Lawrence M. Kelly

Phylogeny and infrageneric classification of Symplocos (Symplocaceae) inferred from DNA sequence data

Symplocos comprises ∼300 species of woody flowering plants with a disjunct distribution between the warm-temperate to tropical regions of eastern Asia and the Americas. Phylogenetic analyses of 111 species of Symplocos based on the nuclear ribosomal internal transcribed spacer (ITS) region and the chloroplast genes rpl16, matK, and trnL-trnF yielded topologies in which only one of the four traditionally recognized subgenera (Epigenia; Neotropics) is monophyletic. Section Cordyloblaste (subgenus Symplocos; eastern Asia) is monophyletic and sister to a group comprising all other samples of Symplocos. Section Palura (subgenus Hopea; eastern Asia) is sister to a group comprising all other samples of Symplocos except those of section Cordyloblaste. Symplocos wikstroemiifolia (eastern Asia) and S. tinctoria (southeastern United States), both of subgenus Hopea, form a clade that groups with S. longipes (tropical North America) and the species of subgenus Epigenia. The remaining samples of subgenus Hopea (eastern Asia) form a clade. Section Neosymplocos (subgenus Microsymplocos; Neotropics) is well nested within a clade otherwise comprising the samples of section Symplocastrum (subgenus Symplocos; Neotropics). Section Urbaniocharis (subgenus Microsymplocos; Antilles) groups as sister to the clade comprising Symplocastrum and Neosymplocos. The data support the independent evolution of deciduousness among section Palura and S. tinctoria. The early initial divergence of sections Cordyloblaste and Palura from the main group warrants their recognition at taxonomic levels higher than those at which they are currently placed. An inferred eastern Asian origin for Symplocos with subsequent dispersal to the Americas is consistent with patterns from other phylogenetic studies of eastern Asian-American disjunct plant groups but contrary to a North American origin inferred from the earliest fossil occurrences of the genus.

Discovery of unusual anatomical and continuous characters in the evolutionary history of Conostegia (Miconieae: Melastomataceae).

Conostegia has been traditionally defined to consist of 42 species in the tribe Miconieae. Recent phylogenetic studies have cast doubt on the monophyly of the genus and highlighted the need for a phylogenetic study focused on Conostegia. The purpose of this study was to test the monophyly of Conostegia and address relationships in the genus. We addressed the evolutionary history of Conostegia using DNA sequences from six loci. Difficulty in finding discrete characters that support clades prompted an anatomical survey of leaves and flowers as well as the exploratory use of some continuous characters. We coded as many species as possible for fifteen characters. Lastly, field work was conducted to document floral traits at anthesis due to the poor quality of preservation of flowers on herbarium specimens. Conostegia was found to be paraphyletic and composed of three main clades. The historically important characters of a calyptrate calyx and pleiostemony were found to have evolved more than once inside the Conostegia clade. Several other characters were found to support the clades we identified. The most unusual characters were mucilage inside the ovary which is here reported in the Melastomataceae for the first time, a stele inside the style which is mostly restricted to one clade of Conostegia and known only in this clade of the Melastomataceae, and herkogamy which has been lost in two clades within Conostegia. A combination of molecular phylogenetic analyses and broad morphological surveys allowed the better understanding of the evolutionary history in a clade of mostly cloud forest Neotropical trees. The need to include anatomical studies and tackle continuous characters is here demonstrated.

Putative floral brood-site mimicry, loss of autonomous selfing, and reduced vegetative growth are significantly correlated with increased diversification in Asarum (Aristolochiaceae).

The drivers of angiosperm diversity have long been sought and the flower-arthropod association has often been invoked as the most powerful driver of the angiosperm radiation. We now know that features that influence arthropod interactions cannot only affect the diversification of lineages, but also expedite or constrain their rate of extinction, which can equally influence the observed asymmetric richness of extant angiosperm lineages. The genus Asarum (Aristolochiaceae; ∼100 species) is widely distributed in north temperate forests, with substantial vegetative and floral divergence between its three major clades, Euasarum, Geotaenium, and Heterotropa. We used Binary-State Speciation and Extinction Model (BiSSE) Net Diversification tests of character state distributions on a Maximum Likelihood phylogram and a Coalescent Bayesian species tree, inferred from seven chloroplast markers and nuclear rDNA, to test for signal of asymmetric diversification, character state transition, and extinction rates of floral and vegetative characters. We found that reduction in vegetative growth, loss of autonomous self-pollination, and the presence of putative fungal-mimicking floral structures are significantly correlated with increased diversification in Asarum. No significant difference in model likelihood was identified between symmetric and asymmetric rates of character state transitions or extinction. We conclude that the flowers of the Heterotropa clade may have converged on some aspects of basidiomycete sporocarp morphology and that brood-site mimicry, coupled with a reduction in vegetative growth and the loss of autonomous self-pollination, may have driven diversification within Asarum.

Phylogenetic relationships in Asarum: Effect of data partitioning and a revised classification

PREMISE OF THE STUDY Generic boundaries and infrageneric relationships among the charismatic temperate magnoliid Asarum sensu lato (Aristolochiaceae) have long been uncertain. Previous molecular phylogenetic analyses used either plastid or nuclear loci alone and varied greatly in their taxonomic implications for the genus. We analyzed additional molecular markers from the nuclear and plastid genomes, reevaluated the possibility of a derived loss of autonomous self-pollination, and investigated the topological effects of matrix-partitioning-scheme choice. METHODS We sequenced seven plastid regions and the nuclear ITS1-ITS2 region of 58 individuals representing all previously recognized Asarum s.l. segregate genera and the monotypic genus Saruma. Matrices were partitioned using common a priori partitioning schemes and PartitionFinder. KEY RESULTS Topologies that were recovered using a priori partitioning of matrices differed from those recovered using a PartitionFinder-selected scheme, and by analysis method. We recovered six monophyletic groups that we circumscribed into three subgenera and six sections. Putative fungal mimic characters served as synapomorphies only for subgenus Heterotropa. Subgenus Geotaenium, a new subgenus, was recovered as sister to the remainder of Asarum by ML analyses of highly partitioned datasets. Section Longistylis, also newly named, is sister to section Hexastylis. CONCLUSIONS Our analyses do not unambiguously support a single origin for all fungal-mimicry characters. Topologies recovered through the analysis of PartitionFinder-optimized matrices can differ drastically from those inferred from a priori partitioned matrices, and by analytical method. We recommend that investigators evaluate the topological effects of matrix partitioning using multiple methods of phylogenetic reconstruction.

A morphological cladistic analysis of Lecythidoideae with emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis

A cladistic analysis was conducted to test the monophyly of Eschweilera and Lecythis as well as to examine the relationships of these two genera and their close relatives Bertholletia and Corythophora. The study included 86 species, representing all four genera and covering the range of taxonomic and morphological variation in the genera. The data matrix included 49 parsimony-informative characters derived from vegetative, floral, fruit, and seed morphology and anatomy. The results based on the consensus of all most parsimonious trees indicate that Bertholletia, Corythophora, Eschweilera, and Lecythis form a clade supported by brachyparacytic stomata, the absence of pedicels (with subsequent reversals in several clades), a two or four-locular ovary, the presence of an aril, and the absence of cotyledons. Within the clade, the monophyly of Corythophora is supported by the presence of inflorescence scales and the absence of nectar. Eschweilera is monophyletic only if E. congestiflora and E. simiorum are excluded. The monophyly of Eschweilera is supported by the presence of a two-locular ovary. Lecythis is not monophyletic, but sections Corrugata, Pisonis, and Poiteaui are monophyletic. Three species of section Lecythis are more closely related to Eschweilera, and other species of section Lecythis along with Bertholletia excelsa remain as unresolved.

Training the next generation: Graduate studies at The New York Botanical Garden, with emphasis on 1996–2015

The purpose of this paper, on the occasion of the 125th anniversary of The New York Botanical Garden, is to provide an overview of NYBG’s graduate studies program along with updated information since the last major review of the program, 25 years ago. Graduate student education has always been, and continues to be, a core mission of The New York Botanical Garden. The program is affiliated six major universities: the City University of New York, Columbia University, New York University, Yale University, Cornell University, and Fordham University. In the 125 year history of the program, the Garden has produced more than 300 graduates, including 216 Ph.D.s and 87 Master’s degrees. Students have focused on floristics, systematics, structural botany, and economic botany; the program has evolved to keep up with the continuous changes in science, with students using the most modern techniques to study plant diversity. Since 1996, the program has produced 93 graduates, including 81 Ph.D.s and 12 Master’s degrees; the Garden’s affiliation with Yale’s School of Forestry and Environmental Studies has produced a dozen graduates; and the Garden’s affiliation with Fordham University, which was reinitiated in 2008, has produced its first graduates. With the growth of the Cullman Program for Molecular Systematics and the Genomics Program in the mid-1990s, student research projects began to incorporate molecular biology (primarily based on DNA sequencing) and to address questions related to genome evolution and evolutionary developmental biology. Technological advances have also changed economic botany research. On the 125th anniversary of NYBG, the hallmark of the graduate studies program is its excellence in teaching and training, with students continuing to integrate data from modern and traditional sources to better understand plant diversity.

Total duplication of the small single copy region in the angiosperm plastome: Rearrangement and inverted repeat instability in Asarum

PREMISE OF THE STUDY As more plastomes are assembled, it is evident that rearrangements, losses, intergenic spacer expansion and contraction, and syntenic breaks within otherwise functioning plastids are more common than was thought previously, and such changes have developed independently in disparate lineages. However, to date, the magnoliids remain characterized by their highly conserved plastid genomes (plastomes). METHODS Illumina HiSeq and MiSeq platforms were used to sequence the plastomes of Saruma henryi and those of representative species from each of the six taxonomic sections of Asarum. Sequenced plastomes were compared in a phylogenetic context provided by maximum likelihood and parsimony inferences made using an additional 18 publicly available plastomes from early-diverging angiosperm lineages. KEY RESULTS In contrast to previously published magnoliid plastomes and the newly sequenced Saruma henryi plastome published here, Asarum plastomes have undergone extensive disruption and contain extremely lengthy AT-repeat regions. The entirety of the small single copy region (SSC) of A. canadense and A. sieboldii var. sieboldii has been incorporated into the inverted repeat regions (IR), and the SSC of A. delavayi is only 14 bp long. All sampled Asarum plastomes share an inversion of a large portion of the large single copy region (LSC) such that trnE-UUC is adjacent to the LSC-IR boundary. CONCLUSIONS Plastome divergence in Asarum appears to be consistent with trends seen in highly rearranged plastomes of the monocots and eudicots. We propose that plastome instability in Asarum is due to repetitive motifs that serve as recombinatory substrates and reduce genome stability.

125 years of Science and Conservation at The New York Botanical Garden: Introduction to the historical issue of Brittonia

Human life depends on plants. Plants provide food, clothing, shelter, medicine, and the raw materials to meet nearly all human needs. Plants make the oxygen we breathe, and they create the rain that sustains the world. The beauty of plants nurtures our souls and inspires our imaginations. The New York Botanical Garden has been exploring the botanical world since 1891, in an effort to describe, document, understand, and preserve plant diversity. A great strength of the institution is the detailed and well-documented knowledge of the plant and fungal kingdoms gained over more than 100 years of intensive study. In a modern world where biodiversity is under threat as never before (see Wilson, 2016), NYBG’s scientific programs contribute to three critical elements of effective conservation: (1) Discover plant biodiversity, through exploration at all scales, from the smallest molecule to the tallest tree; (2) Involve people through capacity building, outreach, and education; and (3) Protect biodiversity by informing the selection and management of protected areas and by identifying species at risk. On the occasion of the 125 anniversary of The New York Botanical Garden, this special issue of Brittonia reports on the history, evolution, and current status of NYBG’s science and conservation programs. The articles in this issue were prepared by staff members active in NYBG’s science programs, and the papers present varied perspectives on the Garden’s research programs, projects, and history. The New York Botanical Garden is characterized by broad-ranging science programs, outstanding collections and facilities, a world-class staff, and a distinguished institutional history. Excellent coverage of the history of the Garden and its programs has been provided by Mikulas (2007) and in the 100 Anniversary issue of Brittonia (Boom, 1996). The purpose of the current issue is to cover topics that were not highlighted in the 100 Anniversary issue and to add information on programs and resources that have developed significantly in the 25 years since the 100 Anniversary issue. Coverage in this issue is as broad as the programs at NYBG, ranging from systematic and economic botany, floristic research, laboratory studies, and conservation activities, to the collections and facilities that underpin this research. Outreach and education are natural extensions of the Garden’s research programs, and these topics are described in a contemporary and historical context as well. These contributions provide evidence of the importance of NYBG’s work and how it has contributed to the ongoing evolution of plant science around the world. While the hallmark of the Garden is its broad range of science, conservation, and education programs, what is more critical is the integration of these programs. Every scientist at NYBG relies on the same collections resources, the same basic data to accurately identify and understand plant diversity, and the same methods and protocols that define science. Modern research projects are multi-dimensional, national and international, and collaboration networks are extensive between researchers and departments at NYBG, for example, economic botanists teaming-up with molecular systematists to explore plant diversity in Vanuatu or Myanmar. Every program depends on every other. The topics of outreach and education are common throughout this issue, and the Graduate Studies Program provides a great example of the benefits of collaboration between departments. The Graduate Program is a key to the Garden’s vitality: it provides a constant infusion of new ideas and fresh perspectives, it is critical to ensure that we have future generations of talented biodiversity specialists, and it bolsters the productivity of our research programs. However, Graduate Studies will only be successful if all of our scientific programs are vibrant and healthy. In turn, the success of all these research programs depends on effective integration with our collections resources, including the Garden’s living collections, the Steere Herbarium, and the Mertz library. Current staff and scientists at the Garden have administered programs during a time of rapid

Vines and Climbing Plants of Puerto Rico and the Virgin Islands

Cajas National Park occurs in central Ecuador, west of the city of Cuenca. It comprises 28,544 hectares, ranging from 3150 m to 4450 m altitude, and consists of humid to wet montane forest to páramo vegetation (a grassland ecosystem occurring above treeline). This handbook is the park’s first guide to 100 wild plants that occur there and is the result of collaborative work involving the Municipality of Cuenca, the Cajas National Park Corporation, botanists from the Universidad del Azuay, and the Missouri Botanical Garden (with grants from the National Geographic Society to C. Ulloa Ulloa and P. M. Jørgensen). The guide is bilingual (Spanish/English) and of a trip size (18.5 × 13.5 cm). It begins with general introductory information about the park, the flora, the species represented in the guide, hints on how to use the guide, and recommendations on how to use and enjoy the park. Then follows the meat of the book: 100 color photos of wild (vascular) plants that occur in the park, arranged principally by color of their flowers or in a few cases as to the color of their most showy parts, and including their scientific name. On the page opposite each photograph, the reader will find the scientific name with its authority, family and vernacular names, a scientific description, the plant’s overall geographical distribution along with that within the park itself, and local uses of the plant when known. Although short, the scientific descriptions of the plants are basic and are written to be informative for identification and differentiation. Endemism of the plants to Ecuador or strictly to the park is also noted. The book ends with a list of useful references, a short glossary of terms used in the descriptions, helpful line-drawing illustrations of leaves, inflorescences and flowers, and finally indices to scientific and common names. This attractive little book is beautifully illustrated with 100 excellent photographs of the plants and a few scenic vistas of the park’s landscape. The authors must be heartily congratulated for what will certainly prove to be a very useful guide to the common plants of this popular national park. The low cost, attractive format, and high quality of paper and photographs used for the book should also add to its popularity and ease of use in the field. A very nice guide indeed!—James L. Luteyn, Mary Flagler Cary Curator of Botany and Senior Curator, Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York, 10458.Cajas National Park occurs in central Ecuador, west of the city of Cuenca. It comprises 28,544 hectares, ranging from 3150 m to 4450 m altitude, and consists of humid to wet montane forest to p&amo vegetation (a grassland ecosystem occurring above treeline). This handbook is the parks first guide to 100 wild plants that occur there and is the result of collaborative work involving the Municipality of Cuenca, the Cajas National Park Corporation, botanists from the Universidad del Azuay, and the Missouri Botanical Garden (with grants from the National Geographic Society to C. Ulloa Ulloa and R M. J0rgensen). The guide is bilingual (Spanish/English) and of a trip size (18.5 x 13.5 cm). It begins with general introductory information about the park, the flora, the species represented in the guide, hints on how to use the guide, and recommendations on how to use and enjoy the park. Then follows the meat of the book: t00 color photos of wild (vascular) plants that occur in the park, arranged principally by color of their flowers or in a few cases as to the color of their most showy parts, and including their scientific name. On the page opposite each photograph, the reader will find the scientific name with its authority, family and vernacular names, a scientific description, the plants overall geographical distribution along with that within the park itself, and local uses of the plant when known. Although short, the scientific descriptions of the plants are basic and are written to be informative for identification and differentiation. Endemism of the plants to Ecuador or strictly to the park is also noted. The book ends with a list of useful references, a short glossary of terms used in the descriptions, helpful line-drawing illustrations of leaves, inflorescences and flowers, and finally indices to scientific and common names. This attractive little book is beautifuIly itlustrated with 100 excellent photographs of the plants and a few scenic vistas of the parks landscape. The authors must be heartily congratulated for what will certainly prove to be a very useful guide to the common plants of this popular national park. The low cost, attractive format, and high quality of paper and photographs used for the book should also add to its popularity and ease of use in the field. A very nice guide indeed!--JAMES L. LUTEYN, MARY FLAGLER GARY CURATOR OF BOTANY AND SENIOR CURATOR, INSTITUTE OF SYSTEMATIC BOTANY, THE NEW YORK BOTANICAL GARDEN, BRONX, NEW YORK, 10458.

Symplocos pachycarpa (Symplocaceae), a new species from southern Mexico

Symplocos pachycarpa is described as new, and an illustration is provided. This species grows in cloud forests and oak-pine forests of Oaxaca and Guerrero, Mexico, and is most similar toS. citrea. A key is provided to distinguishS. pachycarpa from related Mexican species.