Scandianthus gen.nov., Angiosperm Flowers of Gaxifragalean Affinity from the Upper Cretaceous of Southern Sweden

Structurally preserved angiosperm flowers are described fromthe Upper Cretaceous of southern Sweden. They are found in fluviatiledeposits dated, on palynological evidence, as Upper Santonianor Lower Campanian. The material studied includes two speciesof Scandianthus gen.nov. and one unnamed, related taxon. Thefossil flowers represent the best-preserved Cretaceous floralstructures available and the preservation permits detailed studyof organization and arrangement of parts. Comparison with flowersof extant angiosperms indicates a close relationship with membersof the Saxifragales. Scandianthus gen.nov. fossil angiosperm, Upper Cretaceous     

Organelle Sedimentation in Gravitropic Roots of Limnobium is Restricted to the Elongation Zone

Roots of the aquatic angiosperm Limnobium spongia (Bosc) Steud.were evaluated by light and electron microscopy to determinethe distribution of organelle sedimentation towards gravity.Roots of Limnobium are strongly gravitropic. The rootcap consistsof only two layers of cells. Although small amyloplasts arepresent in the central cap cells, no sedimentation of any organelle,including amyloplasts, was found. In contrast, both amyloplastsand nuclei sediment consistently and completely in cells ofthe elongation zone. Sedimentation occurs in one cell layerof the cortex just outside the endodermis. Sedimentation ofboth amyloplasts and nuclei begins in cells that are in theirinitial stages of elongation and persists at least to the levelof the root where root hairs emerge. This is the first modernreport of the presence of sedimentation away from, but not in,the rootcap. It shows that sedimentation in the rootcap is notnecessary for gravitropic sensing in at least one angiosperm.If amyloplast sedimentation is responsible for gravitropic sensing,then the site of sensing in Limnobium roots is the elongationzone and not the rootcap. These data do not necessarily conflictwith the hypothesis that sensing occurs in the cap in otherroots, since Limnobium roots are exceptional in rootcap originand structure, as well as in the distribution of organelle sedimentation.Similarly, if nuclear sedimentation is involved in gravitropicsensing, then nuclear mass would function in addition to, notinstead of, that of amyloplasts.Copyright 1994, 1999 AcademicPress Limnobium spongia, gravitropism, roots, sedimentation, cortex     

The First Record of Fossil Wood of Winteraceae from the Upper Cretaceous of Antarctica

Fossil wood of the Winteraceae from the Upper Cretaceous sedimentsof James Ross Island, Antarctic Peninsula, is described herefor the first time. The specimen is characterized by the absenceof vessels, rays of two distinct sizes and tracheids with one–threerows of circular bordered pits, mainly on the radial walls,grading to horizontally elongate and scalariform. Despite anatomicalconformity to the family Winteraceae, the fossil wood is notidentical to any one extant genus and therefore has been assignedto the fossil organ genus Winteroxylon Gottwald with which thefossil shows greatest similarity. Copyright 2000 Annals of BotanyCompany Antarctica, Cretaceous, angiosperm, wood, anatomy, Winteraceae, Winteroxylon, fossil, palaeoclimate     

A Fossil Wood Flora from King George Island: Ecological Implications for an Antarctic Eocene Vegetation

Early Tertiary sediments of the Antarctic Peninsula region continueto yield a rich assemblage of well-preserved fossil dicotyledonousangiosperm wood. The wood flora under consideration is fromthe Collins Glacier region on Fildes Peninsula, King GeorgeIsland and is derived from tuffaceous sediments of the MiddleUnit of the Fildes Formation. These deposits accumulated ina volcanic setting adjacent to a basic-intermediate stratocone.The fossil assemblage provides further evidence for the existenceof cool temperate forests, similar in composition to those foundtoday in New Zealand, Australia and, in particular, southernSouth America. This paper describes two conifer and five angiospermmorphotypes, four of which are new additions to the Antarcticapalaeoflora records. Cupressinoxylon Goeppert, which is thedominant conifer in terms of numbers, and Podocarpoxylon Gothanrepresent the conifers. The angiosperm component includes twospecies ofNothofagoxylon and two previously undescribed woodmorphotypes that exhibit greatest anatomical similarity to woodsofLuma A. Gray (Myrtaceae) and Eucryphia Cav. (Cunoniaceae).These morphotypes are described and assigned to the organ generaMyrceugenelloxylon Nishida, and Weinmannioxylon Petriella, respectively.A model based on the extant cool temperate Valdivian rainforestsis proposed and ecological reconstructions based on palaeobotanicaland geological evidence suggest that changes in the palaeovegetationreflect natural dynamics following volcanic disturbances.Copyright2001 Annals of Botany Company Fossil wood, Antarctica, Myrtaceae, Eucryphia, Nothofagaceae, Eocene, Valdivia, Tertiary     

Evolution of angiosperm seed disperser mutualisms: the timing of origins and their consequences for coevolutionary interactions between angiosperms and frugivores

The origins of interactions between angiosperms and fruit‐eating seed dispersers have attracted much attention following a seminal paper on this topic by Tiffney (1984). This review synthesizes evidence pertaining to key events during the evolution of angiosperm–frugivore interactions and suggests some implications of this evidence for interpretations of angiosperm–frugivore coevolution. The most important conclusions are: (i) the diversification of angiosperm seed size and fleshy fruits commenced around 80 million years ago (Mya). The diversity of seed sizes, fruit sizes and fruit types peaked in the Eocene around 55 to 50 Mya. During this first phase of the interaction, angiosperms and animals evolving frugivory expanded into niche space not previously utilized by these groups, as frugivores and previously not existing fruit traits appeared. From the Eocene until the present, angiosperm–frugivore interactions have occurred within a broad frame of existing niche space, as defined by fruit traits and frugivory, motivating a separation of the angiosperm–frugivore interactions into two phases, before and after the peak in the early Eocene. (ii) The extinct multituberculates were probably the most important frugivores during the early radiation phase of angiosperm seeds and fleshy fruits. Primates and rodents are likely to have been important in the latter part of this first phase. (iii) Flying frugivores, birds and bats, evolved during the second phase, mainly during the Oligocene and Miocene, thus exploiting an existing diversity of fleshy fruits. (iv) A drastic climate shift around the Eocene–Oligocene boundary (around 34 Mya) resulted in more semi‐open woodland vegetation, creating patchily occurring food resources for frugivores. This promoted evolution of a ‘flying frugivore niche’ exploited by birds and bats. In particular, passerines became a dominant frugivore group worldwide. (v) Fleshy fruits evolved at numerous occasions in many angiosperm families, and many of the originations of fleshy fruits occurred well after the peak in the early Eocene. (vi) During periods associated with environmental change altering coevolutionary networks and opening of niche space, reciprocal coevolution may result in strong directional selection formative for both fruit and frugivore evolution. Further evidence is needed to test this hypothesis. Based on the abundance of plant lineages with various forms of fleshy fruits, and the diversity of frugivores, it is suggested that periods of rapid coevolution in angiosperms and frugivores occurred numerous times during the 80 million years of angiosperm–frugivore evolution.     

A comparative study of the response of the roots and shoots of sunflower and maize to mechanical stimulation

Despite numerous studies of the effects of mechanical stimulationon plant shoots, the response of roots to mechanical stimulationhas largely been neglected. In this study the effects of shootflexure on the morphology and mechanics of two contrasting speciesof herbaceous angiosperm, growing in a glasshouse were compared:maize (Zea mays), a monocot; and sunflower (Helianthus annuusL.) a dicot. Mechanical stimulation affected the root more than the shootcomponents. Root systems of mechanicallystressed sunflowershad a greater angle of spread and increased root number. Aswell as large morphological and weight effects, with increasesover the control of 33% in the length of rigid root and 38%in the dry weight of lateral roots, in sunflowers, there werealso mechanical effects. In both species roots of flexed plantswere more rigid, stronger and composed of stiffer material andtheir root systems also provided greater anchorage strength.In contrast, there was only a small reduction in shoot weightand shoot height in flexed plants and no effects on mechanicalproperties. There were differences in behaviour between species; maize rootmorphology responded less than that of sunflowers to mechanicalstimulation. The basal diameter of roots increased by only 8%compared with 16% in sunflowers, though the roots of both speciesshowed similar increases in material stiffness. This differenceis related to the lack of secondary thickening in the monocotscompared with the dicot sunflowers. Key words: Thigmomorphogenesis, Helianthus annuus L., Zea mays, anchorage, lodging     

First Nuclear DNA C-values for 25 Angiosperm Families

DNA amount is a widely used biodiversity character. As knownDNA C-values represent the global angiosperm flora poorly, bettercoverage of taxonomic groups is needed, including at the familiallevel. A workshop, sponsored byAnnals of Botany , was held atthe Royal Botanic Gardens, Kew in 1997. Its key aim was to identifymajor gaps in our knowledge of plant DNA C-values and recommendtargets for new work to fill them by international collaboration.In 1997 C-values were known for approx. 150 families, meaningthere was no estimate for most angiosperm families (approx 68%).The workshop recommended a goal of complete familial representationby 2002, as a main target for angiosperms. Bennett et al. (Annalsof Botany86: 859–909, 2000) presented a fifth supplementarylist of angiosperm C-values from 70 original sources which includedfirst C-values for 691 species. Only 12 (1.7%) of these werefirst C-values for unrepresented families, so the need to improvefamilial representation was substantially unmet. We began newwork to address this in September 1999, and now report firstDNA C-values for 25 angiosperm families. Such targeting seemsessential to achieve the goal of familial coverage set by the1997 workshop within 5 years. 4C values range from 0.67 pg (similartoArabidopsis thaliana ) in Amoreuxia wrightii(Cochlospermaceae)to 7.49 pg in Deutzia prunifolia(Hydrangeaceae). These datasupport the view that ancestral angiosperms almost certainlyhad small genomes (defined as 1C     

Starch Grain Casts and Moulds in Eocene (Tertiary) Fossil Mangrove Hypocotyls

Further examples of the recently reported fossil starch graincasts and moulds in the fossil mangrove Ceriops cantiensis Chandlerfrom the London Clay (Eocene) of the Isle-of-Sheppey, Kent,England have now been discovered. In addition, similar structureshave been found in the fossil hypocotyls of Palaeobruguieraelongata Chandler from Herne Bay, Kent and also in four recentlycollected hypocotyls of a new species of fossil mangrove fromSheppey. Starch grains from herbarium material of the extantmangrove genera, Bruguiera and Ceriops, have been examined forcomparison with the fossil forms. starch grain casts, fossil, mangrove, hypocotyls, palaeobotany, Eocene, framboids, Rhizophoraceae, Bruguiera sexangula, Ceriops cantiensis, Palaeobruguiera elongata     

Endodermis-like Sheaths in the Submerged Freshwater MacrophyteRanunculus trichophyllusChaix

Light, fluorescence and electron microscopical analysis of therooted freshwater plantRanunculus trichophyllusrevealed a peculiaranatomical feature. In addition to the true endodermis encirclingthe root stele, endodermis-like sheaths occurred around eachvascular bundle of the leaf segments and of the eustelic stemwith its large central cavity, which assumed an anatomical featureresembling that of some pteridophyte stems. These impermeablesheaths, whose cells differentiate suberized walls, can playa major role in hampering the apoplastic leakage of the pressurizedwater solution which flows throughout the plant in xylem vesselsand contains the mineral nutrients taken up by the roots fromthe sediment. Moreover, these sheaths can function in preventingflooding of the aerenchymatic cavities of the submerged organs.In this way the endodermis-like sheaths preserve the correctcirculation of gas and nutrient solution through the entireorganism and assume great significance as a structural mechanismevolved by this species to survive and grow underwater.Copyright1999 Annals of Botany Company. Ranunculus trichophyllus,freshwater macrophyte, submerged angiosperm, anatomy, endodermis, endodermis-like sheaths, light microscopy, fluorescence microscopy.     

Nuclear DNA Amounts in Angiosperms--583 New Estimates

The amount of DNA in the unreplicated haploid nuclear genome(itsC-value) varies over 600-fold between angiosperm species.Information regarding this character is used in a strikinglywide variety of plant biological fields. Moreover, recent studieshave noted a significant need for more information about thisimportant aspect of genome biodiversity. Bennett and co-authorshave published four collected lists of nuclear DNA amounts inangiosperm species, compiled primarily for reference purposes,including the most recent in 1995 (Annals of Botany76: 113–176).Together they list estimates for over 2500 species which representabout 1% of the global angiosperm flora. Interest in angiospermgenome size has remained high, as shown by the recent publicationof many new estimates, creating a need for a fifth compilation.This paper presents a supplementary list of nuclear DNAC-valuesfrom 37 sources for 471 angiosperm species not listed in theaforementioned compilations, plus additional estimates for 113species already listed by them. It contains estimates for palm,orchid and tropical hardwood species which significantly improvesrepresentation of the global flora. Work is in hand to combinethe genome size data compiled in this and the aforementionedpapers into a unified database, and to present the informationin separate lists, with species in alphabetical and systematicorders, respectively. Meanwhile, the availability of DNAC-valuesfor angiosperm species can be checked on the World Wide Web(http://www.rbgkew.org.uk/cval/database1.html). Angiosperm DNA amounts; DNAC-values; nuclear genome sizes; reference lists; plant DNA database     

First Nuclear DNA C-values for Another 25 Angiosperm Families

Nuclear DNA C-value is an important genomic biodiversity characterwith many uses. An international workshop sponsored by Annalsof Botany and held at the Royal Botanic Gardens, Kew, UK, in1997 identified major gaps in our knowledge of plant DNA C-valuesand recommended targets for new work. Improved taxonomic coveragewas highlighted as a key need for angiosperms, especially atthe familial level. In 1997 C-values were known for only approx.32% of angiosperm families; a goal of complete familial representationby 2002 was recommended. A review published in 2000 (Bennettet al.;Annals of Botany86: 859–909) noted poor progresstowards this aim: of the 691 first C-values for species only12 (1.7%) were for unrepresented families. We began new workto address this in 1999, reporting first DNA C-values for 25angiosperm families in 2001 (Hanson et al.;Annals of Botany87:251–258). Here we report first DNA C-values for a further25 angiosperm families, increasing familial coverage in angiospermsto approx. 45%. Such targeting remains essential to approachthe goal set by the 1997 workshop of familial coverage for angiospermswithin 5 years. The 4C DNA amounts presented here range from0.76 pg (similar toArabidopsis thaliana ) in Roridula gorgonias(Roridulaceae)to 29.74 pg in Gunnera manicata(Gunneraceae). 1C values were< 3.5 pg in 23 of the 25 families; these data provide furthersupport for the view that ancestral angiosperms almost certainlyhad small genomes (defined as 1C     

Structural Changes During the in vitro Germination of Vanilla planifolia (Orchidaceae)

A study is reported of histogenesis and organogenesis duringthe processes leading up to seedling formation in cultures ofVanilla planifolia. Prior to germination, all cells of the embryoincreased in size rupturing the seed coat and initiating theprotocorm stage. The cells of the protocorm were heavily ladenwith starch grains. Although all of the cells of the matureembryo were heavily laden with protein bodies, these were confinedto the terminal cell descendents on emergence of the embryofrom the seed coat, and they disappeared during differentiationof the meristem, indicating that some reserves were mobilizedand utilized during germination. The terminal locus of embryonal axis did not differentiate intoa cotyledon and epicotyl as in other angiosperm embryos butformed a thick meristematic layer. Bipolar differentiation withinthe meristem produced the shoot, and after a few leaves hadbeen formed, the first root differentiated endogenously fromthe base of the meristem. Subsequent roots, however, appearedto originate more superficially. The chain of events appearsto be quite unique to Vanilla amongst the angiosperms. Vanilla planifolia, protocorm, starch grains, protein bodies     

Siptionella,a new bivalve genus of Parallelodontidae from the Upper Eocene of Ukraine

A new genus of the family Parallelodontidae (Bivalvia: Arcoidea), Siptionella gen. nov., with four species from the Upper Eocene of Ukraine and Germany and the Maestrichtian of Denmark is described. From the Upper Eocene of Ukraine, two species, the type species of the genus S. prompta (Berezovsky, 2002), previously described by the author within the genus Porterius, and the new species S. demissa sp. nov. are described and figured.     

Was the Early Eocene proboscidean Numidotherium koholense semi-aquatic or terrestrial? Evidence from stable isotopes and bone histology

The Early Eocene deposits of El Kohol, Algeria, have yielded numerous remains of Numidotherium koholense, one of the most primitive and oldest known proboscideans in Africa. The Upper Eocene proboscideans of the Fayum locality (Egypt), Barytherium sp. and Moeritherium sp., were recently interpreted as aquatic or semi-aquatic, according to the stable isotopic compositions (δ¹³C and δ¹⁸O) of their tooth enamel. These data led us to reinvestigate the adaptations of N. koholense. Stable isotopic analysis and observations of histological sections of its long bones reveal that it was essentially terrestrial. According to its position within the phylogenetic tree of Eocene proboscideans, the adaptation to semi-aquatic life appears to have evolved independently in different lineages of Middle and Upper Eocene proboscideans during their adaptive radiation in Africa. Moreover, these new results reopen the debate about the hypothesis that Eocene to Recent proboscideans are derived from semi-aquatic ancestors.     

Nuclear DNA Amounts in Angiosperms

Bennett and Smith (Philosophical Transactions of the Royal Societyof London B274:227-274; B334: 309-345) and Bennett, Smith andHeslop-Harrison (Proceedings of the Royal Society of London,B216: 179-199) published lists of nuclear DNA amounts estimatedfor 1612 angiosperm species collected from 163 sources datedbetween 1951 and 1986. Subsequently, interest in genome sizein angiosperms and its significance has continued, and manynew DNA estimates were published during 1986-1994. Their inaccessibility,and the flow of enquiries for such information, shows that afurther compilation is needed. This paper presents a supplementarylist of nuclear DNA C-values for 105 sources for 899 angiospermspecies not listed in the above-mentioned compilations, plus284 additional estimates for 208 species already listed by them.The data are assembled primarily for reference purposes, withspecies listed in alphabetical order, rather than by any taxonomicscheme. Some advantages and limitations of flow cytometry, nowincreasingly used to quantify DNA C-values in plants, are reviewed.Recent reports regarding the occurrence and extent of intraspecificvariation in genome size are also discussed. While some examplesare real, others reflect technical shortcomings. Work has begunto combine the genome size data compiled in this and the above-mentionedpapers into a unified data base, and to present the informationin separate lists, with species in alphabetical and systematicorders, respectively. DNA C-values are now known for 1% of theworld's angiosperm flora, but improved representation of taxonomicgroups, geographical regions and plant life forms is urgentlyneeded.Copyright 1995, 1999 Academic Press Angiosperm DNA amounts, DNA C-values, nuclear genome sizes, intraspecific variation, flow cytometry, reference lists, genome size database     

Mineral Nutrition of Sandalwood (Santalum spicatum)

Acacia acuminata is a preferred host of the root hemiparasitictree, Santalum spicatum (sandalwood). Comparison between nutrientcontent of adult trees of sandalwood and results for an earlierstudy of the mistletoe, Amyema preissii, on the same host species,A. acuminata, showed similar high levels of K and Na and lowlevels of Zn in both parasites compared with the host plants.Differences in K, Ca, N and Cu levels between parasitized anduninfected Acacias imply that the host plant contributes tothe nutrition of sandalwood. The high K/Ca ratio in sandalwoodconfirms that K uptake in preference to Ca is a general featureof all categories of angiosperm parasites. Patterns of distribution of nutrients between various partsof sandalwood and A. acuminata depend on the type of nutrient,but levels are usually highest in leaves of both species andthe haustoria. Although K, Ca and Na are much lower in the kernelsthan in vegetative parts of the parasite, only seedlings withoutsupplementary Ca in a nutrient omission experiment failed togrow at all in the absence of hosts. Growth is not dependenton the level of K in the unattached plants but other evidenceindicates it may have a role in water uptake in the attachedplant. Calcium supply has a marked effect on internal Ca levelsand growth of unattached plants. Compared with field plants,levels of Ca, and to a lesser extent Zn, were much higher inplants of the Ca/K treatment that produced greatest growth over34 weeks. Haustorial formation is enhanced by the presence of A. acuminataroots. However, competition for nutrients, especially Ca, fromco-planted A. acuminata seedlings results in suppression ofgrowth of young sandalwood compared with their growth in theabsence of the host species. Key words: Mineral nutrients, Santalum spicatum, Acacia acuminata, hemiparasites, K/Ca nutrition, seeds     

Defective Secretion of Mucilage is the Cellular Basis for Agravitropism in Primary Roots of Zea mays cv. Ageotropic

Root caps of primary, secondary, and seminal roots of Z. mayscv. Kys secrete large amounts of mucilage and are in close contactwith the root all along the root apex. These roots are stronglygraviresponsive. Secondary and seminal roots of Z. mays cv.Ageotropic are also strongly graviresponsive. Similarly, theircaps secrete mucilage and closely appress the root all alongthe root apex. However, primary roots of Z. mays cv. Ageotropicare non-responsive to gravity. Their caps secrete negligibleamounts of mucilage and contact the root only at the extremeapex of the root along the calyptrogen. These roots become graviresponsivewhen their tips are coated with mucilage or mucilage-like materials.Peripheral cells of root caps of roots of Z. mays cv. Kys containmany dictyosomes associated with vesicles that migrate to andfuse with the plasmalemma. Root-cap cells of secondary and seminal(i.e. graviresponsive) roots of Z. mays cv. Ageotropic are similarto those of primary roots of Z. mays cv. Kys. However, root-capcells of primary (i.e. non-graviresponsive) roots of Z. mayscv. Ageotropic have distended dictyosomal cisternae filled withan electron-dense, granular material. Large vesicles full ofthis material populate the cells and apparently do not fusewith the plasmalemma. Taken together, these results suggestthat non-graviresponsiveness of primary roots of Z. mays cv.Ageotropic results from the lack of apoplastic continuity betweenthe root and the periphery of the root cap. This is a resultof negligible secretion of mucilage by cells along the edgeof the root cap which, in turn, appears to be due to the malfunctioningof dictyosomes in these cells. Corn, dictyosomes, mucilage, root gravitropism, Zea mays cv. Ageotropic, Zea mays cv. Kys     

Relationship Between Lateral Root Primordia in Different Ranks

Comparisons with theoretical random distributions indicate thatthe longitudinal positional relationships between lateral rootsin different ranks are random in roots of Musa acuminata, Pistiastratiotes and i, species in which lateral root primordia arisein the root tip. In Potentilla palustris, where lateral rootprimordia arise further back in the root, there is a deficiencyof close spacings, which indicates that there is some interactionbetween ranks. Musa acuminata, Pistia stratiotes, Pontederia cordat, Potentilla palustris, roots, lateral roots, pattern     

Some Effects of Sucrose Concentration on Primordium Development in Excised Primary Roots

The effects of varying the amount of sucrose used to supplementthe culture medium maintaining the growth of excised roots ofPisum sativum L., Vicia faba L., Zea mays L. and Phaseolus vulgarisL., on the rates of primordium initiation and subsequent emergenceas lateral roots and on the duration of the interval betweenprimordium inception and emergence as a secondary root throughthe tissues of the primary have been investigated. Variation in the exogenous concentration of sucrose from 0.5to 8 per cent had little effect on the rate of primordium inceptionin Pisum and Vicia and the rates never reached the values obtainedfor the roots of the corresponding intact plants. Moreover,over the 6 day culture period lateral root emergence did notoccur in any of the excised roots of these two species. In contrast,each of the aspects of primordium development examined in theexcised roots of Zea and Phaseolus was markedly affected bythe amount of sucrose used to supplement the culture medium.In addition, in the presence of about 6 per cent sucrose, primordiumdevelopment in these cultured roots was very similar to thatin roots of the corresponding intact plants. The results indicate either that some factor necessary for primordiumdevelopment is present in adequate amounts in excised rootsof Zea and Phaseolus, but not in those of Pisum and Vicia, orthat the factors controlling such development are differentin the former and latter two species. Vicia faba L., Pisum sativum L., Zea mays L., Phaseolus vulgaris L., broad bean, garden pea, maize, dwarf bean, primordium development, sucrose concentration, cultured roots