Floral evolution in Lithospermum (Boraginaceae): independent origins of similar flower types

In Lithospermum (Boraginaceae), floral diversity is quite large, with variation in individual quantitative and qualitative traits as well as suites of floral characteristics. The present study utilizes phylogenetic, morphometric and developmental methodologies to investigate patterns of floral evolution, of individual and suites of traits, in the genus and among related genera. The evolutionary patterns of eight quantitative and five qualitative traits were reconstructed, and morphometrics and the evolution of floral morphospace were examined in the genus. Floral developmental patterns were established with light and scanning electron microscopy. Phylogenetic analyses provided evidence that derived flower types have each evolved multiple times, with differences in the manner in which these flower types arose. Floral morphospace has increased throughout the evolution of Lithospermum. Floral developmental patterns and cell lengths of floral organs at anthesis provided evidence that, in Lithospermum, an increase in the length of the sexual organs of flowers involves both longer cells and a greater number of mitotic divisions, but an increase in the length of the corolla is primarily a result of a larger number of cell divisions. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 213–228.     

A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships,taxonomy, and patterns of character evolution

The angiosperm family Boraginaceae includes ca. 1600 species distributed among ca. 110 genera. Some floral features are constant within the family, but many vegetative, floral, pollen, and nutlet traits vary. Utilizing 224 species of Boraginaceae and related taxa, five matrices were constructed with various combinations of morphological characters, three chloroplast DNA regions, and one nuclear ribosomal DNA region. Phylogenetic analyses were conducted for these matrices, and patterns of character evolution were examined. Boraginaceae is resolved as monophyletic, with Wellstedia as its sister. Codon is sister to Boraginaceae + Wellstedia. Although most of the investigated morphological characters have a low consistency index, particular character states are synapomorphies for large clades in each of the tribes of the family. In Boraginaceae, the breeding system heterostyly evolved at least 12 times, which is the largest number of origins resolved in any family; therefore Boraginaceae can serve as a model for the evolution and development of heterostyly. Nutlet ornamentation is most diverse in Cynoglosseae and Trichodesmeae, while pollen and floral features are most variable in Boragineae and Lithospermeae. Phylogenetic relationships and patterns of character evolution identified in the present study set the stage for future work creating an updated taxonomic system of Boraginaceae.     

Nomenclatural changes in <Emphasis Type="Italic">Lithospermum</Emphasis> (Boraginaceae) and related taxa following a reassessment of phylogenetic relationships

Lithospermum (Boraginaceae) comprises approximately 40 species in both the Old and New Worlds, with a center of diversity in the southwestern United States and Mexico. Using ten cpDNA regions, a phylogeny of Lithospermum and related taxa was reconstructed. Lithospermum (including New World and Old World species) and related New World members of Lithospermeae form a monophyletic group, with Macromeria, Onosmodium, Nomosa, Lasiarrhenum, and Psilolaemus nested among species of Lithospermum. New World Lithospermeae also is a monophyletic group, with Eurasian species of Lithospermum sister to this group. Because Lithospermum is not monophyletic without the inclusion of the other New World genera, species from these genera are transferred to Lithospermum, and appropriate nomenclatural changes are made. New combinations are Lithospermum album, Lithospermum barbigerum, Lithospermum dodrantale, Lithospermum exsertum, Lithospermum helleri, Lithospemum leonotis, Lithospermum notatum, Lithospermum oaxacanum, Lithospermum pinetorum, Lithospermum rosei, Lithospermum trinverium, and Lithospermum unicum; new names are Lithospermum chiapense, Lithospermum johnstonii, Lithospermum macromeria, Lithospermum onosmodium, Lithospermum rzedowskii, and Lithospermum turneri.     

Leaf architecture and anatomy of eleven species of Mortoniodendron (Malvaceae s.l.)

The genus Mortoniodendron is distributed from southeastern Mexico to Colombia and contains about 18 species of trees and shrubs. The main goal of this study is to describe and compare the leaf architecture and the anatomy of 11 species of Mortoniodendron using cleared leaves and paradermal and transverse leaf sections. All of the species of Mortoniodendron share a simple lamina with an entire margin. The differences among the species include the venation pattern, the relative thickness and course of the secondary veins, the presence or absence of the intersecondary veins, the tertiary vein pattern, the number of branches in veinlets, and the marginal ultimate venation. The leaf anatomy is similar to other species of the Malvaceae s.l.: for example, mucilage cells are common in Mortoniodendron. There are unique attributes of the leaf architecture and anatomy, such as the eucamptodromous venation pattern of M. guatemalense, which facilitates the identification of the species. The species M. palaciosii, M. pentagonum, M. ruizii, M. uxpanapense, and M. sp. nov. share features that suggest that they are closely related.     

中国花楸属单叶类群叶脉序特征研究

中国花楸属单叶类群(24种)包含落萼组与冠萼组,亦有学者将其分为两个属(白花楸属和水榆属)或合并成为一个属(白花楸属).由于其花序和果实的高度相似性,长期以来分类地位具有争议,而叶脉序特征在花楸属单叶类群中的分类作用尚不明确.因此,该研究以国产花楸属落萼组的13个物种和冠萼组的7个物种作为研究对象,以清净法制作标本对叶...     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

Marbleseeds are gromwells – Systematics and evolution of Lithospermum and allies (Boraginaceae tribe Lithospermeae) based on molecular and morphological data

Phylogenetic relationships are complex within the Lithospermeae, a large subgroup of the Boraginaceae s.str. The relationships of New World Lasiarrhenum, Macromeria, Nomosa, Onosmodium, Perittostoma, and Psilolaemus to subcosmopolitan and much larger Lithospermum have not been critically investigated in the recent past. No molecular data on the phylogeny of these genera and Lithospermum have so far been published. We investigated the relationships within Lithospermeae using three loci (nuclear ITS plus 5.8S rRNA, chloroplast trnL-F-spacer, and trnS-G-spacer) and micromorphological character traits (pollen, nutlets). Lithospermum s.l. constitutes the sistergroup of Asian Ulugbekia and is monophyletic only when its American segregates “Macromeria”, monotypic Nomosa, and Onosmodium are included. Both the African and the South American species groups of Lithospermum are monophyletic, but North American representatives are not resolved in a single clade. Morphological characters that have been considered as important for generic delimitation in the past (such as large, yellow corollas without faucal scales, particular pollen types, coarsely veined leaves, shrubby habit) have evolved in at least two only distantly related lineages within Lithospermum s.l. The reduction of American “Macromeria”, Nomosa, and Onosmodium as well as Asian Ulugbekia under Lithospermum is proposed to render the latter monophyletic. This redefined Lithospermum s.l. appears to have undergone a type of recent “island radiation” in the Americas, reflected in a morphological diversity far exceeding that found in the Old World.     

贵州小檗属植物(小檗科)叶脉序研究

采用制作叶脉标本和透明叶标本的方法,对贵州产28种2变种小檗属植物叶脉特征进行比较研究。结果表明:贵州小檗属植物的脉序类型有5种:半达缘羽状脉、花环状半达缘羽状脉、简单弓形羽状脉、花环状弓形羽状脉和混合型。叶脉分支一般有五级:1一级脉构架均为羽状脉,粗度有很粗、粗、中等粗细和纤细四种类型,分支方式包括单轴分支和合轴分支;2粗二级脉构架中有分支达缘或分支均不达缘,与中脉夹角变化各异,内二级脉存在或缺失,细二级脉半达缘、真曲行或简单弓形,间二级脉类型复杂多变但频度种间有差异;3三级脉贯串型、结网型或分支型;4四、五级脉网状或自由分支且常混合在一起。脉间区从发育差到良好,小脉从不分支到不均等分支等各种类型均有,叶缘末级脉缺失、不完整、钉状和环状。大部分种类叶缘具齿,每1cm齿数目和齿内腺点的特性等特征在不同种类间有区别,具有鉴定价值,但齿其它特征复杂多变或种间区别较小,同时齿内脉性状也不稳定。此外,齿的有无会对脉序类型产生影响。小檗属植物叶脉类型存在种间差异,具有重要的分类学价值,叶脉类型的变化和复杂程度显示了该属植物的进化特点;叶齿的有无和齿特征具有分类学和系统学意义。基于叶脉特征的研究结果并结合重要的外部形态学特征编制了贵州小檗属植物的分种检索表。研究结果可为小檗属植物分类寻找新的依据并探讨其系统学意义。     

中国吴茱萸属(广义)叶结构特征及其分类学意义北大核心CSCD

该研究以中国产芸香科广义吴茱萸属18种3变种为材料,于体视显微镜下观察叶结构特征。结果发现:(1)吴茱萸属叶脉为羽状脉。(2)二级脉有花环状弓形脉和简单弓形脉两种类型,极少数存在内二级脉,间二级脉存在或缺失。(3)三级脉不规则网状或极少数弱贯穿,边缘三级脉环状。(4)四级脉不规则网状或自由分支。(5)五级脉不规则网状或自由分支,脉间区发育差或中等,游离端小脉末端简单或管状异形,边缘末级脉大多数不完整,极少数环状。(6)基于重要的外部形态学特征和叶结构特征观察结果,编制了广义吴茱萸属植物的分组检索表。研究表明,吴茱萸属这些类群的二级脉与更高级脉序形成的结构极为稳定,同时又存在种间差异,故广义吴茱萸属植物叶结构特征可以为更准确地鉴定一些疑难种和混淆种提供佐证,具有重要的系统分类学价值。     

LIRIODENDRON (MAGNOLIACEAE) FROM THE MIOCENE CLARKIA FLORA OF IDAHO

Miocene Liriodendron carpels, whole fruiting structures and leaves from Clarkia and Oviatt Creek sites in northern Idaho are preserved as imprints and compressed fossils in soft lacustrine clays. The isolated carpels are indistinguishable from those described as L. hesperia Berry from the Spokane Latah flora. Fruit aggregates from the type Clarkia and Oviatt Creek localities and leaves from three Clarkia sites are considered to be within the range of variation of the single species L. hesperia. Comparisons were made regarding leaf architecture, lower leaf epidermal structures, leaf flavonoid and steroid analysis, morphological features of receptacles and carpels, and the venation pattern of carpels of the fossil material to the two extant species, L. tulipifera L. (native to southeastern United States) and L. chinense Sarg. (native to southeastern Asia). Leaf architecture features analyzed by standard statistical and canonical tests and marginal venation patterns near the base of leaves suggest that L. hesperia is more similar to L. tulipifera, whereas the size dimensions of lower epidermal cells and the common presence of two sterane compounds imply that L. hesperia is more similar to L. chinense. The fossil species, however, is a distinct taxon indicated by statistical discriminant and canonical tests, leaf base shape, often smaller epidermal cell dimensions, and the shape of round receptacle carpel scars. Both the fossil and the two living Liriodendron species are associated with comparable mixed mesophytic floras.     

Molecular phylogeny of the diatom genera Amphora and Halamphora (Bacillariophyta) with a focus on morphological and ecological evolution

The taxonomic history of the diatom genus Amphora is one of a broad early morphological concept resulting in the inclusion of a diversity of taxa, followed by an extended period of revision and refinement. The introduction of molecular systematics has increased the pace of revision and has largely resolved the relationships between the major lineages, indicating homoplasy in the evolution of amphoroid symmetry. Within the two largest monophyletic lineages, the genus Halamphora and the now taxonomically refined genus Amphora, the intrageneric morphological and ecological relationships have yet to be explored within a phylogenetic framework. Critical among this is whether the range of morphological features exhibited within these diverse genera are reflective of evolutionary groupings or, as with many previously studied amphoroid features, are nonhomologous when examined phylogenetically. Presented here is a four‐marker molecular phylogeny that includes 31 taxa from the genus Amphora and 77 taxa from the genus Halamphora collected from fresh, brackish, and salt waters from coastal and inland habitats of the United States and Japan. These phylogenies illustrate complex patterns in the evolution of frustule morphology and ecology within the genera and the implications of this on the taxonomy, classification, and organization of the genera are discussed.     

Evolution of skull and body shape in Triturus newts reconstructed from three‐dimensional morphometric data and phylogeny

To explore the relationship between morphological change and species diversification, we reconstructed the evolutionary changes in skull size, skull shape, and body elongation in a monophyletic group of eight species that make up salamander genus Triturus. Their well‐studied phylogenetic relationships and the marked difference in ecological preferences among five species groups makes this genus an excellent model system for the study of morphological evolution. The study involved three‐dimensional imagery of the skull and the number of trunk vertebrae, in material that represents the morphological, spatial, and molecular diversity of the genus. Morphological change largely followed the pattern of descent. The reconstruction of ancestral skull shape indicated that morphological change was mostly confined to two episodes, corresponding to the ancestral lineage that all crested newts have in common and the Triturus dobrogicus lineage. When corrected for common descent, evolution of skull shape was correlated to change in skull size. Also, skull size and shape, as well as body shape, as inferred from the number of trunk vertebrae, were correlated, indicating a marked impact of species' ecological preferences on morphological evolution, accompanied by a series of niche shifts, with the most pronounced one in the T. dobrogicus lineage. The presence of phylogenetic signal and correlated evolutionary changes in skull and body shape suggested complex interplay of niche shifts, natural selection, and constraints by a common developmental system. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 243–255.     

Cryptic diversity,intraspecific phenetic plasticity and recent geographical translocations in Branchiomma (Sabellidae,Annelida)

The importance of identifying biological diversity accurately and efficiently is becoming more evident. It is therefore critical to determine the species boundaries between closely related taxa and to establish diagnostic characters that allow us to define species. This is not an easy task when species exhibit high intraspecific phenotypic plasticity or when distinct evolutionary lineages with an unusually large amount of genetic distinctiveness show no apparent morphological diversity (cryptic species). These phenomena appear to be common in the genus of fan worms Branchiomma (Sabellidae, Annelida), and consequently, taxonomic errors are widespread in the group. Moreover, some Branchiomma species have been unintentionally translocated outside the area where natural range extension is expected, increasing the taxonomic problems. We have performed a range of analytical methods including genetic distances, Bayesian inference, maximum likelihood, maximum parsimony, statistical parsimony analyses and general mixed Yule coalescent model to clarify the taxonomic status and assess the species boundaries of Branchiomma in Australia. This study shows that the traditional diagnostic morphological features are greatly homoplastic. Results also indicate that the diversity of Branchiomma in Australia is higher than previously reported and evidence some cases of high phenetic plasticity (in features previously considered as stable within species), high intraspecific genetic variability, cryptic species and several unexpected cases of translocations.     

CHARLIEA MANZANITANA,N. GEN., N. SP., AND OTHER ENIGMATIC PARALLEL-VEINED FOLIAR FORMS FROM THE UPPER PENNSYLVANIAN OF NEW MEXICO AND TEXAS

Charliea is a new genus (type-species: C. manzanitana), based on pinnately compound leaf material from the richly fossiliferous Virgilian (Upper Pennsylvanian) shales of the Kinney Brick Company quarry near Albuquerque, New Mexico. In several features Charliea resembles Russellites or a zamioid cycad. It has linear-oblong pinnae with broad, oblique attachment and a truncate tip, which is deeply incised to form two to four nearly equal lobes. The venation is simple, parallel, and sparingly dichotomous, each vein ending at the distal margin. The Kinney beds also contain Plagiozamites planchardi, another zamioid form with parallel-veined pinnae, differing from Charliea chiefly in having rounded tips and veins ending in the denticulate margins. An unnamed third form (genus B) in the Kinney beds has long, narrow pinnae with parallel veins and blunt tips; this strongly resembles the Mesozoic conifer Podozamites, but may just as well represent a cycadophyte. Another unnamed taxon (genus A), from an Upper Pennsylvanian deposit in Jack County, Texas, resembles genus B or Russellites in general shape and venation, but the critical distal margins are unknown. In their single-ordered parallel venation, these four foliar types contrast sharply with the two-ordered pinnate venation of most Pennsylvanian fern-like leaves, and seem to foreshadow Mesozoic morphologies. This tendency toward precocious evolution of parallel-veined foliar form in North America is also expressed by a single occurrence of the Asiatic, Permian genus Tingia in the Lower Pennsylvanian of Utah, and by the presence of the predominantly Triassic cycadeoid genus Pterophyllum in the Lower Permian of Texas.     

Phylogenetic Relationships and Morphological Character Evolution of Photosynthetic Euglenids (Excavata) Inferred from Taxon‐rich Analyses of Five Genes

Photosynthetic euglenids acquired chloroplasts by secondary endosymbiosis, which resulted in changes to their mode of nutrition and affected the evolution of their morphological characters. Mapping morphological characters onto a reliable molecular tree could elucidate major trends of those changes. We analyzed nucleotide sequence data from regions of three nuclear‐encoded genes (nSSU, nLSU, hsp90), one chloroplast‐encoded gene (cpSSU) and one nuclear‐encoded chloroplast gene (psbO) to estimate phylogenetic relationships among 59 photosynthetic euglenid species. Our results were consistent with previous works; most genera were monophyletic, except for the polyphyletic genus Euglena, and the paraphyletic genus Phacus. We also analyzed character evolution in photosynthetic euglenids using our phylogenetic tree and eight morphological traits commonly used for generic and species diagnoses, including: characters corresponding to well‐defined clades, apomorphies like presence of lorica and mucilaginous stalks, and homoplastic characters like rigid cells and presence of large paramylon grains. This research indicated that pyrenoids were lost twice during the evolution of phototrophic euglenids, and that mucocysts, which only occur in the genus Euglena, evolved independently at least twice. In contrast, the evolution of cell shape and chloroplast morphology was difficult to elucidate, and could not be unambiguously reconstructed in our analyses.     

Out of Alaska: morphological diversity within the genus <Emphasis Type="Italic">Eurytemora</Emphasis> from its ancestral Alaskan range (Crustacea,Copepoda)

The copepod genus Eurytemora occupies a wide range of habitat types throughout the Northern Hemisphere, with among the broadest salinity ranges of any known copepod. The epicenter of diversity for this genus lies along coastal Alaska, where several species are endemic. Systematic analysis has been difficult, however, because of a tendency toward morphological stasis in this genus, despite large genetic divergences among populations and species. The goals of this study were to (1) analyze patterns of morphological variation and divergence within this genus, focusing on Eurytemora species that occur in North America, and (2) determine patterns of geographic and salinity distribution of Eurytemora species within the ancestral range in Alaska. We applied a comparative multivariate morphological analysis using 16–26 characters from 125 specimens from 20 newly collected sites in Alaska and 15 existing samples predominantly from North America. Results from principal component and hierarchical cluster analyses identified seven distinct morphological species of Eurytemora in North America (E. affinis, E. americana, E. canadensis, E. composita, E. herdmani, E. pacifica, and E. raboti), and identified diagnostic characters that distinguish the species (forming the basis for a new identification key). Several previously named species were regarded as synonyms. The sites we sampled in Alaska were remarkable in the high levels of sympatry of Eurytemora species, to a degree not seen outside of Alaska. Future studies of Eurytemora should shed light on patterns of habitat invasions and physiological evolution within the genus, and yield insights into mechanisms leading to its remarkably broad geographic and habitat range.     

Evolutionary processes and its environmental correlates in the cranial morphology of western chipmunks (Tamias)

The importance of the environment in shaping phenotypic evolution lies at the core of evolutionary biology. Chipmunks of the genus Tamias (subgenus Neotamias) are part of a very recent radiation, occupying a wide range of environments with marked niche partitioning among species. One open question is if and how those differences in environments affected phenotypic evolution in this lineage. Herein we examine the relative importance of genetic drift versus natural selection in the origin of cranial diversity exhibited by clade members. We also explore the degree to which variation in potential selective agents (environmental variables) are correlated with the patterns of morphological variation presented. We found that genetic drift cannot explain morphological diversification in the group, thus supporting the potential role of natural selection as the predominant evolutionary force during Neotamias cranial diversification, although the strength of selection varied greatly among species. This morphological diversification, in turn, was correlated with environmental conditions, suggesting a possible causal relationship. These results underscore that extant Neotamias represent a radiation in which aspects of the environment might have acted as the selective force driving species’ divergence.     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

Gogorevia,a New Monoraphid Diatom Genus for Achnanthes exigua and Allied Taxa (Achnanthidiaceae) Described on the Basis of an Integrated Molecular and Morphological Approach

A new diatom genus, Gogorevia, is described on the basis of an integrated molecular and morphological investigation. Two species are described from Vietnam: the generitype, G. rinatii sp. nov., and G. ovalis sp. nov. These species share many morphological features with Achnanthes exigua and allied taxa that were previously transferred to the genera Achnanthidium or Lemnicola. Our current molecular investigation shows that Gogorevia is distinct and phylogenetically removed from the above-mentioned genera. Morphologically, Gogorevia differs from Achnanthidium and Lemnicola by the shape of the valves, possessing uniseriate striae, raphe morphology, and the presence of an evident sternum. The recently described species Lemnicola uniseriata is transferred to Gogorevia on the basis that it possesses the same morphological features and phylogenetic position as other members of the genus.