The systematic significance of morphological and anatomical variation in fruits of Crotalaria and related genera of tribe Crotalarieae (Fabaceae)

The phylogenetic and taxonomic significance of morphological and anatomical trends in fruits of tribe Crotalarieae has been evaluated, with emphasis on the genus Crotalaria and its seemingly distinctive, inflated and balloon‐shaped pods. In addition to the normal explosive dehiscence as a means of dispersal, several genera (including Crotalaria) show independent evolution of modifications apparently adapted for dispersal by wind, water and gravity. Transverse sections were made of mature pods of 142 species from the 12 currently recognized genera of Crotalarieae. The taxa differ in the orientation of the fibres (related to dehiscence or non‐dehiscence), the overall thickness of the fruit wall, the relative proportions of the pericarp layers, the degree of lignification and the presence or absence of trichomes. Three basic pericarp types can be distinguished: type I, with one, two or three zones of various numbers of cell layers of fibres (almost all genera); type II, with a single cell layer of fibres (only in Rothia, Robynsiophyton, Lebeckia and Lotononis sections Listia and Leobordea); and type III, with one zone of several cell layers of gelatinous fibres and multicellular trichomes associated with the endocarp (only in some species of Calobota and Wiborgiella). Considerable variation was encountered in the tribe, but Crotalaria appears to be rather uniform, with type I predominating. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 84–106.     

Fruit structure of Amborella trichopoda (Amborellaceae)

The indehiscent fruitlets of the apparently basalmost extant angiosperm, Amborella trichopoda, have a pericarp that is differentiated into five zones, a thin one‐cell‐layered skin (exocarp), a thick fleshy zone of 25–35 cell layers (outer mesocarp), a thick, large‐celled sclerenchymatous zone (unlignified) of 6–18 cell layers (middle mesocarp), a single cell layer with thin‐walled (silicified?) cells (inner mesocarp), and a 2–4‐cell‐layered, small‐celled sclerenchymatous zone (unlignified) derived from the inner epidermis (endocarp). The border between inner and outer mesocarp is not even but the inner mesocarp forms a network of ridges and pits; the ridges support the vascular bundles, which are situated in the outer mesocarp. In accordance with previous observations by Bailey & Swamy, no ethereal oil cells were observed in the pericarp; however, lysigenous cavities as mentioned by these authors are also lacking; they seem to be an artefact caused by re‐expanding dried fruits. The seed coat is not sclerified. The fruitlets of Amborella differ from externally similar fruits or fruitlets in other basal angiosperms, such as Austrobaileyales or Laurales, in their histology. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 148 , 265–274.     

A survey of ontogeny pericarp features as contribution to the infratribal characterization of Myrteae (Myrtaceae)

With the aim of correlating the pericarp structure with current phylogenies of Myrteae, this study describes the ontogeny in five species included in five out of the six South American clades of the tribe. In these taxa, the outer and inner ovarian epidermis gives rise to the exocarp and the endocarp, respectively, both with 1 layer. In the mesocarp, derived from the ovarian mesophyll, secretory cavities are arranged into a circle just below the exocarp and near the endocarp in Campomanesia adamantium; only below the exocarp in Eugenia pitanga and Myrcia multiflora; more internally in Myrciaria cuspidata, and below the exocarp and throughout the mesophyll in Myrceugenia alpigena. The promising traits for phylogenetic studies in the group include: direction of elongation of pericarp layers, regions that develop most in relation to the circle of larger vascular bundles, differentiation of spongy and sclerenchymatous tissues and position of secretory cavities.     

Structure of the unusual explosive fruits of the early diverging angiosperm Illicium (Schisandraceae s.l., Austrobaileyales)

All Illicium spp. have explosive fruits, which is a unique character among the basal grade of angiosperms. Illicium fruits consist of several ventrally dehiscing follicles developing from conduplicate carpels, with a prominent, slightly postgenitally fused ventral slit. The closure of the ventral slit is also secured by two mirror‐symmetrical massive longitudinal sclerenchymatous bands in the mesocarp along the edges and by turgor pressure. The pericarp differentiates into a fleshy (or coriaceous) peripheral zone (exocarp and mesocarp) with numerous ethereal‐oil‐containing cells and a sclerenchymatous (single‐layered, palisade) inner zone (endocarp). Dehydration of the fleshy zone of the pericarp and partial compression of the epidermal sclereids with U‐shaped wall thickenings lining the ventral suture are instrumental in explosive fruitlet dehiscence. Generally, the fruit structure of Illicium differs dramatically from those in other early diverging angiosperms. Gynoecium and fruit structure (and a probable early Cretaceous divergence from the Schisandra‐Kadsura clade) provide evidence for treatment of Illicium as separate from Schisandraceae s.s. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 640–654.     

Germination behaviour of Astroloma xerophyllum (Ericaceae), a species with woody indehiscent endocarps

The dispersal unit of many Ericaceae comprises an ovoid drupe with a woody indehiscent endocarp, and diaspores of this type are notoriously difficult to germinate for most members of this widely distributed family. Within the biodiverse south‐west of Western Australia, more than 200 drupaceous species of Ericaceae have been described, more than 50 of which are considered to be rare and threatened, requiring significant conservation action in the near future. In this paper, we investigate the germination ecology of the common Australian endemic, Astroloma xerophyllum, as a proxy for closely related threatened taxa, focusing on the ex situ and in situ germination requirements of seeds and indehiscent endocarps. Each endocarp possessed up to seven locules and means of 2.0–3.4 seeds per endocarp from the two collections used in this study. Seeds were up to 2.74 mm in length and 100% viable. Embryos were linear, differentiated and approximately 1.3 mm in length. Seeds within endocarps imbibed water to 28%, whereas excised seeds became hydrated to 44%. Fifty‐five per cent of seeds extracted from endocarps germinated on water agar alone and 100% germinated when presoaked in gibberellic acid. Seeds remaining inside intact endocarps failed to germinate unless treated with a germination promoter and incubated for more than 20 weeks. Rapid germination of seeds in intact endocarps was promoted by soaking endocarps in gibberellic acid and incubating them in 100% O₂. Embryos grew substantially in length within seeds prior to germination, and thus seeds have morphophysiological dormancy. Seeds under natural conditions required several seasons to germinate to any degree. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 299–311.     

A water drop-shaped slingshot in plants: geometry and mechanics in the explosive seed dispersal of Orixa japonica (Rutaceae)

Background and AimsIn angiosperms, many species disperse their seeds autonomously by rapid movement of the pericarp. The fruits of these species often have long rod- or long plate-shaped pericarps, which are suitable for ejecting seeds during fruit dehiscence by bending or coiling. However, here we show that fruit with a completely different shape can also rely on pericarp movement to disperse seeds explosively, as in Orixa japonica.MethodsFruit morphology was observed by hard tissue sectioning, scanning electron microscopy and micro-computed tomography, and the seed dispersal process was analysed using a high-speed camera. Comparisons were made of the geometric characteristics of pericarps before and after fruit dehiscence, and the mechanical process of pericarp movement was simulated with the aid of the finite element model.Key ResultsDuring fruit dehydration, the water drop-shaped endocarp of O. japonica with sandwich structure produced two-way bending deformation and cracking, and its width increased more than three-fold before opening. Meanwhile the same shaped exocarp with uniform structure could only produce small passive deformation under relatively large external forces. The endocarp forced the exocarp to open by hygroscopic movement before seed launching, and the exocarp provided the acceleration for seed launching through a reaction force.ConclusionsTwo layers of water drop-shaped pericarp in O. japonica form a structure similar to a slingshot, which launches the seed at high speed during fruit dehiscence. The results suggest that plants with explosive seed dispersal appear to have a wide variety of fruit morphology, and through a combination of different external shapes and internal structures, they are able to move rapidly using many sophisticated mechanisms.     

Pericarp formation in early divergent species of Arecaceae (Calamoideae,Mauritiinae) and its ecological and phylogenetic importance

Lepidocaryum tenue, Mauritia flexuosa and Mauritiella armata belong to the subtribe Mauritiinae, one early divergent lineage of the Arecaceae and one of the few of Calamoideae that occur in South America. These species occur in swampy environments and have fruits that are characteristically covered with scales. The objective of this study was to describe the formation of the layers of the pericarp within this subtribe and attempt to correlate fruit structure with the environment where species typically occur. Toward this goal, flowers in pre-anthesis and anthesis and fruits throughout development were analyzed using standard methods for light microscopy. The ontogeny of the layers of the pericarp of all three species was found to be similar. The scales were formed from non-vascularized emergences composed of exocarp and mesocarp. The median mesocarp accumulates lipids only in M. flexuosa and M. armata. The inner mesocarp together with the endocarp becomes papyraceous and tenuous in all species. This internal region of pericarp showed collapsed cells due to seed growth at the end of fruit development. Fruits of Mauritiinae are baccate, and the characters of the pericarp, especially the inner mesocarp and endocarp, help to maintain moisture. On the other hand, many species close to Mauritiinae show pericarp with sclerenchyma adjacent to the seed. This variation can contribute to understand the importance of this striking character in dispersal, germination and colonization in Arecaceae.     

Development of ovule,fruit and seed of Xyris (Xyridaceae,Poales) and taxonomic considerations

The development of the ovule, fruit and seed of Xyris spp. was studied to assess the embryological characteristics of potential taxonomic usefulness. All of the studied species have (1) orthotropous, bitegmic and tenuinucellate ovules, with a micropyle formed by both the endostoma and exostoma; (2) a cuticle in the ovules and seeds between the nucellus/endosperm and the inner integument and between the inner and outer integuments; (3) helobial, starchy endosperm; (4) a reduced, campanulate and undifferentiated embryo; (5) a seed coat formed by a tanniferous endotegmen, endotesta with thick‐walled cells and exotesta with thin‐walled cells; and (6) a micropylar operculum formed from inner and outer integuments. The pericarp is composed of a mesocarp with cells containing starch grains and an endocarp and exocarp formed by cells with U‐shaped thickened walls. The studied species differ in the embryo sac development, which can be of the Polygonum or Allium type, and in the pericarp, which can have larger cells in either endocarp or exocarp. The Allium‐type embryo sac development was observed only in Xyris spp. within Xyridaceae. Xyris also differs from the other genera of Xyridaceae by the presence of orthotropous ovules and a seed coat formed by endotegmen, endotesta and exotesta, in agreement with the division of the family into Xyridoideae and Abolbodoideae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 619–628.     

Two‐tier morpho‐chemical defence tactic in Aethionema via fruit morph plasticity and glucosinolates allocation in diaspores

Fruit dimorphism and the production of glucosinolates (GSLs) are two specific life history traits found in the members of Brassicales, which aid to optimize seed dispersal and defence against antagonists, respectively. We hypothesized that the bipartite dispersal strategy demands a tight control over the production of fruit morphs with expectedly differential allocation of defensive anticipins (GSLs). In dimorphic Aethionema, herbivory by Plutella xylostella at a young stage triggered the production of more dehiscent (seeds released from fruit) than indehiscent fruit morphs (seeds enclosed within persistent pericarp) on the same plant upon maturity. Total GSL concentrations were highest in the mature seeds of dehiscent fruits from Aethionema arabicum and Aethionema saxatile among the different ontogenetic stages of the diaspores. Multivariate analyses of GSL profiles indicated significantly higher concentrations of specific indole GSLs in the diaspores, which require optimal defence after dispersal (i.e., seeds of dehiscent and fruit/pericarp of indehiscent fruit). Bioassays with a potentially coinhabitant fungus, Aspergillus quadrilineatus, support the distinct defensive potential of the diaspores corresponding to their GSL allocation. These findings indicate a two‐tier morpho‐chemical defence tactic of Aethionema via better protected fruit morphs and strategic provision of GSLs that optimize protection to the progeny for survival in nature.     

Ovule,fruit and seed development in Abolboda (Xyridaceae,Poales): implications for taxonomy and phylogeny

Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.     

Gynoecium and fruit histology and development in Eugeissona (Calamoideae: Arecaceae)

The Malesian genus Eugeissona, with six species, is sister to all other Calamoideae, which are in turn sister to all other Arecaceae. The structure of its gynoecium and fruit is thus potentially of great interest in understanding gynoecium evolution in calamoid palms and in Arecaceae as a whole. The wall of the incompletely trilocular gynoecium of Eugeissona is thick and differentiated into several topographic zones, with a well‐developed vascular system even before pollination. During gynoecium and fruit development, the outer and inner epidermises are little specialized and form the exocarp and endocarp (obliterated in the mature fruit), respectively. In contrast, the mesophyll of the carpels differentiates strongly and is markedly specialized: four massive topographic zones are easily distinguished within the mesocarp. The peripheral zone of the mesocarp forms the body of the scales (a synapomorphy for Calamoideae). The second and the fourth zones are multilayered and parenchymatous with a massive derived vascular system in the former. The third zone of the mesocarp comprises a stout sclerenchymatous pyrene, made of fibre‐like sclereids, the innermost bundles of the derived vascular system and dorsal, ventral and lateral vascular bundles. The fruits of all other Calamoideae lack the sclerenchymatous pyrene and thus differ dramatically from Eugeissona fruits. The similarity of the processes of histogenesis during gynoecium and fruit development in Eugeissona with those in Nypa and borassoid palms, suggests these features could be plesiomorphic for the family. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 377–394.     

Gelsemiaceae (Gentianales) expanded to include the enigmatic Asian genus Pteleocarpa

The enigmatic South‐East Asian monotypic genus Pteleocarpa has been considered as a genus incertae sedis among the eudicots for a long time. Molecular data (plastid and nuclear ribosomal regions) from 44 widely sampled species across Lamiidae and phylogenetic analyses have finally clarified its familial relationships, and it is here included in Gelsemiaceae (order Gentianales). Its morphological characteristics support a placement in this family and order as a result of the presence of potential synapomorphies, such as imbricate and commonly yellow corollas, latrorse anther dehiscence, divided styles and compressed seeds. Unique characters for Pteleocarpa in Gelsemiaceae are alternate leaves and indehiscent samaras. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 482–496.     

Fruit micromorphology and anatomy of Valeriana officinalis s. str. (Valerianaceae)

Fruits of two varieties of Valeriana officinalis s. str. (var. officinalis , var. nitida ) are similar in general construction, but differ in details of external and internal structure. The outer cells of the pericarp form a regularly punctuated surface in both taxa. Scanning electron microscopy demonstrates variation in cuticular sculpturing of the outer epidermal cell walls and the presence of epicuticular wax. The surface of fruit hairs varies from micropapillate in var. officinalis to linear warty in var. nitida . In the mature rericarp there occur three distinct histological zones: an outer exocarp, a central mesocarp, and an inner endocarp. The seed is small, enclosed in the indehiscent fruit, with thin seed coat and a straight embryo. Endosperm is absent. The results of this carpological study, especially the SEM characters of pericarp surface, may provide criteria useful for delimitation of V officinalis varieties.     

Pollination by fungus gnats (Diptera: Mycetophilidae) and self-recognition sites in Tolmiea menziesii (Saxifragaceae)

Coleeae (Bignoniaceae) are a tribe almost entirely restricted to Madagascar. Coleeae have previously been placed in neotropical Crescentieae due to species with indehiscent fruits, a character otherwise unusual in Bignoniaceae. A phylogeny based on three chloroplast regions (ndhF, trnT-L spacer, trnL-F spacer) identifies a monophyletic Coleeae that is endemic to Madagascar and surrounding islands of the Indian Ocean (Seychelles, Comores and Mascarenes). African Kigelia is not a member of Coleeae, rather it is more closely related to a subset of African and Southeast Asian species of Tecomeae. The molecular phylogeny indicates that indehiscent fruit have arisen repeatedly in Bignoniaceae: in Coleeae, Kigelia and Crescentieae. The characteristic fleshy fruits of species of Coleeae likely arose autochthonously in Madagascar. Within Coleeae Colea and Ophiocolea are sisters, Phyllarthron is sister to Colea + Ophiocolea, and Rhodocolea is sister to the rest of the tribe.     

Pericarp anatomy of wild roses (Rosa L., Rosaceae)

The pericarp anatomy of representatives of all subgenera and sections of the genus Rosa was studied. All species have the same basic pericarp structure: it is composed of inner and outer endocarps, mesocarp and exocarp formed by the epidermis and hypodermis. The differences concern mainly the thickness of particular layers, and the shape and size of their cells. Cells of the endocarp and mesocarp are thick-walled. The only exception is Rosa rugosa mesocarp, which is composed of rather thin-walled cells with a large lumen. The endocarp structure of Rosa achenes resembles the drupe of the genus Prunus s.l. and drupelets of Rubus species.     

Pericarp anatomy and evolution inCoriaria (Coriariaceae)

The first overall study of pericarp anatomy ofCoriaria is presented to discuss its evolution and relationships within a genus. All 14 species investigated (including 11 narrowly defined species) have somewhat bilaterally flattened mature fruits with five to seven (or more) longitudinal costae. They share a usually nine-(or more-)cell-layered (at intercostal region), stratified mature pericarp, which is basically constructed by an exocarp, an outer, a middle and an inner zone of mesocarp, and an endocarp. While a multi-layered endocarp is composed of circumferentially elongate fibres, a multi-layered inner zone of the mesocarp comprises longitudinally elongate fibres. Despite its uncertain systematic value, the presence of those fibres arranged crisscross is a characteristic feature of the genus. Comparisons among species indicate thatCoriaria terminalis, a species of the Eastern Hemisphere, retains a basic or archaic, well-stratified pericarp structure similar to the one found in all the species investigated of the Southern and Western Hemisphere, and that four species of Asia,Coriaria napalensis, C. sinica, C. intermedia andC. japonica, share a specialized structure (lacking the outer zone of the mesocarp) indicative of their mutual close affinity. Comparisons further suggest distinctness ofCoriaria intermedia, as well as variously derived position ofC. myrtifolia andC. japonica.     

Phylogeography of the whelk genus Cominella (Gastropoda: Buccinidae) suggests long‐distance counter‐current dispersal of a direct developer

The gastropod genus Cominella Gray, 1850 consists of approximately 20 species that inhabit a wide range of marine environments in New Zealand and Australia, including its external territory, the geographically isolated Norfolk Island. This distribution is puzzling, however, with apparently closely‐related species occurring either side of the Tasman Sea, even though all species are considered to have limited dispersal abilities. To determine how Cominella attained its current distribution, we derived a dated molecular phylogeny, which revealed a clade comprising all the Australian and Norfolk Island species nested within four clades of solely New Zealand species. This Australian clade diverged well after the vicariant separation of New Zealand from Australia, and implies two long‐distance dispersal events: a counter‐current movement across the Tasman Sea from New Zealand to Australia, occurring at the origination of the clade, followed by the colonization of Norfolk Island. The biology of Cominella suggests that the most likely method of long‐distance dispersal is rafting as egg capsules. Our robust phylogeny also means that the current Cominella classification requires revision. We propose that our clades be recognized as subgenera: Cominella (s.s.), Cominista, Josepha, Cominula, and Eucominia, with each subgenus comprising only of New Zealand or Australian species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 315–332.     

A systematic histological study of palm fruits. V. Subtribe Archontophoeniciae (Arecaceae)

Pericarp histology in the Archontophoenicinae provides little to characterize the subtribe as a whole, revealing instead two separate trends with parallels in other subtribes of the Areceae. The data support a close relationship among the three genera occurring in New Caledonia:Chambeyronia, Actinokentia, andKentiopsis, in which there is a complex endocarp consisting of short, oblique fibrous bundles embedded in a thick mantle of brachysclereids, and a loose endocarp of heavily fibrous, flattened vascular bundles adjacent to a relatively thin locular epidermis. The data also support a close relationship between the two genera of the New Zealand/Tasman Sea region:Hedyscepe andRhopalostylis, in which the pericarp is more or less fibrous throughout, with purely fibrous bundles in the outer pericarp and heavily fibrous vascular bundles in the inner pericarp. These results confirm relationships revealed by other morphological data.Archontophoenix appears to be most like the New Caledonian genera in its pericarp structure, with a similar mantle of short fibrous bundles embedded in a a mantle of brachysclereids in the outer pericarp, although it differs significantly in other aspects of morphology and anatomy.