Comparative seed morphology of <Emphasis Type="Italic">Leandra</Emphasis> (Miconieae,Melastomataceae)

The seed morphology of 79 species of neotropical Miconieae (Melastomataceae) is presented. These species have been chosen, in majority, from the polyphyletic genus Leandra. A few other species from the polyphyletic genera Miconia, Ossaea, and Clidemia were also sampled, because of potential similarities. Sixteen morphological seed types are defined after analysis through light microscopy and scanning electron microscopy. The seed morphology appears to possess a great deal of variability on the level of the overall shape as well as the structure and the surface of the testa. The different types defined here do not match with genera or sections, but rather are composed of species coming from different genera. In comparison with a preliminary molecular phylogeny study done on Leandra, some types of seeds are related to well supported clades. In some cases seed morphology corresponds with natural groups of species, thus being of high phylogenetic importance.     

西双版纳地区4种榕树果实的资源分配及其种子萌发特性

为了解榕树果实的资源分配和种子萌发特征,对西双版纳地区的垂叶榕(Ficus benjamina)、高山榕(F.altissima)、聚果榕(F.racemosa)和无柄雅榕(F.concinna)的果实营养、繁殖资源分配,以及种子萌发特性进行了研究。结果表明,4种榕果的营养分配存在差异,榕果重的种类果肉亦多,榕果大者果腔较大,且种子数量较多,但雄花和榕小蜂数量不一定多。榕果分配给雌性功能及雄性功能繁殖的小花比例也不相同,聚果榕在雌性功能上的分配显著大于雄性功能,其它3种榕果则分配于雄性功能的资源比雌性功能的多,但程度不同。此外,4种榕树的种子大小及重量存在显著的种间差异,以至于影响到种子的萌发率及根茎生长,果实最大的聚果榕种子最多,萌发率高,但由于种子最小,其根茎生长缓慢且短;果实适中的垂叶榕和高山榕种子较大,不但萌发率较高,其根茎生长也较快;果实和种子都小的无柄雅榕,种子萌发率低,根茎生长慢。生长在水热环境好的榕树比生长在相对贫瘠环境的榕树,由于极少面临水热光照等限制,更容易进化出较小的种子,萌发能力和速度也相对较差。这说明不同生境中榕树进化出了不同的繁殖分配机制及资源利用策略来适应环境变化。     

Effects of ingestion by neotropical bats on germination parameters of native free-standing and strangler figs (Ficus sp., Moraceae)

Fruit-eating animals can influence the germination success of seeds through transportation and handling. We experimentally tested the contribution of ingestion by the common fruit-eating bat, Artibeus jamaicensis (Phyllostomidae, Chiroptera), to the percentage and rate of seed germination of figs (Ficus, Moraceae), which are considered keystone species for many frugivores. We collected fruits from three species of native free-standing figs (subgenus Pharmacosycea: F. insipida, F. maxima and F. yoponensis) and three species of native strangler figs (subgenus Urostigma: F. nymphiifolia, F. obtusifolia and F. popenoei) on Barro Colorado Island, Panama. The germination success of seeds removed from fruit pulp either manually or by ingestion was very high (>92%), while seeds that were not removed from fruit pulp were destroyed by fast-growing fungi within a few days. The dynamics of seed germination were not influenced by ingestion, but differed between the two subgenera of figs. In free-standing figs, germination started significantly earlier (5.3 ± 0.7 days) than in strangler figs (8.6 ± 1.4 days). Furthermore, strangler seeds were covered with a sticky coating and their seedlings developed cotyledons faster than fine roots, in contrast to free-standing figs that showed the opposite pattern. Our results demonstrate that the germination of fig seeds is positively influenced by passage through the gut of A. jamaicensis. Furthermore, free-standing and strangler figs revealed differences in germination parameters that might be adaptive with respect to the suitability of microsites such as tree fall gaps or host trees for establishment.     

Phenology of figs in Budongo Forest Uganda and its importance for the chimpanzee diet

This paper reports on the phenological patterns of figs in Budongo Forest, Uganda, and how it relates to chimpanzee food availability in different seasons. In addition, we analysed the dung of chimpanzees to understand the composition of fruits in their diet. The aim of our study was to assess Ficus phenology and how it affects chimpanzee diet. Fifteen species of figs were monitored for fruit (syconium) and leaf phenology between June 2000 and 2001. Ficus fruit production varied significantly between and within species, and also with tree trunk and crown diameters. Fig fruit production was asynchronous and individual fig trees produced crops from one to five times in a year. In addition to fruits, chimpanzees fed on young leaves of some Ficus species. Shedding of old Ficus leaves coincided with the dry season, followed by appearance of young leaves. The dry season in Budongo is a period of general fruit scarcity. The combination of fig fruits and young leaves make up the most important food in the diet of chimpanzees. From the chimpanzee dung, more than 78% of seeds comprised fig ‘seeds’ (nutlets) and the rest of the diaspores were from other tree species. Our findings suggest that chimpanzees disperse large number of diaspores in their dung, thereby serving as important agents of natural forest regeneration.     

A phylogenetic evaluation of Leandra (Miconieae, Melastomataceae): a polyphyletic genus where the seeds tell the story, not the petals

Melastomataceae is a tropical family of 4500–5000 species divided into nine tribes. The largest tribe, Miconieae, is composed of approximately 2200 species in 30 genera and is found exclusively in the Neotropics. Previous phylogenetic analyses of the Miconieae have suggested that many of the genera are derived from a paraphyletic Miconia. However, these analyses only included six species of the large genus Leandra, so its phylogenetic affinities remained unclear. As currently defined Leandra is characterized by acute petals and terminal inflorescences, but some species of Miconia, Clidemia and Ossaea also have these characters. In this study, we present an analysis of nrITS sequence data for a sample of 63 species of Leandra. The genus is clearly resolved as polyphyletic, but some distinct and well‐supported clades exist. Some of these partially correspond to sections recognized in the nineteenth century by Cogniaux, or to geographic distribution. The distribution of seed structure characters is better correlated with the phylogeny than traditional characters, such as petal morphology. Seed appendages in Leandra have evolved independently at least four times. © The Willi Hennig Society 2007.     

Coevolution of reproductive characteristics in three dioecious fig species and their pollinator wasps

This study investigates dioecious fig species using a pollinator introduction experiment. Our aims were to determine: (1) whether there was a significant difference in foundress distribution between sexes per fig species; (2) whether fig size and foundress number affect reproductive success of dioecious figs; and (3) who is the ‘controlling partner’ in the fig/pollinator mutualism. Three dioecious fig species: Ficus semicordata, Ficus hispida and Ficus tinctoria from Xishuangbanna, China, were selected for this experiment. We found that there was no significant difference of the foundress number in female and male figs of F. semicordata, F. hispida and F. tinctoria. Also, the foundress number did not depend on the fig diameter. The numbers and the proportions of fig seeds and female wasp offspring significantly increased with more foundresses; and fig seed number was significantly higher than female wasp offspring in F. semicordata and F. hispida, but not in F. tinctoria. Our results indicate that figs are generally the ‘controlling partner’ in fig-wasp mutualisms in species with large figs, but not with small figs. Compared with published studies of reproductive success in monoecious figs, the dioecious figs seem to be more efficient in producing both seeds and wasp offspring when there is a high number of foundress.     

Frugivory and seed dispersal in a hyperdiverse plant clade and its role as a keystone resource for the Neotropical fauna

Background and AimsMuch of our understanding of the ecology and evolution of seed dispersal in the Neotropics is founded on studies involving the animal-dispersed, hyperdiverse plant clade Miconia (Melastomataceae). Nonetheless, no formal attempt has been made to establish its relevance as a model system or indeed provide evidence of the role of frugivores as Miconia seed dispersers.MethodsWe built three Miconia databases (fruit phenology/diaspore traits, fruit–frugivore interactions and effects on seed germination after gut passage) to determine how Miconia fruiting phenology and fruit traits for >350 species interact with and shape patterns of frugivore selection. In addition, we conducted a meta-analysis evaluating the effects of animal gut passage/seed handling on Miconia germination.Key Results Miconia produce numerous small berries that enclose numerous tiny seeds within water- and sugar-rich pulps. In addition, coexisting species provide sequential, year long availability of fruits within communities, with many species producing fruits in periods of resource scarcity. From 2396 pairwise interactions, we identified 646 animal frugivore species in five classes, 22 orders and 60 families, including birds, mammals, reptiles, fish and ants that consume Miconia fruits. Endozoochory is the main dispersal mechanism, but gut passage effects on germination were specific to animal clades; birds, monkeys and ants reduced seed germination percentages, while opossums increased it.ConclusionsThe sequential fruiting phenologies and wide taxonomic and functional diversity of animal vectors associated with Miconia fruits underscore the likely keystone role that this plant clade plays in the Neotropics. By producing fruits morphologically and chemically accessible to a variety of animals, Miconia species ensure short- and long-distance seed dispersal and constitute reliable resources that sustain entire frugivore assemblages.     

Prolific fruit output by the invasive tree Bellucia pentamera Naudin (Melastomataceae) is enhanced by selective logging disturbance

Selective logging in tropical rain forests may promote population growth of invasive plants. The ability of invaders to respond, specifically in reproductive traits, to increase in resource abundance may allow them to increase their presence in the seed rain of recipient communities. The invasive pioneer tree Bellucia pentamera (Melastomataceae) is currently spreading within Gunung Palung National Park in West Kalimantan, Indonesia. The park has also experienced periods of illegal, small‐scale, selective logging that seem to have facilitated population growth and spread of this invader. We first used fruiting frequency as a proxy for fruit output to make comparisons between B. pentamera and the native tree community of over 200 genera. We then constructed two Generalized Linear Autoregressive Moving Average (GLARMA) models using 13 months of phenology data to predict both fruiting frequency and crop size of B. pentamera under selective logging versus natural treefall disturbance regimes. Bellucia pentamera fruited at considerably higher frequency than all 200 native genera considered. This invader also responded positively to selective logging with both odds of being in fruit and crop sizes at least doubling in logged plots. Prolific seed output of B. pentamera is especially problematic within the lottery competition of tropical rain forest gap tree communities, in which prominence in the seed rain is particularly important. Furthermore, the reproductive response of B. pentamera to selective logging suggests that this harvesting practice may have a considerable role in facilitating this invasion in Southeast Asia.     

Elucidating the mechanism of poricidal anther dehiscence in Miconia species (Melastomataceae)

Melastomataceae have porate anthers. However, unlike Solanaceae and many monocots, in which the poricidal dehiscence depends on the presence of a mechanical layer (often the endothecium), most members of Melastomataceae have no evident specialized layer related to the poricidal opening. The goal of this study was to characterize the tissues that form the apical pore of the anther in 10 Miconia species, which may help to understand the nearly unknown mechanism of anther dehiscence in this genus, considered to be one of the largest and most diverse New World genera. Before anthesis, the apical pores of all of the species are closed by a uniseriate epidermis, the cells of which lack a cuticle. In contrast, the epidermis of the remainder of the anther is covered by a thick, ornamented cuticle. Among Myrtales, the Melastomataceae form a clade with Alzateaceae, Crypteroniaceae and Penaeaceae, almost all of which have anthers with endothecium lacking wall thickenings. In these families, the endothecium may or may not be present in the mature anther, with degenerating cells in the latter case. Anther dehiscence does not depend on the endothecium as the mechanical layer, and this process is still not well understood. However, in the Miconia species studied here, the cuticle may prevent tissue dehydration, and the pore opening seems to be due to the passive process of dehydration taking place only in the pore region due to the absence of the cuticle.     

Nutritional Values of 14 Fig Species and Bat Feeding Preferences in Panama1

Figs are a critical resource for many tropical frugivores, yet they often are referred to as low quality fruits. To determine their nutritional value, both as a group and for individual species, we analyzed 14 fig species from Barro Colorado Island (BCI), Panama, for fiber, tannins, lipids, protein, carbohydrates, amino acids, and minerals. Seeds and pulp were analyzed separately. Fig fruit pulp consisted of about one‐third digestible components, mostly carbohydrates with some lipids and proteins. Tannin, lignin, and water‐soluble carbohydrates showed considerable variation among species, as did fruit size. Figs contained high amounts of amino acids, such as leucine, lysine, valine, and arginine, and minerals, such as potassium, calcium, magnesium, sodium, and phosphorus. One species, Ficus insipida, contained the highest concentrations of almost all amino acids, many minerals, and protein. Small figs had as much nutritional value per gram as large figs. Free‐standing figs had higher percentages of protein, complex carbohydrates, and ash than strangler figs, which had higher percentages of water‐soluble carbohydrates, tannins, and hemicellulose. The guild of fruit‐eating bats on BCI included ten common species with diets dominated by figs. Fecal analyses and captures at ripe fig trees showed a consistent pattern of resource partitioning. Small bats preferentially ate small‐fruited and strangler figs while large bats consumed mostly large‐fruited and free‐standing figs. Small bats most often ate F. bullenei, which has high levels of lipid and carbohydrates, and F. yoponensis, which has high levels of protein. Medium and large bats most often ate F. insipida, a nutritionally superior species; their second most eaten species was F. obtusifolia, in which the large size may make it efficient to eat. Each bat ate a variety of fig species, supporting the idea that although no single species of fig may be sufficient to sustain frugivores, a mix of fig species can provide a complete set of nutrients.     

Floral constraint resulting from intersexual mimicry in a gynodioecious fig tree

Fig trees (Ficus: Moraceae) are pollinated by female fig wasps (Agaonidae) whose larvae develop inside galled flowers of unusual inflorescences (figs). Most fig trees also support communities of non‐pollinating fig wasps. Figs of different species display great size variation and contain tens to tens of thousands of flowers. Around one‐half the species of fig trees have the gynodioecious breeding system, where female trees have figs that produce seeds and male trees have figs that support development of pollinators. Mutual mimicry between receptive male and female figs ensures that pollinators enter female figs, even though the insects will die without reproducing, but the need to give no sex‐specific cues to the pollinators may constrain differences in size between receptive male and female figs. We compared relationships between inflorescence size and some measures of reproductive success in male and female figs of Ficus montana grown under controlled conditions in the presence of the pollinator Kradibia tentacularis and its main parasitoid Sycoscapter sp. indesc. Female figs that contained more flowers produced more seeds, but male figs did not increase the production of female pollinator K. tentacularis fig wasps in proportion of the flower number. Although more flowers were galled by the pollinators in male figs containing more female flowers, the high larval mortality caused by parasitism and nutritional limitation prevented the increase in the production of adult female offspring. Selection may favor the increase in flower numbers within figs in female plants of F. montana, but contrarily constrain this attribute in male plants.     

Comparative anatomy and morphology of nectar-producing Melastomataceae

Background and Aims

Most neotropical Melastomataceae have bee-pollinated flowers with poricidal anthers. However, nectar rewards are known to be produced in about 80 species in eight genera from four different tribes. These nectar-producing species are pollinated by both vertebrates and invertebrates.

Methods

The floral morphology and anatomy of 14 species was studied in six genera of nectar-producing Melastomataceae (Blakea, Brachyotum, Charianthus, Huilaea, Meriania and Miconia). Anatomical methods included scanning electron microscopy, and serial sections of paraffin-embedded flowers.

Key Results

All vertebrate-pollinated melastome flowers have petals that do not open completely at anthesis, thus forming a pseudo-tubular corolla, while closely related species that are bee pollinated have rotate or reflexed corollas. In most species, nectar secretion is related to stomatal or epidermal nectaries and not filament slits as previously reported. Moreover, the nectar is probably supplied by large vascular bundles near the release area. Blakea and Huilaea have nectary stomata located upon the dorsal anther connective appendages. Brachyotum also has nectary stomata on the anther connectives, but these are distributed lengthwise along most of the connective. Meriania may release nectar through the anther connective, but has additional nectary stomata on the inner walls of the hypanthium. Miconia has nectary stomata on the ovary apex. Charianthus nectaries were not found, but there is circumstantial evidence that nectar release occurs through the epidermis at the apex of the ovary and the lower portions of the inner wall of the hypanthium.

Conclusions

Nectar release in Melastomataceae is apparently related to nectary stomata and not filament slits. The presence of nectary stomata on stamens and on ovary apices in different lineages suggests that the acquisition of nectaries is a derived condition. Nectary location also supports a derived condition, because location is strongly consistent within each genus, but differs between genera.Key words: Blakea, Brachyotum, Charianthus, Huilaea, Meriania, Melastomataceae, Miconia, nectaries, nectary stomata, pollination     

Seasonal variation in seed predation by adult carabid beetles

In many agro‐ecosystems of the temperate region, granivorous ground beetles (Coleoptera: Carabidae) are the most important invertebrate predators of seeds after dispersal. The consumption of a particular species of seed differs among carabid species, but seasonal variation in consumption has not yet been studied. Seasonal differences in the quantity of seed consumed by the carabids Harpalus affinis (Schrank) and Harpalus distinguendus (Duftschmid) were studied in a cafeteria experiment. Adults were collected on five dates between April and November and provided with seeds of 28 species of dicotyledonous herbaceous plants. Proportions eaten of the seeds offered (preferences) were nearly identical in both carabid species. Overall seed consumption increased from early spring until late June, then gradually decreased towards autumn. Despite this variation, the preferences on particular dates were essentially similar. Seasonal variation in carabid physiology and feeding experience thus influenced overall seed consumption, but not their preferences for particular seed species.     

Foraging Patterns of Sympatric Hornbills during the Nonbreeding Season in Arunachal Pradesh, Northeast India

The diet and food resource partitioning of three sympatric hornbills (Great Hornbill Buceros bicornis, Wreathed Hornbill Aceros undulatus, and Oriental Pied Hornbill Anthracoceros albirostris) during the nonbreeding season were studied relative to fruit availability in a foothill semievergreen forest of Arunachal Pradesh, northeast India. Hornbills fed on fruits of 49 plant species that comprised over 95 percent of their diet. Hornbill species partitioned food resources by varying the relative contribution of figs and non‐fig fruits in the diet. Similarity in non‐fig fruit diet was low. Ten species contributed to over 90 percent of the non‐fig diet. The availability of non‐fig fruits was much lower in the nonbreeding season than in the breeding season; however, despite lower fruit availability during the nonbreeding season, hornbills had a wide diet breadth and indirect evidence suggests that Wreathed Hornbills foraged widely in this season for fruit. Great Hornbills relied more on figs that were available year‐round. The diverse diet breadth that included rare and patchy fruit resources underscores the importance of conserving large forest tracts for hornbills.     

Stable isotope analysis indicates trophic differences among forest floor carabids in Japan

Differences in trophic niches among carabid beetles (Coleoptera: Carabidae) co‐occurring on the forest floors of warm temperate forests in central Japan were studied using carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope analyses. Different carabid species showed similar δ¹⁵N values, which were higher than those of their possible invertebrate prey (herbivores and detritivores) collected from the litter layer, indicating that these species were consumers in the same trophic level. In contrast, δ¹³C values differed among carabid species, indicating interspecific differences in prey animals. The variation in the δ¹³C value was larger in summer than in autumn. In summer, δ¹³C values indicated that some carabids depended highly on either grazing (low δ¹³C values) or detrital sources (high δ¹³C values) within the food chain [Chlaenius posticalis Motschulsky and Haplochlaenius costiger (Chaudoir), respectively], although other species with intermediate δ¹³C values likely depended on both. The latter group of species comprised mostly two dominant genera (Carabus and Synuchus). Although congeners might have similar feeding habits, the stable isotope ratios indicated trophic niche differences between adults of different species and between adults and larvae of the same genus.     

Carabid beetle (Coleoptera: Carabidae) assemblages in the intermediate successional stage after Mt. Bandai eruption of 1888: Effects of environmental variables on carabid beetles in the Urabandai area

We studied carabid beetle assemblage structure and species diversity in an intermediate successional stage (seral) forest established in areas affected by the 1888 eruption of Mt. Bandai and a climax forest that had not been affected by the eruption at the Urabandai area, Fukushima Prefecture, Japan. In total, 2,131 carabid beetles representing 31 species were collected using pitfall traps without bait. A comparison of carabid beetle assemblages between the two forest types revealed that the number of species observed was comparable, but their abundance was greater in the seral forest. The assemblage structure clearly differed between the two forest types. In the seral forest, forest generalists, such as Synuchus arcuaticollis and Pterostichus prolongatus, along with forest specialists including Carabus vanvolxemi and Pterostichus asymmetricus, were collected. In the climax forest, forest generalists, such as Synuchus cycloderus and Carabus albrechti tsukubanus, were more abundant than forest specialists. This suggests that the current seral forest in the Urabandai area possesses environmental variables that enable the coexistence of both forest specialist and generalist beetle species. A redundancy analysis showed that six species from the genera Synuchus and Pterostichus were associated with high canopy openness and high understory vegetation cover, whereas species recorded only in the climax forest were associated with deep litter. Therefore, it seems likely that carabid beetles in the Urabandai area were affected by these three environmental variables.     

Two new species of <Emphasis Type="Italic">Miconia</Emphasis> sect. <Emphasis Type="Italic">Sagraea</Emphasis> (Melastomataceae) from the Macaya Biosphere Reserve,Haiti, and twelve relevant new species combinations

The Sagraea clade (Melastomataceae, tribe Miconieae) is briefly characterized, typified, and formally treated as a section within Miconia. In addition, two new species of Miconia sect. Sagraea, endemic to the floristically diverse Massif de la Hotte of southwestern Haiti and discovered during the course of a systematic revision of the Caribbean species of this section, are here described and illustrated. Miconia hottensis and M. navifolia, morphologically similar and possible sister species, are compared to each other and to the widespread Caribbean species M. capillaris and the southwestern Dominican Republic endemic M. tetraptera; these four species share rectangular stems with four low ridges or wings and minute, short-stalked, peltate or pseudopeltate hairs and likely form a clade.     

Dispersal of adult female fig wasps 1. Arrivals and departures

Ficus burtt-davyi, like most other fig species (Ficus, Moraceae), is exclusively pollinated by its own unique species of fig wasp, in this caseElisabethiella baijnathi (Chalcidoidea, Agaonidae). Because fig crop development on any one tree is usually synchronised, the small and short-lived female wasps have to migrate and find other trees bearing figs which are at suitable stage of development for oviposition. However, the likelihood of successful location and subsequent arrival at a new host tree is dependent on distance and the effect of environmental factors such as wind and temperature. This study examines the relationship between ambient temperatures and the timing of fig wasps emergence from their natal figs and the commencement of their dispersal flight. The behaviour of the wasps arriving at figs which were ready to be pollinated was also examined. The female wasps did not appear to distinguish between the figs and other parts of the tree when in flight. However, after landing on the tree their search for figs was more directed as they visited more figs than leaves. Short-range recognition of figs appears to be by contact chemo-reception, but the wasps showed a preference for entering figs which did not already contain a female wasp.     

Una nueva especie de <Emphasis Type="Italic">Miconia</Emphasis> (Melastomataceae: Miconieae) de Colombia

Miconia punctibullata, a new species of Melastomataceae from Colombia, is described and illustrated. The new species can be distinguished by its scandent herbaceous habit, stems and inflorescences with swollen annular projections below the nodes, punctations with sessile, glandular trichomes on the abaxial surface of the leaf, compound dichasial inflorescences, and bright yellow petals. A key to all species of Miconia with annular stem projections is provided.

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Resumen  Se describe e ilustra Miconia punctibullata, una nueva especie colombiana de Melastomataceae, caracterizada por el hábito herbáceo semiescandente, la presencia de proyecciones anulares engrosadas por debajo de los nudos (manicaduras) en tallos y ejes de la inflorescencia, punteaduras con tricomas glandulares sésiles por el envés de la lámina foliar, inflorescencias en dicasio compuesto, y pétalos amarillos. Se incluye una clave para todas las especies de Miconia con nudos manicados.