Batesian mimicry in the non-rewarding orchid Disa pulchra, and its consequences for pollinator behaviour

The non-rewarding flowers of the South African orchid Disa pulchra are remarkably similar in morphology and spectral reflectance to the flowers of a sympatric nectar-producing iris, Watsonia lepida. Field observations indicated that both D. pulchra and W. lepida are pollinated by the long-tongued fly, Philoliche aethiopica (Tabanidae). The hypothesis that D. pulchra is a floral mimic of W. lepida was supported by choice experiments, which showed that, in terms of visits to inflorescences, flies do not discriminate between the two species. However, flies probed fewer flowers and spent less time on D. pulchra inflorescences than on Watsonia inflorescences. Thus the absence of nectar in the flowers of D. pulchra may reduce the risk of geitonogamy and pollen discounting. A breeding system experiment showed that fruits of D. pulchra that arise from self-pollination contain relatively few viable seeds compared with outcrossed fruits. These findings support the hypothesis that deception in orchids evolved under selection for more efficient mating systems.     

Structure and species richness of low-diversity floodplain forest on the Rio Tapajós, Eastern Amazonia, Brazil

Rivers in the Amazonian region have annual water level fluctuations varying from 6 to 20m, and the flooding period ranges from 50 to 270 days between the rising and falling phases. Along the rivers there are many different types of vegetation which are annually flooded. In order to study the variation of the species composition and structure of a forest that is seasonally flooded by a clear water river in Brazilian Amazonia 20km SW of Santarém, Pará State, Brazil, we surveyed three hectares, each divided in ten randomly placed rectangular quadrats (10×50m). The number of species per hectare ranged from 21 to 30. The number of trees varied from 242 to 741. Total basal area ranged from 0.70 to 10.9m2. Leguminosae was the most abundant and dominant family in the three hectares sampled. Couepia paraensis (Chrysobalanaceae), Campsiandra laurifolia (Leguminosae) and Leopoldinia pulchra (Palmae) were the most abundant species in the hectares sampled. Similarity indexes at species level varied from 46 to 60% between the three hectares sampled. The hectares studied have the lowest species richness observed in igapó floodplain forest of the Brazilian Amazonia. This may be correlated with the water type, soil condition and proximity of savanna.     

Frugivory and seed dispersal by chelonians: a review and synthesis

In recent years, it has become clear that frugivory and seed dispersal (FSD) by turtles and tortoises is much more common than previously thought. We here review published and unpublished records of chelonian FSD, and assess the role of chelonians as seed dispersers, from individual species to the community level. We first discuss the distribution of chelonian FSD and the characteristics of the fruit and/or seed species eaten and dispersed by chelonians. We then use the seed dispersal efficiency framework to explore the quantitative and qualitative components of seed dispersal by tortoises and turtles, embarking on a journey from when the fruits and/or seeds are consumed, to when and where they are deposited, and assess how efficient chelonians are as seed dispersers. We finally discuss chelonian FSD in the context of communities and of chelonians as megafauna. A substantial proportion of the world's aquatic and terrestrial turtles and a major part of testudinid tortoises (71 species in 12 families) include fruits and/or seeds in their diet; fruits of at least 588 plant species in 121 families are ingested and/or dispersed by chelonians. For some chelonians, overall or in certain seasons, fruit may even form the largest part of their diet. Contrary to seed dispersal by lizards, the other major reptilian frugivores, chelonian FSD is not an island phenomenon in terms of geographic distribution. Nevertheless, on islands tortoises are often among the largest native terrestrial vertebrates – or were until humans arrived. We synthesise our knowledge of chelonian FSD, and discuss the relevance of our findings for conservation and restoration, especially in relation to rewilding with large and giant tortoises.     

Cryptic clades, fruit wall morphology and biology of Andira (Leguminosae: Papilionoideae)

Andira comprises 29 species distributed throughout tropical America, with two subspecies in Africa. Its fruits are unusual for a papilionoid legume because they are drupes. The majority of species have fruits dispersed by bats, but eight species have larger fruits dispersed by-rodents. Some fruits of both dispersal types are secondarily dispersed by water. Cladistic analysis of chloroplast DNA (cpDNA) restriction site characters discovered four well-supported clades of Andira species. None of these 'cryptic' clades had been recognized by previous workers, because they are not apparently marked by any known morphological innovations. This prompted a search for new characters that might support these groupings. An anatomical study of fruit walls of 25 Andira species revealed the presence of three principal endocarp types, dominated by (1) parenchyma, (2) fibres, or (3) stone cells. These features arc best coded as a single unordered multistate character. When incorporated into a simultaneous cladistic analysis of previously gathered molecular and morphological data, states of this endocarp character are shown to be apomorphies for two of the well-supported clades evident in the cpDNA restriction site data. The most likely plesiomorphic state for the endocarp is parenchyma-dominated. Thicker, stronger endocarps of fibres and stone cells may have evolved in response to the need to protect the seed from predators.     

Multiple drivers of apparent competition reduce re-establishment of a native plant in invaded habitats

Although rarely examined, apparent competition, whereby exotic plants increase consumer pressure on native plants, could play a significant role in affecting native plant establishment in invaded habitats. Moreover, although terrestrial consumer communities often contain many consumer species, little is known about which consumers may generate apparent competition, and whether the strength or mechanism of apparent competition differs among different members of the consumer community. Using consumer-specific experimental exclosures and seed additions in the invaded grasslands of California, we demonstrate that multiple mechanisms of apparent competition are capable of limiting the re-establishment of the native grass Nassella pulchra in the absence of direct competition with exotic plants. The effect of small mammalian consumers (mice and voles) and larger consumers (e.g. rabbits, squirrels, deer) decreased with distance to the exotic forb Brassica nigra , which varied from 0–33 meters from focal N. pulchra . The effect of larger consumers also depended upon characteristics of the plant community directly adjacent (i.e. approx. 1  m) from focal N. pulchra . The effect of large consumers also increased with the richness of the exotic plant community and the degree to which the exotic plant community was dominated by exotic grasses as opposed to exotic forbs. Our finding that apparent competition can be driven by different mechanisms, that the importance of each mechanism depends upon which consumers have access, and that each mechanism has a different spatial extent, suggests that the composition of both the consumer community and the exotic plant community may shape the spatial dynamics of reestablishment, the potential for restoration, and the need for conservation.     

Temperature induced bacterial virulence and bleaching disease in a chemically defended marine macroalga

Host-pathogen interactions have been widely studied in humans and terrestrial plants, but are much less well explored in marine systems. Here we show that a marine macroalga, Delisea pulchra, utilizes a chemical defence - furanones - to inhibit colonization and infection by a novel bacterial pathogen, Ruegeria sp. R11, and that infection by R11 is temperature dependent. Ruegeria sp. R11 formed biofilms, invaded and bleached furanone-free, but not furanone-producing D. pulchra thalli, at high (24°C) but not low (19°C) temperatures. Bleaching is commonly observed in natural populations of D. pulchra near Sydney, Australia, during the austral summer when ocean temperatures are at their peak and the chemical defences of the alga are reduced. Furanones, produced by D. pulchra as a chemical defence, inhibit quorum sensing (QS) in bacteria, and this may play a role in furanone inhibition of R11 infection of furanone-free thalli as R11 produces QS signals. This interplay between temperature, an algal chemical defence mechanism and bacterial virulence demonstrates the complex impact environmental change can have on an ecosystem.     

Induction of settlement of larvae of the sea urchin Holopneustes purpurascens by histamine from a host alga

Larvae of the Australian sea urchin Holopneustes purpurascens are induced to settle and metamorphose (termed settlement herein) by a water-soluble compound produced by the red alga Delisea pulchra, the main host plant of new recruits. The settlement cue for H. purpurascens had previously been identified as a floridoside-isethionic acid complex, and this paper presents new evidence correcting that finding. The actual settlement cue produced by D. pulchra was isolated from the polar extract by cation-exchange chromatography and identified as histamine, using one- and two-dimensional nuclear magnetic resonance spectrometry. The chemical identity of the cue was confirmed by gas chromatography-mass spectrometry (GC-MS) and matrix-assisted laser desorption/ionization-time-of-flight mass spectrometry. Synthetic histamine and histamine at 4.5 microM isolated from D. pulchra both induced rapid settlement in 80%-100% of the larvae of H. purpurascens. Lower concentrations of histamine (0.9-2.3 micro M) induced larval settlement, but this response varied from 0%-90%. The histamine content of two host plants of H. purpurascens--D. pulchra and Ecklonia radiata--and of four other common species was quantified using GC-MS. D. pulchra had the highest histamine content, which is consistent with H. purpurascens recruiting to this species. Histamine was also detected in the seawater surrounding these host algae. This is the first time that a settlement cue has been quantified in the habitat of a marine organism.     

A revision of Gibsoniothamnus L. O. Williams (Schlegeliaceae)

Gibsoniothamnus L. O. Williams is a genus of 11 species native to Mexico, Central America, and northern Colombia. The species are shrubs or small trees with opposite, entire leaves, tubular corollas, four stamens, 2-locular ovary, and fleshy, indehiscent fruits. Calyces are unlobed, or they may have lateral, triangular or winglike lobes. Most species are epiphytes and appear to be bird pollinated and bird dispersed.     

Signal convergence in fruits: a result of selection by frugivores?

The Dispersal Syndrome hypothesis remains contentious, stating that apparently nonrandom associations of fruit characteristics result from selection by seed dispersers. We examine a key assumption under this hypothesis, i.e. that fruit traits can be used as reliable signals by frugivores. We first test this assumption by looking at whether fruit colour allows birds and primates to distinguish between fruits commonly dispersed by birds or primates. Second, we test whether the colours of fruits dispersed by primates are more contrasting to primates than the colours of bird‐dispersed fruits, expected if fruit colour is an adaptation to facilitate the detection by seed dispersers. Third, we test whether fruit colour has converged in unrelated plant species dispersed by similar frugivores. We use vision models based on peak sensitivities of birds’ and primates’ cone cells. We base our analyses on the visual systems of two types of birds (violet and ultraviolet based) and three types of primates (trichromatic primates from the Old and the New Worlds, and a dichromatic New World monkey). Using a Discriminant Function Analysis, we find that all frugivore groups can reliably discriminate between bird‐ and primate‐dispersed fruits. Fruit colour can be a reliable signal to different seed dispersers. However, the colours of primate‐dispersed fruits are less contrasting to primates than those of bird‐dispersed fruits. Fruit colour convergence in unrelated plants is independent of phylogeny and can be better explained by disperser type, which supports the hypothesis that frugivores are important in fruit evolution. We discuss adaptive and nonadaptive hypotheses that can potentially explain the pattern we found.     

浙江忍冬属植物新资料

报道了浙江忍冬科（Caprifoliaceae）忍冬属（Lonicera L.）植物分类研究的若干新发现：排除了L. macrantha在我国的分布和细毡毛忍冬（L. similis）在浙江的分布;恢复了L. guillonii的种级地位,并将异毛忍冬（L. macrantha var. heterotricha）作为其新异名,将灰毡毛忍冬（L. macranthoides）组合为其变种;描述了一新种——华大花忍冬（L. sinomacrantha）和一新变型——红花菰腺忍冬（L. hypoglauca f. pulchra）,编制了浙江产大花忍冬复合群的分类检索表。     

Directed vertebrate-mediated seed dispersal of a fleshy-fruited amaryllid in a heterogenous habitat

Most plants with fleshy fruits have seeds that are ingested by animals, but a less well-understood mode of seed dispersal involves fleshy fruits containing seeds that are discarded by frugivorous animals because they are too large or toxic to be ingested. We studied the seed dispersal biology of Haemanthus deformis, an amaryllid lily species found in a mosaic of bush clumps in a grassland matrix in South Africa. We asked whether seed dispersal is directed in and among bush clumps and whether germination and survival are greater for seeds dispersed to bush clumps than for those dispersed into grassland. Using camera trapping, we found that fruits are consumed mainly by birds and rodents. The pulp was removed from the seeds which were then discarded without ingestion. While many seeds were dispersed close to the parent plant, most (c. 78.5%) were dispersed further than 1 m away from the parent plant. Longer distance dispersal resulted mainly from birds flying off with fruits in their bill or from rodents engaging in scatter-hoarding behavior. Seedling survival was most successful within bush clumps as compared to grasslands and shade was identified as a primary requirement for seedling survival. Seeds from which the fruit pulp had been removed germinated faster than those in intact fruits. Haemanthus deformis deploys a system of directed seed dispersal, whereby both birds and rodents contribute to the dispersal of seeds within patchy bush clumps that are favorable for seedling survival.     

Vertebrate Fruit Removal and Ant Seed Dispersal in the Neotropical Ginger Renealmia alpinia (Zingiberaceae)*

Plants frequently display fruit characteristics that support multiple seed‐dispersal syndromes. These ambiguous characteristics may reflect the fact that seed dispersal is usually a complex process involving multiple dispersers. This is the case for the Neotropical ginger Renealmia alpinia (Zingiberaceae). It was originally suggested that the aromatic fruits of R. alpinia located at the base of the plant are adapted for terrestrial mammal seed dispersal. However, the dark‐purple coloration of the fruits and bright orange aril surrounding the seeds suggest that birds may play a role in R. alpinia seed dispersal. At La Selva Biological Station, Costa Rica, we used camera traps to record vertebrate visits to infructescences of R. alpinia. Most visitors were toucans and aracaris (Ramphastidae). However fruits were also removed by terrestrial mammals (coatis and armadillos). In addition to vertebrate fruit removal, some of the fruits dehisce and the seeds that fall on the ground are dispersed by ants. Fruitfall traps showed that 77 percent of fruits are removed by vertebrates. However, 15 percent of fruits fall to the base of parent plants to be potentially dispersed by ants. Experiments using a laboratory ant colony showed that ants are effective seed dispersers of R. alpinia. Ant seed manipulation increased germination success and reduced time to germination. In conclusion, primary seed dispersal in the Neotropical ginger R. alpinia is mostly performed by birds, additionally ants are effective dispersers at short distances. Seed dispersal in R. alpinia is a complex process involving a diverse array of dispersal agents.     

Standard yet unusual mechanisms of long-distance dispersal: seed dispersal of Corymbia torelliana by bees

Mellitochory, seed dispersal by bees, has been implicated in long-distance dispersal of the tropical rain forest tree, Corymbia torelliana (Myrtaceae). We examined natural and introduced populations of C. torelliana for 4 years to determine the species of bees that disperse seeds, and the extent and distance of seed dispersal. The mechanism of seed dispersal by bees was also investigated, including fruit traits that promote dispersal, foraging behaviour of bees at fruits, and the fate of seeds. The fruit structure of C. torelliana , with seed presented in a resin reward, is a unique trait that promotes seed dispersal by bees and often results in long-distance dispersal. We discovered that a guild of four species of stingless bees, Trigona carbonaria, T. clypearis, T. sapiens , and T. hockingsi, dispersed seeds of C. torelliana in its natural range. More than half of the nests found within 250 m of fruiting trees had evidence of seed transport. Seeds were transported minimum distances of 20–220 m by bees. Approximately 88% of seeds were dispersed by gravity but almost all fruits retained one or two seeds embedded in resin for bee dispersal. Bee foraging for resin peaked immediately after fruit opening and corresponded to a peak of seed dispersal at the hive. There were strong correlations between numbers of seeds brought in and taken out of each hive by bees ( r =  0.753–0.992, P  < 0.05), and germination rates were 95 ± 5%. These results showed that bee-transported seeds were effectively dispersed outside of the hive soon after release from fruits. Seed dispersal by bees is a non-standard dispersal mechanism for C. torelliana, as most seeds are dispersed by gravity before bees can enter fruits. However, many C. torelliana seeds are dispersed by bees, since seeds are retained in almost all fruits, and all of these are dispersed by bees.     

Taxonomic redescriptions of two ciliates, Protogastrostyla pulchra n. g., n. comb. and Hemigastrostyla enigmatica (ciliophora: spirotrichea, stichotrichia), with phylogenetic analyses based on 18S and 28S rRNA gene sequences

The morphology and infraciliature of two stichotrichid ciliates, Gastrostyla pulchra(Perejaslawzewa 1886) Kahl, 1932 and Hemigastrostyla enigmatica(Dragesco and Dragesco-Kernéis 1986) Song & Wilbert, 1997, collected from marine and brackish sediments, were investigated by using living observations and protargol impregnations. Both 18S and 28S rRNA genes of these two species were sequenced. The 18S rDNA show high similarities (98.4%-99.7%) among populations of each species. There is about 94% similarity in 18S rDNA genes between G. pulchra and Gastrostyla steinii, the type species of the genus, which has been confirmed to be an oxytrichid by previous studies. In the phylogenetic trees of 18S, 28S, and combined 18S and 28S rDNA, both G. pulchra and H. enigmatica are consistently placed outside the well-established oxytrichid clade. Based on our analyses and previous ontogenetic data, we conclude that these two species may represent some lower groups in the subclass Stichotrichia, and that G. pulchra should represent a new genus, Protogastrostyla n. g. This new genus, which is morphologically similar to Gastrostyla, differs in its morphogenesis: the apical part of the old AZM is retained combining with the newly built membranelles that develop from the proter's oral primordium; the primary primordia of the dorsal kinety; and marginal primordia commence de novo without a definite contribution from the old structure.     

Fruit size preference in the New Zealand pigeon (Hemiphaga novaeseelandiae)

Abstract We investigated whether the New Zealand pigeon Hemiphaga novaeseelandiae (Columbidae) exhibits size‐based preferences for fruits. We tested the hypothesis that in small‐fruited species, pigeons would prefer larger fruits, but in larger‐fruited species, this preference would reverse as the pigeons become increasingly limited by their gape size. We collected undispersed fruits and bird‐dispersed seeds of 10 plant species, some over several sites or years (13 datasets in total). We estimated the fruit size of dispersed seeds by fitting regressions of fruit diameter to seed diameter in intact fruits. We were able to predict fruit diameter from seed diameter in 12 of the 13 populations, although the relationship was stronger in single‐seeded species than in multi‐seeded species. Seven of the 12 populations tested showed a significant difference in seed diameter among undispersed and dispersed seeds. However, our results showed no consistent pattern in fruit size preference by the New Zealand pigeon and did not support our hypothesis. The large‐bodied New Zealand pigeon is generally not gape limited and fruit size preferences appear to be independent of mean fruit size.     

Density-dependent variation in reproductive success in a terrestrial orchid

 Within population variation in plant density can affect reproductive success and breeding systems. We examined such effects in Listera cordata R. Br., a north temperate terrestrial orchid, in a subalpine population in Colorado, USA. Hand pollinations showed that L. cordata was self-compatible as all pollinations produced fruits. Cross-pollinations differed significantly from self-pollinations, and had a higher number of seeds per capsule and higher % of fertilized ovules. Together they could compensate for the transmission advantage of self-pollinations. Average number of flowers per inflorescence was 4.2–4.4 and average fruit set was 20.7–23.7%. By establishing 36 plots with varying plant densities (1, 4, 8 and 16) and emasculating all flowers in target plants of a set of 20 plots, we estimated density effects on fruit and seed production. Plant density had no effect on the amount of reproduction except in reduced seed production of highly dispersed plants. Our results suggest that inbreeding depression may be an important selective factor in L. cordata. Further studies are needed to define its role in the evolution of mating systems in this species. Received February 21, 2000 Accepted December 26, 2000     

Revealing secondary seed removers: results from camera trapping

This paper reports the results of the first study on secondary seed removal of seeds dispersed by Sykes’ monkeys (Cercopithecus albogularis) using camera traps in Africa. Patterns of primary seed dispersal are often superimposed by secondary conveyance, emphasising the need to study these secondary processes carefully. As the agents and mechanisms of seed dispersal are often concealed, being carried out by cryptic or nocturnal animals in dense vegetation, camera trapping was deemed a viable means to investigate secondary removal of seeds disseminated by C. albogularis in the Western Soutpansberg, South Africa. Camera traps were established at preferred feeding trees of the focal Sykes’ monkey group to identify animal species that remove seeds and fruits spat and dropped to the forest floor and seed removal observations were carried out. This method proved to be effective in identifying seed removers and also allowed to get indications about the quantities of seeds removed. Ten animal species were recorded visiting the trees, of which eight were observed removing seeds and fruits. Overall seed and fruit removal rates were high, indicating that the foraging behaviour of C. albogularis attracts many terrestrial frugivores.     

The Distribution and Evolution of Terrestrial Nemertines

The occurrence of terrestrial nemertines on widely separatedoceanic islands is described: there are three groups of species(in Australia, New Zealand and the Indopacific islands) andthree separate species on Atlantic islands. Parallel evolutionfrom separate marine ancestors is indicated. The evolutionaryrelationship between different species is established on securemorphological grounds, with particular reference to the genusPantinonemertes which comprises marine, semi-terrestrial andfully terrestrial species. The distribution of terrestrial nemertinesis discussed. Most species are extremely localised but a fewhave become widely dispersed, notably Argonemertes dendyi butalso Geonemertes pelaensis and to a lesser extent Leptonemerteschalicophora. At least two of these widespread species are hermaphrodite.     

Latitudinal and elevational variation in fruiting phenology among western European bird-dispersed plants

I try to test the prediction that bird-dispersed plants should produce fruits when fruit-eating birds are most abundant by reviewing some phenological data of fleshy fruit production in western Europe The prediction that fruit ripening dates in populations of the same species should occur later at lower latitudes and elevations, to coincide with the maximum abundance of fruit-eating birds, is not supported by the data The patterns of seasonal variation in the total number and biomass of fruits, but not in the proportion of species in fruit, in communities at different latitudes and elevations do coincide with patterns of seasonal abundances of avian frugivores 1 suggest that this coincidence is due to the greater relative abundance (and contribution to total fruit production) in each locality of species that fruit at times of the year when birds are most abundant These species may have achieved a demographic advantage by getting more seeds dispersed than species that ripen fruits in other seasons     

Functional uniqueness of a small carnivore as seed dispersal agents: a case study of the common palm civets in the Tabin Wildlife Reserve, Sabah, Malaysia

Many carnivorous mammals consume fruits and disperse the intact seeds to specific sites. Few studies have attempted to quantify this seed dispersal or evaluate its effectiveness, despite its potential importance and functional uniqueness. In the study reported here, we found that a frugivorous carnivore, the common palm civet (Paradoxurus hermaphroditus), generated seed shadows that are distinct from those of the sympatric frugivore, the pig-tailed macaque (Macaca nemestrina), and played a unique and important role in the regeneration of Leea aculeata (Leeaceae). We found that macaques dispersed the seeds randomly, while civets dispersed them non-randomly to sites such as the banks of small rivers, rain-flow paths, abandoned trails, and treefall gaps, which are characterized by low stem density and canopy cover. Seeds of L. aculeata that were dispersed by civets to the banks of rivers and gaps had significantly higher survival and growth rates than those dispersed to rain-flow paths or abandoned trails. Seeds dispersed by macaques or to random locations also had low survival. Although the effects of the civets on seed fate were not straightforward, compared with macaques and random dispersal, civets significantly enhanced the survival and growth of L. aculeata seeds after 1 year. These results indicate that non-random dispersal by civets is important for the persistence of L. aculeata. Civets may disperse other plant species and thus could have profound effects on forest dynamics.