Pollination biology and floral scent chemistry of the Neotropical chiropterophilous Parkia pendula

During the past several decades, the pollination biology of Old World plant species pollinated by flying foxes and of New World plants pollinated by highly specialized nectar-feeding glossophagine bats has been studied in detail. However, little is known about Neotropical plants that are pollinated by less specialized phyllostomid bats. Therefore, we studied the pollination biology of Parkia pendula , a tree pollinated by Phyllostomus . Flowers of P. pendula are arranged in capitula, and a capitulum is composed of approximately 800 hermaphrodite flowers and 260 sterile flowers. The sterile flowers produced a total of 7.4 ml nectar per night, with a sugar concentration of 14.95%, and proline as the dominant amino acid. Nectar production is highest at dusk and ends at 03:00 h. The floral scent is dominated by monoterpenoids (97.9%), with ( E )-β-ocimene being the dominant (84.0%) compound. No sulfur compounds were detected. The capitula are heavily visited by four species of phyllostomid bats, of which Phyllostomus discolor is the most abundant (98.9%). Nectar production per capitulum is within the reported range of nectar produced by this pantropical genus (5.0–8.0 ml). This genus-wide range seems to be optimal for attracting non-specialized nectar-feeding bats and forces them to visit capitula of several trees to satisfy their dietary needs, thus increasing the probability of cross-pollination for this plant.     

Predation by a snake of a flower-visiting bat at parkia nitida (Leguminosae: Mimosoideae)

An observation of a snake catching a bat visiting a capitulum of the chiropterophilous treeParkia nitida is described. The implications of predator avoidance on the behavior of pollinators and hence on cross-pollination are discussed. Fear of predation may be the reason why some bats make only brief visits toParkia capitula.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

Relationship between capitulum size and pre-dispersal seed predation by insect larvae in common Asteraceae

The evolution of a showy floral display as an advertisement to pollinators could simultaneously advertise the availability of resources to pre-dispersal seed-predators. The hypotheses tested here are that the incidence of seed predation by bud-infesting insect larvae in capitula of Asteraceae is positively related to (1) capitulum size among species, (2) capitulum size within species, (3) capitulum lifespan, and (4) the degree of flowering asynchrony on individual plants. Three populations of each of 20 common herbaceous species of Asteraceae from disturbed ground and grassland habitats were monitored for the presence of pre-dispersal, seed-eating insect larvae. Mean capitulum size (receptacle width) of each species was measured. In a sub-set of eight species, individual capitula were tagged to determine their flowering phenology and lifespan (from anthesis to seed shedding). From these data an index of flowering synchrony on individual plants was derived. Among species, the incidence of larval infestation increased with capitulum size. Small-flowered species such as Achillea millefolium were largely free of bud-infesting larvae, whilst large-flowered species such as Arctium minus were heavily infested. In three cases investigated in greater detail, bud infestation was found to increase with capitulum size within species, suggesting a potential for natural selection to favour smaller capitula. No relationship was found between infestation levels and either capitulum lifespan or degree of flowering synchrony, and there was no evidence that the relationship between capitulum size and infestation was confounded by correlations with these other features. The results support hypotheses 1 and 2, but not 3 and 4. It is suggested that the characteristic capitulum size of each species may represent a trade-off between the opposing selection pressures of pollinators and pre-dispersal seed predators.     

The antiquity of Madagascar’s grasslands and the rise of C4 grassy biomes

Aim Grasslands and savannas, which make up > 75% of Madagascar’s land area, have long been viewed as anthropogenically derived after people settled on the island c. 2 ka. We investigated this hypothesis and an alternative – that the grasslands are an insular example of the post‐Miocene spread of C₄ grassy biomes world‐wide. Location Madagascar, southern Africa, East Africa. Methods We compared the number of C₄ grass genera in Madagascar with that in southern and south‐central African floras. If the grasslands are recent we would expect to find fewer species and genera in Madagascar relative to Africa and for these species and genera to have very wide distribution ranges in Madagascar. Secondly, we searched Madagascan floras for the presence of endemic plant species or genera restricted to grasslands. We also searched for evidence of a grassland specialist fauna with species endemic to Madagascar. Plant and animal species endemic to C₄ grassy biomes would not be expected if these are of recent origin. Results Madagascar has c. 88 C₄ grass genera, including six endemic genera. Excluding African genera with only one or two species, Madagascar has 86.6% of southern Africa’s and 89.4% of south‐central Africa’s grass genera. C₄ grass species make up c. 4% of the flora of both Madagascar and southern Africa and species : genus ratios are similar (4.3 and 5.1, respectively). Turnover of grasses along geographical gradients follows similar patterns to those in South Africa, with Andropogoneae dominating in mesic biomes and Chlorideae in semi‐arid grassy biomes. At least 16 monocot genera have grassland members, many of which are endemic to Madagascar. Woody species in frequently burnt savannas include both Madagascan endemics and African species. A different woody flora, mostly endemic, occurs in less frequently burnt grasslands in the central highlands, filling a similar successional niche to montane C₄ grasslands in Africa. Diverse vertebrate and invertebrate lineages have grassland specialists, including many endemic to Madagascar (e.g. termites, ants, lizards, snakes, birds and mammals). Grassland use of the extinct fauna is poorly known but carbon isotope analysis indicates that a hippo, two giant tortoises and one extinct lemur ate C₄ or CAM (crassulacean acid metabolism) plants. Main conclusions The diversity of C₄ grass lineages in Madagascar relative to that in Africa, and the presence of plant and animal species endemic to Madagascan grassy biomes, does not fit the view that these grasslands are anthropogenically derived. We suggest that grasslands invaded Madagascar after the late Miocene, part of the world‐wide expansion of C₄ grassy biomes. Madagascar provides an interesting test case for biogeographical analysis of how these novel biomes assembled, and the sources of the flora and fauna that now occupy them. A necessary part of such an analysis would be to establish the pre‐settlement extent of the C₄ grassy biomes. Carbon isotope analysis of soil organic matter would be a feasible method for doing this.     

The capitula of phasmid eggs: an update with a review of the current state of phasmid ootaxonomy

The relationship between egg shape and presence or absence of a capitulum is considered. The distribution of capitula of different types within the order Phasmida is reviewed. These tend to confirm tribal and subfamily groupings but suggest that the suborder Anareolatae is polyphyletic and some suggestions are made for its regrouping. The species of Phasmida which have been recently studied from an ootaxonomic viewpoint are surveyed.     

Variation in floral sex allocation, female success, and seed predation within racemiform synflorescence in the gynomonoecious Ligularia virgaurea (Asteraceae)

Studies of diclinous species have showed that floral sex allocation and female reproductive success were quite variable within inflorescences. However, little attention has been paid to gynomonoecious species, in which individuals produce both female and bisexual flowers. The purpose of this study was to investigate the variations in reproductive patterns at different capitulum positions within racemiform synflorescence in Ligularia virgaurea, and to determine selective mechanisms of variations in reproductive patterns. We conducted observational and experimental studies in natural populations of the gynomonoecious composite L. virgaurea. Floral sex allocation, seed production and pre-dispersal seed predation were quantified in the field. The results showed several patterns of variation from top to bottom capitula, including an increase in bisexual flowers and flower number per capitulum, but a decrease in seed set and size. Removing earlier capitula during bud stage did not change floral sex allocation in later capitula. And no effect was found on seed set under supplemental pollination. Thus, although it has been reported many times in previous studies, the variation of floral sex allocation in L. virgaurea may not result from architectural effect or mating environment, and the variation of seed production could not be fully explained by pollination success. Additionally, our results showed that L. virgaurea was susceptible to high levels of bisexual biased predation, which was greater for top capitula. We therefore suggest that these variations may help to enhance reproductive success of L. virgaurea in the face of bisexual-biased seed predation.     

Diversity and spacio‐temporal distribution of mushrooms in a Nigerian savanna: implication for their conservation

We surveyed the macrofungi of an area dominated by northern Guinea savanna habitats in north‐east Nigeria. A total of 93 different species of mushrooms were found in the study area. These species belong to 29 different families, most species belonging to the family Agaricaceae, Lyophyllaceae, Bolbitiaceae, Pluteaceae and Polyporaceae. A total of 48 species belonged to the Agaricaceae, followed by Lyophyllaceae represented by 6 species. The micro‐habitats of mushrooms in the study area include wood, soil around dead tree stump, waste dump, cow dung, fallow, arable land, and 22 different living trees species. The most important habitats in term of mushroom species diversity were Parkia biglobosa, Tamarindus indica and dead wood. The Parkia biglobosa tree had the highest species richness (45) and species diversity (Shannon diversity index, SDI: 3.6). Tamarindus indica was the second, having 28 species richness and 2.7 SDI. These were followed by dead wood where 22 different mushroom species were recorded and has of 2.1 SDI. Also, these three habitats (Parkia biglobosa, dead wood and Tamarindus indica) had the highest number of mushroom species (14, 8 and 6 respectively) that were confined to them. Most of the other mushroom species were confined to only one microhabitat. There was no statistically significant difference in mushroom abundance between arable land and fallow. Collectively, the indigenous trees were associated by more mushroom species (63) than the exotic trees (20) and the indigenous trees also had significantly higher mushroom abundance than the exotic trees. However, the non‐indegenous trees had more species forming mycorrhiza than the indegenous trees, presumaby because the former were able to form symbioses with both native fungi and introduced species from their origina habitats. Inter‐annual variation of mushroom species diversity was also observed. Human activities contribute positively to mushroom diversity in the study area by creating some unique micro‐habitats that support the growth of certain unique mushroom species. Therefore, for maximum conservation of mushroom diversity, conservation measures excuding all human activities should be avoided.     

FINE STRUCTURE OF MISTLETOE POLLEN. V. MADAGASCAN AND CONTINENTAL AFRICAN VISCUM L. (VISCACEAE)

Pollen characters of Madagascan and continental African Viscum are described and compared to those in Asia and Australia. The subprolate, tricolporate, nonuniformly sculptured pollen of Madagascan taxa is most similar to that of Asian species. Ultrastructurally, however, the completely granular equatorial ektexine of Madagascan Viscum is most similar to that of continental African taxa. Continental African Viscum, in contrast to Madagascan and Asian species, display a wide variation in pollen shape and apertures. Pollen shape ranges from subprolate to oblate, the latter unique to Africa. The most striking feature of continental Viscum is their variability in aperture number and aperture type. Aperture number varies at both the intra- and interpopulational levels with such variation resolvable to the individual flower—a condition unique to the continent. The only simple (colpate) aperture type in the genus is restricted to Africa. The continental species can be divided into two species groups based on pollen characters: Group I (4 spp.) characterized by strictly tricolporate rounded convex pollen with a rodlet/granular equatorial ektexine structure and Group II (most continental species) possessing multiapertures, concave lobate shape, uniform sculpturing and granular equatorial ektexine. The African V. menyharthii, V. fischeri, V. rotundifolium and V. minimum exhibit no clear Group I or II affinities. An analysis of overall pollen characters in Viscum indicates a trend towards spheroidal shape, multiapertures and uniform sculpturing and ektexine organization. Though pollen characters suggest ties between Australia, Asia and Madagascar, they indicate an even stronger relationship between Madagascar and continental Africa, particularly eastern Africa. The relationship of the majority of African Viscum, excluding those with obvious Madagascan affinities, remains obscure. The unique palynological features of Group II species coupled with their inflorescence structure suggest an independently evolving group.     

Cleistogamous capitulum in Centaurea melitensis (Asteraceae): heterochronic origin

Cleistogamous capitula formed by Centaurea melitensis display a number of morphological and functional changes with respect to chasmogamous capitula that ensure self-fertilization. Because no studies have hitherto addressed the evolution of cleistogamy in Asteraceae, it was considered useful to ascertain whether these changes are attributable to one or more of the heterochronic processes reported in the literature. Bivariate allometric analyses were performed, and changes were represented graphically using Gould's clock models for size, shape, and age of several capitulum and floret structures. Results suggest that the partially paedomorphic appearance of cleistogamous with respect to chasmogamous capitula is attributable to three processes: (1) early onset of floral development (predisplacement), (2) decreased growth rate of the whorls studied (except gynoecium width) and (3) early offset time (progenesis). The latter appears to play the most significant role in the origin of the cleistogamous capitulum.     

Partial self‐incompatibility in the polyploid endemic species Scalesia affinis (Asteraceae) from the Galápagos: remnants of a self‐incompatibility system?

Different pollination treatments of capitula were used to examine the breeding system of individuals of the tetraploid endemic species Scalesia affinis from the Galápagos Islands. All types of crossings resulted in approximately 35 achenes per capitulum, but in actively and passively self-pollinated capitula these were mostly without embryos. Among self-pollinated individuals a large variation was found in the production of embryos. Some individuals failed completely to set filled achenes while others produced few or many. Additionally, we found a clear difference in female phenology of florets in self-pollinated capitula compared with florets in cross-pollinated capitula. Female florets in self-pollinated capitula remained receptive for longer time. These results suggest that Scalesia affinis is partly self-incompatible. Outcrossing is assured by the endemic carpenter bee, Xylocopa darwini , which proved to be an important pollinator of Scalesia affinis . Nevertheless, geitonogamy seemed to be considerable in the focal population, for which reason open pollinated capitula produced fewer achenes than controlled outcrossed capitula.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 93–101.     

Taxonomic notes on the genus Otoptera (Phaseoleae, Fabaceae) in southern Africa

A taxonomic study of the genus Otoptera (tribe Phaseoleae) is presented. This distinct genus consists of two species, O. burchellii in southern Africa (Namibia, Botswana, South Africa and Zimbabwe) and O. madagascariensis endemic to Madagascar. The two species differ in the shape and size of leaflets, size and orientation of the spur located at the base of the wing petal, and also by size of the pod. A key to the two species is provided. This study focuses mainly on the southern African O. burchellii DC., which is described and illustrated. The species name is lectotypified and the known geographical distribution is recorded for the first time.     

影响黄帚橐吾种子生产的因素Ⅰ.生境和花序结构

 通过对黄帚橐吾(Ligularia virgaurea)自然种群在不同生境和种群密度下的种子生产的研究,结果显示：1）黄帚橐吾的种子生产无论是在种群水平,还是在总花序上同一部位头状花序水平,均受到生境条件的影响,其种子数量、种子均重和种子总重量等特征在不同生境间表现出沙地＞滩地＞坡地,在同一生境内则为高密度种群＞低密度种群和植被盖度较低＞植被盖度较高的趋势（滩地例外）;2）种子生产中的种子数量和种子总重与总花序大小、头状花序数量以及无性系株大小多呈显著正相关,与个体密度无关,但种子均重与上述因素（除无性系株大小外）的关系不显著;3）在总花序内不同部位头状花序间表现出的种子生产变化趋势（顶部＞基部）不受生境条件、种群密度等因素的影响; 4）种子生产在总花序上所表现出的这种部位依赖性可能是由于不同部位头状花序间开花时序上的差异引起种子从母体获取资源上的不同所致。     

Low local but high beta diversity of tropical forest dung beetles in Madagascar

Aims We have compared local (alpha) and regional (beta) species diversities of dung beetles in wet forests in the main tropical regions including Madagascar. Madagascar is exceptional in lacking native large herbivorous mammals which produce the key resource for dung beetles elsewhere. Location Central and South America, mainland Africa, Madagascar and Southeast Asia. Methods Trapping data on dung beetles and data on mammalian faunas were obtained from published and unpublished studies. We used our original data for Madagascar. Results Species richness of dung beetles and that of large‐bodied (> 15 mm length) species in particular were highly significantly explained by the regional number of large‐bodied (> 10 kg) mammals (R² from 50 to 80%). For a given pairwise spatial distance between two communities, beta diversity was significantly higher in Madagascar than elsewhere, explaining the very high total species richness in Madagascar in spite of low local diversity. Main conclusion The presence and numbers of large herbivorous mammals greatly influence the species richness of dung beetles in tropical wet forests. The lack of native large herbivores rather than a limited species pool explains the low local diversity in Madagascar. Exceptionally high beta diversity in Madagascar suggests a pattern of old radiation involving extensive allopatric speciation.     

Plant–pollinator interactions: comparison between an invasive and a native congeneric species

Plant–pollinator interactions determine reproductive success for animal-pollinated species and, in the case of invasive plants, they are supposed to play an important role in invasive success. We compared the invasive Senecio inaequidens to its native congener S. jacobaea in terms of interactions with pollinators. Visitor guild, visitation rate, and seed set were compared over 3 years in three sites in Belgium. Floral display (capitula number and arrangement) and phenology were quantified, and visiting insects were individually censused, i.e. number of visited capitula and time per visited capitulum. As expected from capitula resemblance, visitor guilds of both species were very similar (proportional similarity = 0.94). Senecio inaequidens was visited by 33 species, versus 36 for S. jacobaea. For both species, main visitors were Diptera, especially Syrphidae, and Hymenoptera. Visitation rate averaged 0.13 visitor per capitulum per 10 min for S. inaequidens against 0.08 for S. jacobaea. However, insects visited more capitula per plant on S. jacobaea, due to high capitula density (886 m⁻² versus 206 m⁻² for S. inaequidens), which is likely to increase self-pollen deposition considerably. Seed set of S. jacobaea was lower than that of S. inaequidens. We suggest that floral display is the major factor explaining the differences in insect visitation and seed set between the two Senecio species.     

Factors Contributing to Variation in Seed Production among Remnant Populations of the Endangered Daisy Gerbera aurantiaca1

The species‐rich mist‐belt grasslands of southern Africa have been severely reduced in extent as a result of commercial afforestation, thus confining many endemic plants and animals to small populations in habitat fragments. We investigated the influence of various environmental factors on seed production and seedling recruitment in remnant populations of the endangered grassland herb Gerbera aurantiaca (Asteraceae). Experiments with color traps showed that Eriesthis beetles, which appear to be the primary pollinators of G. aurantiaca, were most abundant in the two largest y extant populations. Seeds are produced in a very small proportion (typically <10%) of the ca 80 female florets in a capitulum. The mean number of seeds produced in undamaged capitula was found to be significantly lower in small than in large populations. Pollen limitation was evident from a significant overall increase in seed set after supplementalpollination in three populations over two seasons. The proportion of capitula containing seed predators did not differ markedly among populations or years, but lepidopteran larvae, which destroy all of the seeds in a capitulum, weremost abundant in the two largest populations in 2003. The presence of juvenile plants varied markedly amongpopulations, but this could not be linked firmly to estimates of seed production. Clonal growth is likely to contribute to the persistence of small isolated populations of G. aurantiaca, even when seed production is severely compromised by pollen limitation and predation.     

Role of within-individual variation in capitulum size and achene mass in the adaptation of the annual Centaurea eriophora to varying water supply in a Mediterranean environment

To clarify the adaptive value of variation in capitulum size and achene mass, plants of Centaurea eriophora were studied in a glasshouse and in a natural population. C. eriophora plants consist of a basal leaf rosette from which an erect stem grows, with lateral branches of various orders ending in capitula of different orders. Primary, secondary and tertiary capitula are comparable in size and they produce similar numbers of achenes, which are similar in weight (large achenes). These capitula are formed during April, May and early June, and constitute the normal or primary flowering. Following ripening of tertiary capitula, leaves senesce, but, later during June and the first half of July, a secondary flowering of a variable number of smaller capitula may occur if wet conditions persist for longer than usual. Plants that have almost senesced develop small lateral branches 1-2 cm long bearing a few small leaves and ending in a capitulum about half the diameter of capitula from the primary flowering period. The number of achenes produced in these capitula (small achenes) and their weight are 70 and 30% less, respectively, than those of capitula formed during primary flowering. These reductions appear to result from restricted availability of resources. Large and small achenes have similar dispersal characteristics and possess similar germination potential. However, large achenes produce seedlings that are capable of emerging from greater burial depths, providing the resulting plants with a potential advantage. The normal flowering period coincides with the optimum time of year for flowering and fruiting in the south of Spain, and only if rainfall lasts longer than usual does secondary flowering occur. Secondary flowering extends the normal flowering and fruiting periods, thereby providing a supplementary crop of smaller, yet viable, fruits. It can be considered to be an adaptive response to the unpredictable Mediterranean climate, optimizing the use of available resources.     

The biogeography of Miniopterus bats (Chiroptera: Miniopteridae) from the Comoro Archipelago inferred from mitochondrial DNA

The endemic fauna of the Comoro Archipelago is composed of a mixture of taxa originating from Africa and Madagascar. Bats are the only native land dwelling mammals on this archipelago, but the biogeographical origins for the vast majority of species within this group are ambiguous. We report here genetic analyses based on two mitochondrial DNA markers to infer the origin of Comorian bats belonging to a reputed species complex of Miniopterus that is further distributed across Africa and Madagascar. Phylogenetic reconstructions show that east African M. minor are not closely related to the insular Miniopterus of Madagascar and the Comoros (Grande Comore and Anjouan). The latter cluster into two distinct, monophyletic clades (Clade 1 and Clade 2). Representatives of these clades occur sympatrically both on the Comoros and on Madagascar, and are distinguished by a large genetic distance (K2P: 9.9% for cytochrome b). No haplotypes are shared between any islands, suggesting the absence of contemporary gene flow. Populations of the widespread Clade 1 are furthermore characterized by a significant inter‐island structure (Φ_CT = 0.249), and by high haplotype and nucleotide diversities (h = 0.90–0.98, π = 0.04–0.06). Demographic analyses of Clade 1 suggest secondary contact between two distinct phylogroups (Subclade 1 A and 1B) that reached Grande Comore and Anjouan, and a large, stable population with a long evolutionary history on Madagascar. These results and the current distribution of related lineages suggest that the Comoros were colonized independently at least two or three times by ancestors from Madagascar.