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Systematic palynology in Ebenaceae with focus on Ebenoideae: Morphological diversity and character evolution
Abstract
This study examines the diversity and character transformations in pollen and orbicule morphology of Ebenaceae, with a focus on subfamily Ebenoideae (ca. 600 sp.). 62 specimens comprising all three genera of Ebenoideae (Diospyos, Euclea, Royena), were studied using LM and SEM. Bayesian phylogenetic analysis was performed on molecular sequence data to establish an evolutionary hypothesis that was then used as an evolutionary framework to identify synapomorphies and trace evolutionary trends of palynological data with Bayesian posterior mapping and principle component analyses (PCA). Ebenoideae pollen is generally shed as monads (permanent tetrads in two species), medium-sized, prolate-spheroidal to subprolate and tricolporate. A substantial amount of variation is found in pollen size, equatorial outline (lobate, subtriangular, circular and hexagonal) and sexine ornamentation type ((micro)rugulate, striate, granulate and gemmate). Moreover, orbicules were present on the inner locule wall in all specimens examined. Their abundance, degree of fusion with tapetal membrane and aggregation vary considerably. We can conclude that Ebenaceae pollen is more heterogeneous than previously assumed. We traced palynological synapomorphies for groups at different taxonomic levels: subfamily level (pollen size, pollen wall stratification and aperture morphology), generic level (size, equatorial outline and sexine ornamentation types) and subgeneric clades (size, ectocolpus morphology, equatorial outline and sexine ornamentation subtypes) respectively. The granular infratectum and the unique sculpturing pattern on the orbicule walls are the most discriminating pollen features for subfamily Ebenoideae.
... Remarks: The overall shape, aperture configuration and ektexine ornamentation is characteristic for many Diospyros Linné and Royena taxa (e.g., Erdtman, 1969;Geeraerts et al., 2009;Grygorieva et al., 2013, Morton, 1994Morton and Kincaid, 1995), but this is a quite large specimen and only a few species investigated by Geeraerts et al. (2009) fig. 2G), but this species is considerably smaller and has a subspheroidal shape. ...
... Royena fischeri, R. loureiroana, R. austroafricana, Diospyros batocana, D. lotus, D. mespiliformis, D. oubatchensis) and many of these taxa are characterized by exceptionally large pollen grains. The distinction of the two genera on the basis of pollen morphology is difficult (Geeraerts et al., 2009;Morton and Kincaid, 1995), and pollen morphological studies cover only a tenth of the species (e.g., Erdtman, 1969;Geeraerts et al., 2009;Gosling et al., 2013;Grygorieva et al., 2013;Morton, 1994;Morton and Kincaid, 1995;Tissot et al., 1994). The reason we refer the first Ebenaceae as "cf. ...
... Royena fischeri, R. loureiroana, R. austroafricana, Diospyros batocana, D. lotus, D. mespiliformis, D. oubatchensis) and many of these taxa are characterized by exceptionally large pollen grains. The distinction of the two genera on the basis of pollen morphology is difficult (Geeraerts et al., 2009;Morton and Kincaid, 1995), and pollen morphological studies cover only a tenth of the species (e.g., Erdtman, 1969;Geeraerts et al., 2009;Gosling et al., 2013;Grygorieva et al., 2013;Morton, 1994;Morton and Kincaid, 1995;Tissot et al., 1994). The reason we refer the first Ebenaceae as "cf. ...
The 1 m thick intercalated marly sediments of Entrischenbrunn represent a floodplain pond within a braided fluvial system of late Langhian age. The organic rich upper part of the marls that are characterized by the presence of numerous plant macrofossils, are investigated for palynomorphs with SEM. The results reveal that many typical azonal broad leaved forest elements were present (e.g., Fraxinus and Liquidambar) and some of the common macrofossils, such as Hemitrapa, Podocarpium, Platanus sp., Quercus spp. etc. However, because of non-resistant pollen walls, the two dominant components of the macroflora, Daphnogene and Populus are not represented in the palynoflora. In contrast, plants not known from macrofossils or rare for this locality are present: several aquatic plants (e.g., Callitriche sp., Ludwigia sp., Decodon sp.), and a large portion of more subhumid and sclerophyllous plants such as Celtis “tournefortii-type”, two Ephedra spp., two Erica spp. affiliable with Mediterranean species, Quercus sp. section Ilex, a Sapoteae, Sideroxylon sp., Rehderodendron sp., Ulmus sp. section Foliaceae Zelkova “abelicea/serrata-type”, and Ziziphus “mucronata/mauritania/jujube-type”, whereas Diospyros sp. and cf. Royena sp. can be interpreted to present humid to subhumid conditions. The reasons for this mixture might be threefold: 1. localized well-drained substrates and water loss on particularly pebbly, gravelly and sandy deposits of the braided river system or 2. presence of a geographically controlled localized lower humidity in that area, or 3. because of a generally drier period within the Middle Miocene.
... In contrast, flowers of Diospyros possess a genus-diagnostic feature, namely the calyx of the female flower that is leathery, persistent, and often accrescent in fruit. Together with the diagnostic pollen (Geeraerts et al. 2009) this can be used to reliably trace the genus in the fossil record. ...
... -The pollen morphology of Diospyros has been investigated in several studies (e.g. Ng 1971;Cerceau-Larrival et al. 1984;de Franceschi 1993;Morton 1994;Tissot et al. 1994;Kodela 2006;Geeraerts et al. 2009;Sánchez-Dzib et al. 2009;Grygorieva et al. 2010Grygorieva et al. , 2013Li et al. 2011;Miyoshi et al. 2011;Gosling et al. 2013). Pollen of extant Diospyros species display a wide range in size and outline, but many of the investigated species have a distinct (micro)rugulate sculpture (subtype 1b of Geeraerts et al. 2009). ...
... Ng 1971;Cerceau-Larrival et al. 1984;de Franceschi 1993;Morton 1994;Tissot et al. 1994;Kodela 2006;Geeraerts et al. 2009;Sánchez-Dzib et al. 2009;Grygorieva et al. 2010Grygorieva et al. , 2013Li et al. 2011;Miyoshi et al. 2011;Gosling et al. 2013). Pollen of extant Diospyros species display a wide range in size and outline, but many of the investigated species have a distinct (micro)rugulate sculpture (subtype 1b of Geeraerts et al. 2009). The fossil pollen reported from Søby further shares the long colpi and perforations in interapertural areas with pollen of extant Diospyros (Geeraerts et al. 2009, Table 1). ...
Diospyros is a large genus of woody flowering plants with a predominantly subtropical and tropical modern distribution. Fossils attributed to Diospyros are mainly leaf impressions from Cretaceous and Cenozoic strata across the Northern Hemisphere. However, it is difficult to assign such fossils to Diospyros because genus-diagnostic leaf characteristics remain to be identified. Unequivocal fossil records of Diospyros are few, including staminate flowers with in situ pollen from the late Eocene of South Australia and dispersed pollen from Cenozoic strata in the Northern Hemisphere. Here, we investigated dispersed pollen and calyx remains from Miocene deposits of Denmark using a combined scanning electron/light microscopy approach. Tricolporate, relatively large pollen with lalongate pori and long bow-shaped colpi and a distinctive micro/nanorugulate exine ornamentation together with persistent 4-lobed flower calyces allow unambiguous identification of the genus. Based on the large size of the calyx, we describe a new fossil-species of Diospyros. Further, a review of the fossil pollen record of Diospyros shows that, in addition to the Australian record, the genus was present in South China, western North America and Europe during the Eocene and in East and South Africa and Central Asia during the Oligo-Miocene. Although still scanty, the pollen record can contribute vital information for time-calibrated molecular phylogenies to resolve conflicting biogeographic inferences. A thorough description of the historical biogeography of Diospyros is still in its infancy. While we initiate such a study here, development of a comprehensive picture will require further studies of dispersed pollen grains with high taxonomic resolution.
... The > 500 species are distributed among two subfamilies and four genera: the monogeneric subfamily Lissocarpoideae consisting of the South American endemic genus Lissocarpa Benth. with eight species; and the subfamily Ebenoideae consisting of three genera, the African endemic genera Euclea L. (18 species) and Royena L. (17 species), and the widely distributed genus Diospyros L. (500 species) (Duangjai et al., 2006;Geeraerts et al., 2009;Turner et al., 2013;Samuel et al., 2019). Based on the tropical nature of most of the representative taxa and disjunct distribution patterns, Ebenaceae are believed to have an early divergence history during the Cretaceous in western Gondwana (Raven & Axelrod, 1974). ...
... These characters can now be used effectively in phylogenetic studies for reconstructing the divergence time and historical biogeography of plants ( Barreda et al., 2014;He et al., 2016;Grímsson et al., 2017a, b). Detailed palynological studies using SEM imaging techniques have already been conducted on extant and extinct members of Ebenaceae to understand pollen morphological character evolution and the biogeography of the family (Geeraerts et al., 2009;Turner et al., 2013;Hoffmann, 2018;Linan et al., 2019). After reviewing observations on Ebenaceae pollen by many workers (Morton, 1994), Geeraerts et al. (2009) were the first to describe pollen morphology of extant members of Ebenaceae in detail using SEM and LM. ...
... Detailed palynological studies using SEM imaging techniques have already been conducted on extant and extinct members of Ebenaceae to understand pollen morphological character evolution and the biogeography of the family (Geeraerts et al., 2009;Turner et al., 2013;Hoffmann, 2018;Linan et al., 2019). After reviewing observations on Ebenaceae pollen by many workers (Morton, 1994), Geeraerts et al. (2009) were the first to describe pollen morphology of extant members of Ebenaceae in detail using SEM and LM. An evolutionary hypothesis was proposed using molecular sequence data of species of Ebenaceae, and synapomorphies were then identified by mapping some pollen morphological characters (pollen size, ornamentation type, equatorial outline and orbicule size) onto the molecular phylogenetic tree. ...
... Family Ebenaceae Gürke Genus Diospyros L. Diospyros sp. Pollen of extant Diospyros species display a wide range in size and outline, but all have a (micro) rugulate sculpture (Geeraerts et al. 2009). The relatively long colpi and perforations observed in interapertural areas (SEM) of the fossil pollen is shared with pollen of many extant Diospyros taxa (Geeraerts et al. 2009, tables I and II). ...
... Pollen of extant Diospyros species display a wide range in size and outline, but all have a (micro) rugulate sculpture (Geeraerts et al. 2009). The relatively long colpi and perforations observed in interapertural areas (SEM) of the fossil pollen is shared with pollen of many extant Diospyros taxa (Geeraerts et al. 2009, tables I and II). ...
... -The pollen morphology (LM, SEM) and ultrastructure (TEM) of Diospyros has been studied thoroughly by, among others, Erdtman (1952), Ng (1971), Cerceau-Larrival et al. (1984), de Franceschi (1993), Morton (1994), Tissot et al. (1994), Kodela (2006), Geeraerts et al. (2009), Sánchez-Dzib et al. (2009),Grygorieva et al. (2010Grygorieva et al. ( , 2013,Li et al. (2011), Miyoshi et al. (2011), and Gosling et al. (2013. The recent most comprehensive SEM work byGeeraerts et al. (2009) defines four main sculpture types within Ebenoideae (including Diospyros, Euclea, Royena). The sculpture of the fossil pollen type from Lavanttal falls within 'Type 1b' ofGeeraerts et al. (2009) that occurs in various extantDiospyros, but also Euclea and Royena species. ...
The continued investigation of the middle Miocene palynoflora from the Lavanttal Basin reveals numerous additional angiosperm taxa. The Myrtales to Ericales pollen record documented here comprises 46 different taxa belonging to Onagraceae (Ludwigia), Ericaceae (Craigia, Reevesia, Tilia), Anacardiaceae (Pistacia), Rutaceae (Zanthoxylum), Sapindaceae (Acer), Santalaceae (Arceuthobium), Amaranthaceae, Caryophyllaceae, Polygonaceae (Persicaria, Rumex), Cornaceae (Alangium, Cornus, Nyssa), Ebenaceae (Diospyros), Ericaceae (Andromeda, Arbutus, Empetrum, Erica), Sapotaceae (Pouteria, Sideroxylon), Styracaceae (Rehderodendron) and Symplocaceae (Symplocos). Köppen signatures of potential modern analogues of the additional fossil woody elements confirm the hypothesis of a subtropical (Cfa, Cwa) climate at lower elevations and subsequent transition into a temperate climate with altitudinal succession (Cfa → Cfb/Dfa → Dfb; Cwa → Cwb → Dwb-climate). The fossil plants represent different vegetation units, from wetland lowlands to well-drained montane forests. Many of the fossil taxa have potential modern analogues that can be classified as nemoral and/or meridio-nemoral and/or semihumid-meridional vegetation elements. New is the recognition of oreotropical elements, which are direct indicators for a substantial altitudinal gradient.
... The asterid ericalean clade (APG IV, 2016) is morphologically diverse with well-defined families (Schönenberger, Anderberg & Sytsma, 2005), of which several have been recently analysed using molecular phylogenetic techniques: Ericaceae by Kron et al. (1999), Kron, Powell & Luteyn (2002), Gillespie & Kron (2010), Kron & Luteyn (2005) and Kron & Powell (2009), Sapotaceae by Anderberg & Swenson (2003), Swenson, Bartish & Munzinger (2007), Swenson, Richardson & Bartish (2008), Bartish, Antonelli & Richardson (2011), Armstrong et al. (2014) and Richardson et al. (2014), Ebenaceae by Wallnhöfer (2001), Duangjai et al. (2006) and Geeraerts et al. (2009), Styracaceae by Fritsch (2001) and Fritsch et al. (2001Fritsch et al. ( , 2011, and Theaceae by Vijayan, Zhang & Tsou (2009). ...
... The routinely applied SEM investigation of fossil pollen is possible because of previous comprehensive LM and SEM studies on modern pollen. Particularly important for this study were the studies of Sarwar, Ito & Tahahashi (2006), Sarwar & Takahashi (2006, 2014 on Ericaceae, Harley (1991) on Sapotaceae, Geeraerts et al. (2009) on Ebenoideae, Morton & Dickison (1992) on Styracaceae and Zavada & Wei (1993) on Camellia L. ...
... Pollen grains of Euclea resemble those of Diospyros, but are generally much smaller than the normal size range for Diospyros (Morton & Kincaid, 1995: fig. 7;Geeraerts et al., 2009: Christophel et al., 1987: fig. 5) and with at least two types on Hainan, and at the end of the Eocene in the USA, but these latter three pollen types still resembled extant African Diospyros taxa. ...
As scanning electron microscopy (SEM) has not been routinely used in palaeo-palynology, affiliations of fossil pollen to botanical equivalents, based only on light microscopy (LM), are always equivocal. Here 29 taxa of Ericales 55.8-41.2 My in age have been identified using LM and SEM, modifying the evolutionary and palaeo-biogeographical framework of some Ericales. Seven Ericaceae types, 17 Sapotaceae types, three Ebenaceae types, one Styracaceae type and one Theaceae type were identified to tribe or genus level. A 56-My-old Erica-type affiliated with Erica arborea might be the ancestor of African Erica taxa, whereas a younger type with affinities to palaeo-Arctic Erica might be the precursor of northern European heathers. The Kalmia-type suggests a European origin and a wider geographical distribution for this type during the mid-Eocene, and a Rhododendron-type of contemporaneous age corroborates the existence of numerous Rhododendron taxa in Europe. Both the 56-My-old Vaccinium-type, affiliated with temperate continental Asian taxa, and the Gaultheria-type, affiliated with several South American taxa, represent the oldest occurrences of these taxa and suggest a Eurasian origin. The Sapotaceae are represented by Sarcospermatoideae, with two Sarcosperma-types, Chrysophylloideae, with three Chrysophylleae-types affiliated with South American Pouteria/Elaeoluma and one affiliated with Asian Planchonella/Sersalisia, and Sapotoideae with three Mimusopeae/Isonandreae-types with mixed affinities, two Mimusopeae-types affiliated with African/Malegasy Mimusops/Vitellariopsis taxa, one Mimusopeae-type affiliated with South American Manilkara taxa, four Isonandreae-types affiliated with SE Asian Palaquium and one South American Sideroxylon-type. All Sapotaceae types are older than postulated in the literature by molecular dating analyses. Three Diospyros-types affiliated with African taxa corroborate a Gondwanan-African origin of the genus. A Styrax-type resembles extant South American and East Asian taxa and is interpreted as being an early European ancestor of Styrax. The contemporaneous Camellia-type represents the oldest record of Camellia.
... D. ebenum belongs to the family Ebenaceae [26], subfamily Ebenoidae [16] (Table 1). This subfamily contains around 600 species distributed amongst three genera (Diospyros, Euclea, Royena) [16], of which Diospyros is the largest with more than 500 species [12]. ...
... D. ebenum belongs to the family Ebenaceae [26], subfamily Ebenoidae [16] (Table 1). This subfamily contains around 600 species distributed amongst three genera (Diospyros, Euclea, Royena) [16], of which Diospyros is the largest with more than 500 species [12]. The family of Ebenaceae has another subfamiliy, Lissocarpoideae, with a single genus: Lissocarpa. ...
... According to Duangjai et al. [12], approximately 300 species occur in Asia and in the Pacific areas, 98 species in Madagascar and the Comoros, 94 species in the mainland of Africa, around 100 species in the Americas, 15 species in Australia and 31 species in New Caledonia. The four genera of the family Ebenaceae are monophylous while the genus Diospyros is closely related to the African genera Euclea and Royena [16]. However, the authors state that the "current distribution patterns in Diospyros can be explained by dispersal only with no vicariance events", which, according to the authors, indicates that the genus has a capacity for long-distance dispersal. ...
High-quality wood has always been in demand in human societies. It is loved and used extensively by those who can afford it. In Egypt, ebony was most likely used as early as 4500 years B.P. [11], and later, the Roman Empire also developed a high demand for ebony wood. Much of this demand was satisfied by the Indian Diospyros species [41]. The two ebony species mainly traded from India were D. melanoxylon and D. ebenum. However, the latter is the “only one giving a black wood without other streaks or markings …” [13]. The wood was first brought to Rome around 2000 years ago [41] and was traded with rulers of the Europe during the Renaissance. Around 400 years ago, the term “ebenist” was used for the finest carpenters of France [9]. For millennia, ebony has been harvested in India so that by the time of the British arrival a large percentage of the ebony was already cut, mostly by using very destructive methods [37]. Because of this, not much of the species was left on the subcontinent [13], and the still plentiful stocks in Sri Lanka (then Ceylon) were the main source for the British of this much-in-demand wood. Some of these stocks remain today, even though they were heavily exploited. Despite the high value of the species and the potential that the species promises with scientific management, it does not receive much attention from the scientific world. The species is classified as endangered in Sri Lanka [21] and its trade is banned in both India and Sri Lanka. However, in the course of the forest ecosystem restoration work in South India, the species is used and distributed.
... Many individuals have only female flowers and the fruits can arise without fertilization -by parthenocarpy and they are seedless. Information on Diospyros virginiana pollen grain traits occurs scarcely in a few papers (Erdtman 1966;Geeraerts et al., 2009;Grygorieva et al., 2010). Grygorieva, O., Motuleva, S., Nikolaieva, N., Klymenko, S., Schubertová, Z., Brindza, J. Agrobiodiversity, 2017, 151-158 Pollen grains of the Diospyros virginiana occur individually -solitary, defined as monads. ...
... An additional pole is in the colpus middle at the equatorial centre (circular equatorial aperture). This classification correlates with literature data (Sowunmi 1995;Kodela 2006;Geeraerts et al., 2009). ...
... μm. For example Erdtman (1952) in Diospyros virginiana reported the mean value for polar axis length (P) 59.0 μm and the equatorial axis (E) 46.0 μm whereas Geeraerts et al. (2009) for the same species found the mean values for the length P -41.4 μm and E -31.2 μm (Table 2). ...
Morphology of pollen grain was described for nine American persimmons (Diospyros virginiana L.) genotypes (DV-01-DV-09) at the Institute of Biodiversity and Biological Safety of Slovak Agricultural University in Nitra using an electron microscope Carl Zeiss LS 15, and the microphotographs were taken. The measurement of morphometric parameters was carried out on 50 pollen grains from each genotype using the AxioVision Rel. 4.8.2.0 program. The measurements were made in micrometer (μm). The length of polar axis (P) and the equatorial diameter (E) of grain, P/E ratio were measured and their variation was compared among studied genotypes. The polar axis and equatorial diameter of pollen grains values were varied from 51.94 to 78.60 μm and from 23.00 μm to 42.71 μm, respectively. This study showed that there were significant differences among the genotypes in all measured factors. Pollen grains are radial-symmetrical, isopolar, tricolpate (with three sulcus). Exine surface is a bicomponent sculptural type with small perforate-undulate or wrinkled weakly perforate. Percentages of abnormal pollen grains ranged from 2.0 to 19.3% among the genotypes. It was noted that diversity of surface sculpturing of pollen grains in combination with shape and sizes of them enables to use complex of thin morphologic signs for Diospyros virginiana pollen identification.
... Pollen grains 3-zonocolpate-orate, wide or narrowly ellipsoidal, rarely almost spheroidal, in outline from polar view rounded or angular-rounded; polar axis 22,0-35,4 μm, equatorial diameter 18,9-28,5 μm. Colpi long, ends rounded or slightly pointed, narrow (in A. viridis is slit-like); sculpture of colpus membrane of A. aureus granular, granules located mostly along the colpus edges, in other species colpus membrane without ornamentation; apocolpium diameter 6,[5][6][7][8][9][10][11]5 μm,mesocolpium width 11,[5][6][7][8][9][10][11][12][13][14][15][16][17]3,4 х 6,[9][10]5 μm at length. Exine thin (0,7-0,8 μm), columellae layer weakly expressed, almost invisible. ...
... Pollen grains 3-zonocolpate-orate, wide or narrowly ellipsoidal, rarely almost spheroidal, in outline from polar view rounded or angular-rounded; polar axis 22,0-35,4 μm, equatorial diameter 18,9-28,5 μm. Colpi long, ends rounded or slightly pointed, narrow (in A. viridis is slit-like); sculpture of colpus membrane of A. aureus granular, granules located mostly along the colpus edges, in other species colpus membrane without ornamentation; apocolpium diameter 6,[5][6][7][8][9][10][11]5 μm,mesocolpium width 11,[5][6][7][8][9][10][11][12][13][14][15][16][17]3,4 х 6,[9][10]5 μm at length. Exine thin (0,7-0,8 μm), columellae layer weakly expressed, almost invisible. ...
... A. karjaginii Boriss. 30,7 х 27,5 10,[2][3][4][5][6][7][8][9][10][11]5 16,[8][9][10][11][12][13][14][15][16][17] Willd.) 25,6-26,0 х 22,2-23,5 6,5-7,7 16,2-17,0 -/ -reticulate (lumina diameter up to 1,5 μm) A. oleifolius DC. 27,8 х 22,[9][10][11][12][13][14][15][16][17][18][19][20][21][22][23][24]5 3,7 11,[5][6][7][8][9][10][11][12][13] 23,6-24,5 х 18,9-20,5 5,7-6,5 11,5-11,0 -/ -reticulate (lumina diameter up to 1,0 μm) A. uraniolimneus Boiss. ...
This paper is a third part of a series was devoted to the palynological study of woody plants of Armenia. With the help of light (LМ) and scanning electron (SEM) microscopes investigation of pollen morphology of 26 species of Armenian trees and shrubs from the families Ebenaceae Gürke, Elaeagnaceae Juss., Ericaceae Juss. and Fabaceae Lindl. has been carried out.
... Flowering biology complexity is for these species symptomatic, and it brings questions concerning several topics like the pollen grains characteristics, designated in palinology as sporoforms. Information on Diospyros genus pollen grain traits occurs scarcely in a few papers (Erdtman 1966;Kodela 2006;Geeraerts et al. 2009). ...
... Additional pole is in the colpus middle at the equatorial centre (circular equatorial aperture). This classification correlates with literature data (Sowunmi 1995;Kodela 2006;Geeraerts et al. 2009). In some pollen grains were observed annuluses -an area of the exine surrounding a pore with thickened periphery or operculum -part of exine over the pore (Fig. 1-4). ...
... The shape indexes ranging from 1.77 to 1.83 in the genotypes of Diospyros lotus L. species confirmed a certain degree of identity substantiating thus the classification to shape class of prolonged pollen grains, i. e. with the vertical axis longer than equatorial one. In comparison with the literature data by Geeraerts et al. (2009) our results concerning the shape indexes of tested species are generally higher. On the other hand, the data shown in Table 1 and the values of shape indexes (1.28-1.33) ...
Aim of the work was to study the general characteristics and essential morphological traits of pollen grains, as the size, shape of pollen grains and number, form and position of apertures in Diospyros kaki L. f., Diospyros virginiana L., Diospyros lotus L. species and interspecies hybrid of Diospyros virginiana L. × Diospyros kaki L. f. The studies were performed on pollen obtained from genotypes cultivated in Slovakia (D.
kaki L. f. a D. lotus L.) and Ukraine (D. virginiana L., D. virginiana L. × D. kaki L. f.). Pollen grains morphological traits were evaluated using the scanning and transmission electron microscopy. Mean length of polar axis varied between 48.0 μm (D. lotus L.) and 71.19 μm (D. virginiana L. × D. kaki L. f.) and equatorial axis (pollen width) from 23.56 μm (D. virginiana L.) to 33.14 μm (D. virginiana L. × D. kaki L. f.). The highest values of both axes were observed in the pollen grains of interspecies hybrid Diospyros virginiana L. × Diospyros kaki L. Diospyros spp. genotypes tested pollen grains polar axis variation range were classified as medium-sized to large. Regarding the shape index values, which ranged from 1.75 (D. kaki L. f.) to 2.18 (D. virginiana L.), are
the pollen grains predominantly prolate or prolate spheroidal. Significant differences were detected among the tested species and the interspecific hybrid as well as between individual genotypes of D. lotus L., especially in the equatorial axis. The significant differences in the length of polar axis and equatorial diameter axis indicate the interspecific and intraspecific pollen variability of Diospyros species. Generally, the tested
species pollen grains are solitary, isopolar, radially symmetric. According to shape and number of apertures the pollen grains are tricolporate. This study of pollen grains in species of genus Diospyros enlarged the currently sparse knowledge on taxonomical, palinological and practical use in bee-keeping oriented on identification of pollen grains occurring in the corbicular bee-pollen.
... [7] In terms of vascular plant diversity, Mexico ranks fourth in the world, with approximately 23,314 species that are distributed in 2,854 genera, [8] including the genus Diospyros (Ebenaceae). This group of plants comprises 500 species of trees distributed in tropical latitudes, [9] mainly in Asia (300 species) and America (100 species), and 20 species are located in Mexico. [10] The wood of species such as Diospyros ebenum is valued for its hardness and durability, while others with edible fruits such as Diospyros kaki, have commercial and nutritional value. ...
... The PASS online prediction was determined for compounds 1-21 (Table 1). This chemoinformatic approach predicts the (1)(2)(3)(4)(5)(6)(7)(8)(9)(10)(11)(12)(13)(14)(15)(16)(17)(18)(19)(20). Origanal (21), limonin (22), phorate sulfoxide (23) and galantamine (24). ...
This report informs for the first time the chemical constituents of Diospyros xolocotzii and Diospyros digyna , the pesticidal and the acetylcholinesterase (AChE) inhibition potential of some compounds calculated by in silico approaches, the larvicidal activity against Spodoptera frugiperda of available compounds, the AChE inhibition of selected compounds, and the results of the molecular docking of the most active ones with this receptor. From the aerial parts of D. xolocotzii were isolated pentacyclic triterpenes ( 1–4 , 6 , 10 , 11–13 ), phytosterols ( 15–17 ), and isodiospyrin ( 18 ), whereas the analysis of aerial parts of D. digyna conducted to the isolation of pentacyclic triterpenes ( 4 , 5 , 7–9 , 11–14 ), (4 S )‐shinanolone ( 19 ), and scopoletin ( 20 ). For comparison purposes, origanal ( 21 ) was chemically prepared from 11 . The in silico analysis showed that the tested compounds have pesticide potential. The larvicidal activities of 11 > 13 > 12 indicated that the increase of the oxidation degree at C‐28 increases their bioactivity. Compounds 11 and 21 presented the higher inhibition in the acetylcholinesterase assay, and the higher binding energies, and for the interactionswith AChE by molecular docking. Both Diospyros species are sources of triterpenes with pesticidal potential and the molecular changes in lupane triterpenes correlate with the observed bioactivity and molecular docking.
... In Ebenoideae, pollen is generally shed as monads (permanent tetrads in two species) and is medium sized, prolate-spheroidal to subprolate and tricolporate. Palynological synapomorphies at different taxonomic levels such as subfamily, generic and subgeneric level are reported for Ebenoideae (Geeraerts et al., 2009). The granular infratectum and the unique sculpturing pattern on the orbicule walls are the most discriminating pollen features for Ebenoideae (Geeraerts et al., 2009). ...
... Palynological synapomorphies at different taxonomic levels such as subfamily, generic and subgeneric level are reported for Ebenoideae (Geeraerts et al., 2009). The granular infratectum and the unique sculpturing pattern on the orbicule walls are the most discriminating pollen features for Ebenoideae (Geeraerts et al., 2009). ...
In this review article, we provide an overview of the status of research on Old World Ebenaceae with an emphasis on the large genus Diospyros. The well-supported phylogenetic tree obtained from nucleotide sequences of multiple regions of plastid genome gave clear insights into the subfamilial classification of Ebenaceae. It supported inclusion of previously recognized genera such as Cargillia, Gunisanthus, Maba, Macreightia and Tetraclis in Diospyros. Endemic Diospyros spp. of New Caledonia have multiple origins. One of these clades has c. 21 species that are morphologically distinct and occupy different ecological niches, but they exhibit low genetic variation, leading to a lack of phylogenetic resolution. Analyses of whole plastid genome sequences did not greatly increase resolution or support for results of our previous plastid analyses. Geographical clustering of the individuals against a background of lower sequence divergence of the whole plastid genome could be due to transfer of plastid genomes during hybridization and introgression following secondary contact. However, > 8400 filtered single nucleotide polymorphisms (SNPs) from restriction site associated DNA sequencing (RADseq) confirmed species circumscriptions for this clade and produced well-supported phylogenetic relationships, pointing to an early regional clustering among populations and species. This supported allopatric speciation with respect to macrohabitat (i.e. climatic conditions) having had a role in the initial differentiation in the group. A later, more rapid radiation involved divergence with respect to microhabitat (e.g. soil preference). Although chromosome counts indicate that Diospyros spp. are consistently diploids with 2n = 30, extensive variation in genome size has been observed, which is due to an increase of repeat elements, including LTR/gypsy. In Ebenaceae, pollen is heterogeneous, and palynological synapomorphies are traced at different taxonomic levels. Several new Diospyros spp. have recently been identified and documented from India, Thailand, China, Africa and New Caledonia. Taxonomic revisions have been completed for the Australian species, and synonyms are reported for some New Caledonian Diospyros spp.
... "Remarks:" Both the Diospyros sp. 1 and sp. 2 pollen types correspond in aperture configuration and exine sculpture to extant Diospyros pollen (Geeraerts et al., 2009). The form genus Diospyrospollenites Skavińska Ziembińska-Tworzydło is commonly used for fossil dispersed pollen with affinities to extant Diospyros (e.g. ...
The Gleisdorf Formation (Fm.) deposits in the clay pit at Gratkorn, Styria, Austria, are dated to 12.2–12 Ma, and are of late middle Miocene age (late Serravallian or Sarmatian). To reconstruct the paleovegetation and estimate the paleoclimate at this important vertebrate site, the palynoflora close to the boundary between the vertebrate-bearing layers of the Gratkorn Fm. and the overlying limnic clay deposits of the Gleisdorf Fm. was investigated. Using the single-grain method, 140 palynomorphs were identified. The palynoflora suggests that the paleovegetation was characterised by well-drained lowland and upland forests, riparian forest, and swamp forests. Depending on the dominating tree species, lowland and upland forests might have had closed or more open canopies. Open habitats included wet meadows and shrublands. In addition, conifers were present in the swampy lowlands and the forested uplands. The most prominent paleoclimatic signatures of the palynoflora indicate a fully humid warm temperate climate, with hot to warm summers and cool winters (Cfa-, Cfb-climate), and a seasonal climate with cool and drier winters and hot to warm and wetter summers (Cwa-, Cwb-climate). Our results align with existing studies bordering the Styrian Basin and support the presence of subtropical to warm-temperate vegetation around Gratkorn during the Sarmatian.
... Mapping characters on phylogenetic trees is a common approach to understand the evolution of traits Bollback, 2006). However, studies have most commonly focused on morphological characters (e.g., Perret et al., 2003;Geeraerts et al., 2009;Ilves and Taylor, 2009), leaving aside other important traits such as ecology and behavior. In this study, we mapped the evolution of habitat use for a widespread butterfly group and related this to the biogeographic and diversification history of the group. ...
Species losses are increasing and may have an impact on our understanding of patterns of evolutionary pathways and phylogenetic relationships among the groups being lost. The knowledge of such patterns can contribute to preventing future losses by identifying which lineages have higher or lower diversification rates, thus informing conservation strategies. Recent years have seen a significant growth in studies of butterfly systematics, allowing a better understanding of evolutionary relationships among most groups and revealing significant taxonomic chaos in several groups. One of the latter groups is the nymphalid subtribe Euptychiina (Satyrinae), which has been shown to include a number of non-monophyletic genera based on recent molecular phylogenetic analyses. Among others, these genera include Yphthimoides, which is widespread throughout the Neotropical region but particularly diverse in the southeastern Neotropics, and a pair of related genera, Pharneuptychia Forster, 1964 and Moneuptychia Forster, 1964. Using molecular data, this study scope and aims was to provide a phylogenetic hypothesis that corroborates Yphthimoides as presently conceived being non-monophyletic, a result reinforced by a comparative study of the male genitalic morphology. Our results also show that Pharneuptychia and Moneuptychia, plus a species misplaced elsewhere in the Euptychiina, Euptychoides castrensis (Schaus, 1902), form a well supported clade, and that the latter 'species' is a complex of cryptic species. We therefore propose a number of taxonomic rearrangements in the present work to resolve these issues: Yphthimoides eriphule (A. Butler, 1867) will be moved to a new genus; Y. affinis (A. Butler, 1867), Y. maepius (Godart, [1824]), Y. mimula (Hayward, 1954), Y. neomaenas (Hayward, 1967) and Y. mythra (Weymer, 1911) are being transferred to Malaveria Viloria & Benmesbah, 2021; Pharneuptychia innocentia (Godart, [1824]) will be moved to another genus to be described; and Euptychoides castrensis, Pharneuptychia romanina (Bryk, 1953) and Yphthimoides viviana (Romieux, 1927) are being moved to Moneuptychia. The dating of divergences points to a split between the ancestral lineage of Yphthimoides and its sister group, Carminda Ebert and Dias, in Dias 1998, during the last half of the Miocene, around 11.86 Mya, and to the diversification of the Pharneuptychia during the same time 11.35 (± 3.52) Mya. Biogeographic analysis showed that the most recent common ancestor of Yphthimoides started to diversify either in the the Brazilian Cerrado savannas or in a combined area of Cerrado and South Atlantic Forest, with a possible change in the ancestral habitat of Carminda. Furthermore, ancestral character mapping favors a savanna origin hypothesis over a forest origin hypothesis.
... However, according to Erdtman (1971), morphologically comparable tetrads occur also within different taxa included in Empetraceae, Epacridaceae, and Pyrolaceae. Additionally, Geeraerts et al. (2009) reported the occurrence of permanent tetrads in Ebenaceae (i.e. Diospyros mannii and D. longifolia). ...
The Paleocene Fossil-Lagerstätte Menat in France is well known for its wealth of outstandingly well preserved fossil insects and plants. Despite being known for more than a century, the palaeoflora, which is regarded as typical for the late Thanetian by some authors, has largely been neglected since the 1940s. New excavations and surveys yielded exceptionally well-preserved plant material, including a minute, heptamerous flower bearing in situ pollen tetrads, comparable to tetrads of the modern ericacean genus Kalmia L, in its anthers. The only known modern ericacean genus which is characterised by heptamerous flowers is Bejaria Mutis ex L., a basal relative of the tribe Phyllodoceae within Ericaceae, which also includes the genus Kalmia in a relatively basal position. However, heptamerous flowers also occur very rarely (mostly interpreted as teratologies) in a number of other modern Ericaceae, but also in various other modern angiosperm families. Due to the unique combination of a heptamerous flower with Kalmia -type pollen tetrads within the anthers, the new taxon Menatanthus mosbruggeri gen. nov. et sp. nov. is erected. The lack of morphological data from the flower itself and the fact that comparable pollen tetrads can be produced by a number of modern families, however, prevent an assignment of the new taxon to any known angiosperm family.
... Duangjai et al. (2006) dividiu Ebenaceae em duas subfamílias, Lissocarpoideae que abrange as espécies com ovário ínfero, e Ebenoideae que compreende as espécies com ovário súpero. Essa divisão posteriormente foi confirmada por Geeraerts et al. (2009) através de estudos palinológicos: Lissocarpoideae (triporado) e Ebenoideae (tricolporado). ...
Resumo Apresentamos o levantamento florístico das espécies de Ebenaceae para o estado do Ceará, como parte do projeto “Projeto Flora do Ceará: conhecer para conservar”. O estudo foi baseado na análise comparativa dos caracteres morfológicos de espécimes depositados nos Herbários ASE, EAC, HCDAL, HUEFS, HUVA, INPA, MAC e UB, de amostras obtidas em campo, bibliografias especializadas e imagens de coleções-tipo. A família é representada no estado pelo gênero Diospyros, com três espécies: D. coccolobifolia, D. inconstans subsp. obovata e D. sericea. As espécies têm distribuição disjunta, restrita ou amplamente distribuídas em território cearense, ocorrendo em vegetação de Savana (Cerrado), Savana Estépica (Caatinga), Floresta Estacional Decidual (Mata Seca), Floresta Estacional Semidecidual das Terras Baixas (Mata de Tabuleiro), Vegetação com Influência Marinha (Restinga) e Floresta Ombrófila Densa (Mata Úmida). As três espécies foram registradas em Unidades de Conservação no estado.
... Principal component analysis (PCA) as a classical analytical method is widely used for characterization of single-molecule SERS. [56][57][58] For example, the PCA is powerful for decomposition of SERS spectra of a mixture in the bianalyte approach. The decomposition with statistical soundness is usable for characterization of SERS substrates, such as precise estimation of resonance wavelengths of hotspots and evaluation of enhancement efficiencies of SERS substrates. ...
... Pori are mostly well defined, usually lalongate, sometimes with indistinct lateral edges or rarely indistinguishable. The pollen is remarkably constant throughout the family, and the main variations are in size and shape of the grains and the apertures (Erdtman, 1952;Walln€ ofer, 2004;Geeraerts et al., 2009). ...
... Diospyros L. Sharma & Gupta, 1979;Wallnöfer, 2001;Kodela, 2006;Geeraerts et al., 2009;Li et al., 2010 *Diospyros kaki Thunb. (Fig. 4K Muller, 1972Muller, , 1973Tsou, 1994 Couroupita Aubl. ...
This study, the fifth in a series investigating palynological characters in angiosperms, aims to explore the distribution of states for 19 pollen characters on five early diverging orders of Superasteridae (Berberidopsidales, Caryophyllales, Cornales, Ericales, and Santalales) plus Dilleniales. To illustrate the character states found in the pollen of this group, we examined pollen grains of 15 species exemplifying 15 families across all studied orders using light, scanning, and transmission electron microscopy. We reconstructed the phylogeny of the early diverging Superasteridae and related taxa with eight genetic markers for 172 genera, using maximum likelihood (ML) analysis. Nineteen pollen characters were coded for the genera used in this phylogeny and compiled into two morphological matrices using two coding strategies. The characters were then optimized on the newly generated ML tree plus two constrained trees differing in the position of Dilleniales, using three methods of inference. Taxa in this grade show a striking diversity of pollen morphologies, particularly in certain characters such as size, tectum sculpture, and aperture number. The plesiomorphic condition for the early diverging Superasteridae is unambiguously and consistently inferred to comprise monad-dispersed, isopolar, spheroidal, circular in outline, equatorially arranged, tricolpate pollen grains with granular aperture membranes, a smooth tectum, and endexine present. We identify diagnostic character states and synapomorphies for several monophyletic groups, and explore the palynological evidence that may shed light on some unresolved relationships. For example, the hypothesis that Dilleniales is sister to Superrosidae is better supported than alternative hypotheses, being consistent with a number of shared palynological state changes including transitions to presence of costae, reticulate tectum, and columellar infratectum structure. Across this part of the angiosperm phylogeny, most state transitions occur repeatedly, and their frequency varies among both clades and characters. We discuss the impact of optimization method, tree topology, and coding strategy upon ancestral state reconstruction.
... (9.34) T3 -Cross with D. virginiana 50 97.12 (9.85) T4 -Cross with Kandaghat Pink 50 94.68 (9.73) C.D. (0.05) 4.74 Figures in parenthesis are square root transformed valueIn Diospyros mespiliformis the mean values for the length, 32.30 µm and width, 24.80 µm (Geeraerts et al. 2009).The ratio of length and width was 1.36 in case of Kandaghat Pink and 1.34 in Diospyros virginiana. Shape of pollen was prolate in both the genotypes. ...
... Taxonomic inconsistencies have also been found in P. insularis (Nakai) Nakai, an endemic Korean species restricted to Ulleung Island (Nakai, 1918;Oh, 1978; cal studies might help to clarify the generic delimitation in Neillieae. In addition, the morphological details of orbicules, small sporopollenin particles produced by the secretory tapetum, may also provide valuable phylogenetic information (Huysmans et al., 1997(Huysmans et al., , 2010Huysmans, El-Ghazaly & Smets, 2000;Huysmans & Smets, 1998;Geeraerts et al., 2009;Verstraete et al., 2011Verstraete et al., , 2014. Although studies on orbicules are scarce in Rosaceae, recent descriptions of the orbicules in tribe Sorbarieae (subfamily Amygdaloideae) were of systematic importance (Song, Moon & Hong, 2016). ...
Palynological characteristics were examined in Rosaceae tribe Neillieae, comprising the genera Neillia, Physocarpus and Stephanandra. Physocarpus insularis and some taxa of Spiraea were also examined to evaluate the potential usefulness of pollen traits in taxonomic and phylogenetic studies of Rosaceae. Pollen grains of Neillieae are monads, tricolporate, small to medium in size [polar axis (P) = 13.4–45.8 μm, equatorial diameter (E) = 14.3–39.9 μm] and oblate to prolate in shape (P/E = 0.68–1.70). Sexine ornamentation in Neillia and Stephanandra is perforate, whereas Physocarpus and Spiraea have striate pollen. Spherical orbicules with a central perforation (donut-shaped) were consistently found in Neillieae, but these were absent in P. insularis and all taxa of Spiraea. Our palynological results provide strong evidence for the merging of Neillia and Stephanandra, and pollen morphological data supported the recent phylogenetic hypothesis that P. insularis should be a member of Spiraea.
... An understanding of the distribution and morphology of orbicules in this tribe is needed to better assess phylogenetic patterns within the Spiraeeae, since the orbicules may have systematic importance as additional palynological characteristics (Huysmans et al. 1997(Huysmans et al. , 1998(Huysmans et al. , 2000(Huysmans et al. , 2010Geeraerts et al. 2009;Verstraete et al. 2011Verstraete et al. , 2014. Orbicules are small sporopollenin particles that can be produced by species with a secretory tapetum (Verstraete et al. 2014). ...
This study provides pollen data for 38 representative taxa belonging to all nine genera in the current classification of the tribe Spiraeeae (Rosaceae) including the monotypic Korean endemic genus Pentactina, and considers the distribution of orbicules for the first time. Pollen morphology and wall stratification were investigated using light, scanning electron and transmission electron microscopy. Spiraeeae pollen grains are small to medium in size (P = 6.9–34.0 μm, E = 7.1–28.0 μm), oblate to prolate in shape (P/E = 0.66–1.48) and tri-colporate. Spiraeeae pollen is generally characterised by striate sexine ornamentation, but four ornamentation types are recognised based on the length and direction of the ridge patterns. The observed variation in sexine ornamentation is particularly valuable at the generic level. The exine stratification of the representative Spiraeeae studied is similar and characterised by unbranched columellae and a continuous endexine. Orbicules are present in three genera of the tribe (Luetkea, Sibiraea and Xerospiraea). Orbicule distribution patterns indicate that the absence of orbicules is a synapomorphic condition of the more derived clade, comprising Pentactina + Petrophytum + Kelseya + Spiraea.
... Pori are mostly well defined, usually lalongate, sometimes with indistinct lateral edges or rarely indistinguishable. The pollen is remarkably constant throughout the family, and the main variations are in size and shape of the grains and the apertures (Erdtman, 1952;Walln€ ofer, 2004;Geeraerts et al., 2009). ...
The accurate and consistent identification of fossil pollen is essential to allow robust inferences to be drawn with regard to past vegetation and climate change. Identifications are best achieved through the direct inspection of reference material. Most substantial reference collections are located at research institutions in Europe or North America, which restricts access for researchers trying to advance palynology in less developed countries. Due to the development of digital imaging and fast internet access it is now possible to produce high quality images from pollen and spore reference collections and make them globally available. In this pollen and spore atlas we contribute to this growing body of work by presenting images of 240 pollen types (in 202 genera and ∼79 families) and 30 spore types (in 25 genera and ∼17 families) from a wide range of different vegetation types originating from the Kilimanjaro area and tropical East Africa. We provide an overview on the range of pollen and spore types commonly found in Last Glacial and Holocene sedimentary archives in studies from the Kilimanjaro area and tropical East Africa. Besides a pollen key, detailed information is given on pollination, habitus and habitat which all support the interpretation and environmental reconstructions from pollen records. The atlas will serve as useful guide for palynological investigations, aiming at detailed and comprehensive reconstruction of past vegetation, environmental and climate change in tropical East Africa.
... In addition, orbicules, sporopollenin structures like pollen wall, have never been investigated in Sorbarieae. Although the clear function of orbicules is still enigmatic, the orbicules might have phylogenetic signal at the various taxonomic groups (Huysmans et al. 1997(Huysmans et al. , 1998(Huysmans et al. , 2000Vinckier and Smets 2002;Moon et al. 2008a, b, c;Geeraerts et al. 2009;Huysmans et al. 2010;Verstraete et al. 2011Verstraete et al. , 2014. ...
The pollen morphology of the tribe Sorbarieae (Adenostoma, Chamaebatiaria, Sorbaria, and Spiraeanthus) and two related genera Gillenia and Lyonothamnus was investigated by light, scanning electron, and transmission electron microscopy. Sorbarieae pollen was monad, tri-colporate, small to medium in size (P = 10.2–40.0 μm, E = 10.8–32.4 μm), and oblate to prolate in shape (P/E = 0.74–1.88). The sexine ornamentation was striate, but four different types could be divided: striate-psilate, striate-plicate, striate-microechinate, and striate-perforate. The pollen wall stratification typically consisted of unbranched columellae and a continuous endexine. As an additional palynological characteristic, orbicules (small sporopollenin granules) were observed in all taxa and thus a possible synapomorphic character of Sorbarieae as a whole. The pollen characteristics are useful to recognize certain taxa. Gillenia is the only genus with both a pore flap on the aperture and the perforated sexine. Lyonothamnus has a significantly thick exine and relatively larger pollen grains, which is compared to that of Sorbarieae. Some quantitative characteristics (e.g., pollen and colpus size, exine thickness, and ridge width of striae) identified using principal components analysis (PCA) may have diagnostic importance among the taxa in the tribe.
... The results from our previous molecular phylogenetic study of Ebenaceae sensu lato strongly supported monophyly of all four genera, Diospyros, Euclea, Lissocarpa, and Royena (Duangjai et al., 2006), and indicated that the pantropical genus Diospyros is closely related to the smaller African genera Euclea and Royena (Morton et al., 1996;Berry et al., 2001;Duangjai et al., 2006). Diospyros can be distinguished from the closely related genera by seed anatomical and palynological characters (Duangjai et al., 2006;Geeraerts et al., 2009). Royena and Euclea have an ingrowth of the testa surrounding the radicle of the embryo, but members of Diospyros lack of an invagination surrounding the radicle. ...
We aimed to clarify phylogenetic relationships within the pantropical genus Diospyros (Ebenaceae sensulato), and ascertain biogeographical patterns in the New Caledonian endemic species. We used DNA sequences from eight plastid regions (rbcL, atpB, matK, ndhF, trnK intron, trnL intron, trnL-trnF spacer, and trnS-trnG spacer) and included 149 accessions representing 119 Diospyros species in our analysis. Results from this study confirmed the monophyly of Diospyros with good support and provided a clearer picture of the relationships within the genus than in previous studies. Evidence from phylogenetic analyses suggests that Diospyros colonized New Caledonia multiple times. The four lineages of Diospyros in New Caledonia also differ in their degree of diversification. The molecular data indicate that one lineage is paleoendemic and derived from an ancient Australian species. The other three lineages are more closely related to several Southeast Asian species; two of them are neoendemics, and one has radiated rapidly and recently.
Accurate identification is crucial for conserving species, especially in regions such as the Western Ghats, where trade poses a significant threat to endangered and threatened forest species. Traditional morphology-based identification can be challenging and time-consuming, leading to inaccuracies, especially with similar-looking species or dried specimens. Therefore, DNA barcoding offers a potent solution for precise species identification to address illicit trade and address impactful conservation measures. DNA barcoding is a taxonomic technique that uses standardized short DNA sequences to differentiate and classify species. This approach is especially valuable when morphological characteristics alone are insufficient for accurate species identification. In this study, we focused on the development of a DNA barcoding system for the efficient and accurate identification of threatened and endangered important forest species of Western Ghats Karnataka. To develop the DNA barcoding system, a multilocus approach utilizing sixteen standard DNA barcoding markers was used. A total of 47 threatened and endangered forest species from the Western Ghats were selected for this study. Using a larger number of markers to develop DNA barcodes led to the most precise species identification rates. Moreover, the wide availability of DNA barcode databases allows for quick and accurate species identification. In our study, we observed the highest amplification rates for rbcL1 (40 species), psbtrnH2 (36 species), and PsbA-trnH1 (33 species). DNA amplification varied from 11.76–94.11%. Notably, the highest DNA amplification rates were detected for A. wightii (94.11%) and A. hondala ( 92.34%), both of which belong to the Arecaceae and Passifloraceae families, respectively. Sequencing success rates ranged from 37.5–100%. This study will aid in the development of a database of available threatened forest species in western Ghats Karnataka and other regions.
we conducted anatomical analysis of anthers with the aim to establish the differences in the development pattern of microsporophytes and microgametophytes between perfect and imperfect flowers in the tribe Gardenieae (Rubiaceae). The species studied were: Tocoyena formosa (monoecious with perfect flowers), Cordiera concolor, Genipa americana, Randia calycina, and Randia heteromera (dioecious with imperfect flowers). Flowers in successive stages of development were collected and fixed. The material was processed and examined using light microscopy and scanning electron microscopy. The present study revealed the stage when pollen is arrested in the functionally pistillate flowers of the dioecious taxa. Based on these observations an evolutionary sequence of changes towards the reduction of non-functional anthers in Rubiaceae is proposed. In addition, we describe and discuss characters that might be of importance in future phylogenetic studies in Rubiaceae (e.g., pollen morphology and its dispersal unit, the presence of orbicules, and a new type of placentoid).
Persimmon (Diospyros kaki Thunb.) is an ancient fruit tree that originate in East Asia, especially in Southern China. Persimmon is grown in China, Japan and Korea for a long history. Persimmon can be utilized for various purposes with fruits, leaves and its derivates. The genus Diospyros with about 500 species accounts for the largest genus of Ebenaceae. There is a close relationship between D. kaki, D. lotus, D. glandulosa, D. oleifera and ‘Yemaoshi’ based on morphological and molecular evidence. The ancestor of persimmon is not clearly elucidated. Modern hexaploidy persimmon should be evolved from diploid through genome duplication, in which 2n gametes might play an important role. Persimmon cultivars can be horticulturally classified into four types including PCNA (pollination constant and non-astringent), PVNA (pollination variant and non-astringent), PVA (pollination variant and astringent) and PCA (pollination constant and astringent). PCNA is further classified into Chinese PCNA and Japanese PCNA based on their different genetic control of astringency loss traits. Persimmon genotypes with different geographic origins can be distinguished using various molecular markers, while four persimmon cultivar types could not be well-separated by molecular detection. PCNA cultivars from China and Japan exhibit a divergent origin, interestingly, both Chinese PCNA and Japanese PCNA originate in central mountain areas of the two countries, respectively. Most of the genetic resources of Ebenaceae were distributed in tropical and subtropical regions of the world. More than 2000 accessions of persimmon were preserved in China, Japan, Korea, Italy, Spain, etc. Most of the persimmon production was derived from PCA cultivars in China, which is the largest persimmon producer globally. PCNA fruit is more attractive and new orchards prefer to plant PCNA types in Asia, Mediterranean area, Oceania and South America.
A total of 115 palynomorphs recovered from samples collected in the Colônia basin were described. The morphological descriptions were accompanied by illustrations and ecological information. The palynomorphs date back to the penultimate glacial with ages ranging from 135 to 180 ka. The palynomorphs were grouped according to their presence in three forest types represented in the Colônia basin, namely hillside forest, peat forest and swamp forest. The 115 identified taxa represent a total 58 families; one represents the algae, and 57 Embryophyta (ten pteridophytes, two gymnosperms and 45 angiosperms); 52% of the palynomorphs were observed in the three local forest types. 32.4% were observed in other types of vegetation distributed within a radius of 5 to 15 km from the basin. These palynomorphs represent mature forest, secondary forest, cloud forest, grassland, and disturbed environment whereas 15.6% of the palynomorphs were no longer observed in the region of Colônia. The species represented by these palynomorphs most likely disappeared from the region of Colônia during interglacial climate warming. In addition, our study provides reference material for future pollen studies in Colônia since some vulnerable species were identified.
Burada-e-Aabnoos (Diospyros ebenum Koeing) is an important Unani drug for eye diseases belong to the family Ebenaceae. Great Unani scholar Isa bin Kuhal advocated it especially for ophthalmic use in his treatise Tazkira-tul-Kuhalein. The aim of the present review is to collect all the information of abnoos plant available in Unani classical literature, phytochemical and pharmacological studies, and Unani formulations in which it is used as an ingredient also make an effort to prove the strength of action of the drug mentioned in Unani classical literature. In this review, ten Unani books were referred and pharmacological studies were recognized. The drug is having good medicinal properties but is not used for its action mentioned in Unani literature. It may be recommended that this should be taken for further scientific evaluation.
Many plant families lack substantive fossil records, limiting our understanding of their origin and evolution. The abundance and preservation potential of pollen through geological time have helped to overcome such limitations and have provided reliable fossils for reconstructing biogeographical history and character evolution in many angiosperm families. Here, using scanning electron microscopy, we identified six Ebenaceae-type fossil pollen grains from early Palaeogene sediments of western India. Phenetic and phylogenetic analyses using pollen characters of fossil and extant taxa reavealed affinities of these fossils to three genera of Ebenaceae (Euclea, Royena and Diospyros). Furthermore, our divergence dating analysis using these fossils as priors suggested a Gondwanan origin for the family during the mid-Cretaceous [c. 107 Mya, 95% highest posterior density (HPD): 100–112 Mya] and supports the boreotropical and ‘out of India’ dispersal hypotheses as the most probable explanations for the present global distribution of the family. The study also supports the dispersal of the family into India, from Africa, through the Kohistan–Ladakh Arc during the Palaeocene. Finally, comparative phylogenetic analyses suggest significant synapomorphic and phylogenetic signals for a few selected pollen characters in Ebenaceae. Our findings have important implications for understanding the biogeography and evolution of the highly diverse and ecologically and economically important family Ebenaceae.
Two new norbergenin derivatives, 4-O-p-hydroxybenzoylnorbergenin (1) and 11-O-(E)-cinnamoylnorbergenin
(2), were isolated from the leaves of Diospyros gilletii De Wild along with nine known compounds, including
norbergenin (3), 4-O-galloylnorbergenin (4), betulin (5), betulinic acid (6), lupeol (7), ursolic acid (8), corosolic acid (9), β-sitosterol (10) and quercitol (11). Their structures were elucidated from the analysis of their 1D and
2D 1H and 13C NMR spectral data in conjunction with mass spectrometry. Single-crystal X-ray diffraction technique unambiguously established the structure of compound (1). The antimicrobial properties of the eleven compounds isolated were investigated as well as the antioxidant activity of the norbergenin derivatives (1–4).
The results obtained showed that none of those compounds displayed antimicrobial activity at 0.5 μg/ml
whereas norbergenin derivatives exhibited noteworthy antioxidant property with IC50 values ranging from 8.2 μg/ml to 41.6 μg/ml.
The function of orbicules has been a matter of speculation for a long time and until now no satisfactory answer has been put forward. We propose two hypotheses that could contribute to the elucidation of their function: (1) that anemophilous and 'buzz pollination' species have microechinate orbicules and (2) that microechinate orbicules are advantageous for pollen release in both pollination modes. To test the first hypothesis, orbicule morphology of species in which the pollen is released by anther shaking (20 anemophilous and three with buzz-pollination) was analysed. We also conducted a literature review, noting the orbicule surface type and the presence of floral traits suggesting that pollen is released by anther shaking. Ninety-two percent of the species with microechinate orbicules are anemophilous or present 'buzz pollination'. Orbicules without spinules are present in species that lack floral traits linked to anther shaking for pollen release. To test the second hypothesis, a computational simulation that reveals the electric field produced by electrostatically charged orbicules was used. Results showed that the field is increased at the tip of each orbicule spinule. Therefore, an anther loculus surface coated by pointed structures increases the repulsion force of the pollen grains. This is an advantage when pollen is released by shaking the anther.
This study confined to authentication of herbal medicine namely Henna (Lawsonia inermis) the leaves and powdered leaves of which are used for jaundice, hair and skin problems. The study aimed to investigate indigenous medicinal uses, marketing status, macro and microscopical characters (LM & SEM) of pollen, foliar epidermal anatomy, behavior of powdered drug on treatment with different chemical reagents, fluorescence analysis (under visible & UV light) and preliminary phytochmeical tests to differentiate the genuine source from its adulterant. Such investigation may provide basis for authentication, standardization and characterization of genuine drug. The study concludes with authentication of Lawsonia inermis from its adulterant Mirabilis jalapa based on taxonomic and pharmacognostic characterization. These studies are useful especially for traded herbal drugs for their originality which leads to safe and quality assured herbal formulations for global acceptance.
Next to pollen, stamens of flowering plants often produce microstructures, called orbicules, lining the locules. Although the existence of orbicules has been known since 1865, their function still remains enigmatic. This paper surveys orbicule distribution throughout angiosperms, including +1,500 entries. We show that orbicules are found all over of flowering plants with an evolutionary trend towards orbicule absence in more derived clades. Orbicules are common in the ANITA-grade and 85 % of the monocots studied produce orbicules, with Orchidaceae, Commelinales and Zingiberales as notable exceptions. Within eudicots, asterids are most densely sampled with 61 % orbicule presence. Asteraceae and the majority of Lamiaceae lack orbicules. For 17 angiosperm orders orbicule distribution data are lacking entirely. We demonstrate that the hypothesized correlation of orbicule presence with non-amoeboid tapetum types holds true. The presence of orbicules is therefore a convenient proxy for tapetum characterization. The potential of orbicules as an a-cellular model system for patterned sporopollenin polymerization is discussed and suitable model plants for future functional orbicule-research are identified.
To clarify phylogenetic relationships among New Caledonian species of Diospyros, sequences of four plastid markers (atpB, rbcL, trnK-matK and trnS-trnG) and two low-copy nuclear markers (ncpGS and PHYA) were analysed. New Caledonian Diospyros species fall into three clades, two of which have only a few members (1 or 5 species); the third has 21 closely related species for which relationships among species have been mostly unresolved in a previous study. Although species of the third group (NC clade III) are morphologically distinct and largely occupy different habitats, they exhibit little molecular variability. Diospyros vieillardii is sister to the rest of the NC clade III, followed by D. umbrosa and D. flavocarpa, which are sister to the rest of this clade. Species from coastal habitats of western Grande Terre (D. cherrieri and D. veillonii) and some found on coralline substrates (D. calciphila and D. inexplorata) form two well-supported subgroups. The species of NC clade III have significantly larger genomes than found in diploid species of Diospyros from other parts of the world, but they all appear to be diploids. By applying a molecular clock, we infer that the ancestor of the NC clade III arrived in New Caledonia around nine million years ago. The oldest species are around seven million years old and the youngest ones probably much less than one million years.
Small, glabrous trees without latex. Leaves alternate, simple, exstipulate, finely and pinnately veined, with entire margins; extrafloral nectaries often present on abaxial leaf surface. Inflorescences axillary, racemose. Flowers with prophylls, articulated at base, sessile or shortly pedicellate, actinomorphic, 4-merous, unisexual (plants probably dioecious); calyx campanulate with 4 imbricate, retuse lobes, persistent but not enlarging in fruit; corolla sympetalous, isomerous with calyx, with lobes contorted sinistrorsely; tube prominent, bearing distally a corona of 8 lobes which, however, is missing in some species; male flowers with 8 stamens in one whorl, the filaments shortly connate and attached to the corolla tube below the middle; anthers linear, erect, basifixed, with an apiculate-prolonged connective, 4-sporangiate, dehiscing by longitudinal slits; locules and ovules not developed; female flowers: staminodes resembling stamens, but anthers collapsed and without pollen; gynoecium syncarpous, 4-carpellate; ovary inferior, 4-locular; style terminal; stigma shallowly 4-lobed; ovules 2 per locule, anatropous; placentation apical. Fruit an ovoid berry with a persistent calyx; seeds 1–2 by abortion, pendulous; hilum relatively small, apical; testa smooth, thin, coriaceous; endosperm horny, abundant, smooth; embryo straight, with two small foliaceous cotyledons and a strongly developed radicle.
The field of palynology is reviewed in terms of its contributions to angiosperm systematics and phylogeny. Principal pollen characters which are phylogenetically useful at higher taxonomic levels (including aperture type, pollen wall architecture, pollen-unit, polarity, symmetry, shape, and grain size), and their evolutionary trends are examined. Many palynological characters and concepts are subjected to re-examination, particularly in an evolutionary-phylogenetic context. An attempt is made to show how pollen characters correlate with various higher categories of the Takhtajan and Cronquist systems of angiosperm classification and to outline certain phylogenetic tends observed in the pollen of different groups of angiosperms. With some exceptions, pollen morphology is consistent with the levels of relative advancement and the relationships postulated in the Takhtajan and Cronquist systems. Angiosperm pollen grains are clearly divisible into two fundamentally different types (each with its own derivatives): heteropolar, bilateral, boat-shaped monosulcate pollen versus isopolar, radiosymmetric, globose tricolpate pollen. "Gymnospermous" monosulcate pollen and derivative types (ulcerate, disulculate, etc) characterize both the putatively primitive dicotyledonous subclass Magnoliidae and the monocotyledons. The six non-magnoliid dicotyledonous subclasses, on the other hand, are characterized by tricolpate pollen and derivative types (tricolporate, triporate, rugate etc.). Relatively primitive tricolpate pollen is retained by many Ranunculidae, Caryophyllidae, and "lower' Hamamelididae. The Dilleniidae (except Dilleniaceae) and Rosidae are somewhat more advanced in having basically compound-aperturate tricolporate pollen. The subclass Asteridae, which retains indications of a rosid ancestry, exhibits the greatest array of specialized pollen types. The most important palynological contradiction of the Takhtajan and Cronquist systems is the fact that the highly specialized, basically triporate pollen of the "higher" Hamamelididae (Amentiferae) can be more directly related to triangular tricolporate pollen of the Rosidae than to the tricolpate pollen of the "lower" Hamamelididae. Briefer sections of the paper deal with pollen technique and the major reference works of systematic palynology.
Molecular support for monophyly of Ebenales and its phylogenetic relationships to other dicot families was investigated using parsimony analysis of nucleotide sequences of the large subunit of ribulose-1,5-biphosphate carboxylase (rbcL). All families of Ebenales were examined except for the remote monogeneric family Lissocarpaceae, due to the lack of material. In addition we examined 93 rbcL sequences of 34 other families. All analyses strongly support monophyly of the asterid III grouping (identified in an analysis of 500 rbcL sequences by Chase et al.). The rbcL tree identified Ebenales to be distributed among several major subclades of the asterid III clade. Sapotaceae are sister to Symplocaceae and more closely related to Lecythidaceae/Scytopetalaceae than to other families of Ebenales. Ebenaceae are sister to Primulales. Styracaceae are polyphyletic, with Diapensiaceae and some Styracaceae sister to Polemoniaceae, and this clade is then sister to members of Clethraceae and Styrax. Although the phylogenetic hypothesis appears to be poorly resolved between groups, there is considerable support, as indicated by decay and jackknife support, for many of the inter-familial relationships.
A revised and updated classification for the families of flowering plants is provided. Many recent studies have yielded increasingly detailed evidence for the positions of formerly unplaced families, resulting in a number of newly adopted orders, including Amborellales, Berberidopsidales, Bruniales, Buxales, Chloranthales, Escalloniales, Huerteales, Nymphaeales, Paracryphiales, Petrosaviales, Picramniales, Trochodendrales, Vitales and Zygophyllales. A number of previously unplaced genera and families are included here in orders, greatly reducing the number of unplaced taxa; these include Hydatellaceae (Nymphaeales), Haptanthaceae (Buxales), Peridiscaceae (Saxifragales), Huaceae (Oxalidales), Centroplacaceae and Rafflesiaceae (both Malpighiales), Aphloiaceae, Geissolomataceae and Strasburgeriaceae (all Crossosomatales), Picramniaceae (Picramniales), Dipentodontaceae and Gerrardinaceae (both Huerteales), Cytinaceae (Malvales), Balanophoraceae (Santalales), Mitrastemonaceae (Ericales) and Boraginaceae (now at least known to be a member of lamiid clade). Newly segregated families for genera previously understood to be in other APG-recognized families include Petermanniaceae (Liliales), Calophyllaceae (Malpighiales), Capparaceae and Cleomaceae (both Brassicales), Schoepfiaceae (Santalales), Anacampserotaceae, Limeaceae, Lophiocarpaceae, Montiaceae and Talinaceae (all Caryophyllales) and Linderniaceae and Thomandersiaceae (both Lamiales). Use of bracketed families is abandoned because of its unpopularity, and in most cases the broader circumscriptions are retained; these include Amaryllidaceae, Asparagaceace and Xanthorrheaceae (all Asparagales), Passifloraceae (Malpighiales), Primulaceae (Ericales) and several other smaller families. Separate papers in this same volume deal with a new linear order for APG, subfamilial names that can be used for more accurate communication in Amaryllidaceae s.l., Asparagaceace s.l. and Xanthorrheaceae s.l. (all Asparagales) and a formal supraordinal classification for the flowering plants.
Quantitative data play an important role in palynological research. With the advent of digital imaging in light and electron microscopy, palynologists now have the opportunity to perform measurements faster and more precisely than ever before. Several image analysis software packages already exist for these tasks, but they are often expensive, difficult to use or not adapted to the specific needs of palynologists. After studying the daily workflow of a palynologist, we designed CARNOY, an image analysis application written from the ground up for use in palynology and morphology. CARNOY offers an easy-to-use interface and several features to make measuring easier and faster. The program can export measurements to almost every other software package for further analysis and is available for free on the Internet.
Coding quantitative data has always been one of the most difficult steps in morphological phylogenetics. A robust and theoretically well-supported method for coding quantitative data was proposed by Thiele (1993). This method relies on differential weighting of gaps between coded states within one character. The downside of gap weighting is that it requires many calculations. In order to provide an easy way of integrating quantitative data into phylogenetic analyses, we have created MorphoCode, an open-source software project. MorphoCode implements a variation on Thiele's method and offers the user the possibility to import data as tab-separated text, to choose the number of character states and to export the newly coded data to a NEXUS file.
In the course of a revision of Ebenaceae for "Flora Neotropica", "Flora of Ecuador", "Flora of the Guianas", and while preparing a corresponding treatment for Kubitzki's "Families and Genera of Vascular Plants", a study of the available literature was undertaken. Data pertaining to vegetative morphology and anatomy, indumentum, extrafloral nectaries, sexual polymorphism, inflorescence and flower structure, pollen mor-phology, pollination, embryology, fruits, seeds, seed dispersal, germination, seedlings, caryology, phyto-chemistry, ecology, geographical distribution, phylogenetic affinities, paleobotany, and use and economic importance are here compiled, together with relevant bibliography. In addition, a revised description of the family and its genera, and a key for the identification of Diospyros and Euclea are included. Zusammenfassung Im Rahmen einer Revision der Ebenaceae für "Flora Neotropica", "Flora of Ecuador", "Flora of the Guianas", sowie der Vorbereitung eines entsprechenden Beitrages für Kubitzkis "Families and Genera of Vascular Plants", wurde die verfügbare Literatur ausgewertet. Eine Zusammenfassung der relevanten Daten betreffend vegetative Morphologie und Anatomie, Behaarung, extraflorale Nektarien, Sexualpolymorphis-mus, Infloreszenz-und Blütenstruktur, Pollen, Bestäubung, Embryologie, Früchte, Samen und deren Ver-breitung, Keimung, Keimlinge, Karyologie, Phytochemie, Ökologie, geographische Verbreitung, Verwandt-schaftsverhältnisse, Paläobotanik, Verwendung und ökonomische Bedeutung wird hier mit entsprechender Bibliographie vorgestellt. Aktualisierte Beschreibungen der Familie und der beiden akzeptierten Gattungen Diospyros und Euclea, sowie ein Bestimmungsschlüssel für die Gattungen sind gleichfalls inkludiert.
Abstract— Data scored for cladistic analyses may be quantitative or qualitative, continuous or discrete, and show overlapping or non-overlapping values between taxa. Quantitative and qualitative are modes of expression of data, while continuous or discrete refer to properties of the set of numbers that express the data; both these pairs of terms have been confused with overlapping and non-overlapping. The degree of overlap of values between taxa is often used to filter characters in cladistic analyses: if a minimum amount of overlap is exceeded, or a minimum amount of disjunction not reached, characters are rejected as “not cladistic". However, this rests on a confusion between features of taxa and features of individual organisms (attributes). Cladistic characters are features of taxa, and comprise frequency distributions of attribute values over individuals of a taxon. Cladistic characters logically cannot overlap, although taxa may have overlapping attribute values. Thus, a priori rejection of characters that have overlapping attribute values is non-sensical. Such data may still be rejected from consideration for cladistic analysis if it could be demonstrated that they contain little recoverable phylogenetic signal. Few published analyses have empirically tested this. An analysis of overlapping morphometric data from three series of Banksia suggests that, at least in these cases, they map phylogeny almost as accurately as more conventional, qualitative morphological data. While more such tests are required, morphometric data should not be rejected a priori from cladistic analyses.
Orbicules, or Ubisch bodies, are sporopollenin particles lining the inner tangential and sometimes also the radial tapetal cell walls. They occur only in species with a secretory tapetum. The surface ornamentation of orbicules and pollen of the same species is often strikingly similar. Although orbicules were discovered more than a century ago, these structures remain enigmatic since their function is still obscure. Proposed hypotheses about their possible function are discussed. We also deal here with topics such as the possible allergenicity of orbicules and their representation in the fossil record. The use of orbicule characters for systematics is reviewed.
The distribution of orbicules throughout the angiosperms, based on a literature review from the first report until today, is shown in a list with 314 species from 72 families. Those species found in the literature without orbicules are presented together with their tapetum type. We plotted this information on a dahlgrenogram to visualize the distribution of orbicules. Orbicules occur in all subclasses of the angiosperms. Their occurrence is not correlated with certain modes of pollination or habitats.
The program MRBAYES performs Bayesian inference of
phylogeny using a variant of Markov chain Monte Carlo.
Availability: MRBAYES, including the source code, documentation,
sample data files, and an executable, is available at http://brahms.biology.rochester.edu/software.html.
Contact: johnh{at}brahms.biology.rochester.edu
Phylogenetic relationships of the pantropical family Ebenaceae s.l. were investigated using plastid DNA sequence data from six regions: atpB, matK, ndhF, trnK intron, trnL intron, and trnL-trnF spacer. Sampling included representatives of all currently recognized genera of Ebenaceae, Diospyros, Euclea, and Lissocarpa, and nearly all taxa that were previously recognized at the generic level, e.g., Cargillia, Gunisanthus, Maba, Macreightia, Royena, and Tetraclis. Our results strongly support monophyly of Ebenaceae s.l. and demonstrate that the previous infrafamilar classifications of the family do not circumscribe monophyletic groups. A new infrafamilial classification based on a phylogenetic approach is proposed here and consists of two subfamilies, Lissocarpoideae and Ebenoideae, and four genera, Lissocarpa, Euclea, Royena, and Diospyros. Relationships and potential synapomorphic characters are discussed and summarized. This study supports a western Gondwanan origin of family and indicates that both vicariant and long-distance dispersal events played an important role in attaining current distributions.
The program MODELTEST uses log likelihood scores to establish the model of DNA evolution that best fits the data.
AVAILABILITY: The MODELTEST package, including the source code and some documentation is available at http://bioag.byu. edu/zoology/crandall_lab/modeltest.html.
The recent development of Bayesian phylogenetic inference using Markov chain Monte Carlo (MCMC) techniques has facilitated the exploration of parameter-rich evolutionary models. At the same time, stochastic models have become more realistic (and complex) and have been extended to new types of data, such as morphology. Based on this foundation, we developed a Bayesian MCMC approach to the analysis of combined data sets and explored its utility in inferring relationships among gall wasps based on data from morphology and four genes (nuclear and mitochondrial, ribosomal and protein coding). Examined models range in complexity from those recognizing only a morphological and a molecular partition to those having complex substitution models with independent parameters for each gene. Bayesian MCMC analysis deals efficiently with complex models: convergence occurs faster and more predictably for complex models, mixing is adequate for all parameters even under very complex models, and the parameter update cycle is virtually unaffected by model partitioning across sites. Morphology contributed only 5% of the characters in the data set but nevertheless influenced the combined-data tree, supporting the utility of morphological data in multigene analyses. We used Bayesian criteria (Bayes factors) to show that process heterogeneity across data partitions is a significant model component, although not as important as among-site rate variation. More complex evolutionary models are associated with more topological uncertainty and less conflict between morphology and molecules. Bayes factors sometimes favor simpler models over considerably more parameter-rich models, but the best model overall is also the most complex and Bayes factors do not support exclusion of apparently weak parameters from this model. Thus, Bayes factors appear to be useful for selecting among complex models, but it is still unclear whether their use strikes a reasonable balance between model complexity and error in parameter estimates.
Marcgraviaceae are a rather small family of seven genera and approx. 130 neotropical species. This study aims to present a detailed palynological survey of the family in order to comment on the intrafamily relationships and possible correlations with pollinators.
In total, 119 specimens representing 67 species and all genera are observed using light microscopy and scanning electron microscopy. Furthermore, eight species from five genera are studied with transmission electron microscopy.
Our results show that pollen grains of Marcgraviaceae are small (20-35 microm), have three equatorial apertures, granules on the colpus membrane, oblate spheroidal to prolate spheroidal shapes, mainly psilate to perforate ornamentations, and lalongate colpus-shaped thinnings at the inner layer of the exine, and show the presence of orbicules. Based on our fragmentary knowledge of the pollination biology of the family, there are no clear correlations between pollinators and pollen features.
The genus Marcgravia has a high percentage of reticulate sexine patterns and a relatively thin nexine. Sarcopera can be defined by the presence of an oblate spheroidal to even suboblate shape, while Ruyschia and Souroubea typically show prolate spheroidal to subprolate pollen grains. The presence of a thick foot layer in the pollen wall is characteristic of the genera Norantea, Sarcopera and Schwartzia. Pollen features that are taxonomically useful within the family are the shape, sexine sculpturing, and ultrastructure of the pollen wall.
Character mapping on phylogenies has played an important, if not critical role, in our understanding of molecular, morphological, and behavioral evolution. Until very recently we have relied on parsimony to infer character changes. Parsimony has a number of serious limitations that are drawbacks to our understanding. Recent statistical methods have been developed that free us from these limitations enabling us to overcome the problems of parsimony by accommodating uncertainty in evolutionary time, ancestral states, and the phylogeny.
SIMMAP has been developed to implement stochastic character mapping that is useful to both molecular evolutionists, systematists, and bioinformaticians. Researchers can address questions about positive selection, patterns of amino acid substitution, character association, and patterns of morphological evolution.
Stochastic character mapping, as implemented in the SIMMAP software, enables users to address questions that require mapping characters onto phylogenies using a probabilistic approach that does not rely on parsimony. Analyses can be performed using a fully Bayesian approach that is not reliant on considering a single topology, set of substitution model parameters, or reconstruction of ancestral states. Uncertainty in these quantities is accommodated by using MCMC samples from their respective posterior distributions.
Numerous flowers and a diverse assemblage of leaves are mummified in clay lenses in the base of the Demons Bluff Formation overlying the Eastern View Coal Measures, in the Alcoa of Australia open cut near Anglesea, Victoria, Australia. Flowers are tubular, less than 10 mm long, and about 5 mm wide. Flowers all unisexual and staminate. Pollen subprolate, tricolporate, and about 32 mu m in diameter. Exine smooth to slightly scabrate. A rudimentary ovary occurs in some flowers. One of the more common leaf types found associated with the flowers is characterized by the same peculiar cuticular thickenings and overarching papillae on subsidiary cells that occur on sepals, which indicates that flowers and leaves represent a single taxon. Leaves are highly variable in size and shape but are consistently entire margined, with pinnate, brochidodromous venation. The suite of features characterizing the flowers is unique to the Ebenaceae. Flowers and leaves of many extant species of Diospyros (Ebenaceae) closely resemble the fossil flowers and leaves. Both flowers and leaves are considered conspecific and are named Austrodiospyros cryptostoma gen. et sp. nov. The Anglesea fossils represent one of the earliest well-documented occurrences of the Ebenaceae and are the earliest known remains of Ebenaceae from Australia. They support the hypothesis of a Gondwanan origin for the family with late Tertiary diversification in the Malesian region. -after Authors
A plastid rbcL sequence was obtained from a recent collection of Lissocarpa benthamii Gürke, and was analysed with two earlier molecular datasets. The results strongly support Lissocarpa as the sister group of Diospyros. We propose that Lissocarpa should be included in a broader concept of Ebenaceae. The number of genera that should be recognized in the family will depend upon more extensive molecular sampling.
The Sapotaceae is a well-defined pantropical family of about 900 species. Within it, however, there has been much controversy and little agreement over generic limits. The pollen of ca. 480 species have been examined in a worldwide morphological, palynological, and phytochemical generic survey of the family, carried out in collaboration with T. D. Pennington (New York Botanical Garden) and P. Waterman (University of Strathclyde). The pollen roughly separates into two groups, the first of which aligns with subfamilies Sideroxyloideae and Omphalocarpoideae, the second of which aligns with Mimu-sopoideae and Madhucoideae. Although pollen structure has been useful in helping to confirm the taxonomic position of one or two genera which were in doubt, it does not appear at present that it will be very effective in solving difficult taxonomic problems. Nevertheless the study has disclosed that the pollen possesses some interesting and characteristic structural elements which should prove useful in some instances for specific identification.
Pollen representing twenty-three species in all eleven genera of Styracaceae was examined by combined light, scanning and transmission electron microscopy. A comparative examination of the morphological features of the grains was conducted in an attempt to evaluate the importance of pollen morphology to the systematics of the family. Pollen morphology of the Styracaceae is rather uniform, with few variable features. Grains of nearly all examined species are solitary, mediumsized, radially symmetrical, tricolporate with lalongate endoapertures each with an equatorial bridge, and a tectate-pcrforate-columellatc exine. The stratified exine has four consistent layers, including a well-developed tectum and an endexine that is often internally thinly fragmented along the mesocolpi and thickened under the colpi. Although the pollen of the family is relatively uniform, subtle variation is present, particularly in sculpturing type, where intertwined rugulate-reticulate, transitional fossulatc-foveolate, finely rugulate, reticuloid, and weakly crotonoid exines occur. Outer tcctal surfaces are smooth, granulose, microspinulose, beaded and banded. The large genus Styrax encompasses much of the pollen morphological variation observed within the family. Grains of Parastyrax are occasionally 4-aperturate and have especially thick exines. Pollen morphology of the Styracaceae is similar to that of Theaceae, with both families possessing basically tricolporate grains with a finely perforate or reticulate sculpturing pattern and a smooth outer tectal wall. It is suggested that within both families specialization has led to similar fossulate-finely rugulate and foveolate-fossulate sculpturing patterns with beaded or banded outer tectal walls.
A revised and updated classification for the families of the flowering plants is provided. Newly adopted orders include Austrobaileyales, Canellales, Gunnerales, Crossosomatales and Celastrales. Pertinent literature published since the first APG classification is included, such that many additional families are now placed in the phylogenetic scheme. Among these are Hydnoraceae (Piperales), Nartheciaceae (Dioscoreales), Corsiaceae (Liliales), Triuridaceae (Pandanales), Hanguanaceae (Commelinales), Bromeliacae, Mayacaceae and Rapateaceae (all Poales), Barbeuiaceae and Gisekiaceae (both Caryophyllales), Geissolomataceae, Strasburgeriaceae and Vitaceae (unplaced to order, but included in the rosids), Zygophyllaceae (unplaced to order, but included in eurosids I), Bonnetiaceae, Ctenolophonaceae, Elatinaceae, Ixonanthaceae, Lophopyxidaceae, Podostemaceae (Malpighiales), Paracryphiaceae (unplaced in euasterid II), Sladeniaceae, Pentaphylacaceae (Ericales) and Cardiopteridaceae (Aquifoliales). Several major families are recircumscribed. Salicaceae are expanded to include a large part of Flacourtiaceae, including the type genus of that family; another portion of former Flacourtiaceae is assigned to an expanded circumscription of Achariaceae. Euphorbiaceae are restricted to the uniovulate subfamilies; Phyllanthoideae are recognized as Phyllanthaceae and Oldfieldioideae as Picrodendraceae. Scrophulariaceae are recircumscribed to include Buddlejaceae and Myoporaceae and exclude several former members; these are assigned to Calceolariaceae, Orobanchaceae and Plantaginaceae. We expand the use of bracketing families that could be included optionally in broader circumscriptions with other related families; these include Agapanthaceae and Amaryllidaceae in Alliaceae s.l., Agavaceae, Hyacinthaceae and Ruscaceae (among many other Asparagales) in Asparagaceae s.l., Dichapetalaceae in Chrysobalanaceae, Turneraceae in Passifloraceae, Erythroxylaceae in Rhizophoraceae, and Diervillaceae, Dipsacaceae, Linnaeaceae, Morinaceae and Valerianaceae in Caprifoliaceae s.l. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141, 399–436.
The glossary of pollen and spore terminology was first presented to the international palynological community as the final outcome of the Working Group on Palynological Terminology at the 8th International Palynological Congress in Aix-en-Provence in 1992. It became widely accepted as reference guide for palynologists to assist in the preparation of accurate and consistent descriptions of their material. It also serves as a practical source of information for non-specialists who wish to understand the meaning of the large number of existing palynological terms.The history of the glossary began in 1972 at the 3rd IPC at Novosibirsk when the working group on palynology was established. Throughout its history the project has been a collaborative effort with contributions from many palynologists, representing all branches of the discipline. Only through this long and elaborate procedure, with input from many people, it has been possible to produce the glossary.The entries are arranged alphabetically and are accompanied by simple schematic illustrations where appropriate. These contain the minimum amount of information needed to explain the feature. Moreover, to simplify the recognition of pollen and spore wall layers, colours have been used to indicate the corresponding layers.The first edition had 547 terms of which 339 have been accepted and recommended for use. In the second edition, a further 41 terms have been added with their appropriate illustrations. Of these, 10 have been accepted and 31 rejected for a variety of reasons. Where necessary, illustrations have been revised. An extensive list of consulted literature has been added.
The effect of various chemicals on the size of recent pollen grains of Corylus avellana L. and Quercus robur L. was studied. The size of acetolysed grains was affected by the treatment prior to acetolysis and moreover by the duration of acetolysis. Preparation methods, which produce comparable sizes and shapes, are given for both fresh and dried polliniferous material. It is concluded that size and shape are valuable for the determination of types. A new method is described for isolating anthers from herbarium sheets without causing great damage to the flower. Glycerin jelly proved to be a good mounting medium, provided that the cover glass is supported by granules of modelling clay.
Typescript. Thesis (Ph. D.)--City University of New York, 1994. Includes bibliographical references (p. 318-330). Photocopy.
This article is based on the address given by Miss Thompson at the Africa Day Conference held at the Royal Society of Arts on November 17, 1956.
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A taxonomic study of the Ebenaceae with special reference to Malesia. Thesis at the University of Oxford
- F S P Ng
Ng, F.S.P., 1971. A taxonomic study of the Ebenaceae with special reference to Malesia.
Thesis at the University of Oxford, 221 pp. (microfilmed by British lending library
division: no. D 183904).
PalDat-A Palynological Database: Descriptions, Illustrations, Identification, and Information Retrieval
- R Buchner
- M Weber
- Onwardss
Buchner, R., Weber, M., 2000 onwardss. PalDat-A Palynological Database: Descriptions,
Illustrations, Identification, and Information Retrieval. (http://www.paldat.org/).