Structure and evolution of the pod in Indigofera (Fabaceae) reveals a trend towards small thin indehiscent pods

Pod morphology and anatomy have been studied in 28 species and four varieties of Indian Indigofera. Pods of Indigofera spp. differ with respect to size, the relative thickness of the fruit wall, the number of sclerenchymatous layers in the endocarp, the presence/absence of hypodermis and trichomes, and the presence of separation tissue. Anatomically, the pericarp is broadly characterized into three types: type I (thin pericarp and three to five sclerenchymatous layers in the endocarp), type II (intermediary pericarp thickness and six to eight sclerenchymatous layers), and type III (thick pericarp and more than eight sclerenchymatous layers). The distribution of these types across the tribe is not congruent with the current phylogenetic analyses. Type III pericarp (present in the early diverging lineages of the tribe) represents the most primitive state, whereas type I and type II pericarps are derived. Fruits of Indigofera generally show normal explosive dehiscence as a means of dispersal of seeds, although some species show adaptations for dispersal by wind. In Indigofera, dehiscence is caused by a separation layer present at the dorsal and ventral sutures except in some species (Indigofera hochstetteri, Indigofera karnatakana, Indigofera glandulosa var. sykesii, and Indigofera trita var. scabra) in which no separation tissue is present; these species show delayed dehiscence or an indehiscent condition. The indehiscent pod type is considered to be apomorphic. The taxonomic, functional, and evolutionary significance of morphological and anatomical features in fruits of the genus Indigofera has been evaluated. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 260–276.     

The systematic value of flower structure in Crotalaria and related genera of the tribe Crotalarieae (Fabaceae)

Flowers in the tribe Crotalarieae of the family Fabaceae are generally adapted to bee pollination mechanisms. Molecular systematics have recently provided a major step towards a profound insight into generic relationships, thereby creating the opportunity to re-evaluate the taxonomic and functional significance of flower structure in the tribe, with emphasis on the large genus Crotalaria. A representative sample of flowers from 211 species was dissected to record morphological character states. These data were supplemented from the literature to allow for generalizations for the tribe as a whole. Six structural–functional flower types were identified: (1) pump; (2) gullet; (3) hugging; (4) saddle; (5) tunnel and (6) brush (saddle and tunnel types described here for the first time). Crotalaria uniquely has the brush type, characterized by a rostrate keel, highly dimorphic anthers, stylar trichomes and elaborate callosities on the standard petal. Remarkably, Crotalaria and Bolusia are the only genera of the tribe Crotalarieae with callosities present in all of the species. In other genera, callosities are generally absent or infrequent. Trends towards specialization of pollination syndromes are apparent as assemblages of apomorphic states that co-occur in what we refer to here as “specialized flowers”; individual characters are labile or non-homologous (e.g. callosities) and diagnostically less valuable. Unique combinations of flower characters are often useful to support current generic concepts in Crotalarieae.     

Crystal types and their distribution in the bark of African genistoid legumes (Fabaceae tribes Sophoreae,Podalyrieae, Crotalarieae and Genisteae)

The occurrence and distribution of seven crystal types in 114 bark samples from 25 genera and 91 species, representing all four tribes of African genistoid legumes, are reported. The seven types are prismatic, druse (including irregular crystal clusters), navicular (including truncated navicular, here reported for the first time), spherical crystal cluster, styloid, crystal sand and acicular crystals in sheaf‐like aggregates. Unlike most studies, the elemental composition of the crystals was examined using X‐ray microanalyses. With the exception of acicular crystals, all crystals showed the typical peaks of calcium (sometimes accompanied by small amounts of magnesium). Acicular crystals in sheaf‐like aggregates were composed only of carbon and oxygen, indicating that they are organic and precipitate during drying or after fixation in alcohol. These crystals are found only in the two early‐diverging lineages of Podalyrieae (Cyclopia and Virgilia+Calpurnia clades), consistent with the phylogenetic pattern in the tribe (indicating a secondary loss). Navicular crystals are restricted to Podalyrieae and Crotalarieae. Prismatic crystals in bark are proposed to be the ancestral condition, with multiple losses (or reversals) in Podalyrieae, Crotalarieae and Genisteae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 620–632.     

Leaf epidermal character variation and evolution in Gaultherieae (Ericaceae)

The taxonomic value and evolutionary significance of 30 leaf epidermal characters from 238 samples representing 127 species of all seven genera in the tribe Gaultherieae (Ericaceae) and two outgroup genera were investigated by scanning electron microscopy. The character states were coded and optimized onto a maximum‐likelihood tree based on previous molecular data with Fitch parsimony and hierarchical Bayesian analysis to trace the evolution of character states throughout all internodes in the phylogenetic tree for Gaultherieae. Leaf epidermal characters were found to be largely consistent within species, but highly variable at interspecific and higher taxonomic levels. The most recent common ancestral states of 15 characters diagnosed various lineages recovered from prior studies, some with no prior morphological support. Relatively high frequencies of state change occur in the eastern Asian clade Gaultheria series Gymnobotrys + Diplycosia, the American clade G. subsection Dasyphyta p.p., the core East Asian clade and the Australia/New Zealand clade. The characters with the highest frequencies of state change are the outer stomatal ledge ornamentation type, the stomatal apparatus level, stomatal density and area, and the type of abaxial trichomes. These character state change patterns may provide insight into the ecological adaptions of Gaultherieae during their evolutionary history. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 686–710.     

Systematic and phylogenetic value of wood anatomy in Heteromorpheae (Apiaceae,Apioideae)

The wood anatomy of all four woody genera of the tribe Heteromorpheae (Apiaceae, subfamily Apioideae) has been described and compared, based on 40 wood samples (representing nine species of Anginon, one species of Glia, three species of Heteromorpha and two species of Polemannia). The four genera were found to be relatively similar in their wood anatomy. Helical thickenings on the vessel walls occur in all species investigated and appear to represent an ancestral character state and a symplesiomorphy for the tribes Bupleurieae and Heteromorpheae. Each of four genera has a diagnostically different combination of character states relating to the diameter of vessels, size of intervessel pits, length of fibres, presence and arrangement of banded axial parenchyma, size of rays and ray cells, and presence of septate fibres and crystals in the ray cells. The occurrence of marginal axial parenchyma in Anginon and Glia may be an additional synapomorphy for these taxa. Variation in the wood anatomy of 31 samples from nine species of Anginon is not correlated with habitat (Fynbos or Succulent Karoo Biomes), but instead appears to reflect adaptations to seasonal aridity found in both ecosystems. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 569–583.     

Structure and distribution of floral trichomes in Lycaste and Sudamerlycaste (Orchidaceae: Maxillariinae s.l.)

Phylogenetic analysis indicates that Lycastinae should be incorporated into a more broadly defined Maxillariinae. This is supported by several anatomical features, including the presence of sunken, glandular trichomes in both Lycastinae and Maxillariinae s.s. Until recently, these were known only from vegetative organs, but have since been reported from flowers of Maxillaria dichroma. One character currently used to distinguish between Lycaste and Sudamerlycaste is the distribution of floral trichomes. In this article, we test the reliability of this character, describe the floral micromorphology of Lycaste and Sudamerlycaste and investigate whether their flowers bear sunken hairs. Their floral micromorphology is compared with that of other genera currently assigned to Maxillariinae s.l. Flowers of Lycaste and Sudamerlycaste bear conical or obpyriform papillae and unbranched and unequally branched multicellular trichomes. Contrary to previous reports that trichomes are confined to the column in Sudamerlycaste, they also occur in the tepal axils. Labellar trichomes, although often present in Lycaste, are lacking in Sudamerlycaste. In Lycaste sections Lycaste and Aromaticae, floral trichomes tend to be unbranched, whereas section Intermediae has both unbranched and branched hairs. Branched hairs are more common in Sudamerlycaste. Some hairs are tracheoidal, pitted and lignified. These mainly occur in section Lycaste and, to a degree, in section Intermediae, but are absent from section Aromaticae and most species of Sudamerlycaste. Branched column hairs, present in Sudamerlycaste, are absent from all sections of Lycaste, and tracheoidal column hairs occur only in Sudamerlycaste. Sunken floral hairs are absent from both genera. Trichome structure and distribution may prove useful in distinguishing between these taxa and in elucidating the intergeneric relationships of Maxillariinae s.l.© 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 409–421.     

Role of trichomes and pericarp in the seed biology of the desert annual Lachnoloma lehmannii (Brassicaceae)

In many angiosperms, the fruit rather than the seed is the dispersal/germination unit, and this is the case with Lachnoloma lehmannii, a desert annual ephemeral in central southwestern Asia with indehiscent nonmucilaginuous silicles covered with trichomes. The primary aim of this study was to assess the role of trichomes and pericarp in dispersal, anchorage of diaspores, and seed germination of this species. Mature silicles are dispersed by wind and gravity, and trichomes not only significantly increased their dispersal distance, adherence to sandy soil particles, mass of water imbibed and moisture content, but also decreased the rate of water loss and moisture content of seeds. A significantly higher percentage of seeds within silicles than of isolated seeds retained viability after exposure to 60 °C for 24 h. Seed dormancy is due to the pericarp and to nondeep physiological dormancy, as shown by the increase in germination percentage of isolated seeds following dry storage and treatment with GA₃. Removal of pericarp increased germination of 6-month-old seeds from 0 to 80–90 %, and leachate from both pericarp and trichomes significantly inhibited germination of isolated seeds. Ninety-five percent of seeds within silicles buried in soil for 2 years were viable, but only 28 % of them germinated in light at 15/2 °C; thus L. lehmannii forms a persistent soil seed bank. The pericarp and its trichomes may maximize plant fitness by determining the settlement location of silicles, thus helping to ensure that seeds germinate during the cool season for seedling survival in the desert environment.     

Floral evolution and phylogeny in the tribeThelymitreae (Orchidaceae: Neottioideae)

The Australian orchid tribeThelymitreae, composed ofCalochilus, Epiblema, andThelymitra, is unique in theOrchidaceae because of the presence of a mitra or staminodal complex. Evidence from floral structure suggests thatEpiblema andThelymitra are sister genera and thatCalochilus is derived from aThelymitra ancestor. A Gene Pool Vortex model and a hypothetical phylogeny illustrate that introgressive hybridization, allopatry, and long distance dispersal have played a major role in the evolution of the tribe. Pollination and hybridization in the tribe are discussed with major emphasis on floral mimicry.     

Dehiscence of Fruit in Oilseed Rape (Brassica napus L.): I. ANATOMY OF POD DEHISCENCE

The development sequence of anatomical changes taking placewithin the pericarp tissues of Brassica napus siliquae havebeen studied at a fine- and ultra-structural level. Tissue differentiationoccurred during the initial 20 d after anthesis (DAA) and allowedthe identification of dehiscence zone cells. This descrete tissuewas subsequently further delineated by extensive lignificationof adjacent valve edge and replar vascular cells. Concomitantwith the onset of pericarp lignification, cytoplasmic contentsof the thin-walled dehiscence zone cells exhibited progressivesenescence and degradation. Wall breakdown, initially evidentin pods by 60 DAA, exclusively affected cells within the dehiscencezone, and eventually extended throughout this tissue from theepidermal suture to the locule, thus precipitating valve detachment.Ultrastructural examination confirmed that this loss of cellularcohesion was primarily attributable to middle lamella degradationand, furthermore, the dissolution of wall material was apparentlydependent on rupture of the dehiscence zone protoplast. Thesignificance of dehiscence zone cell modifications in relationto autolytic cell wall breakdown, together with possible implicationsfor the regulation of pod shatter, are discussed. Key words: Oilseed rape, Brassica napus, pod shatter, dehiscence zone, cell wall breakdown     

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.     

Gynoecium, fruit and seed structure of Paullinieae (Sapindaceae)

Despite an emphasis on fruit characters in Paullinieae taxonomy, few detailed morphological and anatomical studies of the gynoecia, fruits and seeds exist. The aims of the present study were (1) to provide a detailed documentation of gynoecium, fruit and seed structure and ontogeny in selected Paullinieae taxa; (2) to determine whether the gynoecium, seed and seedling provide additional characters of systematic value within the tribe; and (3) to relate the structural findings to mechanisms of fruit dehiscence and dispersal within these taxa. Newly described characters of systematic value within Paullinieae are shape and surface of the obturator, type of pollen tube transmitting tract, indumentum of the inner and outer surface of the carpels, ovary wall anatomy, aril anatomy, pseudo-hilum form, seedling germination mode and structure of first leaves. The fruits of Paullinia are septifragal, and conspicuous colour contrasts between the pericarp, aril and seed in most species of this genus are suggestive of a bird dispersal syndrome. Interestingly, it appears that relatively minor structural changes are associated with switches to rodent dispersal in Paullinia sphaerocarpa and water dispersal in P. clathrata and P. hystrix. Anemochorous fruits are septifragal ( Cardiospermum and Urvillea ) or schizocarpic ( Houssayanthus , Lophostigma , Serjania ). They are structurally similar and Cardiospermum with septifragal capsules may also show septicidal dehiscence.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 159–189.     

Investigations of pod characters in the Vivieae

A study of the fruits of modern members of the Vicieae (vetch tribe) has been undertaken to seek morphological, micromorphological and anatomical criteria which may assist in the identification of the archaeological remains of pods and in particular ones that may be useful for the diagnosis of their wild or domesticated status. The study included wild and domesticated Pisum (peas), Lens (lentils) and Vicia (common vetch) and some of their wild relatives found in the Old World. The results show that the micromorphological features of the pod surface tend to be highly variable, and do not distinguish the taxa. Most importantly, the pods of less-developed domesticated taxa show no reduction in the number of fibrous layers in the pod wall compared with those of their closest wild relatives, and no morphological or anatomical evidence has been found to indicate the dehiscent or indehiscent status of a pod. This appears to challenge some of our accepted wisdom on the mechanics of legume dehiscence. Received August 8, 2001 / Accepted January 15, 2002     

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.     

The uncommon cavitated secretory trichomes in Bauhinia s.s. (Fabaceae): the same roles in different organs

Cavitated secretory trichomes are characterized by a short or absent stalk that is connected to a secretory hollow head. They are rare structures in angiosperms; in Fabaceae, they have been recorded in only seven genera, including Bauhinia s.s. Because B. curvula and B. rufa exhibit glands that are responsible for attracting pollinators to flowers, this study aimed to test whether the cavitated secretory trichomes present in the flowers of these species have an attraction function. As leaf trichomes are commonly related to plant defence, comparative analyses of the morphology, ontogeny, ultrastructure and chemical profile of the secretory trichomes present in flowers and leaves were conducted. It was found that cavitated secretory trichomes are similar in their external morphology and development, regardless of the organ or species analysed. However, interspecific differences were found in the secretion process and chemical profile of the exudate. The differences found in the cavitated secretory trichomes between species indicate that they secrete distinct compounds, whereas the similarities found in these structures between vegetative and reproductive organs indicate that the cavitated trichomes have equivalent ecological functions within a species, probably in plant defence during organ development. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 104–122.     

Phylogenetic relationships in tribe Spiraeeae (Rosaceae) inferred from nucleotide sequence data

Tribe Spiraeeae has generally been defined to include Aruncus, Kelseya, Luetkea, Pentactina, Petrophyton, Sibiraea, and Spiraea. Recent phylogenetic analyses have supported inclusion of Holodiscus in this group. Spiraea, with 50-80 species distributed throughout the north temperate regions of the world, is by far the largest and most widespread genus in the tribe; the remaining genera have one to several species each. Phylogenetic analyses of nuclear ITS and chloroplast trnL-trnF nucleotide sequences for 33 species representing seven of the aforementioned genera plus Xerospiraea divided the tribe into two well supported clades, one including Aruncus, Luetkea, Holodiscus, and Xerospiraea, the second including the other genera. Within Spiraea, none of the three sections recognized by Rehder based on inflorescence morphology is supported as monophyletic. Our analyses suggest a western North American origin for the tribe, with several biogeographic events involving vicariance or dispersal between the Old and New Worlds having occurred within this group.     

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.