Drosophyllum lusitanicum, an endangered West Mediterranean endemic carnivorous plant: threats and its ability to control available resources

Drosophyllum lusitanicum (L.) Link (Droseraceae), Erva pinheira orvalhada, an endangered carnivorous plant, is a local endemic of clearings in pine, cork oak and oak forests or their successional shrublands in the western Iberian Peninsula and Morocco. The conservation status, distribution and population dynamics of this species are only partially known, both for Spanish and Portuguese occurrences. Portuguese distribution data from herbarium and bibliographic sources were collected for this study. Field work on 50 populations was undertaken in order to improve knowledge on their conservation status and possible threats. Natural causes, infrastructure and housing construction are stressed as the most important threats. Germination tests were carried out and the distribution patterns, plant height and the number of flowers were studied in an arbitrarily chosen population. A relatively high light requirement and low competitive ability to gain light are considered factors likely for the microdistribution pattern. Drosophyllum lusitanicum cannot compete for light in habitats with intense competition. Seeds from D. lusitanicum have reduced probability of germination when adult plants are already growing in the area. It is hypothesized that seed germination shows a similar pattern to flowering, viz over a considerable time rather than in a sudden flush. This strategy may be a key factor for the species' survival, representing attempts to take advantage of available resources, preventing intraspecific competition and, finally, preventing sudden total disappearance of a population, resulting from any catastrophic events due to natural causes or to human activity. As a result of this study, some conservation measures are suggested.   © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 383–390.     

Is Drosera meristocaulis a pygmy sundew? Evidence of a long-distance dispersal between Western Australia and northern South America

Background and aims

South America and Oceania possess numerous floristic similarities, often confirmed by morphological and molecular data. The carnivorous Drosera meristocaulis (Droseraceae), endemic to the Neblina highlands of northern South America, was known to share morphological characters with the pygmy sundews of Drosera sect. Bryastrum, which are endemic to Australia and New Zealand. The inclusion of D. meristocaulis in a molecular phylogenetic analysis may clarify its systematic position and offer an opportunity to investigate character evolution in Droseraceae and phylogeographic patterns between South America and Oceania.

Methods Drosera meristocaulis

was included in a molecular phylogenetic analysis of Droseraceae, using nuclear internal transcribed spacer (ITS) and plastid rbcL and rps16 sequence data. Pollen of D. meristocaulis was studied using light microscopy and scanning electron microscopy techniques, and the karyotype was inferred from root tip meristem.

Key Results

The phylogenetic inferences (maximum parsimony, maximum likelihood and Bayesian approaches) substantiate with high statistical support the inclusion of sect. Meristocaulis and its single species, D. meristocaulis, within the Australian Drosera clade, sister to a group comprising species of sect. Bryastrum. A chromosome number of 2n = approx. 32–36 supports the phylogenetic position within the Australian clade. The undivided styles, conspicuous large setuous stipules, a cryptocotylar (hypogaeous) germination pattern and pollen tetrads with aperture of intermediate type 7–8 are key morphological traits shared between D. meristocaulis and pygmy sundews of sect. Bryastrum from Australia and New Zealand.

Conclusions

The multidisciplinary approach adopted in this study (using morphological, palynological, cytotaxonomic and molecular phylogenetic data) enabled us to elucidate the relationships of the thus far unplaced taxon D. meristocaulis. Long-distance dispersal between southwestern Oceania and northern South America is the most likely scenario to explain the phylogeographic pattern revealed.     

Minor pollinator–prey conflict in the carnivorous plant, Drosera anglica

We studied the physical and temporal isolation of two arthropod guilds interacting with Drosera anglica Huds., a terrestrial carnivorous plant. Flowers are separated from basal trap leaves by a leafless stalk. Since arthropods are potentially employed both as prey and pollinators, we asked whether separation of traps from flowers reduces the frequency with which flower visitors are captured by the leaves. Plants captured prey throughout the season, with peak trapping activity occurring before flowering began. The diverse prey spectrum included at least 109 species in 94 genera in 26 of 37 identified families representing 11 arthropod orders. The most common prey were adult flies of Nematocera, particularly Ceratopogonidae (50%) and Chironomidae (42%). The following taxa were periodically abundant: Acarina, Diptera–Cecidomyiidae, Chloropidae, Sciaridae, Hemiptera nymphs and Thysanoptera–Thripidae. Flies (Diptera) were chief flower visitors (95%), dominated by Syrphidae (66%), Bombyliidae and Muscidae (10% each), Calliphoridae (7%), Tachinidae and Dolichopodidae (3% each). Additionally, visitors were a bee (Hymenoptera–Halictidae) and thrips (Thysanoptera–Thripidae). Four families were common to both guilds: Diptera–Dolichopodidae, Muscidae, Tachinidae; and Thysanoptera–Thripidae. However, direct comparisons of identified taxa within these families showed that overlap between flower visitors and prey occurred for Thrips sp. larvae alone, which comprised only 3% of all flower visitors and 0.5% of prey. Drosera anglica exploits distinct guilds of insects for pollinators and prey.     

Uniformity of organellar DNA in Aldrovanda vesiculosa, an endangered aquatic carnivorous species, distributed across four continents

Organellar DNA from the widely distributed but rare and critically endangered aquatic carnivorous plant Aldrovanda vesiculosa (Droseraceae) was examined. Six chloroplast intergenic regions (3700 nt in total) were sequenced before analyzing the Southern-RFLP (Restriction Fragment Length Polymorphism) of 2 mt gene flanking regions. Only two different chloroplast haplotypes among 15 A. vesiculosa accessions from Africa, Australia, Europe, and Japan were found, generally distinguishing European and non-European plants, with two exceptions. Genetic variation observed in A. vesiculosa appears to be even lower than in other aquatic species with a similar world-wide distribution. A recent bottleneck followed by long-distance dispersal by water birds or low mutation rates could be responsible for the observed genetic uniformity. Estimation of genetic distances based on six chloroplast intergenic regions led to the conclusion that the chloroplast genome of A. vesiculosa matches more closely to that of Drosera regia than Dionaea muscipula, a sister genus sharing snapping traps. The inconsistency between genetic distance estimates based on nuclear and cytoplasmic markers may reflect a chloroplast capture. In A. vesiculosa, a four amino acids substitution (TGWS) in the amino acid sequence of ATP synthase alpha subunit (ATP1), highly conserved mitochondrial protein, was discovered, unique among all organisms based on current knowledge.     

Micropropagation of <Emphasis Type="Italic">Drosophyllum lusitanicum</Emphasis> (Dewy pine), an endangered West Mediterranean endemic insectivorous plant

In this work, in vitro clonal propagation of Drosophyllum lusitanicum (Dewy pine) was obtained from seedlings germinated in vitro. Seeds were collected in various populations identified in the Algarve region and germinated in vitro on MS medium supplemented with 0.5 mg l^–1 BA (6-benzyladenine) and 0.1 mg l^–1 GA₃ (gibberellic acid). The obtained shoots were used in several multiplication assays. The best results were observed in MS medium supplemented with 0.2 or 0.5 mg l^–1 zeatin. The highest rooting frequency (83%) was observed on 1/4MS medium supplemented with 0.2 mg l^–1 IBA (indole-3-butyric acid). Fifty percent of the plantlets were successfully acclimatized to ex vitro conditions, exhibiting normal development. Plans are underway to reintroduce the in vitro produced plants from this study in selected locations in their natural habitat.     

Floral and reproductive biology of Drosophyllum lusitanicum (L.) Link (Droseraceae)

We studied the floral and reproductive biology of Drosophyllym lusitanicum (Droseraceae), a species endemic to the western Iberian Peninsula and northwest Morocco. Flowering lasted from March to August, peaking in April. The species is clearly homogamous, with pollen germination and stigma receptivity occurring even in preanthesis. This was reflected in the high rates of fruit set and seed set in bagged inflorescences. Fructification did not differ significantly between different pollination treatments, although we did find differences between some treatments in the numbers of viable seeds per flower and fruit. However, seed weight did not differ significantly. The importance of self-pollination over cross-pollination was also supported by the low percentages of fructification (24.67%) after emasculation and open pollination. Metameric deviations were seen in flowers, although this abnormality was not translated as any reproductive benefit or disadvantage (fructification, number or weight of seeds) in comparison with normal flowers.     

Effects of competition and litter on a carnivorous plant, Drosera capillaris (Droseraceae)

Although carnivorous plants are widely recognized as being generally restricted to open habitats, tests of the effects of competition on individual performance are extremely rare. In this study, I examined the effects of the removal of herbaceous and shrub canopies on seedling density and growth, survival, and reproduction of phytometers of a small insectivorous plant, Drosera capillaris (pink sundew). I also examined the distribution of this species in relation to the occurrence of woody species in a frequently burned wet savanna in southeastern Mississippi. Killing plants and removing dead biomass increased seedling density in both open areas and shrub thickets. The removal of dead biomass following herbicide application was critical to increasing densities of seedlings. Killing plants with herbicide without also clearing residual litter and standing dead was not sufficient to increase seedling densities in shrub thickets. Although the removal of the groundcover canopy strongly influenced the density of seedlings, it had very little effect on survival, growth, and reproduction of small phytometers during a single growing season. Survival of phytometers was greater in open areas than in shrub thickets, regardless of whether the groundcover canopy was removed. Densities of both seedlings and adults were greater in open areas away from shrub thickets than beneath the woody canopies of thickets and were negatively correlated with the leaf area index of groundcover vegetation. Results of this study show that the establishment of this carnivorous plant species is limited in part by the effects of litter on seedling density in both open areas and shrub thickets.     

Naphthalene glucoside and other phenolics from the shoot and callus cultures of Drosophyllum lusitanicum

The callus and, for the first time established, shoot cultures of Drosophyllum lusitanicum Link. (Droseraceae) yielded new naphthalene glucoside-5-hydroxy-4-methoxy-2-naphthalenecarboxylic acid 5-O-beta-glucoside (drosophylloside) and 5-hydroxy-4-methoxy-2-naphthalenecarboxylic acid methyl ester besides other phenolics like naphthalenes-5-hydroxy-4-methoxy-2-naphthalenecarboxylic acid (ancistronaphthoic acid B), hydroplumbagin 4-O-glucoside, naphthoquinones-plumbagin and 3-chloroplumbagin, C-glycosylflavones- vitexin, isovitexin, orientin and isoorientin. The pattern of phenolics found supports affinity of Drosophyllum to the families-Droseraceae, Ancistrocladaceae and Dioncophyllaceae.     

The homogentisate ring-cleavage pathway in the biosynthesis of acetate-derived naphthoquinones of the droseraceae

Photosynthesis experiments with ¹⁴CO₂ established that of 16 Droseraceae species tested Drosophylum lusitanicum incorporated the highest amount of label into plumbagin (2-methyl-5-hydroxy-1,4-naphthoquinone). Tyrosine-[β-¹⁴C] fed to Drosophyllum was shown to label plumbagin efficiently (20% incorporation). Extensive chemical degradation of the labeled naphthoquinone showed, however, that the incorporation of tyrosine was indirect, the label being distributed throughout the molecule. It was established that plumbagin and the closely related 7-methyljuglone are biosynthesized via the acetate-polymalonate pathway. Tyrosine is broken down to acetate in this tissue via the homogentisate pathway, which was demonstrated by feeding and incorporation of label into plumbagin of intermediates such as homogentisate-[¹⁴C], maleyl- and fumarylacetoacetate-[¹⁴C]. Simultaneous application of tyrosine-[β-¹⁴C] and α,α′-bipyridyl, an inhibitor of the homogentisate oxigenase, led to an accumulation of homogentisate-[¹⁴C] within the tissue. The degradation of tyrosine to acetate by Drosophyllum is not due to epiphytic bacteria since ring cleavage of tyrosine and formation of plumbagin from breakdown products occurred both within sterile grown plants and sterile cell suspension cultures. In tissue kept in darkness, plumbagin undergoes a slow turnover with a half life of about 400 hr.