Anatomical characterization of Strobilanthes (Acanthaceae) species from the northern Western Ghats of India and its implication for identification at vegetative state

A comprehensive study of stem, leaf and petiole anatomy of 10 species of Strobilanthes from northern Western Ghats of India was carried out to identify characteristics which would enable species identification when flowering material is unavailable. In Strobilanthes, some species bloom annually, others are plietesials, i.e. they grow without blooming for several years and then produce huge quantities of flowers, release seeds and die. Therefore, alternative methods, such as anatomical characters, are essential to distinguish Strobilanthes species in their vegetative stage. We collected ten species of Strobilanthes for anatomical characterization. Under the bright‐field microscope, stem cross‐sections of different species were found to be undulate, quadrangular, quadrangular‐winged or terete. Study of the stem revealed a distinct outer and inner cortex, the distribution of cystoliths (CaCO₃ crystals), raphides (CaC₂H₂O₅ crystals) and sclereids which varied from species to species. Study of leaf anatomy showed structural variation and vascular bundle shapes that differed between the species. Leaf epidermal characters under light and scanning electron microscopy exhibited variation in characters such as stomatal index, stomatal length and width, stomatal type and presence of glandular and non‐glandular trichomes. The petiole anatomy was species‐specific, especially with respect to vascular bundle structure and the distribution of structures such as sclereids, cystoliths, sphaeraphides and tannin cells varied. Hence, unique anatomical features of the stem, leaf and petiole could be used as taxonomic characters to identify Strobilanthes species in a vegetative state.     

The coralline genera Sporolithon and Heydrichia (Sporolithales,Rhodophyta) clarified by sequencing type material of their generitypes and other species

Interspecific systematics in the red algal order Sporolithales remains problematic. To re‐evaluate its species, DNA analyses were performed on historical type material and recently collected specimens assigned to the two genera Sporolithon and Heydrichia. Partial rbcL sequences from the lectotype specimens of Sporolithon ptychoides (the generitype species) and Sporolithon molle, both from El Tor, Egypt, are exact matches to field‐collected topotype specimens. Sporolithon crassum and Sporolithon erythraeum also have the same type locality; material of the former appears to no longer exist, and we were unable to PCR amplify DNA from the latter. A new species, Sporolithon eltorensis, is described from the same type locality. We have not found any morpho‐anatomical characters that distinguish these three species. No sequenced specimens reported as S. ptychoides from other parts of the world represent this species, and likely reports of S. ptychoides and S. molle based on morpho‐anatomy are incorrect. A partial rbcL sequence from the holotype of Sporolithon dimotum indicates it is not a synonym of S. ptychoides, and data from the holotype of S. episporum confirm its specific recognition. DNA sequences from topotype material of Heydrichia woelkerlingii, the generitype species, and isotype material of Heydrichia cerasina confirm that these are distinct species; the taxon reported to be H. woelkerlingii from New Zealand is likely an undescribed species. Type specimens of all other Sporolithon and Heydrichia species need to be sequenced to confirm that they are distinct species; morpho‐anatomical studies have proved inadequate for this task.     

Consistent and variable leaf anatomical characters in Carex (Cyperaceae)

The anatomy and morphology of leaves in Carex have the potential to be taxonomically useful. However, studies on the variability of leaf characteristics in the genus are sparse. Researchers therefore risk using leaf anatomical characters without the knowledge of whether they are consistent in a species. We examined 22 qualitative and seven quantitative leaf anatomy characters from transverse leaf sections to test their consistency across 11 Carex spp. The characters were clearly described and primarily microscopic. Some characters were found to exhibit high levels of intraspecific variation, whereas other characters exhibited high levels of consistency in a species, including the shape of the leaf section, the density of papillae and the size of epidermal cells. Caution must be applied when choosing leaf anatomy to delimit taxa because of the intraspecific variability found in some characters, but sufficient numbers of invariant characters exist to provide useful taxonomic separation. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 371–384.     

Crusticorallina gen. nov., a nongeniculate genus in the subfamily Corallinoideae (Corallinales,Rhodophyta)

Molecular phylogenetic analyses of 18S rDNA (SSU) gene sequences confirm the placement of Crusticorallina gen. nov. in Corallinoideae, the first nongeniculate genus in an otherwise geniculate subfamily. Crusticorallina is distinguished from all other coralline genera by the following suite of morpho‐anatomical characters: (i) sunken, uniporate gametangial and bi/tetrasporangial conceptacles, (ii) cells linked by cell fusions, not secondary pit connections, (iii) an epithallus of 1 or 2 cell layers, (iv) a hypothallus that occupies 50% or more of the total thallus thickness, (v) elongate meristematic cells, and (vi) trichocytes absent. Four species are recognized based on rbcL, psbA and COI‐5P sequences, C. painei sp. nov., the generitype, C. adhaerens sp. nov., C. nootkana sp. nov. and C. muricata comb. nov., previously known as Pseudolithophyllum muricatum. Type material of Lithophyllum muricatum, basionym of C. muricata, in TRH comprises at least two taxa, and therefore we accept the previously designated lectotype specimen in UC that we sequenced to confirm its identity. Crusticorallina species are very difficult to distinguish using morpho‐anatomical and/or habitat characters, although at specific sites, some species may be distinguished by a combination of morpho‐anatomy, habitat and biogeography. The Northeast Pacific now boasts six coralline endemic genera, far more than any other region of the world.     

Dust accumulation due to anthropogenic impact induces anatomical and photochemical changes in leaves of Centranthus ruber growing on the slope of the Vesuvius volcano

• In Mediterranean ecosystems, some natural areas are exposed to severe anthropogenic impact. Especially in summer, the considerable number of tourists visiting such areas, often with vehicles, causes deposition of dust over the vegetation due to formation of powder clouds, also favoured by wind erosion, high temperature, low precipitation and incoherent soil structure. The main aim of this study was to analyse whether the deposition of dust can induce changes in leaf anatomical functional traits and in the efficiency of photosynthetic apparatus in Centranthus ruber, a species widespread in Mediterranean ecosystems.
• Leaf morpho‐functional traits were quantified in plants growing at sites characterised by high (HD) and low (LD) dust deposition, in periods with high anthropogenic impact. Analyses included quantification of chlorophyll fluorescence emission parameters, photosynthetic pigment concentration as well as stomatal size and frequency, leaf lamina thickness, quantification of intercellular spaces and phenolics in the mesophyll through microscopy.
• The overall analysis suggested that the different conditions of dust deposition induced different adjustment of morpho‐functional traits in leaves of C. ruber. High dust deposition shielded the leaf lamina, protecting the photosynthetic apparatus from excess light and favoured plant photochemical efficiency. Leaves exposed to low dust deposition showed higher accumulation of phenolic compounds, protecting chloroplast membranes and characterised by high thermal dissipation of excess light.
• Such adaptive phenomena can affect vegetation dynamics due to possible different species‐specific plant responses, resulting in different plant competitiveness under the limiting conditions of Mediterranean environments.

     

Correlation of drought traits and the predictability of osmotic potential at full leaf turgor in vegetation from New Zealand

Scientists do not know precisely how severe will be the impact of climate change on species. Evidence suggests that for some species, their future distributions might be jeopardized by local extinctions and drought‐induced tree mortality. Thus, we require models capable of estimating drought tolerance across many species. We can approach this goal by assessing functional traits. The trait osmotic potential at full turgor, π_O, is potentially a good drought indicator; however, few studies address its importance as a drought‐tolerance predictor and it is difficult to measure in the field with accuracy. In this work, we aim to answer the questions: which drought traits correlate with π_O?; do morpho‐anatomical traits correlate with π_O?; and which trees and shrubs are more (or less) vulnerable to drought? To achieve this aim, we assessed physiological and morpho‐anatomical traits for 14 native species from New Zealand forests. We included leaf‐ and wood‐related traits, π_O, water potential and stomatal conductance. We examined how these traits correlate with π_O and sought to generate models to predict π_O as a function of other traits. We tested 33 different models and evaluated them using Akaike's information criterion. Unfortunately, none of the morpho‐anatomical traits correlated well with π_O. Instead, water potential correlated most strongly with π_O. None of the models using only morpho‐anatomical traits produced plausible results. The model with the best predictive performance incorporated the effects of both morpho‐anatomical and physiological traits: water potential and wood saturated water content. Of the species analysed, and based on their π_O response, Lophozonia menziesii was considered the most vulnerable to drought stress, whereas Plagianthus regius was the least vulnerable. Our findings imply that it is potentially valuable to keep exploring the use of π_O as a drought indicator and that the effort required to measure some physiological traits, such as water potential, may be essential to consider plant drought responses and to predict π_O.     

Climate change alters leaf anatomy,but has no effects on volatile emissions from arctic plants

Biogenic volatile organic compound (BVOC) emissions are expected to change substantially because of the rapid advancement of climate change in the Arctic. BVOC emission changes can feed back both positively and negatively on climate warming. We investigated the effects of elevated temperature and shading on BVOC emissions from arctic plant species Empetrum hermaphroditum, Cassiope tetragona, Betula nana and Salix arctica. Measurements were performed in situ in long‐term field experiments in subarctic and high Arctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography‐mass spectrometry. In order to assess whether the treatments had resulted in anatomical adaptations, we additionally examined leaf anatomy using light microscopy and scanning electron microscopy. Against expectations based on the known temperature and light‐dependency of BVOC emissions, the emissions were barely affected by the treatments. In contrast, leaf anatomy of the studied plants was significantly altered in response to the treatments, and these responses appear to differ from species found at lower latitudes. We suggest that leaf anatomical acclimation may partially explain the lacking treatment effects on BVOC emissions at plant shoot‐level. However, more studies are needed to unravel why BVOC emission responses in arctic plants differ from temperate species.     

Using leaf anatomy to solve taxonomic problems within the Anemopaegma arvense species complex (Bignonieae,Bignoniaceae)

Anemopaegma is a monophyletic lineage included in the tribe Bignonieae. The genus is taxonomically problematic, especially in the Anemopaegma arvense species complex, a group whose taxa have been delimited on the basis of leaf external morphology. Here we study the leaf anatomy of all three species and nine varieties currently included in this species complex (A. acutifolium DC., A. arvense (Vell.) Stellf. ex de Souza and A. glaucum Mart. ex DC.) in search for additional characters that may help circumscribe taxa within this group. For comparison, this study also investigated the leaf anatomy of two species that are morphologically similar to representatives of the species complex but currently placed outside it (A. album Mart. ex DC. and A. scabriusculum Mart. ex DC.). All taxa were analyzed using standard anatomical procedures and light microscopy. In addition, leaves of A. acutifolium and Anemopaegma album were analyzed with scanning electron microscopy (SEM). Overall, stomata position, composition of the vascular system of the petiole, midrib, lateral veins and margin constitution represented the most important anatomical features for the recognition of the species and varieties. The study also demonstrated that A. scabriusculum presents leaf anatomical traits that differ from those encountered in species of the Anemopaegma arvense complex, corroborating earlier anatomical and molecular phylogenetic results and indicating that A. scabriusculum is indeed best placed outside the A. arvense species complex. Leaf anatomical data also support the synonymization of some varieties of A. mirandum with A. arvense, as well as varieties of A. glaucum into this species.     

Leaf morphology and anatomy of Camellia section Camellia (Theaceae)

The delimitations of species in Camellia section Camellia have been disputed for many years, resulting from uncertain relationships among species. Leaf morphological and anatomical characters for 54 species and three varieties in this section were investigated to reveal the relationships. Principal component analysis and cluster analysis were conducted using the transformed data for quantitative and qualitative characters from leaf morphology and anatomy. Combining the results of statistical analysis with comparative leaf characters of morphology and anatomy, we discussed the taxonomic treatment of section Camellia by Chang compared with that of Ming and we conclude that section Camellia consists of c. 50 species. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 456–476.     

Comparative vegetative anatomy and classification of Diseae (Orchidaceae)

The leaf, stem, root, tuber and dropper anatomy of the orchid tribe Diseae (including the subtribes Satyriinae, Disinae, Brownlecinac, Huttonaeinae and Coryciinae) is reviewed. The study is largely based on investigations of 123 species, and data from several previous publications have also been incorporated. Two characters were identified as being taxonomically valuable: (1) the presence of sclerenchyma caps associated with leaf vascular bundles, and (2) the degree of dissection of the siphonostele of the tuber (‘polystelic’ or ‘monostelic’). The phylogenetic analysis shows that anatomical characters do not change the basic structure of a cladogram that is based on morphological characters. The taxa of Diseae are discussed on the basis of anatomical data. Subtribes Satyriinae (excluding the anatomically unusual genus Pachites), Brownleeinae, Huttonaeinae, and Coryciinae are uniform in. critical anatomical characters. However, subtribe Disinae is rather diverse in vegetative anatomy. Disa sect. Micranthae differs from the rest of the genus in its leaf anatomy. The occurrence of foliar sclerenchyma bundle caps and ‘polystelic’ tubers supports the incorporation of Herschelianthe in Disa sect. Stenocarpa.     

Leaf anatomy of Ficus subsection Urostigma (Moraceae)

Species of Ficus subsection Urostigma show much overlapping variation in vegetative morphology, which often precludes correct identification of the species. The aim of this study was to describe the leaf anatomical characters and their variation and to check their suitability for identification. Included were 41 samples belonging to 25 species of subsection Urostigma, four samples belonging to two species of section Leucogyne and one specimen of Ficus glaberrima subsp. siamensis of subsection Conosycea. Transverse sections of lamina, midrib and petiole and cuticular macerations were used, and the observed anatomical characters are described for each species. On the basis of a limited number of studied samples, leaf anatomy shows little variation within each species and each species has a unique combination of character states, facilitating identification. Ficus arnottiana shows some leaf anatomical characters that are quite different from those of other members of subsection Urostigma, including a multiple epidermis and enlarged lithocysts on both sides of the leaf. Both characters are generally considered as typical for Ficus subsection Conosycea. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 259–281.     

A new combination in Aulonemia (Poaceae: Bambusoideae: Bambuseae) based on floral analysis, anatomical features, and distribution

Recent bamboo collections from Brazil, and the study of specimens from several Brazilian herbaria revealed that the species described as Eremocaulon setosum, subtribe Guaduineae, in fact fits within the Arthrostylidiinae genus Aulonemia, based on floral and anatomical characters. Several characters found in this species, such as paniculate determinate synflorescences, solitary spikelets, an abaxial marginal green stripe on the foliage leaf blades, absence of stomata on the entire adaxial side of the leaf blade, and intercostal fibers in the mesophyll are typical features of the subtribe Arthrostylidiinae. The new combination Aulonemia setosa is here proposed, an epitype is designated, and a broader distributional range of this species is documented.     

Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (g_s) and xylem cavitation vulnerability. In addition, maximum g_s was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum g_s increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ₅₀) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ₅₀ values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.     

The systematic importance of anatomical data in Gagea (Liliaceae) from the Flora Iranica area

Anatomical features of basal leaves, pedicels and tepals of 22 species of Gagea belonging to four sections (Platyspermum, Plecostigma, Gagea and Didymobulbos) are investigated. Anatomical characters are mapped onto a molecular phylogenetic tree and their evolution is evaluated. The systematic importance of the anatomical characters is discussed. Anatomical characters are more systematically useful in the Irano‐Turanian taxa than in Euro‐Siberian taxa. The presence of collenchyma and/or sclerenchyma and the pentagonal outline of the transverse section of the basal leaf is found in Irano‐Turanian taxa of section Platyspermum and is mostly absent in Euro‐Siberian taxa. A diagnostic key based on combined anatomical characters is provided. The level of variation in anatomical characters is greater in the basal leaf than the pedicel and lowest in the tepals. Convergent evolution in anatomical characters is associated with ecological shifts between sunny, open, dry habitats and closed, humid habitats. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 155–177.     

Comparative anatomy of leaves and stems in some species of the South American genus Chrysolaena (Vernonieae,Asteraceae) and taxonomic implications

Vegetative anatomical features are poorly known in the South American genus Chrysolaena. In this study, leaves and stems of six Chrysolaena species were described and compared morphologically and anatomically using diaphanization, microtome serial sectioning and scanning electron microscopy. The species differed in leaf epidermis, type of stomata, shape of anticlinal walls of epidermal cells, trichome density, and presence or absence in stems of small air spaces in the cortical parenchyma and of druse‐shaped oxalate crystals. Furthermore, glandular trichomes and three types of non‐glandular trichomes with different number of basal cells were identified on leaves and stems. Collectively, these features proved instrumental to discriminate among the six studied species, suggesting that leaves and stems of Chrysolaena can represent a source for taxonomically useful characters. We also discuss anatomical features in relation to the environmental conditions in the species’ habitats.     

Variations in leaf morpho‐anatomy and photosynthetic traits between sun and shade populations of Eurya japonica (Pentaphylacaceae) whose seeds are dispersed by birds across habitats

Eurya japonica occurs in diverse light environments through seed dispersal by birds. As the seed size is extremely small, we hypothesized that newly germinated seedlings with restricted depth of roots and length of the hypocotyl would suffer high mortality due to increased transpiration in sunny habitats and low light in shady habitats. We also expected that surviving seedlings would differ in leaf traits between habitats as a result of selection. We aimed to determine how photosynthetic traits differ between habitats and how leaf structure is related to this difference. We examined photosynthesis and leaf morpho‐anatomy for plants cloned from cuttings collected from the forest understory (shade population) and neighboring roadsides and cut‐over areas (sun population) and then grown under two irradiances (18.5% and 100% sunlight) in an experimental garden. Under growth in 100% sunlight, cloned plants from the sun population exhibited significantly greater area‐based photosynthetic capacity compared to cloned plants from the shade population at a comparable stomatal conductance, which was attributable to a higher area‐based leaf nitrogen concentration. On the other hand, mean values of photosynthetic capacity did not significantly differ between the two populations. Cloned plants from the sun population had significantly thicker leaf laminas and spongy tissue and lower stomatal density compared to cloned plants from the shade population. Thickened leaf lamina might have increased leaf tolerance to physical stresses in open habitats. The variation in leaf morpho‐anatomy between the two populations can be explained in terms of the economy of leaf photosynthetic tissue.