A new species of Encephalartos (Zamiaceae) in Sudan

Encephalartos mackenziei Newton is described as a new cycad species occurring in south-east Sudan. It is characterized by its branching at the base to produce up to ten trunks per plant and by its broad leaflets, as well as some details of the cone scales. The population is large and widespread, and the presence of small seedlings indicates successful regeneration.  © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 187–192.     

Comparative anatomy and morphology of fertile complexes of Prumnopitys and Afrocarpus species (Podocarpaceae) as revealed by histology and NMR imaging, and their relevance to systematics

Fertile complexes (individual reproductive units of ovulate cones) of three Prumnopitys species and Afrocarpus falcatus (Podocarpaceae) were subjected to histological examination and non-destructive NMR imaging. The latter technique allowed the display, frame-by-frame analysis and electronic 'dissection' of internal structures such as the number and courses of vascular traces and resin canals and their morphology. Characters of these internal structures distinguished all three Prumnopitys species from each other and thus were shown to be taxonomically diagnostic. Fertile complexes of Prumnopitys andina and P. taxifolia were most similar, possessing simple vascular traces and few unbranched resin canals. Those of P. ferruginea were very different and possessed an interconnected network of resin ducts within the sarcotesta. These findings are congruent with relationships inferred from molecular phylogenetic studies, in which two subclades were recovered within Prumnopitys . The anatomy of the female fertile complexes of Afrocarpus falcatus was very distinct from all Prumnopitys species analysed. Its most distinctive feature was the existence of a complex network of radial vascular strands originating from within the outer layers of the sarcotesta and penetrating the inner layers of the fertile complex. The surface texture and morphology of the sclerotesta of the seed was also unique to each species.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 295–316.     

Facultative apomixis and hybridization in arctic Potentilla section Niveae (Rosaceae) from Svalbard

The degree of sexual seed set and the ability to cross were investigated in three taxa of Potentilla section Niveae ( P. chamissonis , P. insularis and P. nivea ) from the Svalbard Archipelago. Emasculated and bagged flowers had little or no seed set, while 71% of the emasculated, bagged and pollinated flowers had some. The taxa are interpreted as pseudogamous apomicts. Parental plants and offspring were subjected to isozyme analysis. Most of the offspring were clones of the mother plant, but 1.7% had bands from the pollen donor. The sexual offspring were all produced by mother plants of P. insularis , which is interpreted as a facultative apomict. Surprisingly, crosses between different taxa gave higher seed to ovule ratios and numbers of sexually produced offspring than crosses within taxa. Some of the sexually produced hybrid offspring had the same kind of hairs on the petioles as their mother taxon, showing that hybrids may not be intermediate in hair characters, which are considered important in the delimitation of these taxa. The results of the present study indicate very close relationships among the investigated plants.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 373−381.     

Shoot and compound leaf comparisons in eudicots: dynamic morphology as an alternative approach

Recent developmental studies suggest that the compound leaf is a more or less incompletely developed shoot. Instead of treating compound leaves and shoots as non-homologous, this interpretation draws a continuum between them. The present work considers the plant as a hierarchical series of units on which similar developmental processes are at work, and where each level (shoot, compound leaf, leaflet) is 'repeated' by the next higher level. Measurements related to the expression of developmental processes operating on leaves at the shoot level and on leaflets at the compound leaf level were used to determine if similar processes are at work at these different levels during early stages of organogenesis. Plants with compound leaves showing acropetal leaflet inception, representing a total of 16 species from ten eudicot families, were studied. Based on several types of quantitative analyses, there appears to be a continuum between so-called shoots, compound leaves and leaflets in the species studied. This perspective, qualified as dynamic morphology, both parallels and complements the classical interpretation.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143, 219−230.     

Expressed sequence tag analysis in <Emphasis Type="Italic">Cycas</Emphasis>, the most primitive living seed plant

Background

Cycads are ancient seed plants (living fossils) with origins in the Paleozoic. Cycads are sometimes considered a 'missing link' as they exhibit characteristics intermediate between vascular non-seed plants and the more derived seed plants. Cycads have also been implicated as the source of 'Guam's dementia', possibly due to the production of S(+)-beta-methyl-alpha, beta-diaminopropionic acid (BMAA), which is an agonist of animal glutamate receptors.

Results

A total of 4,200 expressed sequence tags (ESTs) were created from Cycas rumphii and clustered into 2,458 contigs, of which 1,764 had low-stringency BLAST similarity to other plant genes. Among those cycad contigs with similarity to plant genes, 1,718 cycad 'hits' are to angiosperms, 1,310 match genes in gymnosperms and 734 match lower (non-seed) plants. Forty-six contigs were found that matched only genes in lower plants and gymnosperms. Upon obtaining the complete sequence from the clones of 37/46 contigs, 14 still matched only gymnosperms. Among those cycad contigs common to higher plants, ESTs were discovered that correspond to those involved in development and signaling in present-day flowering plants. We purified a cycad EST for a glutamate receptor (GLR)-like gene, as well as ESTs potentially involved in the synthesis of the GLR agonist BMAA.

Conclusions

Analysis of cycad ESTs has uncovered conserved and potentially novel genes. Furthermore, the presence of a glutamate receptor agonist, as well as a glutamate receptor-like gene in cycads, supports the hypothesis that such neuroactive plant products are not merely herbivore deterrents but may also serve a role in plant signaling.

     

Genetic diversity and structure of the cycad Zamia loddigesii Miq. (Zamiaceae): implications for evolution and conservation

The genetic diversity and structure of four populations of the cycad Zamia loddigesii were studied throughout its range in Mexico. Allozyme electrophoresis of 15 loci was conducted. The mean number of alleles per locus was 1.80 ± 0.09, the percentage of polymorphic loci was 66.6 ± 5.4, and the expected heterozygosity was 0.266 ± 0.02. The results indicated that the genetic diversity was relatively higher, with respect to tropical tree species and other cycads. The genetic variation explained by differences among populations was 18%. On average, gene flow between paired populations was similar ( Nm  = 1.6) to other tropical forest trees and cycad species. Our results indicated that the geographical isolation among populations of Z. loddigesii generated allele loss, as well as a clinal variation in the frequencies of two loci ( MDH and MNR2 ), in relation to the latitudinal distribution of populations. The populations have become fragmented due to increasingly higher pressure of habitat conversion and disturbance. The importance of the establishment of sanctuaries and protected areas and a reduction in deforestation is highlighted in this research as a way of preserving the high genetic diversity of this and other endemic species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 533–544.     

A new species of Swertia L. (Gentianaceae) from Nepal

Swertia barunensis P. Chassot sp. nov. from Nepal is described and illustrated. It was collected in 1997 in an alpine meadow in the Makalu Barun National Park at an elevation of 4200 m. It belongs to Swertia section Macranthos T.-N. Ho & S.-W. Liu and resembles S. pseudohookeri H. Smith, from which it differs mainly by the shape of the nectary and the exomorphic seed structure. A key to all the species of sect. Macranthos is provided. The affinities of S. barunensis with some other taxa in the subtribe Swertiinae (Griseb.) Rchb. are also briefly discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141, 389–394.     

Pollination ecology of Arum italicum (Araceae)

The pollination ecology of Arum italicum was studied in south-western France. This plant attracts olfactory dung-breeding flies through deceit. These insects are principally represented by Diptera, all belonging to saprophyte families. The volatilization of the odouriferous compounds, responsible for their attraction, is achieved through the production of heat by the appendix. The insects are trapped for 24 h in order to participate in both sexual phases of the protogynous inflorescence. The male flowers produce three heat events during flowering. These peaks of heat seem to be involved in the spathe movements, since they occur during the opening of the inflorescence and the liberation of the insects. The last male heat event may be linked with the liberation of pollen and its dispersion by stimulating trapped flies. According to their frequency and pollen-load, two Psychoda species appear to be the most efficient pollinators ( P. crassipenis and P. pusilla ). Nevertheless, each of the other attracted species could play a significant role under different spatio-temporal conditions. Experiments on self-pollination have shown that obligate cross-pollination is necessary for A. italicum to set seeds. Moreover, hand- and natural-pollinated plants showed similarly high abortion frequencies suggesting that seed set may be more constrained by resources rather than by pollination limitation.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 205–214.     

Two new species from Gabon show the need to reduce Commitheca to the synonymy of Pauridiantha (Rubiaceae, Pauridiantheae)

Two new Pauridiantha species (Pauridiantheae, Rubiaceae) from Gabon are described and illustrated. Pauridiantha pleiantha is characterized by a combination of entirely glabrous stems, stipules with a narrow base, and large inflorescences. Pauridiantha smetsiana is characterized by entirely glabrous stems, narrow, early deciduous stipules, leaves red-tinged when dry, and small inflorescences. The two new species, especially P. smetsiana , close the morphological gap between Commitheca and Pauridiantha in combining typical Commitheca characters, such as glabrous stems and a narrow stipular base, with Pauridiantha characters, such as eucamptodromous venation. A detailed morphological comparison (including ovary structure, seed morphology, exotesta anatomy, and pollen morphology) between the two genera is given; in conclusion, Commitheca is relegated to the synonymy of Pauridiantha . The necessary new specific combinations are provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 105–117.     

Seed ecology of a Mediterranean perennial herb with an exceptionally extended flowering and fruiting season

Lobularia maritima is a Mediterranean short-lived herb with a flowering and fruiting season that lasts for ten months. Previous studies have shown that recruitment in periods other than autumn of the flowering season has few demographic implications; that is contributes little to the population growth rate. Since environmental conditions in periods other than autumn would allow recruitment, we examined to what extent the seed ecology of L. maritima accounts for recruitment shortage for the greater part of the year. To this end, we studied the effects of selfing and outcrossing on seed production and germination, within- and between-year variation in seed mass and germination, seed characteristics in the soil seed bank throughout the year, and the effect of temperature as a factor controlling seed germination. Results indicate that selfing does not decrease recruitment, and thus the observed changes in visitation rate and pollinator composition throughout the year cannot account for differences in recruitment. Germinability decreases throughout the year, suggesting a possible cost in reproduction associated with extended flowering. L. maritima has a transient seed bank whose seeds also experience a decrease in their germination throughout the year. Finally, temperature affects seed germination patterns, indicating the existence of quiescence mechanisms that prevent germination in the months prior to the summer drought. Overall, the results obtained support and, at least partly, explain the recruitment patterns of L. maritima observed in the field.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 273–280.     

Seed dormancy and germination behaviour in two Euphrasia species (Orobanchaceae) occurring in the Swiss Alps

The optimal time for germination of a seed depends on environmental conditions of its habitat, the life cycle of the germinating plant, and the conditions for successful establishment, growing and reproduction. We studied the germination behaviour of the alpine annual Euphrasia minima and an alpine ecotype of E. salisburgensis in a lowland garden experiment. Seeds of both species and their hybrids germinated at constant (5 °C) and at varying temperatures (3–10 °C), and never before spring after seed ripening. Germination was spread over 3 years, which suggests that the seeds formed a persistent seed bank. The two species together with E. minima and E. minima hybrids differed significantly in the germination rate in the first and second spring.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 649–656.     

Trichomes of the seed fern Blanzyopteris praedentata: implications for plant–insect interactions in the Late Carboniferous

The fronds and compound tendrils of the Stephanian (Late Carboniferous) seed fern Blanzyopteris praedentata possess several types of trichomes, two of which may, based on their morphology, have functioned as deterrents against herbivores. Bands of upwardly curved trichomes, occurring on the adaxial surfaces of tendrils, frond- and pinna rachides, are also known from extant plants, where they create mechanical obstacles. Other trichomes that are glandular occur on most parts of the foliage and tendrils and represent a different form of defence mechanism. These trichomes apparently possessed a touch-sensitive mechanism that opened the secretory cell when touched. They are interpreted as functionally similar to the so-called 'explosive' trichomes of certain extant Cucurbitaceae and Solanaceae. Studies of living Oleander aphids ( Aphis nerii ) on Sicana odorifera (Cucurbitaceae) demonstrate the effectiveness of the physical component of this defence. When touched and ruptured by an aphid, the trichomes of S. odorifera rapidly release a sticky exudate, which adheres to the animal's legs; the accumulation of exudate on the legs eventually impedes the aphid. Based on these studies, hypotheses are presented on the types of animals that might have been deterred by the glandular trichomes of B. praedentata in the Late Carboniferous.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 133–149.     

Water lilies and scarabs: faithful partners for 100 million years?

Night-flowering water lilies (Nymphaeaceae) in South America are pollinated by Cyclocephala scarab beetles (Scarabaeidae: Cyclocephalini) in a specialized relationship involving synchronized flowering movements, strong floral scent, food tissues, and heat-producing flowers. We report that a similar and closely related association exists in West Africa between Nymphaea lotus L. and Ruteloryctes morio Fabricius (Cyclocephalini). This finding strongly supports a late Early Cretaceous origin of a symbiosis between the night-flowering water lilies and pollinating Cyclocephalini beetles. We believe that this is the first unambiguous evidence of a plant-pollinator relationship of this age.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 539–543.     

Influence of selfing and maternal effects on life-cycle traits and dispersal ability in the herb Hypochaeris radicata (Asteraceae)

The ecological and evolutionary implications of dispersal are many. Pollination type and maternal effects may affect plant fitness traits, including life-cycle traits as well as dispersal ability. This study investigated the joint influence of pollination type and maternal effects on both life-cycle traits and dispersal ability in the herb Hypochaeris radicata . We conducted experimental crosses to obtain selfed and outcrossed progeny. Individual seeds and their pappuses were measured to determine seed terminal velocity. Seed size was also used to assess maternal effects. Selfing dramatically decreased seed set, indicating that H. radicata is self-incompatible. However, the few selfed seeds produced outperformed outcrossed seeds in seed size and flowering probability, surely as a result of an effective reallocation of resources among selfed seeds. None of the life-cycle traits was affected by seed size, the estimate of maternal effects. Selfed seeds were larger and bore a smaller pappus than outcrossed seeds. As a result, dispersal ability was lower for selfed than outcrossed seeds. Several factors, such as the low proportion of plants that produced selfed seeds, the low number of selfed seeds produced per plant, and the lack of self-fertility mechanisms might act in concert to prevent the evolution of selfing in H. radicata .  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 163–170.     

A new species of Valantia (Rubiaceae) from Spain

A new species of Valantia , V. lainzii Devesa & Ortega-Olivencia, endemic to the coastal zone of Granada (southern Spain) is described. The species recalls V. muralis L. in its general aspect, and V. deltoidea Brullo in the type of fructiferous body.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 331–335.     

Stomatal length and frequency as a measure of ploidy level in black wattle, Acacia mearnsii (de Wild)

The length and frequency of stomata on leaf surfaces were examined as rapid techniques for future identification of ploidy level of Acacia mearnsii (de Wild). Diploid (2 n  = 2 x  = 26) and tetraploid (2 n  = 4 x  = 52) plants were germinated from chipped seed at 25°C and grown under nursery conditions. After one month, measurements showed that the mean stomatal length was 27.17 ± 0.474 µm for diploids and 40.24 ± 0.521 µm for tetraploids and these differed significantly from each other ( P  < 0.001). The frequency of stomata per leaf surface was shown to decrease significantly ( P  < 0.001) as the ploidy level increased, with a mean of 22.11 ± 0.495 for diploids and 10.26 ± 0.495 for tetraploids. It was concluded that stomatal length and stomatal frequency are rapid indirect methods to identify ploidy level in black wattle.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 177–181.     

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.     

Evolution probing for semiochemicals based on secondary metabolites in the cuticles of three species of Halobates (Heteroptera: Gerridae)

We developed methods for characterizing semiochemicals in the cuticles of the sea-skater Halobates (Gerridae). Such substances may be involved in chemical communication at the sea surface and have been shown to act as sexual pheromones for a related marine insect Trochopus plumbeus (Veliidae). We identified 86 volatile compounds in extracts from three Halobates species ( H. micans Eschscholtz, H. hawaiiensis Usinger and H. sobrinus White) and used them to reconstruct the phylogeny of the clade. Sixty-seven of the compounds were found to be parsimony-informative. Suspected semiochemicals appear to be specific autapomorphies in the phylogram, and our results indicated that chemical profiles may be used to predict the phylogenetic history of Halobates . Moreover, the phylogenetic tree derived from our data is congruent with a previously published one based on morphology and mitochondrial DNA sequence (780 bp in the COI gene). The potential role of each compound to be used for chemical communication was also reviewed. At appropriate concentrations, some of the cuticular compounds, being partly hydrophobic and partly hydrophilic, could disperse on the sea surface and thereby play an energetically efficient role in chemical communication, e.g. mate location. Less dispersible components might act as 'sun-screens' and provide protection from UV-irradiation during daylight hours.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 671–688.