Trichome complement of Turnera and Piriqueta (Turneraceae)

The indumentum of Piriqueta and Turnera is made up of nine different types of trichomes, which broadly can be divided into glandular and non-glandular. Taking into account foot shape, head size and pedicle size, five variants of glandular trichomes are recognized: microcrapitate, stipitate-capitate, sessile-capitate and setiform. The non-glandular trichomes can be simple (unicellular or pluricellular-uniseriate), stellate or porrect-stellate. The setiform glandular hairs are present in most species of Piriqueta . Simple unicellular hairs are the most widespread type, frequently being found in combination with other trichomes. Stellate trichomes show a restricted distribution in both genera. Within Piriqueta , section Africana has only simple trichomes, whereas section Piriqueta has also porrect-stellate trichomes; groups of species can be set up according to the presence and type of glandular trichomes. Within Turnera the stipitate-capitate trichomes are exclusive to series Papilliferae ; sessile-capitate trichomes are found in series Microphyllae , Annulares and in some species of Salicifoliae ; clavate trichomes are found only in series Turnera ; setiform glandular hairs are exclusive to T. collotricha , whereas the microcapitate trichomes are widely distributed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 85–97.     

The uncommon cavitated secretory trichomes in Bauhinia s.s. (Fabaceae): the same roles in different organs

Cavitated secretory trichomes are characterized by a short or absent stalk that is connected to a secretory hollow head. They are rare structures in angiosperms; in Fabaceae, they have been recorded in only seven genera, including Bauhinia s.s. Because B. curvula and B. rufa exhibit glands that are responsible for attracting pollinators to flowers, this study aimed to test whether the cavitated secretory trichomes present in the flowers of these species have an attraction function. As leaf trichomes are commonly related to plant defence, comparative analyses of the morphology, ontogeny, ultrastructure and chemical profile of the secretory trichomes present in flowers and leaves were conducted. It was found that cavitated secretory trichomes are similar in their external morphology and development, regardless of the organ or species analysed. However, interspecific differences were found in the secretion process and chemical profile of the exudate. The differences found in the cavitated secretory trichomes between species indicate that they secrete distinct compounds, whereas the similarities found in these structures between vegetative and reproductive organs indicate that the cavitated trichomes have equivalent ecological functions within a species, probably in plant defence during organ development. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 104–122.     

A new species of Lepidostrobus from the Upper Devonian of Xinjiang, China and its bearing on the phylogenetic significance of the order Isoëtales

Lepidostrobus xinjiangensis sp. nov. is described from Upper Devonian rocks of the eastern Junggar Basin, Xinjiang, north-west China. It provides new insights into the reproductive diversification and phylogenetic relationships of lycopsids in the Late Devonian. The strobilus shares more characteristics with arborescent lycopsids than any herbaceous forms, and falls within the diagnosis of Lepidostrobus . Each sporophyll consists of a deltoid pedicel and a triangular lamina. Sporophylls are horizontally inserted into the strobilus axis in low spirals. The pedicel, with lateral alations and an abaxial keel, extends distally into the upturned lamina and downturned heel, producing a peltate appearance. A single sporangium with terminal longitudinal dehiscence is a radially elongate, dorsiventrally flattened ovoid and is attached along its length to the adaxial surface of the pedicel. A column-like subarchesporial pad is found in the sporangium. A possible ligule occurs on the adaxial surface of the pedicel distal to the sporangium. The strobilus is microsporangiate, containing Lycospora -type spores with granulate ornamentation and an equatorial flange. Based on this new species, the reproductive diversification and evolutionary pattern in arborescent lycopsids from the Devonian through the Carboniferous are discussed in a phylogenetic framework. We suggest that the reproductive strategies represented by bisporangiate- and monosporangiate-strobili had proliferated by the Late Devonian, which implies that phylogenetically advanced arborescent lycopsids bearing Lepidostrobus strobili had an earlier origin than previously thought.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 55−67.     

Transgenic Nicotiana TabacumL. with enhanced trichome exudate cembratrieneols has reduced aphid infestation in the field

Glandular trichomes of many plants secrete natural products that influence plant/insect interactions. For example, tobacco varieties having relatively high cembratrieneols (CBTols) are known to have enhanced aphid resistance in the field. CBTols and corresponding CBTdiols comprise 1.4 and 62%, respectively, of trichome exudate in the aphid susceptible tobacco variety, T.I. 1068. Using this cultivar we suppressed the CYP71D16 gene that encodes the enzyme that converts CBTols to CBTdiols. In suppressed plants CBTols and CBTdiols accounted for about 27 and 35% of exudate weight, respectively. Total CBTols plus CBTdiols was not changed substantially. Here we studied the relationship between aphid infestation and increased exudate CBTols in the field using self progeny derived from 5 independent primary transgenic T. I. 1068 plants having suppressed CYP71D16 activity. Two hundred individual plants were scored for aphid infestation, and their trichome exudate compositions were determined by gas chromatography. A significant negative correlation was found between high CBTols levels in trichome exudates and aphid infestation. No aphid infestation was observed on the majority of plants with CBTols/CBTdiols ratios of >1.49, which represents a >20-fold increase in CBTols, and a >40% decrease in CBTdiols over control T.I. 1068 exudate. In contrast, aphid infestation occurred on most plants with CBTols/CBTdiols ratios <0.201, which is similar to that of the untransformed control T.I. 1068. These results demonstrate the feasibility of using metabolic engineering of glandular trichomes to enhance natural product-based pest resistance.     

Atypical pseudopollen-forming hairs in Maxillaria (Orchidaceae)

Pseudopollen is found on the labella of several species of Maxillaria and is formed by the fragmentation of uniseriate, multicellular, moniliform trichomes. The resultant component cells are rich in protein. Since flowers of pseudopollen-forming species generally lack nectar, it is probable that pseudopollen gradually replaced nectar as the pollinator reward. However, direct evidence for this hypothesis is lacking. The present paper examines the labellar micromorphology of five Brazilian species of Maxillaria assigned to the M. discolor alliance. The flowers of two of these species, namely M. violaceopunctata and M. villosa , produce both food hairs and a lipoidal, labellar secretion which is rich in aromatic amino acids. Moreover, in the case of M. violaceopunctata , the secretion was found to contain reducing sugars. This may represent an intermediate stage in the transition from nectar-producing to pseudopollen-forming flowers. SEM studies indicate how pseudopollen-forming, moniliform trichomes, such as those found in the Maxillaria grandiflora complex, may have evolved from simple, uniseriate, multicellular hairs.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 151−158.     

Heritable genetic variation and potential for adaptive evolution in asexual aphids (Aphidoidea)

Aphid life cycles can encompass cyclical parthenogenesis, obligate parthenogenesis, obligate parthenogenesis with male production and an intermediate 'bet-hedging' strategy where an aphid genotype will over-winter by continuing to reproduce by parthenogenesis and by investment in sexually produced eggs. In this paper, we focus on aphid lineages that reproduce entirely parthenogenetically (asexual aphids), in contrast to those that have any sexual forms in the annual cycle. Using modern molecular techniques, aphid biologists have made many empirical observations showing that asexual lineages are widespread both geographically and temporally. Indeed, we are collectively beginning to gather data on the evolution and persistence of these lineages through time. Here we review aphid karyology and parthenogenesis, both essential for interpretation of the molecular and ecological evolution of aphid asexual lineages. We describe the growing list of studies that have identified aphid genotypes that are both temporally and geographically widespread. We then collate examples of molecular and chromosomal evolution in asexual aphids and review the literature pertaining to phenotypic evolution and ecological diversification of asexual aphid lineages. In addition, we briefly discuss the potential of bacterial endosymbionts and epigenetic effects to influence the evolution of asexual aphid lineages. Lastly we provide a list of aphid taxa believed to be obligately asexual. This will be a useful resource for those seeking parthenogenetic animals as study systems. In conclusion, we present guidelines for the use of the term clone in aphid biology and stress the need for well-designed and well-executed studies examining the potential of asexual aphid lineages for adaptive evolution.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 115–135.     

A new polypterid fish: Polypterus faraou sp. nov. (Cladistia, Polypteridae) from the Late Miocene, Toros-Menalla, Chad

Polypterus faraou sp. nov. (Cladistia, Polypteridae) from the Late Miocene of Toros-Menalla (western Djourab, Chad) is described on the basis of a subcomplete articulated skeleton preserved in three dimensions. This is the first time such a complete fossil polypterid skeleton has been described. It is the only verifiable fossil record for the genus Polypterus . P. faraou closely resembles P. bichir and P. endlicheri , extant fish found in the Chad–Chari system. Intrarelationships among the polypterids are not yet resolved: however, P. faraou shows a primitive shape of the body and a primitive shape of the opening of the lateral line on the scales, similar to that of three living species ( P. bichir , P. endlicheri and P. ansorgii ).  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 146 , 227–237.     

Genetic changes within an aphid clone: homogenization of rDNA intergenic spacers after insecticide selection

A single asexual maternal lineage (i.e. clone) of the greenbug aphid, Schizaphis graminum (Rondani) was repeatedly selected with the insecticide disulfoton ( O , O -diethyl S- [2-(ethylthio)ethyl] phosphorodioate). A parallel colony of the non-selected clone was also maintained. After approximately 200 generations (4 years) of continuous selection, both the selected and non-selected clones were assayed for changes in intergenic spacer (IGS) length variants of the rRNA cistron. No changes in sets of IGS variants were detected in the non-selected clone. However, the selected clone was found to have lost three variants present in the non-selected clone. This probably occurred by unequal cross-over between sister chromatids, whereby the cistron became homogenized by an increase of frequency of two smaller variants. This documents a large-scale genetic change occurring within the rRNA cistron in a parthenogenetically reproducing aphid.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 101–105.     

A phylogeny of the families of fossil and extant tetraodontiform fishes (Acanthomorpha, Tetraodontiformes), Upper Cretaceous to Recent

A data matrix is presented of 210 morphological characters (mostly osteological, some external) for 20 extant taxa of the ten Recent families of tetraodontiform fishes and 36 fossil tetraodontiforms. The oldest of these are from the Upper Cretaceous (95 Mya); most are from the Lower to Middle Eocene (50–58 Mya). There are two outgroup taxa (a zeiform and a caproid). A cladistic analysis of this matrix for only the extant taxa produced two equally parsimonious trees that call into question the monophyly of some of the previously accepted major higher-level tetraodontiform clades. Inclusion in the analysis of the large number of available fossil taxa helps to resolve relationships between family level clades. The new phylogenetic hypothesis, together with stratigraphic and biogeographical data, is used to discuss scenarios of the origin and evolution of the major clades of the order.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 565−617.     

Trichomes of Lycopersicon spp. and their effect on Myzus persicae (Sulzer) (Hemiptera: Aphididae)

Abstract The green peach aphid ( Myzus persicae ) is becoming resistant to many of the pesticides used in the tomato ( Lycopersicon esculentum ) industry. A potential alternative method of control is host-plant resistance, which may be mediated by trichomes. Resistance to the green peach aphid was assessed for six accessions of L. hirsutum, three of L. pennellii and a L. esculentum control. Mortality, entrapment and number of aphids unaffected by trichomes were recorded 24, 48 and 72 h after placement. Using multiple regression against counts of glandular and non-glandular trichomes, the role of foliar pubescence in resistance to the aphid was established. Lycopersicon pennellii possessed the highest level of resistance after 72 h and type IV trichome density accounted for 26.5% of variance in mortality ( P  < 0.001). Low densities of type IV and high densities of type VII trichomes increased the number of aphids 'unaffected' by trichomes, a relationship accounting for 29% of variance ( P  < 0.001). Information on which types of trichomes most strongly affect resistance to insect pests will aid future breeding and L. pennellii shows potential for use in breeding programs to increase levels of host-plant resistance in L. esculentum.     

Leaf morphology of hosts and nonhosts of the thrips Heliothrips haemorrhoidalis (Bouché)

This study highlights the morphological differences between two groups of plants; those that are recorded as hosts of the thrips Heliothrips haemorrhoidalis (Bouché) and those that remain free from infestation. Scanning electron microscope techniques were used to describe the morphology of the leaf surface of 19 species in order to help identify which morphological features could play a role in the selection of plant species by thrips from among a diverse botanical collection. H. haemorrhoidalis had a preference for species with leaves that were coriaceous, with one or both surfaces being smooth. Plants evading these thrips commonly possessed glandular trichomes. Thus morphology may have a role in deterring thrips from the leaf surface, as well as influencing the behaviour of predators that control thrips. Further detailed studies into the chemistry of leaves of hosts and nonhost species may help to further our understanding of selection mechanisms.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 109–130.     

Killer sperm whale: a new basal physeteroid (Mammalia, Cetacea) from the Late Miocene of Italy

Zygophyseter varolai , a new genus and species of Physeteroidea (Cetacea, Odontoceti), is based on an almost complete skeleton from the Late Miocene (Tortonian) in southern Italy. The extreme elongation of the zygomatic process of the squamosal and the circular supracranial basin (probably for housing the spermaceti organ) delimited by a peculiar anterior projection of the supraorbital process of the right maxilla are the most distinctive features of this bizarre sperm whale. Large body size, large teeth present in both lower and upper jaw, and anteroposteriorly elongated temporal fossa and zygomatic process of the squamosal indicate that this cetacean (for which we suggest the English common name killer sperm whale) was an active predator adapted to feeding on large prey, similarly to the extant killer whale ( Orcinus orca ). A phylogenetic analysis reveals that Zygophyseter belongs to a Middle–Late Miocene clade of basal physeteroids, together with Naganocetus (new genus for the type of ' Scaldicetus ' shigensis ). Moreover, the phylogenetic analysis shows evidence of a wide physeteroid radiation during the Miocene and that the extant Physeter and Kogia belong to two distinct families that form a clade representing the crown-group Physeteroidea.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 103–131.     

Ontogenesis of internal secretory cells in leaves of Lantana camara (Verbenaceae)

Internal secretory cells may be morphologically indistinguishable from their neighbours except for the presence of secreted material, or they may differ to such an extent that could be classified as secretory idioblasts. Several authors have reported the presence of glandular trichomes secreting essential oil in Verbenaceae, including Lantana . However, none have reported internal secretory cells. Anatomical and histochemical methods applied to Lantana camara leaves revealed the occurrence of internal secretory cells whose ontogenesis and chemical nature are described in this paper. According to leaf developmental analysis, L. camara secretory cells originated from the ground meristem, started to differentiate in the third node leaves, and were actively secreting in the fourth node leaves. The content of the secretory cells was of a lipidic nature, and a terpenoid essence of their secretion was also identified. Based upon differences in size and shape from neighbouring cells and on detection of nonvolatile terpenes, they were confirmed as true internal secretory idioblasts.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 427–431.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

Function of the siphunculi in aphids with particular reference to the sycamore aphid, Drepanosiphum platanoides

Sycamore aphids with occluded siphunculi survive, give birth to as many offspring and maintain their weight as well as aphids with normal siphunculi and it is therefore unlikely that siphunculi have an excretory function. Blocking the siphunculi of sycamore aphids does not affect the way they space out. Siphunculi are used in defence against predators most effectively when the aphid and predator are similar in size.The smell of siphuncular exudate causes aphids like Myzus persicae to scatter but in the sycamore aphid the smell has to be accompanied by the sight or vibration of a struggling aphid to induce an alarm response. Prodding the abdomen and pinching the legs of a sycamore aphid induces it to produce siphuncular exudate.     

Structure,Ontogeny, Classification and Organographic Distribution of Trichomes in Some Cucurbitaceae

The structure, ontogeny, classification and organographic distribution of trichomes in 9 genera and 12 species of the Cucurbitaceae are presented. The vegetative and floral trichomes are classified into glandular and non-glandular. In all 13 main types and 26 subtypes of glandular and nonglandular trichomes are observed. All the types of trichomes originate from a single papillate hair initial. The trichomes are classified on the basis of form, structure and contents.     

The fossil history of palms (Arecaceae) in Africa and new records from the Late Oligocene (28–27 Mya) of north-western Ethiopia

The African palm fossil record is limited but the data provide an outline of palm evolution from the Late Cretaceous through the Neogene. Pollen attributed to palms is reported from the Aptian (125–112 Mya), but the earliest unequivocal record in Africa is Campanian (83.5–70.6 Mya). Palms diversified 83.5–65.5 Mya and became widespread, although most records are from the west and north African coasts. Many taxa were shared between Africa and northern South America at that time, but a few were pantropical. Extirpations occurred throughout the Palaeogene, including a notable species turnover and decline at the Eocene–Oligocene boundary (33.9 Mya), a change that resulted in the elimination of nypoid palms from Africa. The Neogene plant macrofossil record is better sampled than the Palaeogene, although few palms are documented. Thus, the low diversity of African palms today is more likely the result of Palaeogene, rather than Neogene extinctions. Newly discovered palm fossils of leaves, petioles and flowers from the Late Oligocene (27–28 Mya) of north-western Ethiopia document the abundance and dominance of palms in some communities at that time. The fossils represent the earliest records of the extant genera Hyphaene (Coryphoideae) and Eremospatha (Calamoideae).  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 69–81.     

Some remarks on the genus Astragalus sect. Incani in Iran

New findings are given for the Astragalus sect. Incani , which occurs within the area covered by the Flora of Iran. Astragalus wojciechowskianus Ranjbar is described as a new species. A. procerus Boiss. & Hausskn . , A. cottonianus Aitch. & Baker, A. achundovii Grossh., A. orduabadensis Grossh. and A. glaucophyllus Bunge are reported as new records for the flora of Iran .   © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 443–447.     

Role of leaf glandular trichomes of melon plants in deterrence of Aphis gossypii Glover

External characteristics of the leaf epidermis and their effects on behaviour of Aphis gossypii Glover were evaluated in two Cucumis melo L. genotypes, ‘Bola de Oro’ (aphid susceptible) and TGR‐1551 (aphid resistant) in order to explore their role in the early rejection of TGR‐1551 by this aphid. No differential effects of epicuticular waxes on aphid behaviour were observed. The type, distribution and number of trichomes on melon leaves were also studied. Pubescence in melon, measured as the number of non‐glandular trichomes per cm², was not sufficient to prevent aphid settling. However, there was a high density of type I glandular trichomes on leaves of the aphid‐resistant genotype. According to microscopic observations and stain testing, these trichomes store and secrete phenols and flavonoids. Free‐choice tests were conducted to determine the effect of these glandular trichomes on A. gossypii preference, revealing that aphids reject leaf disks of TGR‐1551 from the onset of the experiment. Additional experiments after removal of leaf type I glandular trichome exudates showed that A. gossypii preferred washed TGR‐1551 leaf disks over unwashed disks, while this effect was not observed in experiments using washed and unwashed ‘Bola de Oro’ leaf disks. These results suggest that a high density of glandular trichomes and chemicals secreted by them deter A. gossypii and disturb aphid settling on TGR‐1551.