Artificial Defoliation Induces Trichome Production in the Tropical Shrub Cnidoscolus aconitifolius (Euphorbiaceae)1

Induction of plant defenses and their spatial variability are key subjects in the field of ecology and evolution of defensive traits in plants. Nevertheless, induction has been more commonly studied under controlled environments, ignoring other factors that might influence this process in natural settings. The main goal of this study was to determine if artificial defoliation induces trichome production in three natural populations of the tropical shrub Cnidoscolus aconitifolius. First, we performed trichome counts for each population before imposing artificial defoliation to assess differences in trichome loads between populations. Trichome densities (trichomes/cm²) were quantified for leaf blades, petioles, and flower stalks. To determine if defoliation induced trichome production, three defoliation treatments (0% leaves defoliated or controls, 50% of total leaves defoliated, and 100% defoliation) were applied once at the beginning of the reproductive season. Trichome counts were performed on each structure every ～20 d during a 3‐mo period after the application of treatments. Trichome counts showed significant differences in trichome densities between populations for all three structures. In turn, artificial defoliation increased trichome density. Significant differences among treatments were found for trichome densities on leaf blades and petioles. In both these structures, the 100 percent defoliation treatment differed significantly from control plants, presenting higher trichome densities. In addition, the treatment × population interaction was not significant for leaf blades and petioles, indicating that induction is a generalized response in this species, at least at the study sites. These results indicate that trichomes in C. aconitifolius are inducible due to defoliation.     

Scaling of Petiole Dimensions with Respect to Leaf Size for a Tropical Tree, Scaphium macropodum (Sterculiaceae), in Borneo

Scaphium macropodum (Miq.) Beumee ex Heyne (Sterculiaceae) in a tropical rain forest in West Kalimantan (Indonesia) was analyzed from the viewpoint of statics. The petiole diameter must increase with increasing leaf size to retain enough mechanical stability and a sufficient amount of conductive vessels. The petiole's cross-sectional area at its base was found to be proportional to the leaf blade's dry mass, which indicates that Shinozaki's pipe model is applicable to leaves with different sizes. Although larger leaves produce greater bending moments on the petiole's cross-section as a result of their greater weights, the bending stresses at the petiole's base caused by the leaf's weight were constant at ca. 76,900 g cm⁻² regardless of leaf size. Thicker petioles increase the leaf's mechanical stability, but require sizable energy investments for their construction. It is hypothesized that the constant value for petiolar stress indicates an optimal balance between energy economy and the mechanical stability of S. macropodum leaves. To keep bending stress constant, the leaf blade's center of gravity shifts to a more proximal position and the cross-sectional area of the petiole increases. Received 8 December 1997/ Accepted in revised form 1 December 1998     

Recruitment Distance from the Nearest Reproductive Conspecific Increases with Tree Size in Tropical Premontane Wet Forests

Studies have shown that median distances of plants from the nearest reproductive conspecific (recruitment distance) shifts outward with increasing age or size class, and that plant spatial distribution changes over time in a predictable manner. However, observations and empirical evidence for such predictable changes are limited, and underlying mechanisms explaining such patterns for a wide range of individual sizes are not fully explored. In Costa Rican premontane wet forests, I empirically tested whether recruitment distance changes in a predictable manner with increasing size for five animal‐dispersed tree species by considering all post‐germination sizes. Specifically, I tested the Janzen–Connell hypothesis and the colonization hypothesis by considering distance, density, size, herbivory, biotic infection, and light availability simultaneously. Recruitment distance increased with increasing size (16–22 m) for the four non‐pioneer species, suggesting eventual regeneration success for seeds dispersed away from reproductive conspecifics. During the 2 years of this study, I found positive distance‐dependent survivorship and light availability were important for post‐seedling survivorship, in agreement with the Janzen–Connell hypothesis and the colonization hypothesis, respectively, but only for seedlings. However, seedlings did not escape aboveground herbivory or biotic infection better when the seeds were dispersed greater distances. Results highlight the importance of seed dispersal for successful regeneration, and suggest that changes in spatial distribution over time may be predictable in the vicinity of maternal trees for some non‐pioneer tree species.     

The mis-measure of manioc (Manihot esculenta,Euphorbiaceae)

The distinction between “bitter” and “sweet” (toxic and non-toxic) varieties of the cyanide-containing food crop manioc (Manihot esculenta, Euphorbiaceae) has a long tradition in the tropical forest areas of South and Central America where it was first cultivated. Yet this distinction has no taxonomic basis. The levels of cyanogenic glucosides found in manioc varieties not only vary widely, but do not correspond with any other known morphological or ecological feature. Nonetheless, these two “varieties” are commonly reported to have distinct geographical and cultural distributions and each is associated with a particular traditional food complex. This paper reviews the literature regarding the nature, distribution, and traditional uses of manioc varieties and concludes that (1) the geographical and cultural distribution of bitter and sweet varieties of manioc may not be as distinct as has been thought; (2) traditional categories of bitter and sweet manioc may stem more from culturally derived belief systems than from actual known levels of toxicity; and (3) the choice of complex and labor intensive processing methods usually associated with bitter manioc may not be required for detoxification but rather for the derived food products, particularly manioc flour.     

Microbial Biomass,Respiration, and Decomposition of Hura crepitans L. (Euphorbiaceae) Leaves in a Tropical Stream1

The processing of leaves in temperate streams has been the subject of numerous studies but equivalent tropical ecosystems have received little attention. We investigated leaf breakdown of a tropical tree species (Hura crepitans, Euphorbiaceae), in a tropical stream using leaf bags (0.5 mm mesh) over a period of 24 days. We followed the loss of mass and the changes in adenosine triphosphate (ATP) concentrations and respiration rates associated with the decomposing leaves. The breakdown rate was fast (k=?0.0672/d, k_d=?0.0031/degree‐day), with 81 percent loss of the initial mass within 24 days. This high rate was probably related to the stable and high water temperature (22°C) favoring strong biological activity. Respiration rates increased until day 16 (1.1 mg O₂/h/g AFDM), but maximum ATP concentrations were attained at day 9 (725 nmol ATP/g AFDM) when leaf mass remaining was 52 percent. To determine the relative importance of fungi and bacteria during leaf decomposition, ATP concentrations, and respiration rates were determined in samples treated with antibiotics, after incubation in the stream. The results of the samples treated with the antifungal or the bacterial antibiotic suggest a higher contribution of the fungal community for total microbial biomass and a higher contribution of the bacterial community for microbial respiration rates, especially during the later stages of leaf decomposition. However, these results should be analyzed with caution since both antibacterial and antifungal agents did not totally eliminate microbial activity and biomass.     

Patterns of Seed Size,Germination and Seed Viability of Tropical Tree Species in Southern India1

Seed weight, days to germination and seed viability were observed for 99 species growing in the Western Ghats of Karnataka, India. Seed size was strongly correlated with days to germination; smaller seeds germinated faster than larger seeds. Species which flowered during the rainy season had lighter seeds than species which flowered during the dry season. It was also found that seed size and viability of seeds were related to the season of fruiting. Species which fruit during the rainy season had heavier seeds and shorter viability than species which fruit during the dry season. These flowering and fruiting patterns and varying seed sizes are argued to be adaptations to the time of dispersal, time of moisture availability in the habitat and seedling survival.     

Synopsis of the tribe Hureae (Euphorbioideae, Euphorbiaceae)

A synopsis of the tribe Hureae is presented with nomenclatural updates, a discussion of diagnostic features, and summaries of geographical distributions. This study is based on the analysis of approximately 300 voucher specimens, including collections and photographs of types, in addition to bibliographic documentation. Seventeen species distributed in three genera were recognized: Algernonia (11 species), Hura (2), and Ophthalmoblapton (4). All species are American with the majority distributed within the Atlantic Forest, particularly in southeastern Brazil. A key for the identification of genera and species is provided along with illustrations, information on the geographic distributions and conservation status. Lectotypification for Algernonia leandrii is proposed.     

Seed Size,Seed Germination,and Seedling Survival of Brazilian Tropical Tree Species Differing in Successional Status1

Seed germination and seedling establishment patterns have been used to classify species as shade tolerant or intolerant. The main objective of this research was to investigate, under controlled conditions, seed germination of species from different successional positions as well as to follow seed germination and seedling survival under natural shade in the field. The species studied were Solarium granuloso‐leprosum, Trema micrantha, Cecropia pachystachya, Croton piptocalyx, Bauhinia forficata subsp. pruinosa. Senna macranthera, Schizolobium parahyba, Piptadenia gonoacantha, Chorisia speciosa, Pseudobombax grandiflorum, Ficus guaranitica, Esenbeckia leiocarpa, Pachystroma longifolium, Myroxylon peruiferum, and Hymenaea courbaril. Field trials were carried out at Santa Genebra Municipal Reserve, Campinas, SP, Brazil, at the forest edge and in the understory. No significant correlations were detected between successional status and seed size or seed water content. Light‐regulated germination was present only in small‐seeded species. In field experiments, most species, including the light‐sensitive ones, were able to germinate under the canopy, where a low red/far‐red ratio predominates. Most species, mainly those of early‐ and intermediate successional positions, presented low seedling survival rates under shade. Myroxylon peruiferum was the most shade tolerant species, while 5. granuloso‐leprosum, C. speciosa, P. gonoacantha, F. guaranitica, T. micrantha, and 5. parahyba were the most shade intolerant. These latter species showed little or no survival under the shade conditions.     

Pollen morphology of the Flueggeinae (Euphorbiaceae,Phyllanthoideae)

A total of 129 species from the subtribe Flueggeinae of the tribe Phyllantheae (Euphorbiaceae, Phyllanthoideae) were investigated using light and scanning electron microscopy, and 10 species using transmission electron microscopy, in order to evaluate the relationships between the eight constituent genera: Breynia, Flueggea, Glochidion, Margaritaria, Phyllanthus, Reverchonia, Richeriella, and Sauropus. Of these genera, Flueggea, Margaritaria and Richeriella share pollen with a prolate spheroidal meridional outline and a 3-colporate aperture system. Pollen of Reverchonia is also 3-colporate, but differs from that of the Flueggea alliance by its clearly prolate shape, tilioid ornamentation and absence of costae endopori. Breynia and Sauropus have 4–12 and 3–16-colporate pollen, respectively, with diploporate colpi. Two pollen types are recognised in Breynia, and four in Sauropus, one of which supports the recognition of Sect. Hemisauropus. Glochidion pollen is 3–6-colporate, and similar to that of Breynia in having reticulate sculpture with Y-shaped sexine structures, but it has monoporate colpi. Of the genus Phyllanthus, only species with pollen with diploporate colpi have been studied. Seven types are described. Diploporate Phyllanthus pollen can be distinguished from that of Breynia and Sauropus by its distinct colpus margins consisting of parallel muri. Colpal irregularities and endoaperture configurations in the subtribe are discussed, and pollen morphological trends are hypothesised. Placed in the successiform aperture series, the Flueggea alliance and Reverchonia form a basal group. Glochidion is considered intermediate, giving rise to the Breynia-Sauropus group. The relationship with Phyllanthus remains unclear.     

The Effect of Rust Infection on Reproduction in a Tropical Tree (Faramea occidentalis)1

Fungal pathogens that infect reproductive structures of plants (e.g., flowers and fruits) can reduce the seed production and seedling recruitment of host plants. We report here on the effects of a rust, Aecidium farameae, that infects the ovaries and pedicels of mature flowers on Faramea occidentalis (Ruhiaceae), a small tree common on Barro Colorado Island, Republic of Panama. Rust infection of ovaries reduced the number of maturing fruit on infected trees. Trees with low rust incidence in June of 1992 had 68 percent fruit survival, compared to 17 percent fruit survival for those with high rust incidence. Infected fruits developed abnormally and were usually aborted long before uninfected fruits were mature. One hundred percent of the infected ovaries marked in July were diseased or missing in August. We conclude that infection by A. farameae has the potential to seriously decrease the reproductive output of Faramea occidentalis and may represent an important source of variation in the relative fitness of individual plants.     

Anchorage Mechanics of the Tap Root System of Winter-sown Oilseed Rape (Brassica napus L.)

The anchorage mechanics of mature winter-sown oilseed rape (‘Envol’)were investigated by combining a morphological and mechanicalstudy of the root system with anchorage tests on real and modelplants. Oilseed rape plants were anchored by a rigid tap root;the few laterals all emerged below the centre of rotation ofthe root system (approx. 30 mm below the soil surface). Whenplants were pulled over, the tap root bent and the top 30 mmmoved in the soil towards the direction of pull, creating acrevice on the opposite side. The maximum anchorage moment was2.9 ± 0.36 N m. Two main components of anchorage wereidentified: the bending resistance of the tap root and the resistanceof the soil on the near side to compression. The relative importanceof these components was determined by measuring both the bendingresistance of the tap root, and the resistance of metal tubesof varying diameter, inserted to various depths in the soil,to being pulled over. These tests showed that the tap root bendingmoment at failure could account for around 40% of anchoragemoment, while soil resistance could account for around 60%.The model tests on the tubes also help to shed light on theway in which the dimensions of tap roots will influence theiranchorage capability. Copyright 2001 Annals of Botany Company Anchorage, lodging, root bending resistance, mechanical properties, oilseed rape, Brassica napus L     

The genusPicris (Asteraceae,Lactuceae) in Tropical Africa

Three species of the genusPicris L. are native in Tropical Africa:P. abyssinica Sch. Bip. (Ethiopia),P. xylopoda Lack, spec. nova (Nigeria, Ethiopia) andP. humilis DC. (Senegal, Mali). There are indications that the two perennial species,P. abyssinica andP. xylopoda, are related to and have evolved from a primitive Central Asiatic stock in a manner parallel to many African species ofCrepis L.P. humilis, on the other hand, is a small annual plant with a high number of derived characters. The introduced species of European origin growing south of the Sahara are briefly mentioned.     

Isozyme Variation in a Tropical Pioneer Tree Species (Cecropia obtusifolia,Moraceae) with High Contents of Secondary Compounds1

We define 48 allozyme loci for a tropical pioneer tree species, Cecropia obtusifolia Bertol, which has high contents of secondary compounds. Our goals were to find the effects of extraction procedures on artifacts and variation in resolution of enzyme banding patterns; to explore the relationship among the variation of the loci sampled and the enzymes' molecular structure, metabolic function and substrate; to obtain estimates of the genetic variation in this species at Los Tuxtlas rain forest (México) and to explore the variation of allelic frequencies in six successive life-history stages of the species. The resolution of the isozymes bands and the actual banding pattern varied with the type and age of tissue, the collection and storage procedures, the extraction buffer, and other loading and running procedural details. However, artifactual variation was eliminated with a new extraction buffer for species with high contents of secondary compounds. Of the 26 enzymes resolved for C. obtusifolia, we found that enzymes with a greater number of substrates and an oligomeric quaternary structure tended to be more variable than their counterparts, but the relationship was not statistically significant. The proportion of polymorphic systems varied significantly with the metabolic pathway and the function of the enzymes. Enzymes involved in starch synthesis are significantly more variable (p < 0.05) than all others, except those involved in amino acids metabolism and the proportion of polymorphic enzymes is also significantly associated with the hnction of the enzyme, the hydrolases and isomerases are significantly more variable than lyases and oxidoreductases enzymes. The percentage of polymorphic loci for C. obtusifolia was estimated at P = 27.1%. The effective number of alleles was estimated at ne = 1.3 and ne = 2.4 for all loci and only polymorphic ones respectively and the average heterozygosity (H) for all 48 loci was estimated at H = 0.05. Allele frequencies varied throughout the life-cycle of the species, with significant differences for some alleles and loci among some life-cycle stages. “Tree seeds” allele frequencies differ significantly (P < 0.05) from “rainy dispersed seeds” in 7 of 8 loci and from “soil seeds” in Six of eight loci. Allele frequencies of all three seed categories (“tree seeds”, “rainy dispersed seeds”, and “soil seeds”) differed strongly from established individuals (seedlings, juveniles and adults), while allele frequencies of established individuals are relatively similar to one another. Seedling allele frequencies at most loci were also significantly different from those found in seeds collected from trees, seed-rain, and soil. Two alleles (at GOT-2 and FE-2) were only found in soil seeds and one allele (at LAP-2) was only found in seedlings.     

The Species and Size Composition of Tropical Freshwater Zooplankton with Special Reference to the Oriental Region (South East Asia)

A contrast between temperate and tropical Cladocera, Copepoda and Rotifera zooplankton fauna shows a markedly different size composition and species diversity. There are fewer species, and these are generally smaller in the tropics than in either the North or South Temperate Zones. Larger species of the genera Daphnia and Simocephalus are relatively rare in the tropics, while members of Eurycerus and Saycia are absent. Members of the families Holopedidae, Leptodoridae and Polyphemidae increase in abundance toward the temperate regions. Two small cyclopoid copepods are common in tropical zooplankton, while many of the common larger species of other regions are absent or very rare. The chief rotifer components are Keratella tropica and species of Brachionus.     

Root Morphology and Anchorage of Six Native Tree Species from a Tropical Montane Forest and an Elfin Forest in Ecuador

Root architecture of tree species was investigated at two different altitudes in tropical forests in Ecuador. Increasing altitude was accompanied by higher wind speeds and more shallow soils, while slope angles of both sites were comparable (20–50°). Three tree species typical for the montane forest at 1900 m (Graffenrieda emarginata (Ruiz & Pav.) Triana (Melastomataceae), Clethra revoluta (Ruiz & Pav.) Spreng. (Clethraceae), Vismia tomentosa Ruiz & Pav. (Clusiaceae)) and for the elfin forest at 3000 m (Weinmannia loxensis Harling (Cunoniaceae), Clusia spec. (Clusiacaea) Styrax foveolaria Perkins (Styraceae)) were examined. At 1900 m, 92% of the trees grew upright, in comparison to 52% at 3000 m. At 3000 m, 48% of the trees were inclined, lying or even partly uprooted. At this altitude, all trees with tap roots or with shoots connected by coarse rhizomes, 83% of the trees with stilt roots, and 50% of the trees in which stems or roots were supported by other trees grew upright, suggesting that these characteristics were relevant for tree stability. Root system morphology differed markedly between altitudes. In contrast to 1900 m, where 20% of structural roots originated in the deeper mineral soil, root origin at 3000 m was restricted to the forest floor. The mean ratio of root cross sectional area to tree height decreased significantly from 6.1 × 10⁻³ m² m⁻¹ at 1900 m to 3.2 × 10⁻³ m² m⁻¹ at 3000 m. The extent of root asymmetry increased significantly from 0.29 at 1900 m to 0.62 at 3000 m. This was accompanied by a significantly lower number of dominant roots at 3000 m (2.3 compared to 3.8 at 1900 m). In conclusion, native tree species growing in tropical montane and elfin forests show a variety of root traits that improve tree stability. Root system asymmetry is less important for tree stability where anchorage is provided by a deep and solid root–soil plate. When deep rooting is impeded, root traits improving the horizontal extension of the root–soil plate are more pronounced or occur more frequently. Furthermore, mutual mechanical support of roots and stems of neighboring trees seems to be an appropriate mechanism to provide anchorage in soils with low bulk density and in environments with high wind speeds.     

Reiteration in the Monodominant Tropical Tree Dicymbe corymbosa (Caesalpiniaceae) and its Potential Adaptive Significance

The tropical monodominant tree Dicymbe corymbosa reiterates via epicormic shoots and roots, resulting in multistemmed trees with complex pseudotrunks and root mounds. In 2 ha of primary forest on the Guiana Shield, we quantified the reiterative structure and aboveground soil development of 307 D. corymbosa individuals ≥ 10 cm dbh and investigated the potential adaptive significance of reiteration in terms of genet persistence and root exploitation of aboveground soil accumulations. We also investigated the incidence of the heart rot fungus Phellinus robustus in D. corymbosa and examined its relationship to the reiteration process. Large trees contained more and larger reiterations, greater trunk, root mound, and organic soil volumes, and a higher incidence of Phellinus than smaller trees. Roots and ectomycorrhizas were abundant in aboveground soils on the trees, occurred at higher densities than those of the surrounding forest floor, and may be important in recycling mineral nutrients. Stem turnover and reiteration were associated with Phellinus heart rot and appeared to be cumulative over time, resulting in persistent, structurally complex trees of indeterminate lifespan. Dicymbe corymbosa provides a rare example of a tree species that exploits both persistence and recruitment niches, as it successfully recruits through mast fruiting.     

Hybridization, polyploidy, and the evolution of sexual systems in Mercurialis (Euphorbiaceae)

Hybridization and polyploidy are widely believed to be important sources of evolutionary novelty in plant evolution. Both can lead to novel gene combinations and/or novel patterns of gene expression, which in turn provide the variation on which natural selection can act. Here, we use nuclear and plastid gene trees, in conjunction with morphological data and genome size measurements, to show that both processes have been important in shaping the evolution of the angiosperm genus Mercurialis, particularly a clade of annual lineages that shows exceptional variation in the sexual system. Our results indicate that hexaploid populations of M. annua, in which the rare sexual system androdioecy is common (the occurrence of males and hermaphrodites) is of allopolyploid origin involving hybridization between an autotetraploid lineage of M. annua and the related diploid species M. huetii. We discuss the possibility that androdioecy may have evolved as a result of hybridization between dioecious M. huetii and monoecious tetraploid M. annua, an event that brought together the genes for specialist males with those for hermaphrodites.