New species of Polyalthia (Annonaceae) from Borneo and their relationship to Polyalthia cauliflora

A group of seven Polyalthia species from Borneo allied to P. cauliflora Hook. f. & Thomson is revised. Three of these are described as new. Polyalthia myristica sp. nov. and P. trochilia sp. nov. are similar to P. stenopetala (Hook. f. & Thomson) Finet & Gagnep., particularly in having seeds with a deep circumferential groove. Polyalthia kinabaluensis sp. nov., endemic to Mount Kinabalu, is similar to P. cauliflora, but differs in having petals slightly connate at the base and larger monocarps. Polyalthia ichthyosma sp. nov. is the fourth new species described. It bears some similarity to one of the P. cauliflora group, P. dictyoneura Diels, but differs in not being cauliflorous. Two synonyms of Polyalthia cauliflora, Unona desmantha Hook. f. & Thomson and Unona pycnantha Hook. f. & Thomson, are lectotypified.     

A new species of Polyalthia (Annonaceae) from Meghalaya, India

A new species, Polyalthia meghalayensis (Annonaceae) is described with illustrations from the Garo Hills of Meghalaya, India.     

Floral ontogeny of five species ofTalinum and of related taxa (Portulacaceae)

The floral development of five species ofTalinum is studied. Each flower is surrounded by two involucral bracts. The perianth consists of five tepals initiated in a 2/5 phyllotaxis. In all species studied a first whorl of 10–13 stamens is initiated, except inT. napiforme where this whorl is reduced to five stamens. In multistaminate androecia, additional whorls develop centrifugally. InT. paniculatum, T. portulacifolium andT. napiforme the first stamens are initiated in pairs opposite the outer tepals. In several flowers ofT. paniculatum andT. portulacifolium ten stamens are incepted in spiral sequence resembling diplostemony. Similar ontogenetic patterns are present in several species ofPhytolacca. However, within the genusTalinum the ontogenetic pattern of the firstly initiated stamens is not consistent with traditional diplostemony. InT. triangulare the firstly initiated stamens are incepted in sectors on a ring meristem, resembling the early inception in several species ofAnacampseros andPortulaca. The nectaries are associated with the filament bases and can be defined as caducous nectaries of the staminal type. The development of the tricarpellate, syncarpous gynoecium is very similar in all species studied; it is characterised by a leptate carpel-form.     

Floral ontogeny of Cneorum tricoccon L. (Rutaceae)

The floral ontogeny of the Spurge olive (Cneorum tricoccon L.) is studied by means of scanning electron microscopic observations. Special attention is paid to the sequence of initiation of the floral parts, the occurrence of septal cavities, and the development of the nectariferous tissue. The nectary disc arises as a receptacular outgrowth below the ovary and independently from stamen development. By the extensive growth of this voluminous androgynophore, stamen filaments become enclosed by nectary tissue and as a result, they are seated in pits between the lobes of the disc. Between ovary and style, three lobes are present, which are covered with stomata – their function is unknown. The significance of the unusual trimery of the flower is discussed. Floral developmental evidence supports a Rutalean affinity, although more ontogenetic investigations are needed in Rutaceae, subfamily Spathelioideae.     

Floral traits, pollinators and breeding systems in Syncolostemon (Lamiaceae)

Differentiation in floral traits among clusters of related species may reflect a process of pollinator-driven evolution. Pollination systems in the morphologically diverse southern African genus Syncolostemon (Lamiaceae) were investigated by means of field observations of floral visitors and analysis of their pollen loads. Among the five study species, those with short corolla tubes (S. parviflorus, S. ramulosus) were pollinated solely by bees, while those with long corolla tubes were pollinated by a broader array of visitors, primarily long-proboscid flies in S. rotundifolius and S. macranthus and sunbirds in S. densiflorus. The predominately insect-pollinated taxa have lax inflorescences, but S. densiflorus has a compact terminal inflorescence, which facilitates feeding by sunbirds from a single perching position. Experimental hand-pollinations involving three taxa (S. macranthus, S. densiflorus, and S. rotundifolius) showed that these possess a genetic self-incompatibility system. Production of fruits and seeds per fruit was pollen-limited in S. densiflorus and S. rotundifolius, but not in S. macranthus.     

Floral ontogeny, pattern formation, and evolution in Hibbertia and Adrastaea (Dilleniaceae)

Floral development was compared with scanning electron microscopy in 12 Australian species of Hibbertia representing most of its morphological variation, and in the related Adrastaea (Dilleniaceae). Calyx and corolla arise in quincuncial helices in radially symmetrical species, while the petals initiate unidirectionally from one side in zygomorphic species. Stamen number (3-200+) proliferates by centrifugal addition of individual primordia or by innovations of common primordia and ring meristems. Common primordia arise in single-stamen positions alternately with petals, and each produces one to several stamens centrifugally that remain attached to a shared base and form a stamen fascicle. A ring meristem in Adrastaea initiates a whorl of five stamens, alternate with the first stamens but outside their whorl. In radially symmetrical species of Hibbertia, a first ring of stamens is supplemented centrifugally by additional stamens on a meristem ring. The first stamens in zygomorphic species of Hibbertia initiate as a terminal ridge on the floral apex, with subsequent stamens added centrifugally on one side and two carpels initiated on the opposite side. The carpels arise as a simultaneous ring in radially symmetrical flowers, or as a simultaneous pair in zygomorphic species. Staminodial presence is viewed as of minor significance. Four pollinator syndromes are proposed for Hibbertia, related to differing floral architecture.     

The breeding systems of six species of Arabis (Brassicaceae)

The ability of organisms to produce genetic variation for any trait, including resistance to pathogens, is partially determined by breeding system. I used enzyme electrophoresis, crossing experiments, and cytology to assess the breeding systems of cooccurring Arabis species that are often infected by rust fungi. The Arabis holboellii surveyed were pseudogamous apomicts with relatively high population-level ailozyme variation, and variable chromosome numbers (2n, 3n, 4n). Arabisgunnisoniana (3n) and Arabis lignifera (2n) were also pseudogamous but showed no allozyme variation either at the population level, or within progeny arrays. Arabis hirsuta may be an autogamous polyploid or it may be a pseudogamous apomict; more work is needed to clarify the breeding system of this species. Arabis drummondii and Arabis crandallii were sexual, but exhibited little genetic variability due to a predominance of self-fertilization. The use of several techniques was necessary to evaluate these breeding systems. Insect exclusion indicated when pollen was necessary for seed set, but could not differentiate between sexual reproduction and pseudogamy. Electrophoresis yielded information on the degree of selling (as evidenced by homozygosity) and apomixis (fixed heterozygosity), but could not differentiate between autogamy and apomixis in polyploids without ailozyme variation. Pseudogamy was confirmed when crosses between dissimilar genotypes yielded only the maternal genotype, and cytologically by irregular meiosis.     

异果芥的花部综合征及其繁育系统

异果芥Diptychocarpus strictus是十字花科中为数不多的在同一花序上具有两种不同形态果实的短命植物。对其花部综合征及繁育系统的研究结果表明：（1）具有白色花与紫红色花两种不同花色的植株,且其花色是稳定遗传的。两种植株的比例基本上接近1：1。（2）两种植株花序上、下部发育成不同形态果实的花在形态特征上存在差异,同时不同花色植株的花在形态特征上也存在差异。（3）紫红色和白色花植株花序上、下部花的单花花粉量分别为7589.07±976.64、5428.93±900.45、7044.73±557.37和5138.60±813.48;胚珠数分别为31.83±3.36、25.03±3.72、32.03±3.43和25.70±2.61;P/O值分别为241.31±41.85、222.33±52.51、222.19±27.80和201.84±37.52。花粉败育率很低且两种花色植株花序上、下部花的长、短雄蕊之间的花粉败育率均差异显著（P〈0.05）。同一花中短雄蕊花粉量比长雄蕊的多。（4）两种花色植株花序上、下部花之间的长、短雄蕊每花药花粉数、单花花粉数、胚珠数均存在极显著差异（P〈0.01）,均是上部花比下部花的大。（5）两种植株花中长、短雄蕊花粉活力动态变化曲线相似,花粉寿命与柱头可授期具高度的同步性,其花粉活力最高的时期正是柱头的最佳可授期。（6）P/O值、套袋试验、传粉媒介观察和花粉萌发的荧光显微观察结果表明,该物种是以自交为主、异交为辅,具有兼性自交的繁育系统。     

Floral bract function, flowering process and breeding systems ofSarcandra andChloranthus (Chloranthaceae)

Structure and function of floral bracts, anthesis and breeding systems were investigated in greenhouse plants ofSarcandra chloranthoides, Sarcandra glabra, andChloranthus spicatus (Chloranthaceae). In early developmental stages the floral bract replaces the lacking perianth as a protective structure by enclosing the floral organs completely in a pocket-like structure, which is formed by the u-shaped attachment zone of the floral bract to the spike axis. The floral bract has a tip with an epithem tissue, from which traces of secretion seem to be released by the stomata. All species investigated have protogynous and longlasting flowers. The female phase begins five to seven days earlier than the male phase. It continues during the male phase and for some time thereafter. The male phase lasts between one day (Chloranthus spicatus) and seven days (Sarcandra chloranthoides). This is the first study on breeding systems of Chloranthaceae. They are of special interest in such a basal family of the angiosperms. Although the investigatedSarcandra andChloranthus species have similar flowers with entomophilous features, surprisingly their breeding systems are diverse:Sarcandra glabra is self-compatible,S. chloranthoides is agamospermous andChloranthus spicatus is self-incompatible.     

Floral ontogeny of Ruteae (Rutaceae) and its systematic implications

Floral development was investigated in Ruta graveolens and Psilopeganum sinense, representing two genera in the tribe Ruteae. Special attention was paid to the sequence of initiation of organ whorls in the androecium and gynoecium. The antepetalous stamens arise at the same level as the antesepalous stamens in both species. The carpels are antepetalous in both taxa, indicating the androecium in both genera is obdiplostemonous. Compared with floral ontogeny of the ancestral genus Phellodendron (Toddalioideae), the obdiplostemonous androecium is a derived condition. The floral apex in P. sinense is quadrangular before initiation of the two carpels. Additionally, there are four dorsal and four ventral traces in the ovary. Integrated morphological and anatomical evidence indicates that the bicarpellate gynoecium in Psilopeganum most likely evolved from a tetracarpellate ancestor. Considering the similarities in morphological, geographical and chromosomal features, the ancestor may be Ruta‐like. Further molecular phylogenetic and genetic studies are needed to verify this assumption.     

Contrasting breeding systems in twoEriotheca (Bombacaceae) species of the Brazilian cerrados

The pollination biology and breeding systems ofEriotheca pubescens andE. gracilipes have been studied. These two species occur as trees in cerrado vegetation, the neotropical savannas of Central Brazil, with partially sympatric distributions. They have similar phenology and floral structure, although the flowers ofE. pubescens are larger. Both species have nectar flowers pollinated by largeAnthophoridae bees but the main pollinators of each species differ in size. The species have markedly different breeding systems: late-acting self-incompatibility inE. gracilipes and apomixis stimulated by pollination inE. pubescens.     

Floral organogenesis and floral evolution of the Lecythidoideae (Lecythidaceae)

The subfamily Lecythidoideae of Lecythidaceae (Brazil nut family) is a dominant group in neotropical forests, especially those of Amazonia. New World members of the family have large showy flowers that are either polysymmetric or monosymmetric. In this study, floral organogenesis of all 10 neotropical genera was examined using SEM. Our observations of floral development are put into the context of a molecular phylogeny based on sequences of the ndhF and trnL-F genes (Am. J. Bot. 94: 289-301). Floral evolution of the subfamily is explained as having undergone four different levels of complexity in regard to floral symmetry. The basal most genera, Grias and Gustavia, have polysymmetric flowers. At level two, represented only by Couroupita, monosymmetry is established through the expression of abaxial dominance and the development of an androecial hood; at this level, abaxial dominance impacts the perianth and androecium, but not the gynoecium. At the third level, monosymmetry is developed in groups of Couratari and Cariniana domestica; but, in the Allantoma/Cariniana decandra lineage, a reversal back to polysymmetric flowers, resulting from a gradual weakening of abaxial dominance, and the loss of the hood has occurred. Finally, in level four, including Bertholletia, Corythophora, Eschweilera, and Lecythis, monosymmetry is so strongly expressed that the gynoecium is also influenced by abaxial dominance. In this group, the hood is complicated in both structure and function, and the floral axis is changed from straight to slightly inclined. This study demonstrates that the development of floral abaxial dominance is the proximate cause of monosymmetry in the Lecythidoideae. We suggest that monosymmetric flowers are more efficiently pollinated, and therefore the bees and bats that pollinate the monosymmetric flowers in this group are ultimately responsible for the monosymmetry.     

A new species of Annona (Annonaceae) from the northeastern Guayana Shield

Annona prevostiae, a new species from the northeastern margin of the Guayana Shield, is described and illustrated.     

Floral ontogeny in Sophoreae(Leguminosae: Papilionoideae). III. Radial symmetry and random petal aestivation in Cadia Purpurea

Floral organogeny and development are described in Cadia purpurea, a legume with radial symmetry, unstable petal aestivation, and free organs, all unusual features among papilionoids. Flowers are usually solitary or, rarely, in few-flowered racemes. No bracteoles are formed. The order of organ initiation is unidirectional in each whorl, and the carpel initiates directly after petal initiation.The petal primordia remain small until all other floral organs have initiated, enlarged, and differentiated. Petal aestivation is variable, unlike the great majority of papilionoid flowers. Petal margins of Cadia grow straight outward rather than some petal margins curving inward inside the adjacent petal as is the case in other papilionoids. When adjacent growing petal margins of Cadia meet, chance determines which petal becomes positioned inside the other. Hence, the pattern of petal aestivation is random.     

Floral biology of the dioecious species Laurus nobilis L. (Lauraceae)

The present study examines the cytological, physiological, chemical and ecological characteristics of pollen and nectar offered by male and female flowers of the dioecious plant Laurus nobilis. The various phases of floral phenology and the insect pollinators were observed. We used cytological methods to determine anther, pollen and nectary structure. Nectar sugar composition was evaluated by HPLC. Pollen viability in time was compared with cytoplasmic and intine water content. Pollen presentation was found to be reversible by opening and closing of anther valves, determined by hydration of the mechanical layer of the anther. Pollen, covered by pollenkitt, was presented for dispersal for 3 consecutive days and during this time the intine and cytoplasm lost water and pollen viability diminished. At germination exine ruptured together with the outermost layer of the intine. Nectaries of male flowers were observed on the anther filament and on staminodes of female flowers. The nectar consisted almost entirely of sucrose and was more concentrated in male flowers. Secreted through stomata, nectar was presented in a thin layer. In the study area, the main pollinators (about half the total number of all visits) were hymenopterans. Pollen is of the recalcitrant type due to its high water content (>30%) but its viability is long-lasting because the intine is thick and stores water, keeping the cytoplasm of the vegetative cell hydrated and viable, and because anther valves may close under adverse conditions, protecting the pollen. Insects are attracted by male and female flowers similarly, males offer nectar and pollen, whilst females only nectar.     

Comments on flower evolution and beetle pollination in the generaAnnona andRollinia (Annonaceae)

Dynastid scarab beetle pollination appears basic within the genusAnnona. Those species ofAnnona which are more morphologically derived, as well as allRollinia spp. possess reduced floral chambers and attract small beetles likeNitidulidae orStaphylinidae. Pollination of the primitive species ofAnnona byDynastinae would imply that the genus had not evolved before the Tertiary. The fossil record is in congruence with this hypothesis. Once again it is stressed that the cantharophilous syndrome, as it is found in theAnnonaceae, Magnoliaceae, Eupomatiaceae andCalycanthaceae, with beetles being exclusive pollinators, is a secondary and derived condition and obviously different from the expected basic entomophily of the original angiosperms.