Publications by authors named "Selena Y Smith"

21 Publications

  • Page 1 of 1

Fossil palm reading: using fruits to reveal the deep roots of palm diversity.

Am J Bot 2021 03 23;108(3):472-494. Epub 2021 Feb 23.

Department of Earth and Environmental Sciences & Museum of Paleontology, University of Michigan, Ann Arbor, MI, 48109, USA.

Premise: Fossils are essential for understanding evolutionary history because they provide direct evidence of past diversity and geographic distributions. However, resolving systematic relationships between fossils and extant taxa, an essential step for many macroevolutionary studies, requires extensive comparative work on morphology and anatomy. While palms (Arecaceae) have an excellent fossil record that includes numerous fossil fruits, many are difficult to identify due in part to limited comparative data on modern fruit structure.

Methods: We studied fruits of 207 palm species, representing nearly every modern genus, using X-ray microcomputed tomography. We then developed a morphological data set to test whether the fossil record of fruits can improve our understanding of palm diversity in the deep past. To evaluate the accuracy with which this data set recovers systematic relationships, we performed phylogenetic pseudofossilization analyses. We then used the data set to investigate the phylogenetic relationships of five previously published fossil palm fruits.

Results: Phylogenetic analyses of fossils and pseudofossilization of extant taxa show that fossils can be placed accurately to the tribe and subtribe level with this data set, but node support must be considered. The phylogenetic relationships of the fossils suggest origins of many modern lineages in the Cretaceous and early Paleogene. Three of these fossils are suitable as new node calibrations for palms.

Conclusions: This work improves our knowledge of fruit structure in palms, lays a foundation for applying fossil fruits to macroevolutionary studies, and provides new insights into the evolutionary history and early diversification of Arecaceae.
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http://dx.doi.org/10.1002/ajb2.1616DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC8048450PMC
March 2021

C plant carbon isotope discrimination does not respond to CO concentration on decadal to centennial timescales.

New Phytol 2021 03 22;229(5):2576-2585. Epub 2020 Nov 22.

Department of Earth and Environmental Sciences, University of Michigan, Ann Arbor, 1100 N University Avenue, Ann Arbor, MI, 48109, USA.

Plant carbon isotope discrimination is complex, and could be driven by climate, evolution and/or edaphic factors. We tested the climate drivers of carbon isotope discrimination in modern and historical plant chemistry, and focus in particular on the relationship between rising [CO ] over Industrialization and carbon isotope discrimination. We generated temporal records of plant carbon isotopes from museum specimens collected over a climo-sequence to test plant responses to climate and atmospheric change over the past 200 yr (including Pinus strobus, Platycladus orientalis, Populus tremuloides, Thuja koraiensis, Thuja occidentalis, Thuja plicata, Thuja standishii and Thuja sutchuenensis). We aggregated our results with a meta-analysis of a wide range of C plants to make a comprehensive study of the distribution of carbon isotope discrimination and values among different plant types. We show that climate variables (e.g. mean annual precipitation, temperature and, key to this study, CO in the atmosphere) do not drive carbon isotope discrimination. Plant isotope discrimination is intrinsic to each taxon, and could link phylogenetic relationships and adaptation to climate quantitatively and over ecological to geological time scales.
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http://dx.doi.org/10.1111/nph.17030DOI Listing
March 2021

Cornalean affinities, phylogenetic significance, and biogeographic implications of Operculifructus infructescences from the Late Cretaceous (Campanian) of Mexico.

Am J Bot 2018 11 25;105(11):1911-1928. Epub 2018 Oct 25.

Department of Geosciences, University of Rhode Island, Kingston, Rhode Island, 02881, USA.

In English: Premise of the Study Cretaceous Cornales provide a crucial record of the early history of asterids. Most lineages of the order are well represented in the fossil record, but South African families of Curtisiaceae and Grubbiaceae remain poorly understood. Seventy-three specimens of a fossil infructescence belonging to the genus Operculifructus Estrada-Ruiz & Cevallos-Ferriz emend. Hayes & Smith from the Late Cretaceous (Campanian) El Gallo Formation, Baja California, Mexico bear previously undescribed characters that suggest a relationship to Grubbiaceae. Methods Microstructures of the fossils were examined through light microscopy and x-ray microcomputed tomography (microCT) scanning. Modern Grubbia tomentosa (Thunb.) Harms fruits were scanned for comparison to the fossil material. Phylogenetic analyses using the 77 fruit characters of Atkinson () were performed to test relationships of the fossil to major lineages of the order. Several analyses applied topological constraints to the extant taxa, based on various genetically supported hypotheses of relationship within Cornales. Key Results Novel structures of Operculifructus newly observed here include (1) anatropous ovules, (2) drupaceous fruits, (3) an epigynous disc, (4) and a stylar canal in the center of the disc aligned with the micropylar protrusion of the seed. Phylogenetic analysis consistently resolves Operculifructus as sister to Grubbiaceae. Conclusions Operculifructus provides direct evidence for the occurrence of Grubbiaceae in the Late Cretaceous, much older than previous Eocene evidence. The phylogeny of Atkinson () indicates that the new phylogenetic position recovered for Operculifructus also establishes the presence of the most basal drupaceous cornalean fruits in North America by the Campanian. RESUMEN EN ESPAÑOL Hipótesis de la Investigación Cornales cretácicos representan un registro esencial en la historia de los astéridos. Casi todos los linajes del orden están bien representados en el registro fósil, pero las familias africanas sureñas Curtisiaceae y Grubbiaceae permanecen pobremente entendidas. Setenta y tres ejemplares de una infrutescencia fósil perteneciente al género Operculifructus Estrada-Ruiz & Cevallos-Ferriz emend. Hayes & Smith de la formación campaniana (Cretácico Tardío) El Gallo, Baja California, México, poseen caracteres no descritos previamente y sugieren una relación con Grubbiaceae. Metodología Microestructuras de los fósiles fueron examinadas con microscopio de luz y microtomografía computarizada (micro-CT) de rayos X. Frutos actuales de Grubbia tomentosa (Thunb.) Harms fueron escaneados para su comparación con el material fósil. Se realizaron análisis filogenéticos usando los 77 caracteres de frutos de Atkinson () para probar las relaciones de los fósiles con los linajes principales del orden. En algunos análisis se aplicaron restricciones topológicas a los taxa actuales basándose en varias hipótesis, soportadas genéticamente, de las relaciones dentro de Cornales. Resultados Centrales (Cruciales) Las estructuras novedosas de Operculifructus, recientemente observadas aquí, incluyen (1) óvulos anátropos; (2) frutos drupáceos; (3) disco epígino; y (4) un canal estilar en el centro del disco alineado con la protuberancia micropilar de la semilla. Los análisis filogenéticos consistentemente dan como resultado que Operculifructus es hermano de Grubbiaceae. Conclusiones Operculifructus proporciona evidencia directa de la presencia de Grubbiaceae en el Cretácico Tardío, mucho antes que la previa evidencia en el Eoceno. La filogenia de Atkinson () indica que la nueva posición filogenética recuperada para Operculifructus también resalta la presencia de frutos drupáceos cornaleanos más basales en Norteamérica durante el Campaniano.
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http://dx.doi.org/10.1002/ajb2.1179DOI Listing
November 2018

Building the monocot tree of death: Progress and challenges emerging from the macrofossil-rich Zingiberales.

Am J Bot 2018 08 2;105(8):1389-1400. Epub 2018 Aug 2.

School of Integrative Plant Sciences, Section of Plant Biology and the Bailey Hortorium, Cornell University, Ithaca, NY, 14853, USA.

Premise Of The Study: Inclusion of fossils in phylogenetic analyses is necessary in order to construct a comprehensive "tree of death" and elucidate evolutionary history of taxa; however, such incorporation of fossils in phylogenetic reconstruction is dependent on the availability and interpretation of extensive morphological data. Here, the Zingiberales, whose familial relationships have been difficult to resolve with high support, are used as a case study to illustrate the importance of including fossil taxa in systematic studies.

Methods: Eight fossil taxa and 43 extant Zingiberales were coded for 39 morphological seed characters, and these data were concatenated with previously published molecular sequence data for analysis in the program MrBayes.

Key Results: Ensete oregonense is confirmed to be part of Musaceae, and the other seven fossils group with Zingiberaceae. There is strong support for Spirematospermum friedrichii, Spirematospermum sp. 'Goth', S. wetzleri, and Striatornata sanantoniensis in crown Zingiberaceae while "Musa" cardiosperma, Spirematospermum chandlerae, and Tricostatocarpon silvapinedae are best considered stem Zingiberaceae. Inclusion of fossils explains how different topologies from morphological and molecular data sets is due to shared plesiomorphic characters shared by Musaceae, Zingiberaceae, and Costaceae, and most of the fossils.

Conclusions: Inclusion of eight fossil taxa expands the Zingiberales tree and helps explain the difficulty in resolving relationships. Inclusion of fossils was possible in part due to a large morphological data set built using nondestructive microcomputed tomography data. Collaboration between paleo- and neobotanists and technology such as microcomputed tomography will help to build the tree of death and ultimately improve our understanding of the evolutionary history of monocots.
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http://dx.doi.org/10.1002/ajb2.1123DOI Listing
August 2018

Reinvestigating an enigmatic Late Cretaceous monocot: morphology, taxonomy, and biogeography of .

PeerJ 2018 6;6:e4580. Epub 2018 Apr 6.

Nikon Metrology, Inc., Brighton, MI, USA.

Angiosperm-dominated floras of the Late Cretaceous are essential for understanding the evolutionary, ecological, and geographic radiation of flowering plants. The Late Cretaceous-early Paleogene Deccan Intertrappean Beds of India contain angiosperm-dominated plant fossil assemblages known from multiple localities in central India. Numerous monocots have been documented from these assemblages, providing a window into an important but poorly understood time in their diversification. One component of the Deccan monocot diversity is the genus , known from anatomically preserved infructescences. was first collected over a century ago and has been the subject of numerous studies. However, resolution of its three-dimensional (3D) morphology and anatomy, as well as its taxonomic affinities, has remained elusive. In this study we investigated the morphology and taxonomy of genus , combining traditional paleobotanical techniques and X-ray micro-computed tomography (μCT). Re-examination of type and figured specimens, 3D reconstructions of fruits, and characterization of structures in multiple planes of section using μCT data allowed us to resolve conflicting interpretations of fruit morphology and identify additional characters useful in refining potential taxonomic affinities. Among the four species previously recognized, we consider two to be valid ( and ), and the other two to be synonyms of these. Furthermore, we found that permineralized infructescences of from the late Campanian of Mexico correspond closely in morphology to . We argue that and are congeneric and provide the new combination, (Cevallos-Ferriz, Estrada-Ruiz & Perez-Hernandez) Matsunaga, S.Y. Smith, & Manchester comb. nov. The significant geographic disjunction between these two occurrences indicates that the genus was widespread and may be present in other Late Cretaceous assemblages. exhibits character combinations not present in any extant taxa and its affinities remain unresolved, possibly representing an extinct member of Alismatales. The character mosaic observed in and the broad distribution of the genus provide new data relevant to understanding early monocot evolution and suggest that the (thus far) largely invisible Late Cretaceous monocot diversification was characterized by enigmatic and/or stem taxa.
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http://dx.doi.org/10.7717/peerj.4580DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5890723PMC
April 2018

Crowds Replicate Performance of Scientific Experts Scoring Phylogenetic Matrices of Phenotypes.

Syst Biol 2018 Jan;67(1):49-60

Department of Evolution, Ecology and Organismal Biology, The Ohio State University, 43210, USA.

Scientists building the Tree of Life face an overwhelming challenge to categorize phenotypes (e.g., anatomy, physiology) from millions of living and fossil species. This biodiversity challenge far outstrips the capacities of trained scientific experts. Here we explore whether crowdsourcing can be used to collect matrix data on a large scale with the participation of nonexpert students, or "citizen scientists." Crowdsourcing, or data collection by nonexperts, frequently via the internet, has enabled scientists to tackle some large-scale data collection challenges too massive for individuals or scientific teams alone. The quality of work by nonexpert crowds is, however, often questioned and little data have been collected on how such crowds perform on complex tasks such as phylogenetic character coding. We studied a crowd of over 600 nonexperts and found that they could use images to identify anatomical similarity (hypotheses of homology) with an average accuracy of 82% compared with scores provided by experts in the field. This performance pattern held across the Tree of Life, from protists to vertebrates. We introduce a procedure that predicts the difficulty of each character and that can be used to assign harder characters to experts and easier characters to a nonexpert crowd for scoring. We test this procedure in a controlled experiment comparing crowd scores to those of experts and show that crowds can produce matrices with over 90% of cells scored correctly while reducing the number of cells to be scored by experts by 50%. Preparation time, including image collection and processing, for a crowdsourcing experiment is significant, and does not currently save time of scientific experts overall. However, if innovations in automation or robotics can reduce such effort, then large-scale implementation of our method could greatly increase the collective scientific knowledge of species phenotypes for phylogenetic tree building. For the field of crowdsourcing, we provide a rare study with ground truth, or an experimental control that many studies lack, and contribute new methods on how to coordinate the work of experts and nonexperts. We show that there are important instances in which crowd consensus is not a good proxy for correctness.
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http://dx.doi.org/10.1093/sysbio/syx052DOI Listing
January 2018

Open data and digital morphology.

Proc Biol Sci 2017 Apr 12;284(1852). Epub 2017 Apr 12.

School of Earth Sciences, University of Bristol, Life Sciences Building, Tyndall Avenue, Bristol BS8 1TQ, UK

Over the past two decades, the development of methods for visualizing and analysing specimens digitally, in three and even four dimensions, has transformed the study of living and fossil organisms. However, the initial promise that the widespread application of such methods would facilitate access to the underlying digital data has not been fully achieved. The underlying datasets for many published studies are not readily or freely available, introducing a barrier to verification and reproducibility, and the reuse of data. There is no current agreement or policy on the amount and type of data that should be made available alongside studies that use, and in some cases are wholly reliant on, digital morphology. Here, we propose a set of recommendations for minimum standards and additional best practice for three-dimensional digital data publication, and review the issues around data storage, management and accessibility.
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http://dx.doi.org/10.1098/rspb.2017.0194DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5394671PMC
April 2017

Species diversity driven by morphological and ecological disparity: a case study of comparative seed morphology and anatomy across a large monocot order.

AoB Plants 2016 27;8. Epub 2016 Oct 27.

Swiss Light Source, Paul Scherrer Institut, 5232 Villigen PSI, Switzerland.

Phenotypic variation can be attributed to genetic heritability as well as biotic and abiotic factors. Across Zingiberales, there is a high variation in the number of species per clade and in phenotypic diversity. Factors contributing to this phenotypic variation have never been studied in a phylogenetic or ecological context. Seeds of 166 species from all eight families in Zingiberales were analyzed for 51 characters using synchrotron based 3D X-ray tomographic microscopy to determine phylogenetically informative characters and to understand the distribution of morphological disparity within the order. All families are distinguishable based on seed characters. Non-metric multidimensional scaling analyses show Zingiberaceae occupy the largest seed morphospace relative to the other families, and environmental analyses demonstrate that Zingiberaceae inhabit both temperate and tropical regions, while other Zingiberales are almost exclusively tropical. Temperate species do not cluster in morphospace nor do they share a common suite of character states. This suggests that the diversity seen is not driven by adaptation to temperate niches; rather, the morphological disparity seen likely reflects an underlying genetic plasticity that allowed Zingiberaceae to repeatedly colonize temperate environments. The notable morphoanatomical variety in Zingiberaceae seeds may account for their extraordinary ecological success and high species diversity as compared to other Zingiberales.
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http://dx.doi.org/10.1093/aobpla/plw063DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5091906PMC
October 2016

X-rays and virtual taphonomy resolve the first Cissus (Vitaceae) macrofossils from Africa as early-diverging members of the genus.

Am J Bot 2016 09 19;103(9):1657-77. Epub 2016 Sep 19.

Imaging and Analysis Centre, Natural History Museum, London, SW7 5BD, UK.

Premise Of The Study: Fossilized seeds similar to Cissus (Vitaceae) have been recognized from the Miocene of Kenya, though some were previously assigned to the Menispermaceae. We undertook a comparative survey of extant African Cissus seeds to identify the fossils and consider their implications for the evolution and biogeography of Cissus and for African early Miocene paleoenvironments.

Methods: Micro-computed tomography (µCT) and synchrotron-based X-ray tomographic microscopy (SRXTM) were used to study seed morphology and anatomy. Virtual taphonomy, using SRXTM data sets, produced digital fossils to elucidate seed taphonomy. Phylogenetic relationships within Cissus were reconstructed using existing and newly produced DNA sequences for African species. Paleobiology and paleoecology were inferred from African nearest living relatives.

Key Results: The fossils were assigned to four new Cissus species, related to four modern clades. The fossil plants were interpreted as climbers inhabiting a mosaic of riverine woodland and forest to more open habitats. Virtual taphonomy explained how complex mineral infill processes concealed key seed features, causing the previous taxonomic misidentification. Newly sampled African species, with seeds most similar to the fossils, belong to four clades within core Cissus, two of which are early diverging.

Conclusions: Virtual taphonomy, combined with X-ray imaging, has enabled recognition of the first fossil Cissus and Vitaceae from Africa. Early-divergent members of the core Cissus clade were present in Africa by at least the early Miocene, with an African origin suggested for the Cissus sciaphila clade. The fossils provide supporting evidence for mosaic paleoenvironments inhabited by early Miocene hominoids.
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http://dx.doi.org/10.3732/ajb.1600177DOI Listing
September 2016

Revisiting the Zingiberales: using multiplexed exon capture to resolve ancient and recent phylogenetic splits in a charismatic plant lineage.

PeerJ 2016 21;4:e1584. Epub 2016 Jan 21.

Department of Plant and Microbial Biology, Department of Integrative Biology and the University and Jepson Herbaria, University of California, Berkeley , Berkeley, CA , United States.

The Zingiberales are an iconic order of monocotyledonous plants comprising eight families with distinctive and diverse floral morphologies and representing an important ecological element of tropical and subtropical forests. While the eight families are demonstrated to be monophyletic, phylogenetic relationships among these families remain unresolved. Neither combined morphological and molecular studies nor recent attempts to resolve family relationships using sequence data from whole plastomes has resulted in a well-supported, family-level phylogenetic hypothesis of relationships. Here we approach this challenge by leveraging the complete genome of one member of the order, Musa acuminata, together with transcriptome information from each of the other seven families to design a set of nuclear loci that can be enriched from highly divergent taxa with a single array-based capture of indexed genomic DNA. A total of 494 exons from 418 nuclear genes were captured for 53 ingroup taxa. The entire plastid genome was also captured for the same 53 taxa. Of the total genes captured, 308 nuclear and 68 plastid genes were used for phylogenetic estimation. The concatenated plastid and nuclear dataset supports the position of Musaceae as sister to the remaining seven families. Moreover, the combined dataset recovers known intra- and inter-family phylogenetic relationships with generally high bootstrap support. This is a flexible and cost effective method that gives the broader plant biology community a tool for generating phylogenomic scale sequence data in non-model systems at varying evolutionary depths.
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http://dx.doi.org/10.7717/peerj.1584DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4727956PMC
January 2016

Seed morphology and anatomy and its utility in recognizing subfamilies and tribes of Zingiberaceae.

Am J Bot 2015 Nov 27;102(11):1814-41. Epub 2015 Oct 27.

Advanced Light Source, Lawrence Berkeley National Laboratories, Berkeley, California 94720 USA.

Premise Of The Study: Recent phylogenetic analyses based on molecular data suggested that the monocot family Zingiberaceae be separated into four subfamilies and four tribes. Robust morphological characters to support these clades are lacking. Seeds were analyzed in a phylogenetic context to test independently the circumscription of clades and to better understand evolution of seed characters within Zingiberaceae.

Methods: Seventy-five species from three of the four subfamilies were analyzed using synchrotron based x-ray tomographic microscopy (SRXTM) and scored for 39 morphoanatomical characters.

Key Results: Zingiberaceae seeds are some of the most structurally complex seeds in angiosperms. No single seed character was found to distinguish each subfamily, but combinations of characters were found to differentiate between the subfamilies. Recognition of the tribes based on seeds was possible for Globbeae, but not for Alpinieae, Riedelieae, or Zingibereae, due to considerable variation.

Conclusions: SRXTM is an excellent, nondestructive tool to capture morphoanatomical variation of seeds and allows for the study of taxa with limited material available. Alpinioideae, Siphonochiloideae, Tamijioideae, and Zingiberoideae are well supported based on both molecular and morphological data, including multiple seed characters. Globbeae are well supported as a distinctive tribe within the Zingiberoideae, but no other tribe could be differentiated using seeds due to considerable homoplasy when compared with currently accepted relationships based on molecular data. Novel seed characters suggest tribal affinities for two currently unplaced Zingiberaceae taxa: Siliquamomum may be related to Riedelieae and Monolophus to Zingibereae, but further work is needed before formal revision of the family.
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http://dx.doi.org/10.3732/ajb.1500300DOI Listing
November 2015

Reading the leaves: A comparison of leaf rank and automated areole measurement for quantifying aspects of leaf venation.

Appl Plant Sci 2014 Aug 12;2(8). Epub 2014 Aug 12.

School of Forestry and Environmental Studies, Yale University, 195 Prospect Street, New Haven, Connecticut 06511 USA.

The reticulate venation that is characteristic of a dicot leaf has excited interest from systematists for more than a century, and from physiological and developmental botanists for decades. The tools of digital image acquisition and computer image analysis, however, are only now approaching the sophistication needed to quantify aspects of the venation network found in real leaves quickly, easily, accurately, and reliably enough to produce biologically meaningful data. In this paper, we examine 120 leaves distributed across vascular plants (representing 118 genera and 80 families) using two approaches: a semiquantitative scoring system called "leaf ranking," devised by the late Leo Hickey, and an automated image-analysis protocol. In the process of comparing these approaches, we review some methodological issues that arise in trying to quantify a vein network, and discuss the strengths and weaknesses of automatic data collection and human pattern recognition. We conclude that subjective leaf rank provides a relatively consistent, semiquantitative measure of areole size among other variables; that modal areole size is generally consistent across large sections of a leaf lamina; and that both approaches-semiquantitative, subjective scoring; and fully quantitative, automated measurement-have appropriate places in the study of leaf venation.
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http://dx.doi.org/10.3732/apps.1400006DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4141712PMC
August 2014

Seed fertilization, development, and germination in Hydatellaceae (Nymphaeales): Implications for endosperm evolution in early angiosperms.

Am J Bot 2009 Sep 13;96(9):1581-93. Epub 2009 Aug 13.

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK.

New data on endosperm development in the early-divergent angiosperm Trithuria (Hydatellaceae) indicate that double fertilization results in formation of cellularized micropylar and unicellular chalazal domains with contrasting ontogenetic trajectories, as in waterlilies. The micropylar domain ultimately forms the cellular endosperm in the dispersed seed. The chalazal domain forms a single-celled haustorium with a large nucleus; this haustorium ultimately degenerates to form a space in the dispersed seed, similar to the chalazal endosperm haustorium of waterlilies. The endosperm condition in Trithuria and waterlilies resembles the helobial condition that characterizes some monocots, but contrasts with Amborella and Illicium, in which most of the mature endosperm is formed from the chalazal domain. The precise location of the primary endosperm nucleus governs the relative sizes of the chalazal and micropylar domains, but not their subsequent developmental trajectories. The unusual tissue layer surrounding the bilobed cotyledonary sheath in seedlings of some species of Trithuria is a belt of persistent endosperm, comparable with that of some other early-divergent angiosperms with a well-developed perisperm, such as Saururaceae and Piperaceae. The endosperm of Trithuria is limited in size and storage capacity but relatively persistent.
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http://dx.doi.org/10.3732/ajb.0900033DOI Listing
September 2009

Elucidating the affinities and habitat of ancient, widespread Cyperaceae: Volkeria messelensis gen. et sp. nov., a fossil mapanioid sedge from the Eocene of Europe.

Am J Bot 2009 Aug 20;96(8):1506-18. Epub 2009 Jul 20.

Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey TW20 0EX, UK.

The sedges (family Cyperaceae) are an economically and ecologically important monocot group dating back at least to the Paleocene. While modern genera are mostly unknown before the Oligocene, several extinct taxa are recognized as the earliest sedges. Their affinities have been unclear until now, because they are found as isolated, often abraded fruits or endocarps. Exceptionally preserved sedge fossils from the Middle Eocene of Messel, Germany yield more characters for identification. Fossil cyperacean infructescences with in situ pollen are recognized for the first time and show features of the early-divergent mapanioid sedges. Pollen resembles that of tribe Hypolytreae. Comparisons with extant taxa suggest the closest affinities with Hypolytrum and Mapania. However, the Messel fossils represent a distinct taxon, Volkeria messelensis gen. et sp. nov. Without the additional characters of infructescence and pollen, the Messel fruits would have been placed in the extinct genus Caricoidea, a typical Eocene sedge that was widespread across Eurasia. Similarities of fruit structure suggest that Caricoidea was also a mapanioid sedge. Mapanioid sedges are found today in tropical wet forests and swamps, a distribution suggesting that early sedges occupied a similar habitat, unlike many modern sedges, and were not precursors to open grassland vegetation.
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http://dx.doi.org/10.3732/ajb.0800427DOI Listing
August 2009

Virtual taphonomy using synchrotron tomographic microscopy reveals cryptic features and internal structure of modern and fossil plants.

Proc Natl Acad Sci U S A 2009 Jul 2;106(29):12013-8. Epub 2009 Jul 2.

Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey TW20 0EX, United Kingdom.

While more commonly applied in zoology, synchrotron radiation X-ray tomographic microscopy (SRXTM) is well-suited to nondestructive study of the morphology and anatomy of both fossil and modern plants. SRXTM uses hard X-rays and a monochromatic light source to provide high-resolution data with little beam-hardening, resulting in slice data with clear boundaries between materials. Anatomy is readily visualized, including various planes of section from a single specimen, as clear as in traditional histological sectioning at low magnifications. Thus, digital sectioning of rare or difficult material is possible. Differential X-ray attenuation allows visualization of different layers or chemistries to enable virtual 3-dimensional (3D) dissections of material. Virtual potential fossils can be visualized and digital tissue removal reveals cryptic underlying morphology. This is essential for fossil identification and for comparisons between assemblages where fossils are preserved by different means. SRXTM is a powerful approach for botanical studies using morphology and anatomy. The ability to gain search images in both 2D and 3D for potential fossils gives paleobotanists a tool--virtual taphonomy--to improve our understanding of plant evolution and paleobiogeography.
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http://dx.doi.org/10.1073/pnas.0901468106DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2715531PMC
July 2009

Scanning electron microscopy and synchrotron radiation x-ray tomographic microscopy of 330 million year old charcoalified seed fern fertile organs.

Microsc Microanal 2009 Apr;15(2):166-73

Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey TW200EX, UK.

Abundant charcoalified seed fern (pteridosperm) pollen organs and ovules have been recovered from Late Viséan (Mississippian 330 Ma) limestones from Kingswood, Fife, Scotland. To overcome limitations of data collection from these tiny, sometimes unique, fossils, we have combined low vacuum scanning electron microscopy on uncoated specimens with backscatter detector and synchrotron radiation X-ray tomographic microscopy utilizing the Materials Science and TOMCAT beamlines at the Swiss Light Source of the Paul Scherrer Institut. In combination these techniques improve upon traditional cellulose acetate peel sectioning because they enable study of external morphology and internal anatomy in multiple planes of section on a single specimen that is retained intact. The pollen organ Melissiotheca shows a basal parenchymatous cushion bearing more than 100 sporangia on the distal face. Digital sections show the occurrence of pollen in some sporangia. The described ovule is new and has eight integumentary lobes that are covered in spirally arranged glandular hairs. Virtual longitudinal sections reveal the lobes are free above the pollen chamber. Results are applied in taxonomy and will subsequently contribute to our understanding of the former diversity and evolution of ovules, seeds, and pollen organs in the seed ferns, the first seed-bearing plants to conquer the land.
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http://dx.doi.org/10.1017/S1431927609090126DOI Listing
April 2009

Fossil cyclanthus (cyclanthaceae, pandanales) from the eocene of Germany and England.

Am J Bot 2008 Jun;95(6):688-99

Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey TW20 0EX, UK.

The first known fossil fruits and seeds of Cyclanthaceae are described here. Cyclanthus messelensis sp. nov., from the Middle Eocene of Messel, Germany, has discoidal fruiting cycles up to 6 cm in diameter, with a central hole, radiating fiber strands, a thickened outer rim, and paratetracytic stomata. In situ seeds are up to 2 mm long, with an elongate micropylar end, a chalazal neck, and adpressed bands. The Messel fruits and seeds are nearly identical to those of Cyclanthus, differing in minor details of cuticular structure and seeds. The exceptional preservation at Messel (including in situ and isolated seeds) has also allowed us to establish the taxonomic affinity of isolated seeds ('Scirpus' lakensis) that are spatially and temporally widespread in the late Early and early Middle Eocene of southern England. Cyclanthus lakensis comb. nov. is described here as a morphotaxon for isolated fossil Cyclanthus seeds, preserved only as cuticular envelopes. Cyclanthus is another example of links between Eocene Europe and Recent South American floras because it is found today only from Mexico to South America. This material represents the first fossil fruits and seeds of Cyclanthus, which clearly was once growing in the Paleogene of the Old World.
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http://dx.doi.org/10.3732/ajb.2007390DOI Listing
June 2008

Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov. (Saururaceae) from the Middle Eocene Princeton Chert.

Am J Bot 2007 Oct;94(10):1642-57

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada.

Investigations of small permineralized flowers from the Middle Eocene Princeton Chert, British Columbia, Canada have revealed that they represent an extinct species of Saururus. Over 100 flowers and one partial inflorescence were studied, and numerous minute perianthless flowers are borne in an indeterminate raceme. Each flower is subtended by a bract, and flowers and bracts are borne at the end of a common stalk. Five stamens are basally adnate to the carpels. Pollen is frequently found in situ in the anthers. Examined under SEM and TEM, pollen grains are minute (6-11 μm), monosulcate, boat-shaped-elliptic, with punctate sculpturing and a granulate aperture membrane. The gynoecium is composed of four basally connate, lobed carpels with recurved styles and a single ovule per carpel. Flower structure and pollen are indicative of Saururaceae (Piperales), and in phylogenetic analyses using morphological characters, the fossils are sister to extant Saururus. The fossil flowers are described here as Saururus tuckerae sp. nov. These fossil specimens add to the otherwise sparse fossil record of Piperales, represent the oldest fossils of Saururaceae as well as the first North American fossil specimens of this family, and provide the first evidence of saururaceous pollen in the fossil record.
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http://dx.doi.org/10.3732/ajb.94.10.1642DOI Listing
October 2007

Cretaceous and Eocene poroid hymenophores from Vancouver Island, British Columbia.

Mycologia 2004 Jan-Feb;96(1):180-6

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada.

Two fossil poroid hymenophore fragments, one from the Cretaceous Period and the other from the Eocene Epoch, are described. The permineralized specimens were obtained from marine calcareous concretions on Vancouver Island, British Columbia, Canada, and were studied using the cellulose acetate peel technique. Size and distribution of pores in the hymenophores, as well as the hyphal anatomy of the dissepiments and some hymenial elements, were examined. In the Cretaceous specimen, Quatsinoporites cranhamii sp. nov., pores are round to elliptical, three per mm, and 130-540 μm diam. Dissepiments consist of narrow, simple septate, hyphae. Neither basidia nor basidiospores are present, but acuminate hymenial cystidia, up to 54 μm in length, are common. The Eocene specimen, Appianoporites vancouverensis sp. nov., has a pore density of six per mm and pores are 130-163 μm in diam. Dissepiments consist of narrow, simple septate, thin-walled hyphae. Neither basidia nor basidiospores are present, but acuminate, thick-walled hymenial cystidia, up to 32 μm in length, are common. The poroid hymenophore is a characteristic of a number of extant basidiomycete taxa, including the Boletales, Polyporales and Hymenochaetales. It is unlikely that the fleshy, ephemeral, terrestrial basidiomata of the Boletales would be preserved in a marine environment, and thus the specimens are interpreted as belonging to basidiomycete lineages, with persistent, leathery or corky basidiomata. The simple septate hyphae, the minute pores and presence of cystidia most closely resemble taxa of the Hymenochaetales. These fossils unequivocally push back the minimum age of homobasidiomycetes and extend their paleogeographical range.
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October 2012

Cyathea cranhamii sp. nov. (Cyatheaceae), anatomically preserved tree fern sori from the Lower Cretaceous of Vancouver Island, British Columbia.

Am J Bot 2003 May;90(5):755-60

Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9 Canada;

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous.
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http://dx.doi.org/10.3732/ajb.90.5.755DOI Listing
May 2003