Comparative capitular morphology and anatomy of Coreopsis L. and bidens L. (compositae), including a review of generic boundaries

The capitular and floral morphology and anatomy ofBidens L. andCoreopsis L. were studied. All the North American species ofCoreopsis were studied. Selected species ofBidens from North and South America andCoreopsis from South America were included. The results were compared with previous observations on African species ofBidens (incl.Coreopsis). Emphasis was given to character states of the ray florets, paleae, stylearm apices, outer phyllaries, achenes, and pollen grains. Some of the character states are unique features ofCoreopsis, e.g., globular and elongately conical receptacles, deltoid outer phyllaries, truncate and indistinctly 3–5-dentate, 3–4-lobed ray florets, narrowly spathulate paleae, subulate paleae with linear-filiform upper half, hairy and apically 3-cleft paleae, truncate, convex or shallowly conical stylearm apices with the sweeping hairs limited to the area above the stigmatic surfaces and the orbicular to circular achenes. The cylindric setaceous pappus bristles so commonly encountered inBidens are unknown inCoreopsis. The pappus bristles inCoreopsis are paleaceous but similar, though thicker ones are also found in African species ofBidens (incl.Coreopsis) with winged achenes. Twin-celled hairs (setulae) with differing degrees of wall thickness are found on the achenes ofCoreopsis sect.Pseudoagarista (Mexico and South America),Coreopsis sect.Pugiopappus (California), AfricanBidens with winged achenes (e.g.,B. prestinaria, B. macroptera) and some North AmericanBidens (e.g.,B. aristosa). Similar sclerotic parenchyma make up the achenial wings of species in both genera. These may be interpreted as homologous structures, indicating the underlying similarity of these taxa and their derivation from a common ancestral stock.     

Genetic components of heterocarpy inMicroseris hybrid B87 (Asteraceae,Lactuceae)

Microseris B87 is derived from a single hybrid specimen betweenM. pygmaea with few, weakly hairy peripheral achenes and aM. bigelovii with many, strongly hairy peripheral achenes. Offspring through the F4 and F5 generations obtained by spontaneous selfing were analyzed for the segregation of quantitative and qualitative characters relating to achene dimorphism. The phenotypic effects of two previously identified unlinked genes determining the relative number of outer achenes are characterized in partially and completely homozygous sublines. We show that two morphological markers genetically linked to one of these genes are themselves regulated by the system inducing heterocarpy. Not more than two more unlinked genes are involved in the genetic basis of the heterocarpic response. The interaction of these genes in determining the heterocarpy phenotypes is discussed in the framework of a model postulating genes for a morphogen gradient across the capitulum and genes responding to this gradient.     

Nest site selection by fire ant queens

Summary We examined nest site selection by foundresses of the polygyne form ofSolenopsis invicta. In the laboratory, foundresses were allowed to nest in control soil vs. soil inoculated with refuse from a colony ofPheidole dentata, a natural enemy. In a second experiment foundresses chose between control andS. invicta soil. More foundresses (100%) avoided the soil of a natural enemy than the soil of a conspecific colony (79%) that may accept them into the nest. Foundresses from this polygyne ant appear to avoid other colonies in general, but avoid heterospecific colonies more.     

Sexual tetraploid and apomictic pentaploid populations ofHieracium pilosella (Compositae)

Five species are recognized inHieracium subgen.Pilosella sect.Pilosellina Fries. Four are diploid (2x, 2n = 18), one (H. pilosella L.) is highly variable morphologically and cytologically (from 2x to 10x), in its mode of reproduction (self-incompatibility, agamospermy, amphimixis, apo-amphimixis) and in its hybridization pattern. A part of this huge agamic complex was analysed by comparing sexual 4x and apomictic 5x plants (crossing and germination experiments, measurements of vegetative reproduction by stolons etc.). In the experimental garden apomictic 5x produced more stolons than the sexual 4x plants and the total length of the stolons per rosette was greater. However, in nature, the competitive potential of the sexual plants seems to be higher, presumably as a result of the higher mortality of ramets in 5x. Sexual 4x plants often grow in dense and grazed grass vegetation, whereas 5x apomicts often occur in dunes with patchy vegetation. Apomicts produce more capitula per rosette, and sexual rosettes form only about 60% of the number of viable achenes as compared to apomictic ones. Therefore, apomicts appear to be characterized by a greater colonizing ability than sexual plants. Apomictic plants produce equal numbers of viable achenes under conditions of both open pollination and isolation. Sexual plants do not form any viable achenes after isolation and produce a somewhat lower percentage of achenes after open pollination than do apomictics. 5xreproduce exclusively apomictically. Apo-amphimixis was never observed in pentaploids and only very rarely in tetraploids. Addition hybrids are very rare. The cross sexual 4x × apomictic 5x failed in 70% of the attempts, but the recombination of genomes carrying genes for apomixis is possible and results in apomictic 4x and sexual 5x, both with a reduced number of viable achenes. In nature sexual and apomictic plants may occur in close proximity. In such cases the germination rate of the achenes of 4x and 5x is lower; this may indicate that apomictic plants fertilize sexual plants in nature (unidirectional gene-flow). 5x plants form euploid gametes carrying two or three genomes. The results of the crossing experiments can be explained in terms ofNogler's theory of monogenic inheritance of apospory.Variation and evolution inHieracium subg.Pilosella sect.Pilosellina I.     

Ant-seed interactions: combined effects of ant and plant species on seed removal patterns

Seed dispersal by ants (i.e. myrmecochory) is usually considered as a mutualism: ants feed on nutritive bodies, called elaiosomes, before rejecting and dispersing seeds in their nest surroundings. While mechanisms of plant dispersal in the field are well documented, the behaviour of the ant partner was rarely investigated in details. Here, we compared in laboratory conditions the foraging behaviour of two ant species, the omnivorous Lasius niger and the insectivorous Myrmica rubra to which seeds of two European myrmecochorous plants (Chelidonium majus and Viola odorata) were given. Ant colonies were simultaneously presented three types of items: entire seeds with elaiosome (SE), seeds without elaiosome (S) and detached elaiosomes (E). The presence of elaiosomes on seeds did not attract workers from a distance since ants first contact equally each type of items. Although ants are mass-recruiting species, we never observed any recruitment nor trail-laying behaviour towards seeds. For ants having contacted seed items, their antennation, manipulation and seed retrieval behaviour strongly varied depending on the species of each partner. Antennation behaviour, followed by a loss of contact, was the most frequent ant-seed interaction and can be considered as a “hesitation” clue. For both plant species, insectivorous Myrmica ants removed items in larger number and at higher speed than Lasius. This fits with the hypothesis of a convergence between odours of elaiosomes and insect preys. For both ant species, the small Chelidonium seeds were retrieved in higher proportion than Viola ones, confirming the hypothesis that ants prefer the higher elaiosome/diaspore-ratio. Thus, in these crossed experiments, the ant-plant pair Myrmica/Chelidonium was the most effective as ants removed quickly almost all items after a few antennations. The presence of an elaiosome body increased the seed removal by ants excepting for Myrmica which retrieved all Chelidonium seeds, even those deprived of their elaiosome. After 24 h, all the retrieved seeds were rejected out of the nest to the refuse piles. In at least half of these rejected items, the elaiosome was discarded by ants. Species-specific patterns and behavioural differences in the dynamics of myrmecochory are discussed at the light of ant ecology. Received 10 September 2007; revised 5 February 2008; accepted 5 March 2008.     

Lipochaeta and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): establishing their natural limits and a synopsis

We restrict the genusLipochaeta to the allopolyploid species of the typical section.Lipochaeta s.str. is interpreted to be the result of an intergeneric hybridization betweenMelanthera and a presently unknown taxon, perhaps of the genusWedelia. Lipochaeta is characterized, in addition to its allopolyploidy (n=26), by having both flavonols and flavones, disk corollas with 4 lobes, achenes tuberculate at maturity, the disk achenes flattened to slightly biconvex, and ray achenes obcompressed.Lipochaeta sect.Aphanopappus andWollastonia are here reduced to synonymy underMelanthera. We transfer 14 HawaiianLipochaeta and one New Caledonian species as well as the AsianWedelia prostrata toMelanthera. These transfers, along with the species in Africa and North America, bring the number of species in the genus to 35.Melanthera is delimited by an abruptly narrowed to truncate and flattened top of the achene, (0−)1–15(−20) often unequal, ciliate or barbellate, caducous pappus bristles immediately surrounding the corolla, involucral bracts and receptacular paleae with many veins forming longitudinal striations, and n=15. The florets are 5-merous, the corollas are yellow or white, and rays are absent (in white-flowered species) or present and neutral or fertile. In dealing with species formerly placed inLipochaeta, the GalapagosL. laricifolia is here transferred from the illegitimate generic nameMacraea toTrigonopterum and the BrazilianL. goyazensis is transferred toAngelphytum. We maintain the earlier reduction ofEchinocephalum underMelanthera and reduce all three taxa originally described in it to one,M. latifolia.     

Comparison between the allergenic airborne pollen in Trieste and at Lozzo di Cadore (Italy) in 1989

Summary A comparative study was carried out between aeroallergenic pollen spectra in Trieste and at Lozzo di Cadore. The two localities were found to be different on a five-day running mean basis as to the 8 pollen taxa monitored by the National Aerobiological Monitoring Network managed by the Italian Association of Aerobiology (AIA). In the zone of Lozzo the pollination curve of the spring flowering taxa showed a lag of about one month due to the colder climate. Lozzo di Cadore showed a poor airborne pollen content,Corylus and Graminaceae being prevailing. Trieste has a higher airborne pollen diversity and longer pollination times because of its higher floristic and vegetational complexity. In Trieste the allergenic pollen spectrum showed great quantities ofOstrya, Quercus, Oleaceae, Graminaceae and of ruderal taxa, widespread over the area, due to man's influence.     

Pappus part number in annual species ofMicroseris (Compositae,Cichoriaceae)

All North American annual species of the genusMicroseris have a five-part pappus, the one South American annual,M. pygmaea, has ten pappus parts. The pappus develops over a constant number of ten provascular bundles with or without inhibition between alternate sites of pappus development. Each natural population contains a predictable proportion of achenes with aberrant pappus part numbers. Hybridization betweenM. bigelovii (5 parts) andM. pygmaea results in F 1 and F 2 plants with many aberrant achenes. In each plant either five or ten can be shown to be the basic number with aberrant numbers following a Poisson distribution for numbers added to 5 or deleted from 10. Occasional plants show no basic number but have a random distribution of numbers about an intermediate mean. The evolutionary genetics of this character is discussed.     

Yeasts and filamentous fungi carried by the gynes of leaf-cutting ants

Insect-associated microbes exhibit a wide range of interactions with their hosts. One example of such interactions is the insect-driven dispersal of microorganisms, which plays an essential role in the ecology of several microbes. To study dispersal of microorganisms by leaf-cutting ants (Formicidae: Attini), we applied culture-dependent methods to identify the filamentous fungi and yeasts found in two different body parts of leaf-cutting ant gynes: the exoskeleton and the infrabuccal pocket. The gynes use the latter structure to store a pellet of the ants’ symbiotic fungus during nest founding. Many filamentous fungi (n = 142) and yeasts (n = 19) were isolated from the gynes’ exoskeleton. In contrast, only seven filamentous fungi and three yeasts isolates were recovered from the infrabuccal pellets, suggesting an efficient mechanism utilized by the gynes to prevent contamination of the symbiotic fungus inoculum. The genus Cladosporium prevailed (78%) among filamentous fungi whereas Aureobasidium, Candida and Cryptococcus prevailed among yeasts associated with gynes. Interestingly, Escovopsis, a specialized fungal pathogen of the leaf-cutting ant-fungus symbiosis, was not isolated from the body parts or from infrabuccal pellets of any gynes sampled. Our results suggest that gynes of the leaf-cutter ants Atta laevigata and A. capiguara do not vertically transmit any particular species of yeasts or filamentous fungi during the foundation of a new nest. Instead, fungi found in association with gynes have a cosmopolitan distribution, suggesting they are probably acquired from the environment and passively dispersed during nest foundation. The possible role of these fungi for the attine ant–microbial symbiosis is discussed.     

Inter- and intraspecific alarm response in the termiteMacrotermes subhyalinus (Rambur)

Summary In laboratory studies we analysed the reaction of aMacrotermes subhyalinus (Rambur) colony in contact with a small group of test termites placed at the head of a gallery system. As test termites we usedMacrotermes subhyalinus of the same nest,Macrotermes subhyalinus of a foreign nest and the related sympatric speciesMacrotermes bellicosus (Smeathman).Contact with an introduced group of foragers evokes a significant recruitment of workers and minor soldiers. The intensity of the response depends on the test groups we introduced. The highest recruitment results withM. bellicosus andM. subhyalinus from a foreign nest. During the recruitment the increase in the number of minor soldiers is greater than the increase in the number of workers, and as a result, the relative proportion of the soldiers in the population leaving the nest is significantly higher than before.Direct observations at the place of contact show thatM. bellicosus individuals are immediately attacked and killed. In the case ofM. subhyalinus from a foreign nest, not all individuals are killed; some are only intensively examined with the antennae or seized with the mandibles.M. subhyalinus from the same nest are rarely attacked. Mostly they are antennally inspected or enter the gallery system unhindered.The recruitment of termites from the nest is released by major workers which, after contact with the test termites, run back into the nest and provoke an alarm by tactile stimuli. On their way back into the nest the major workers, in all probability, lay a pheromone trail which leads the outgoing alarmed termites towards the place of contact.     

Site selection by a social wasp in a nesting association (Hymenoptera: Vespidae)

The tropical social waspMischocyttarus immarginatus is a nesting associate ofPolybia occidentalis in northwestern Costa Rica. We show that an activeP. occidentalis colony is a necessary component of aM. immarginatus nesting site in this part of the latter’s range.M. immarginatus foundresses actively seekP. occidentalis colonies to initiate nests at a modal distance of 8 cm from them. There is an 18-mm lower limit to how closeM. immarginatus can nest in relation toP. occidentalis. At distances of less than this,P. occidentalis workers can reach theM. immarginatus nest while standing on their own nest and robM. immarginatus of its brood when the adults are absent.     

Overcoming seed dormancy in ex situ plant germplasm conservation programmes; an example in the endemic Argyranthemum (Asteraceae: Anthemideae) species from the Canary Islands

Germplasm of 21 diverse Argyranthemum taxa was collected from contrasting ecological zones in the Canary Islands. Seed dormancy was considerable in the majority of taxa. Extensive investigations, based on a germination test procedure algorithm for Asteraceae, with achenes from ray and disc florets of five contrasting taxa identified a procedure to promote full (85%) germination of the seeds from both ray and disc florets of all five taxa; viz, excision of the seeds from the achenes, followed by testing at 15°C with 2.6×10^-3 m GA₃ co-applied. Subsequent tests showed that this regime was effective in promoting full germination in seeds from both ray and disc florets of the remaining 16 taxa. The results are discussed in the context of ex situ plant germplasm conservation.     

Effects of pollen donors on seed formation inEspeletia schultzii (Compositae) populations at different altitudes

The influence of different pollen donors on seed formation was investigated in three populations ofEspeletia schultzii that differ in environmental conditions and life history characteristics. Self pollen and pollen from different donors (< 15m apart) within each population was used in a diallel design in order to test the genetic base of seed set variation. Three measures of seed formation were used: (1) achene number; (2) proportion of filled achenes (fruits) that distinguishes between achenes with seeds and empty achenes; (3) proportion of aborted seeds that distinguishes between viable and aborted seeds. Self-pollinations resulted in empty achenes. Achene number did not vary between the different pollen donors. A bimodal pattern of filled achenes was found in two populations in two consecutive years. On the other hand, a unimodal pattern was found in crosses between more distant donors (> 30m). These patterns seems to be the results of a sporophytic incompatibility system. Seed abortion was highest at the higher elevations and seems to be correlated with elevation rather than with any genetic effect.     

Social parasitism of <Emphasis Type="Italic">Polistes dominulus</Emphasis> by <Emphasis Type="Italic">Polistes nimphus</Emphasis> (Hymenoptera,Vespidae)

Summary Interspecific facultative social parasitism is well known in ants and in bumble-bees, but it is rarer in wasps. This form of parasitism is traditionally considered to be an intermediate stage in the evolution of obligate interspecific parasitism, where the parasites are no longer able to nest alone. We report field and experimental observations of a newly discovered facultative parasitic relationship between two closely related free-living Polistes species: P. nimphus and P. dominulus. P. nimphus foundresses sometimes usurp the nests of the larger P. dominulus before worker emergence. The invading queen takes over the nest with abundant abdomen stroking on the nest surface and is accepted by workers if they emerge 6 or more days after usurpation. Morphometric comparisons show that the usurper species, though smaller than its victims, has morphological adaptations consisting of larger heads, mandibles and front femora relative to their body size that may give it an advantage during nest invasion. This strategy is likely to be taken only after the foundress loses her original nest because invading P. nimphus queens have lower reproductive success than they would have had on their own nest. Overall, we found that P. nimphus usurpers use strategies of invasion similar to those of two obligate parasites, suggesting that this may be an example of one of the pathways by which social parasitism evolved.Received 4 April 2003; revised 8 August 2003; accepted 14 September 2003.     

Differences in polypeptide expression in thermoinhibited and germinating achenes of Tagetes minuta L.

When imbibed at their optimum germination temperature of25°C, achenes of Tagetes minutaL.germinate over a period of 48 h. At a temperature of36°C, the achenes become thermoinhibited and do notgerminate. This elevated temperature does not harm the achenes, however, and assoon as the temperature is reduced below 35°C,germination occurs, but is spread over a reduced period of only 24h. The polypeptide patterns of germinating and thermoinhibitedachenes were compared, to test whether thermoinhibition is the result ofdifferential gene expression or repression. Ten polypeptides which weredifferentially expressed in thermoinhibited achenes were observed. Thesepolypeptides were all extremely specific to thermoinhibited achenes anddeclinedrapidly when the incubation temperature was decreased, in a manner whichcorrelated with the increase in germinability of the achenes. The tightlyregulated nature of the expression of these polypeptides suggests thatthermoinhibition in T. minuta may be under positivecontrol. These proteins may therefore result in the prevention of radicleemergence at unfavourable temperatures.     

Phylogenetic relationships of the monotypic Peruvian genus Laccopetalum (Ranunculaceae)

Laccopetalum giganteum (Ranunculaceae) is a rare endemic buttercup from the montane regions of the Peruvian Andes. The systematic position of this monotypic genus within Ranunculaceae was investigated using cpDNA matK sequence data. Our findings indicate that L. giganteum forms a highly supported clade together with Krapfia. Several morphological characters are shared by these genera; large subglobose and fleshy flowers, presence of androgynophore with a separated staminal and carpellate region and many tiny achenes. The Laccopetalum-Krapfia clade is sister to the core Ranunculus group with a high bootstrap support. The number of sepals and similar characteristics of achene morphology support an affinity of Laccopetalum with Ranunculus s.s tr.     

The protective function of the compact silk nest of social Stegodyphus spiders (Araneae,Eresidae)

Summary This study investigates the suggested protective function of the compact silken nest of two species of social Stegodyphus spiders (S. dumicola and S. mimosarum), as a possible ultimate factor for their sociogenesis. Being inhabitants of African dry thornbush country, these spiders are endangered by overheating and desiccation. In the laboratory, both species were found to avoid temperatures >40° C. In the field, temperature in the nest between 7:00 and 21:00 h tends to be higher than air-temperature outside, and between 13:00 and 15:00 h even tends to exceed 40° C. The nest thus is of no value in temperature regulation. Nor is it effective in protecting the spiders from desiccation: Both species have a body water content of 65.2±3.8% and are extremely desiccation resistant; they survived 9 days at 37°C in an exciccator with P₂O₅, with a daily average % liveweight reduction of 4.94(±1.1). In the field, relative humidity inside the nest between 11:00 and 19:00 h tends to be lower than that of the air outside. Inspection of burnt areas showed that the nest does not protect the spiders from bushfire. The nest does protect from wind, hail, and direct sun-radiation; but available natural retreats, or a simple silken shield, or the funnel-shaped silk tube inhabited by solitary Stegodyphus, have the same effect.     

Variability of the inflorescence ofMicroseris laciniata (Compositae: Cichorieae)

Quantitative characters of the flowering head of a garden population ofMicroseris laciniata were scored during the second, third, and fourth season of growth. Number of achenes per head, number of phyllaries per head and the average number of pappus parts per achene in single heads show significant plant to plant variation. Achenes per head and pappus parts per achene were scored in identical plants in two subsequent seasons. The number of pappus parts per achene varies freely between five and ten. This contrasts with annual species ofMicroseris in which either five or ten pappus parts are found, depending on the species. In spite of a clear plant-specific average of pappus parts, both high and low pappus part determination can be demonstrated in all specimens. The number of pappus parts depends on the position of an achene on the receptacle, marginal achenes usually having fewer pappus parts than central ones. This gradient is not closely correlated with the position of an achene on the genetic spiral.     

How plants may benefit from their consumers: leaf-cutting ants indirectly improve anti-herbivore defenses in <Emphasis Type="Italic">Carduus nutans</Emphasis> L

Although herbivores often have a negative impact on plant fitness, sometimes plants may benefit from their consumers. However, these positive interactions usually occur as a result of plant damage (e.g., overcompensation, defense induction). I present evidence of a novel way by which plants may benefit from their consumers without being eaten. Plants of Carduus nutans increased their physical defenses when grown in external refuse dumps of the leaf-cutting ant Acromyrmex lobicornis. Seedlings planted in refuse exhibited longer spines and tougher leaves than those planted in control soils. Pick-up assays with entire leaves and leaf discs demonstrated that these enhanced physical defenses prevented leaf-cutting ant harvest. Additionally, plants established in refuse dumps showed fewer insect herbivory than those in non-nest soils. The nutrient-rich refuse dump appeared to reduce the stage at which leaves are tender and thus more vulnerable to herbivory. This is the first case where plants may benefit from insect herbivores via waste products without the cost of being eaten. This illustrates how plants may plastically respond to reliable cues of the risk of herbivory.