the Willdenowia clade is either the sterilization of two carpels in an originally trimerous gynoecium, followed by the loss of the anterior carpel, or the sudden loss of the anterior carpel, preceeding the sterilization of one lateral carpel. The concordant development of the taxa of the Willdenowia clade supports a one-time loss of a carpel and the homogeneity of the clade. Received 12 March 2001/ Accepted in revised form 29 May 2001     

New hyphomycetes from Restionaceae in fynbos: <Emphasis Type="Italic">Parasarcopodium ceratocaryi</Emphasis> gen. et sp. nov., and <Emphasis Type="Italic">Rhexodenticula elegiae</Emphasis> sp. nov.

Two new hyphomycetous anamorphs were isolated from plant hosts of the Restionaceae in the fynbos of the Cape Floral Kingdom of South Africa. Parasarcopodium ceratocaryi gen. et sp. nov. on Ceratocaryum decipiens produces aseptate, cylindrical conidia with amorphous mucoid appendages at both ends in rows or whorls of phialides on verruculose, mononematous conidiophores. Rhexodenticula elegiae sp. nov. on Elegia capensis produces pale brown, fusiform or broadly fusiform, 3-septate conidia on denticles located in the upper part of smooth, geniculate conidiophores. The morphological characteristics of both taxa are described, illustrated, and contrasted to other morphologically similar fungi. To elucidate the phylogenetic relationships of Parasarcopodium, partial large subunit rRNA (28S) gene sequence was generated from the ex-type strain, which revealed it to be a member of the Bionectriaceae (Hypocreales).     

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     

CHARACTER DIAGNOSIS, FOSSILS AND THE ORIGIN OF TETRAPODS

I. The traditional view of the origin of tetrapod vertebrates is that they are descendants of fossil osteolepiform fish, of which Eusthenopteron is best known. In recent years both that conclusion and the methodology by which it has been reached have been challenged by practitioners of cladistic analysis. Particularly a recent review by Rosen et al. (1981) claims that Dipnoi (lungfish) are the sister-group of the Tetrapoda, that Osteolepiformes is a non-taxon and that Eusthenopteron is more distant from tetrapods than are Dipnoi, coelacanths and probably the fossil Porolepiformes. We attempt to refute all these concludions by use of the same cladistic technique. 2. We accept that all the above-mentioned groups, together with some less well-known taxa, can be united as Sarcopterygii by means of shared derived (apomorph) characters. We also agree that Porolepiformes and Actinistia (coelacanths) can be characterized as valid taxa. The primitive and enigmatic fossil fish Powichthys is accepted as representing the plesiomorph sister-group of true porolepiforms. 3. Only two apomorph features, the course of the jaw adductor muscles and the position of incurrent and excurrent nostrils, appear to unite all the fish, living and fossil, currently regarded as Dipnoi. The characteristic tooth plates and the presence of petrodentine both exclude important primitive fossil forms. 4. Contrary to the opinion of Rosen et al., Osteolepiformes can be characterized — by the arrangement of bones forming the cheek plate, the presence of basal scutes to the fins and by the unjointed radials of the median fins. However, if these are true autapomorphies they exclude any osteolepiform from direct tetrapod ancestry. 5. Tetrapoda is a monophyletic group characterized by ten or more autapomorphies, including the bones of the cheek plate, a stapes and fenestra ovalis, and a series of characters of the appendicular skeleton. 6. Tetrapods have a true choana (internal nostril). We accept that the posterior (excurrent) nostril of Dipnoi is the homologue of the tetrapod choana. However, we assert that the posterior nostril of all bony fish is the homologue of the choana. This assertion would be refuted if any fish showed separate posterior nostril and choana. We reject the claim that this ‘three nostril condition’ occurred in porolepiforms and osteolepiforms. The evidence for a choana in porolepiforms is inadequate. Osteolepiforms had a true choana, characterized as in tetrapods by its relationship to the bones of the palate, but no third nostril. Dipnoans are not choanate. 7. Following cladistic practice, the relationship of the extant taxa is established first. Dipnoi are thus shown to be the living sister-group of tetrapods, but only on ‘soft anatomy’ characters unavailable in fossils. Coelacanths are the living sister-group of the taxon so formed. 8. The relationship of the fossil taxa to the extant sarcopterygians is then considered. The synapomorphy scheme proposed by Rosen et al. is discussed at length. Virtually all the characters they use to exclude close relationship of Eusthenopteron (and hence all osteolepiforms) to tetrapods, in favour of coelacanths and dipnoans, are invalid. 9. A series of synapomorphies uniting osteolepiforms and tetrapods is proposed, including a true choana (hence the taxon Choanata), the histology of the teeth, and a number of characters of the humerus. The recently discovered fossil Youngolepis, which lacks a choana, represents the sister-group of the Choanata, and is not uniquely close to Powichthys. The latter, as a porolepiform (s.l.) is a member of the sister-group to Choanata plus Youngolepis. 10. Our cladistic analysis suggests that all the extinct taxa considered are more closely related to tetrapods than are the Dipnoi. Moreover fossil evidence suggests that Dipnoi, considered as an extant taxon, may not even be the living sister-group of Tetrapoda. Early fossil dipnoans appear to have been marine fish without specific adaptations for air breathing. If so the apparent synapomorphies of Dipnoi and Tetrapoda may be homoplastic — the insistence on grouping extant taxa first would then have yielded an invalid inference.     

Stimulation of seed germination in South African species of Restionaceae by plant-derived smoke

The Restionaceae is one of the three major families defining fynbos, the characteristic vegetation type of the Cape Floristic Region. Periodic fires with a frequency of 5 to 40 years are a natural phenomenon in fynbos vegetation. Fire-stimulated seed germination has been reported for a variety of fynbos species, and species in the Asteraceae, Ericaceae, Proteaceae and Restionaceae have shown a germination response to smoke and/or aqueous smoke extracts. In the present study seed of 32 species was screened to obtain an indication of how important the smoke cue is for germination in the South African Restionaceae. The results of the present study represents the first occasion that comparative germination data for South African species in this family have ever been obtained. Twenty-five of the 32 species tested showed a statistically significant improvement in germination following smoke treatment. Untreated seeds of 18 of the species responding, showed a high degree of dormancy with only 0.1% to 2.0% germination. These results suggest that under natural conditions smoke from fynbos fires may provide an important cue for triggering seed germination in this family. The degree of improvement in germination following smoke treatment ranged from 147% in the case of Restio festuciformis to 25300% in the case of Rhodocoma capensis. It is suggested that the 16 species which showed a 1000% or more increase in germination following smoke treatment form a group in which smoke is likely to be the major cue for germination. In those species in which there is a lesser response, smoke may be one of a number of germination cues which include heat, and possibly alternating high and low incubation temperatures. The four species that did not germinate were all myrmecochorus, nut-fruited species. More information is needed concerning the fire survival strategy of South African Restionaceae species and many more will have to be investigated in the nursery and in the field, before the full pattern of response within the family and its significance can be elucidated.