Fruit and seed ontogeny related to the seed behaviour of two tropical species of Caesalpinia (Leguminosae)

Caesalpinia echinata and C. ferrea var. ferrea have different seed behaviours and seed and fruit types. Comparison of the seed ontogeny and anatomy partly explained the differences in seed behaviour between these two species of Brazilian legumes; some differences were also related to fruit development. The seed coat in C. ferrea consisted of two layers of osteosclereids, as well as macrosclereids and fibres, to form a typical legume seed coat, whereas C. echinata had only macrosclereids and fibres. In C. echinata , the developing seed coat had paracytic stomata, a feature rarely found in legume seeds. These seed coat features may account for the low longevity of C. echinata seeds. The embryogeny was similar in both species, with no differences in the relationship between embryo growth and seed growth. The seeds of both species behaved as typical endospermic seeds, despite their different morphological classification (exendospermic orthodox seeds were described for C. echinata and endospermic orthodox seeds for C. ferrea ). Embryo growth in C. ferrea accelerated when the sclerenchyma of the pericarp was developing, whereas embryonic growth in C. echinata was associated with the conclusion of spine and secretory reservoir development in the pericarp. Other features observed included an endothelial layer that secreted mucilage in both species, a nucellar summit, which grew up into the micropyle, and a placental obturator that connected the ovarian tissue to the ovule in C. ferrea . © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 57–70.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

Morphology of fruits, seeds and embryos of Argentinian Capparis L. (Capparaceae)

Embryos, seeds and fruits of four species of Capparis L. (Capparaceae) from Argentina are described in this paper: C. flexuosa (L.) L. sensu lato, C. retusa Griseb., C. speciosa Griseb. and C. tweediana Eichl. Chlorophyllous embryos are found in C. flexuosa and C. retusa , species with thin seed coats, dehiscent siliques, and somewhat fleshy red endocarps. Nonchlorophyllous embryos are found in the berry-fruited C. speciosa and C. tweediana , the former with leathery indehiscent fruit wall, the latter with thin-walled dehiscent berries with massive fleshy pulp. The pulp is permeated by seed coat hairs in C. tweediana . The embryos of C. flexuosa , C. retusa and C. tweediana belong to the 'Axile Division, Foliate Subdivision, and Folded Type' of the classification devised by A. C. Martin in 1946. Some features of the embryo of C. speciosa (which are not considered in Martin's classification), support a new subdivision (Massive) and a new morphological type (Unequal cotyledons), characterized by profound anisocotyly, globose shape of the major cotyledon, presence of suberized cells and an undeveloped radicle. Anatomical studies of embryos and seedlings confirm the anisocotyly.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 209−218.     

Abortive seed development in Ulmus minor (Ulmaceae)

Three genotypes of field elm ( Ulmus minor ) were studied to determine the structural basis of seed abortion in this species. In the non-abortive control, P-VV1, the pattern of seed development is similar to many flowering plants. The embryo progresses through defined morphological stages leading to developmental arrest as the seed matures. Storage products are abundant within embryo cells. Endosperm development is similar to the nuclear type; however, a more extensive cellularization of the endosperm occurs prior to it being crushed by the expanding embryo. For the abortive genotypes, M-SF1 and V-JR1, abnormalities in endosperm development are found. This is judged by the early cellularization and the massive synthesis of the PAS-positive material in the cellular endosperm. In these abortive genotypes, embryo development is delayed and storage products failed to accumulate within embryo cells. After seed desiccation, no living embryo tissue remains within the seed coat in the abortive genotypes.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 455–467.     

An unusual case of epigeal cryptocotylar germination in Rollinia salicifolia (Annonaceae)

An unusual case of epigeal cryptocotylar germination found in Rollinia salicifolia Schltdl. (Annonaceae) is described and is discussed in relation to the terminology used for germination studies. The terms cryptocotylar and phanerocotylar are used to describe the permanence of cotyledons within the seed coat or endocarp; the terms epigeal and hypogeal are used to refer the position of cotyledons in relation to ground level. The terms to describe position about ground level and permanence of cotyledons within the seed coat or endocarp are not used simultaneously in many papers, and some authors consider the terms cryptocotylar and hypogeal to be synonyms. Cases of epigeal cryptocotylar germination seem to be scarce in the literature. The morphological characters shown by R. salicifolia seeds and seedlings − a non-chlorophyllous embryo, abundant endosperm, thick woody seed coat, thickened hypocotyl and non-photosynthetic haustorial cotyledons − are closely related to its epigeal cryptocotylar germination. This is the first record of this type of germination in Argentina.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 53–56.     

Seed dormancy in relation to seed storage behaviour in Acer

Dormancy in seeds of Acer opalus is shown to be mainly caused by the seed coats, although a slight embryo dormancy exists in fresh seeds. The ability to germinate after drying indicates that seed storage behaviour is orthodox. Recalcitrant seeds were heavier than orthodox seeds not only within section Acer but also within the whole genus after statistical control of phylogeny, through a phylogenetic ANOVA with data from two different Acer phylogenies. An evolutionary change from orthodox to recalcitrant behaviour is postulated for genus Acer , but this change appears not to have been accompanied by a change in seed dormancy, at least in the taxonomic section in which Acer opalus belongs.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 203–208.     

Seed micromorphology in the genus Neotinea Rchb.f. (Orchidaceae, Orchidinae)

The ultrastructure of testa seed in the genus Neotinea (Orchidaceae, Orchidinae) was examined for the first time. The morphology of the seed and of the anticlinal and periclinal walls was analysed by scanning electron microscopy. Quantitative data concerning the length and width of the seed and embryo, seed and embryo volume, free air space, and number of cells along the longitudinal axis are presented. In all species, the seeds are fusiform in shape with transverse ridges on the inner periclinal walls. This ornamentation pattern is characteristic for the genus Neotinea . It is a good diagnostic value supporting the monophyly of this genus, which has recently been proposed by several authors.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 133–140.     

Seed size variation within Zostera marina L. (Zosteraceae)

In 1997 and 1998 seeds were collected from the seagrass, Zostera marina L. (Zosteraceae) from Long Island, New York, USA (var. typica ), Shaw Island, Washington, USA (var. latifolia ) and Izembek Lagoon, Alaska, USA (var. izembekensis ) to determine if seed size varied between the varieties, if seed size varied within the same variety, and if the relationship between seed length and seed weight was linear. Measurements of seed length revealed statistically significant differences in this metric between varieties associated with smaller intertidal plants (var. typica and var. izembekensis ) and larger plants from the subtidal region (var. latifolia ). It was also found that seed length varied within the same variety and verified that the cross-sectional shape of these seeds was primarily elliptical. Using seed cross-sectional area and weight from a randomly selected batch of seeds from the three locations sampled in 1998, a linear model was created to determine seed weight and seeds from var. latifolia were found to be heavier than seeds from var. typica . The potential advantage of heavy seeds for subtidal seeds in search of safe sites is discussed, together with the merit of using the model to calculate the weight of other elliptical seeds within the Zosteraceae and other seagrass families.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 281–288.     

Frequency-dependent seed size selection on Cryptocarya alba (Mol.) Looser (Lauraceae): testing the effect of background

Seed size affects the probability of seed predation. Large seeds should be preferred over small seeds but the selective responses of seed predators may also be frequency-dependent. Seed predators may prefer the most common seed sizes (apostatic selection), the rarest ones (antiapostatic selection) or even be unresponsive to the size of seeds. Moreover, seed density may further modify the selective responses of seed predators. We expect that at a low seed density seed consumption should be concentrated on common seed sizes, and at high seed density it should be concentrated on rare seed sizes, as common ones act as a background that makes rare phenotypes more conspicuous (the effect of background). We tested this prediction in a field experiment with seeds of Cryptocarya alba (Lauraceae) at La Campana National Park, central Chile. We presented large and small seeds in two contrasting seed densities (ten and 100 seeds per m²) and at five frequencies of large seeds: 0.1, 0.25, 0.5, 0.75 and 0.9. After 28 days we measured the proportions of the two size classes in the remaining seeds. Large seeds were always preferred to small seeds, even when they were at low frequency. Contrary to the predictions, at low seed density consumption was antiapostatic whereas at high seed density selection was independent of frequency. We discuss the causes and consequences of such selective responses expressed by seed predators.  © 2005 The Linnean Society of London , Biological Journal of the Linnean Society , 2005, 84 , 137–142.     

Flowering phenology and reproductive biology of Drosera anglica (Droseraceae)

Prompted by the sparse knowledge of the reproductive biology of carnivorous plants, compared with studies of their trapping habits, we investigated the flowering phenology and pollination biology of Drosera anglica Huds. in two fens in mid-western Canada. Seed set and germination were used to compare the effectiveness of a series of pollination treatments, including single insect visits to virgin flowers. Flowers opened during mid-morning but closed by early afternoon, and exhibited pseudo-cleistogamic behaviour in cool, overcast weather. D. anglica was found to be self-compatible, and able to self-pollinate and self-fertilize. Geitonogamy was an uncommon mode of self-reproduction because plants typically possessed a lone inflorescence upon which a single, short-lived flower opened, a few days before the next bud reached anthesis. Insect visits to the fragrance-lacking, nectarless flowers, chiefly by flies (Diptera: Syrphidae), were infrequent (one visit per 1 h 40 min of observation), and the low frequency of seed set and low numbers of seeds per fruit in pollination treatments involving insects, suggest the species does not rely on insects to effect pollination. Self-pollination, with or without the aid of a vector (insects, wind) was as effective as natural pollination; ultimately, autogamy is chiefly responsible for natural seed set. Thus, the species exhibits characteristics of facultative autogamy.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 417–426.     

Seed coat morphology of Aizoaceae–Sesuvioideae, Gisekiaceae and Molluginaceae and its systematic significance

Macro- and micromorphological characters of the seeds of 32 species belonging to Aizoaceae (26 species), Gisekiaceae (one species) and Molluginaceae (five species) were analysed for their taxonomic value. Seed morphology is found to be of considerable taxonomic value within the taxa investigated, although additional criteria are usually necessary for distinguishing the species. However, a subgroup of Trianthema , consisting of T. cussackiana , T. megasperma , T. pilosa , T. rhynchocalyptra and T. oxycalyptra var. oxycalyptra (Aizoaceae) shares the synapomorphy of seeds with scattered idioblast cells (papillae). There is little infrageneric variation within Sesuvium and Zaleya in contrast to Trianthema , whose seeds are considerably polymorphic. The fairly different seed structures found in Aizoaceae s.s . and Molluginaceae support their treatment as two distinct families. However, the position of Lineum (Molluginaceae) remains uncertain. Moreover, seed characters support the independence of Gisekia (Gisekiaceae) in a family of its own.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 189–206.     

Fruits and seeds of Ruppia (Potamogetonaceae) from the Pliocene of Yushe Basin, Shanxi, northern China and their ecological implications

Numerous fruits and seeds of Ruppia are reported from the Upper Pliocene (2.3–3.5 Myr ago) Zhangcun Formation in Yushe Basin, Shanxi, northern China. They are the first fossil Ruppia from China and demonstrate the importance of fruit and seed fossils in recording genera not represented by fossil leaves. These Ruppia are characterized by possessing a small oval endocarp, smooth endocarp surface, distinct elliptical external depressions, distinct apical mucro, slightly curved seed shape and conspicuous globose hilum. A new species, R. yushensis Zhao, Collinson and Li, is described from these endocarp and seed features. Comparison with the two European Miocene species, R. palaeomaritima Negru and R. maritime-miocenica Szafer, indicates the existence at that time of three different geographical and stratigraphical species. R. yushensis constitutes the first Pliocene record of Ruppia and extends the range of fossils of this genus from Europe to eastern Asia. R. yushensis is the only aquatic plant in the uppermost middle part of the Zhangcun Formation. This monotypic occurrence indicates a brackish, clear, tranquil and shallow lake in this region in the Late Pliocene. The smooth endocarp surface further suggests a warm temperate or temperate palaeoclimate.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 317–329.     

Multiple pleurograms in Chamaecrista Moench (Leguminosae, Caesalpinioideae)

According to the literature, the seeds of Chamaecrista lack pleurograms and have pitted testas aligned in vertical lines. Preliminary observations have revealed that these small structures consist of superficial alterations of the testa that are coloured differently to the rest of the seed coat and stand out as pits. However, no anatomical inspection has yet determined the precise composition of these pits. The present work examined the morphology, surface micromorphology, anatomy, and histochemistry of the seed coat of mature seeds of C. desvauxii var. latistipula , C. flexuosa , and C. nictitans var. patellaria in order to elucidate the structural nature of the seed coat pits. For this, seed coats were studied using scanning electron microscopy and light microscopy. Structural modifications were examined in the seed coats of the studied species, and variations were noted in the mucilaginous layer, in the thickness of the palisade layer, and in the presence of larger hypodermic cells. Based on the anatomical features observed, and comparing these with published data on pleurograms in the Caesalpinioideae, it was determined that the structures referred to previously as tegumental pits on Chamaecrista seeds are actually small and numerous pleurograms, reported here for the first time.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 487–492.     

Phylogeny of the Afro-Madagascan Aristea (Iridaceae) revisited in the light of new data on pollen morphology

Scanning and electron microscopic studies of the pollen grains of several species of the sub-Saharan African and Madagascan genus Aristea of c . 50 species, supplement data from an earlier study and enable us to include these species in a phylogenetic analysis. TEM examination of a range of pollen grains of representative pollen types in Aristea makes it possible to reinterpret past SEM results so that apertures covered with exine masses and previously called sulculate, are now seen to be either zonasulculate or dizonasulculate. Revised and expanded data are combined in a matrix together with new data for seed morphology in the genus, which like the capsules is remarkably diverse, including angular seeds with reticulate sculpturing, lamellate or triangular-columnar seeds with smooth or reticulate surfaces and with smooth or papillate or areolate margins. This new phylogenetic analysis suggests that Aristea should remain treated as comprising three subgenera. In subgenus Eucapsulares we add a new section Latifolieae for A. latifolia , which seems an isolated relic restricted to the mountains of the Cape Region of South Africa, an area of winter rainfall. Species with similar unspecialized capsule and seed morphology occur only in eastern south and tropical Africa and Madagascar, but they have derived pollen morphology whereas the pollen of A. latifolia is plesiomorphic. This species appears to be the closest living ancestor of the genus. In subgenus Aristea we continue to recognize three sections, sect. Racemosae , sect. Singulares and sect. Aristea. Nesting of subgenus Pseudaristea within section Racemosae of subgenus Aristea is probably an artefact of the analysis and does not receive bootstrap support, and hence we continue to recognize this group of species at subgeneric rank.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 41–68.     

Two new species from Gabon show the need to reduce Commitheca to the synonymy of Pauridiantha (Rubiaceae, Pauridiantheae)

Two new Pauridiantha species (Pauridiantheae, Rubiaceae) from Gabon are described and illustrated. Pauridiantha pleiantha is characterized by a combination of entirely glabrous stems, stipules with a narrow base, and large inflorescences. Pauridiantha smetsiana is characterized by entirely glabrous stems, narrow, early deciduous stipules, leaves red-tinged when dry, and small inflorescences. The two new species, especially P. smetsiana , close the morphological gap between Commitheca and Pauridiantha in combining typical Commitheca characters, such as glabrous stems and a narrow stipular base, with Pauridiantha characters, such as eucamptodromous venation. A detailed morphological comparison (including ovary structure, seed morphology, exotesta anatomy, and pollen morphology) between the two genera is given; in conclusion, Commitheca is relegated to the synonymy of Pauridiantha . The necessary new specific combinations are provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 105–117.     

Hesperis turkmendaghensis (sect. Hesperis) (Cruciferae/Brassicaceae), a new species from the Central Anatolia region, Turkey

Hesperis turkmendaghensis A.Duran & A.Ocak sp. nov. (Cruciferae) is described and illustrated from Anatolia, Turkey. The species grows under mixed forest, open forest and shady slopes in Türkmen Dağı (B3 Eskişehir) in Central Anatolia. It is closely related to H. matronalis L. ssp. matronalis , an endemic confined to Central Anatolia. Diagnostic morphological characters are discussed. Notes are also presented on its ecology, biogeography and conservation status. In addition, the pollen characteristics and seed coat surface of H. turkmendaghensis and H. matronalis are examined by SEM.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 239–247.     

Imbibition of Swietenia macrophylla (Meliaceae) seeds: the role of stomata

BACKGROUND AND AIMS: The occurrence of stomata in seed coats is uncommon and there is limited information about their function(s). The aim of this study was to verify the distribution of stomata in seed coats of Swietenia macrophylla and to relate it to the imbibition process and aspects of the structure of the outer integument layers. METHODS: For the structural and ultrastructural studies, the seeds were processed using the usual techniques and studied under light and scanning electron microscopes. Histochemical tests were employed to identify the cell wall composition in the different seed coat portions. To assess the role of the stomata in the imbibition, non-impervious seeds were compared with partially impervious ones, in which only the embryo, median or hilar regions were left free. Further, the apoplastic pathway marker was employed to confirm the role of the stomata as sites of water passage during imbibition. KEY RESULTS: A positive relationship was observed between seed coat thickness and stomata density. The stomata were devoid of movement, with a large pore. They occurred in large numbers in the embryo region and extended with lower frequency towards the wing. Imbibition rates were related to stomata density, suggesting that the stomata act as preferential sites for water entry in the S. macrophylla seeds. CONCLUSIONS: At maturity, the stomata in the seed coat play a significant role in seed imbibition. The data may also infer that these permanently opened stomata have an important role in gas exchange during seed development, aiding embryo respiration.     

A new species of Veronica L. (Scrophulariaceae) in the Cantabrian Range (Spain)

A new species of Veronica vadiniense in section Veronica is described from material from the Cantabrian Range, in the north-east of the province of León (north-west Spain). Several morphological characters related to stems, leaves, racemes, flowers and capsules are given, and also those related to the indumentum of different parts of the plant, especially calyx lobes and capsule. Indications about distribution and habitat with reference to bioclimatic, biogeographical and phytosociological aspects are provided. Diagnostic features are also given to enable comparison with the most similar species, Veronica officinalis and V. allionii , particularly in the indumentum and consistency of leaves, and in the indumentum and shape of the capsule.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 119–124.     

A new species of Achetaria (Plantaginaceae) from south-eastern Brazil

A new species, Achetaria latifolia V.C.Souza, is described and illustrated. It is characterized by its broadly ovate leaves, indument of stems and leaves of noncapitate hairs, and pubescent capsule. Achetaria latifolia is known only in the coastal dune-area of Cabo Frio region, Rio de Janeiro State, Brazil.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 73–75.