Simon J. Mayo

Phylogenetic relationships of aroids and duckweeds (Araceae) inferred from coding and noncoding plastid DNA

Familial, subfamilial, and tribal monophyly and relationships of aroids and duckweeds were assessed by parsimony and Bayesian phylogenetic analyses of five regions of coding (rbcL, matK) and noncoding plastid DNA (partial trnK intron, trnL intron, trnL-trnF spacer) for exemplars of nearly all aroid and duckweed genera. Our analyses confirm the position of Lemna and its allies (formerly Lemnaceae) within Araceae as the well-supported sister group of all aroids except Gymnostachydoideae and Orontioideae. The last two subfamilies form the sister clade of the rest of the family. Monophyly of subfamilies Orontioideae, Pothoideae, Monsteroideae, and Lasioideae is supported, but Aroideae are paraphyletic if Calla is maintained in its own subfamily (Calloideae). Our results suggest expansion of the recently proposed subfamily Zamioculcadoideae (Zamioculcas, Gonatopus) to include Stylochaeton and identify problems in the current delimitation of tribes Anadendreae, Heteropsideae, and Monstereae (Monsteroideae), Caladieae/Zomicarpeae, and Colocasieae (Aroideae). Canalization of traits of the spathe and spadix considered typical of Araceae evolved after the split of Gymnostachydoideae, Orontioideae, and Lemnoideae. An association with aquatic habitats is a plesiomorphic attribute in Araceae, occurring in the helophytic Orontioideae and free-floating Lemnoideae, but evolving independently in various derived aroid lineages including free-floating Pistia (Aroideae).

Population differentiation and species cohesion in two closely related plants adapted to neotropical high-altitude ‘inselbergs’, Alcantarea imperialis and Alcantarea geniculata (Bromeliaceae)

Isolated granitic rock outcrops or ‘inselbergs’ may provide a window into the molecular ecology and genetics of continental radiations under simplified conditions, in analogy to the use of oceanic islands in studies of species radiations. Patterns of variability and gene flow in inselberg species have never been thoroughly evaluated in comparison to related taxa with more continuous distribution ranges, or to other species in the same kingdom in general. We use nuclear microsatellites to study population differentiation and gene flow in two diploid, perennial plants adapted to high‐altitude neotropical inselbergs, Alcantarea imperialis and Alcantarea geniculata (Bromeliaceae). Population differentiation is pronounced in both taxa, especially in A. imperialis. Gene flow in this species is considerably lower than expected from the literature on plants in general and Bromeliaceae in particular, and too low to prevent differentiation due to drift (Nem < 1), unless selection coefficients/effect sizes of favourable alleles are great enough to maintain species cohesion. Low gene flow in A. imperialis indicates that the ability of pollinating bats to promote gene exchange between inselbergs is smaller than previously assumed. Population subdivision in one inselberg population of A. imperialis appears to be associated with the presence of two colour morphs that differ in the coloration of rosettes and bracts. Our results indicate a high potential for inselbergs as venues for studies of the molecular ecology and genetics of continental radiations, such as the one that gave rise to the extraordinary diversity of adaptive strategies and phenotypes seen in Bromeliaceae.

Relationships within the Araceae: comparison of morphological patterns with molecular phylogenies.

PREMISE OF THE STUDY The first family-wide molecular phylogeny of the Araceae, a family of about 3800 published species in 120 genera, became available in 1995, followed by a cladistic analysis of morpho-anatomical data in 1997. The most recent and comprehensive family-wide molecular phylogeny was published in 2008 and included species from 102 genera. We reanalyzed the molecular data with a more complete genus sampling and compared the resulting phylogeny with morphological and anatomical data, with a view to contributing to a new formal classification of the Araceae. METHODS We analyzed 113 aroid genera and 4494 aligned nucleotides that resulted from adding 11 genera to the 2008 molecular matrix. We also analyzed 81 morphological characters in the context of the molecular phylogeny, using an extended version of the 1997 morpho-anatomical data set. KEY RESULTS The resulting maximum-likelihood phylogeny is well resolved and supported, and most of the 44 larger clades also have morphological or anatomical synapomorphies as well as ecological or geographic cohesion. Of the 44 clades, 16 are here newly circumscribed and informally named. However, some relationships remain poorly supported within the Aroideae subfamily. The most problematic placement is Calla within Aroideae, which conflicts with the distribution of morphological, anatomical, and palynological character states. CONCLUSIONS The comparison of the molecular analysis with morphological and anatomical data presented here represents an important basis for a new formal classification for the Araceae and for the understanding of the evolution of this ancient family, a monocot group known in the fossil record from the early Cretaceous.

Anthocyanin pigments and leaf flavonoids in the family araceae

Anthocyanins, variously identified in inflorescence, fruit, leaf or petiole of 59 representative species of the Araccae, are of a simple type. The most common pigment is cyanidin 3-rutinoside, while pelargonidin 3-rutinoside and cyanidin 3-glucoside are regularly present. Two rare pigments are: cyanidin 3-gentiobioside in Anchomanes and Rhektophyllum, both in the subfamily Lasioideae; and delphinidin 3-rutinoside in Schismatoglottis concinna. In a leaf survey of 144 species from 58 genera, flavone C-glycosides (in 82%) and proanthocyanidins (in 35–45%) were found as the major flavonoids. In the subfamily Calloideae, subtribe Symplocarpeae, flavonols replace glycoflavones as the major leaf components but otherwise flavonols are uncommon in the family (in 27% of the sample) and more usually co-occur with flavone C-glycosides. Two new flavonol glycosides were characterized from Lysichiton camtschatcense: kaempferol 3-(6-arabinosylgalactoside)and kaempferol 3-xylosylgalactoside. Simple flavones, luteolin and chrysoeriol (in 6%) were found only in the subtribes Arinae and Cryptocoryninae, subfamily Aroideae. Flavonoid sulphates were identified in only four taxa: glycoflavone sulphates in two Culcasia species and Philodendron ornatum and a mixture of flavone and flavonol sulphates in Scindapsus pictus. Caffeic ester sulphates were more common and their presence in Anthurium hookeri was confirmed. These results show that the Araceae are unusual amongst the monocots in their simple and relatively uniform flavonoid profile; no one subfamily is clearly distinguished, although at tribal level some significant taxonomic patterns are observed. The best defined groups are the subfamilies Lasioideae and Monsteroideae, and the tribes Symplocarpeae and Arophyteae, and the subtribe Arinae. The greatest chemical diversity occurs in Anthurium and Philodendron, but this may only reflect the fact that these are the two largest genera in the family. The origin and relationship of the Araccae to other monocot groups are discussed in the light of the flavonoid evidence.

Within-population spatial genetic structure in four naturally fragmented species of a neotropical inselberg radiation, Alcantarea imperialis, A. geniculata, A. glaziouana and A. regina (Bromeliaceae)

Studies of organisms on ‘terrestrial islands’ can improve our understanding of two unresolved issues in evolutionary genetics: the likely long-term effects of habitat fragmentation and the genetic underpinnings of continental species radiations in island-like terrestrial habitats. We have addressed both issues for four closely related plant species of the adaptive radiation Bromeliaceae, Alcantarea imperialis, A. geniculata, A. regina and A. glaziouana. All four are adapted to ancient, isolated inselberg rock outcrops in the Brazilian Atlantic rainforest and are thus long-term fragmented by nature. We used eight nuclear microsatellites to study within-population spatial genetic structure (SGS) and historical gene dispersal in nine populations of these species. Within-population SGS reflected known between-species differences in mating systems. The strongest SGS observed in A. glaziouana (Sp=0.947) was stronger than literature estimates available for plants. Analysis of short- and long-distance components of SGS identified biparental inbreeding, selfing and restricted seed dispersal as main determinants of SGS, with restricted pollen dispersal by bats contributing in some localities. The ability of Alcantarea spp. to colonize isolated inselbergs probably stems from their flexible mating systems and an ability to tolerate inbreeding. Short-ranging gene dispersal (average sigma=7–27 m) is consistent with a loss of dispersal power in terrestrial island habitats. Population subdivision associated with sympatric colour morphs in A. imperialis is accompanied by between-morph differences in pollen and seed dispersal. Our results indicate a high potential for divergence with gene flow in inselberg bromeliads and they provide base-line data about the long-term effects of fragmentation in plants.

Alpha e-taxonomy: responses from the systematics community to the biodiversity crisis

SummaryThe crisis facing the conservation of biodiversity is reflected in a parallel crisis in alpha taxonomy. On one hand, there is an acute need from government and non-government organisations for large-scale and relatively stable species inventories on which to build major biodiversity information systems. On the other, molecular information will have an increasingly important impact on the evidential basis for delimiting species and is likely to result in greater scientific debate and controversy on their circumscription. This paper argues that alpha-taxonomy built on the Internet (alpha e-taxonomy) can provide a key component of the solution. Two main themes are considered: (1) the potential of e-taxonomic revisions for engaging both the specialist taxonomic community and a wider public in gathering taxonomic knowledge and deepening understanding of it, and (2) why alpha-species will continue to play an essential role in the conventional definition of species and what kinds of methodological development this implies for descriptive species taxonomy. The challenges and requirements for sustaining e-taxonomic revisions in the long-term are discussed, with particular reference to models being developed by five initiatives with botanical exemplar websites: CATE (Creating a Taxonomic E-Science), Solanaceae Source, GrassBase and EDIT (European Distributed Institute of Taxonomy) exemplar groups and scratchpads. These projects give a clear indication of the crucially important role of the national and regional taxonomic organisations and their networks in providing both leadership and a fruitful and beneficial human and technical environment for taxonomists, both amateur and professional, to contribute their expertise towards a collective global enterprise.

Venation pattern analysis of leaf images

The work investigates pattern recognition methods to detect venation patterns on leaves. An automated technique that involves scale space analysis and an automated technique that includes a combination of edge detectors are compared with a manual technique. A small data set of five images is considered in this initial exploratory work and the results are qualitatively evaluated. The results show that the technique involving scale-space analysis is demonstrated to be a promising research direction to pursue.

Comparative morphology of populations of Monstera Adans. (Araceae) from natural forest fragments in Northeast Brazil using elliptic Fourier Analysis of leaf outlines

SummaryA comparative study of the leaf outline morphometrics of Monstera adansonii var. klotzschiana, M. adansonii var. laniata and M. praetermissa was carried out. The study focused on populations in isolated montane humid (brejo) forests of Ceará state in Northeast Brazil and compared them with populations from Amazonia and the Brazilian Atlantic Forest. Digitised outlines were prepared from a total of 1,695 field-collected leaf images from 20 populations, and elliptic Fourier analysis was used to generate matrices of coefficients, from which six shape variables (principal components) were extracted using Principal Components Analysis. Intra-population variability and inter-population differences were analysed with multivariate distance methods. Separate analyses were carried out for each of three leaf size classes (juvenile, submature, mature) because of the strong heteroblasty typical of this genus. Juvenile leaves were the least variable size class within populations of M. adansonii var. klotzschiana. The shape variables expressed very similar types of variation in all three size classes. The Ceará brejo populations of M. adansonii var. klotzschiana showed significant differences between mature leaf outlines in all pairwise comparisons; the Pacatuba population was the most distinct. The Ceará populations did not cluster together exclusively. In all three size classes, populations clustered together into their taxonomic groups, most clearly so in mature leaves. No correlation between morphological and geographic distance matrices was found, nor between morphological and molecular distance. The study showed that leaf outline shape is a practicable and useful quantitative trait for studying morphological variability at species, varietal and population levels.

How deep learning extracts and learns leaf features for plant classification

Abstract Plant identification systems developed by computer vision researchers have helped botanists to recognize and identify unknown plant species more rapidly. Hitherto, numerous studies have focused on procedures or algorithms that maximize the use of leaf databases for plant predictive modeling, but this results in leaf features which are liable to change with different leaf data and feature extraction techniques. In this paper, we learn useful leaf features directly from the raw representations of input data using Convolutional Neural Networks (CNN), and gain intuition of the chosen features based on a Deconvolutional Network (DN) approach. We report somewhat unexpected results: (1) different orders of venation are the best representative features compared to those of outline shape, and (2) we observe multi-level representation in leaf data, demonstrating the hierarchical transformation of features from lower-level to higher-level abstraction, corresponding to species classes. We show that these findings fit with the hierarchical botanical definitions of leaf characters. Through these findings, we gained insights into the design of new hybrid feature extraction models which are able to further improve the discriminative power of plant classification systems. The source code and models are available at: https://github.com/cs-chan/Deep-Plant .

Aspectos da evolução e da geografia do gênero Philodendron Schott (Araceae)

The genus Philodendron can be divided into three subgenera which are very distinct in vegetative and floral morphology, floral anatomy and distribution. Phylogenetic and phenetic analysis of floral characters indicate that subgenus Meconostigma, which unlike subgenera Philodendron and Pteromischum has a predominantly southeastern range in South America, is highly apomorphic but cladistically primitive in the genus. Phenetic analyses show that the genus can be viewed as distinct, although no definitive diagnostic character is known to exist. These analyses also show that Philodendron is phenetically closer to certain west African genera (Culcasia, Cercestis, Rhektophyllum) than to the mainly Asiatic genera with it has been traditionally associated. Gynoecial morphology in subgen. Meconostigma varies considerably, with simpler forms predominating in southeastern Brazil, and the most elaborate in Amazonia. Outgroup comparison suggests that the simpler types are more primitive and indicates that the rupicolous species P. adamantinum Schott and P. leal-costae Mayo & G.M.Barroso possess the most primitive gynoecial type. Based on gynoecial morphology it is suggested that the subgenus evolved initially in eastern Brazil as a group adapted to open habitats, arriving only later in the Amazon basin. The other two subgenera, Philodendron aud Pteromischum, arose later, as epiphytes of humid forestes and today are most diverse in the northeast of South America