Regional interpretation of environmental gradients which influence Trachypogon savannas in the Orinoco Llanos

Multivariate structural data describing Trachypogon savanna relationships were used to characterize the sources of variation among savannas as a function of the environmental characteristics of the Orinoco Llanos. Twenty-six savannas including major physiognomic types of the Orinoco Llanos ranging from herbaceous to bush island savannas were ordinated on a regional scale. Results of Detrented Canonical Correspondence Analysis (DCCA) using floristics variables indicated that savannas were ordinated along two complex gradients. A soil physical and chemical gradient as expressed by changes in bulk density and magnesium concentration were evident after the analysis of the first DCCA axis. The second axis was a climatic gradient of decreasing annual precipitation and incresing monthly precipitation during the dry season. The impact of human disturbance on the savanna composition was also an explanatory variable of the second DCCA axis. Savanna sites overlapped considerably in composition and most species were widely distributed, with aboveground phytomass abundance depending upon the site.     

Restoration of Midwestern U.S. Savannas: One Size Does Not Fit All

Lowland savannas are a rare variant of Midwestern United States savanna occurring on alluvial soils, for which reference information is sparse. To evaluate the appropriateness of using upland savanna as a surrogate source of reference information for lowland savanna, we studied a pre‐Euro‐American lowland savanna using original U.S. Public Land Survey data and other historical records. Historical vegetation was reconstructed and compared among upland savannas, lowland savannas, and lowland forests; we also evaluated potential disturbance dynamics maintaining these systems. We found that all three communities were dominated by members of the genus Quercus but also had extensive representation by many other tree species, especially notable for savannas in this region. There were no clear size–density relationships for species in the genus Quercus, indicating that these historical savannas were not characterized exclusively by large, scattered oak trees but rather by trees of many oak species and nonoak species in a wide range of size classes. Both upland and lowland savannas also contained a substantial shrub component. We found no evidence that lowland savannas were maintained by flooding, although the uneven‐aged canopy structure suggested that periodic disturbance occurred. Restoration of lowland savanna in this region should include provisions for maintaining nonoak species and shrubs, with disturbance timed to maintain an uneven‐aged canopy structure. Although the appropriateness of historical data in the face of climate change may be questionable, in this region, a warmer climate may actually help promote the “oak parkland” that was present from 8,000 BP up to Euro‐American settlement.     

The external morphology and distribution of cuticular hair organs on the claws of the American lobster, Homarus Americanus (Milne-Edwards)

Four classes of microscopic cuticular hair organs were found on the chelae of Homarus americanus (Milne-Edwards). Type I and Type II organs possess long single sensillae, 30–60 μm and 70–130 μm, respectively. Type III organs are toroid bumps 20–30 μm in diameter, with a small tuft of fibers projecting from the center. Type IV organs are small conical hairs ≈ 1 μm in length.Type IV organs were uniformly distributed over the claws. The distributions of Types I, II, and III organs were analysed through multivariate analysis of variance (MANOVA) with respect to claw, side, and area. Type I organs showed significance of the three-way interaction only. Type II organs showed significance of side, area, and the claw by area interaction. Type III organs showed significance of area, and all interactions. Changes in number and density of Types I, II and III organs, and in number of Type IV organs, were found with growth.     

Effects of fire and defoliation on the life history of native and invader C4 grasses in a Neotropical savanna

African grasses, introduced into Neotropical savannas to improve forage quality, have spread successfully and displaced native plants. To understand their competitive relationships, we compared biomass production and allocation, plant architecture and phenology, net photosynthesis (Pn), water relations, and nutrient content under fire and simulated herbivory between two C₄ grasses, the native Trachypogon plumosus and the introduced Hyparrhenia rufa from a seasonal savanna in Venezuela. All variables were strongly influenced by the rainfall regime. Hyparrhenia produced bigger plants (in mass and size) with a large proportion of mass (>75%) allocated to leaves and culms. Its biomass production was more affected by fire than by defoliation. In contrast, Trachypogon was more affected by defoliation than by fire which promoted a flush of leaf growth even in the dry season. Fire caused up to 85% mortality in Hyparrhenia but none in Trachypogon where it increased inflorescence production. However, fire promoted abundant seed germination and fast seedling growth in Hyparrhenia, enabling it to colonize new areas. During the growing season Trachypogon had higher Pn and lower leaf water potential (Ψ) than Hyparrhenia but differences among treatments were not significant for either grass. Pn of Trachypogon ceased at a lower Ψ (−3.0 MPa) than in Hyparrhenia (−2.0 MPa), indicating its higher tolerance to water stress. During the dry season, Trachypogon leaves remained alive and retained low Pn. Leaf nutrient content was higher during the rainy season in both species. Differences in Pn could not explain the higher seasonal biomass production of Hyparrhenia. However, its water stress evasion strategy, larger biomass allocated to leaves, abundant germination and fast seedling growth appeared to be responsible for the success of Hyparrhenia as an invader of Neotropical savannas. Received: 17 August 1998 / Accepted: 3 March 1999     

Dynamics of energy and nutrient concentration and construction cost in a native and two alien C4 grasses from two neotropical savannas

In Venezuela, the alien grasses Melinis minutiflora Beauv. and Hyparrhenia rufa (Nees.) Stapf tend to displace the native savanna plant community dominated by Trachypogon plumosus (Humb. and Bonpl.) Nees. This occurs in either relatively wetter and fertile highland savannas or in drier and less fertile lowland savannas. Although the native and aliens are perennial C₄ grasses, higher net assimilation leaf biomass per plant and germination rate of the latter are some causes for their higher growth rates and for their competitive success. The objective of this study is to compare seasonal tissue energy, N, P and K concentrations and the calculated construction costs (CC) between the native grass and either one of the alien grasses from lowland and highland savannas. We predict that, in order to out-compete native plants, alien grasses should be more efficient in resource use as evidenced by lower tissue energy and nutrient concentrations and CC.Tissue energy and nutrient concentration were measured throughout the year and compared between M. minutiflora and the co-occurring local population of T. plumosus in a highland savanna and between H. rufa and its neighbor local population of T. plumosus in a lowland savanna. CC was calculated from energy, N and ash concentrations considering ammonium as the sole N source. Differences between co-occurring species, T. plumosus populations, seasons, and organs were analyzed with ANOVA.Highland and lowland grasses differed in concentration and allocation of energy and nutrients whereas the differences between alien and native grasses were specific for each pair considered. Highland grasses had higher energy, N, P and CC than lowland grasses. These variables were always lowest in the culms. In the more stressed lowland site, tissue energy and nutrient concentrations decreased significantly during the dry season except in the roots of both grasses which had the highest energy and nutrients concentrations during the drought. This seasonal response was more marked in the local lowland population of T. plumosus in which maximum CC alternated seasonally between leaves and roots. Energy and nutrient concentrations and CC were the lowest in H. rufa. In the lowland savannas, the higher efficiency of resource use in the invader grass contributes to its higher competitive success through increased growth rate. In the highlands, overall tissue energy concentration and CC, but not N nor P concentration, were lower in the fast growing M. minutiflora but seasonal differences were lacking. The higher leaf CC in T. plumosus can be attributed to the higher proportion of sclerenchyma tissue which is more expensive to construct. Considering CC, both fast growing alien grasses are more efficient in resource use than the co-occurring native grass. However, the role of CC explaining the competitive success of the former, through higher growth rates, is more evident in the more stressful environment of the lowland savanna.     

Spatial and temporal variability of fires in relation to ecosystems,land tenure and rainfall in savannas of northern South America

Fire is a predominant factor forcing global terrestrial biomass dynamics, with more than 30% of the land surface showing frequent burning, particularly in the tropics, where it mostly affects savannas ecosystems annually. Savannas, which cover approximately 269 million ha in South America, play a major role in the global carbon cycle. They are affected by increasing human pressures and global climate change. Using satellite data, this study quantifies vegetation burning in the Colombian Llanos savannas for the period 2000–2008, and analyzes how fire spatial pattern, frequency and extent vary with ecosystem type, land tenure and rainfall. On average 2.75±0.5 million ha (24±4.2%) of the savannas burn each year. Burned area is highly variable, with 3.4 million ha burned in 2002–2003 and <1.9 million ha in 2005–2006. However, during the 2000–2008 period near of 3.7 million ha (33.5%) of the savannas never burned. Compared with the average 8–10 years of fire return time for the tropics and subtropics, these savannas burn twice as often. In addition, the average burn size figure for tropical and subtropical grassland savannas (with <5% trees) of 7000 ha (median 5000 ha), is about seven times the average burned patch size we found in our study. Fires predominate in the well‐drained high plain savannas, lowest figures occurring along the Andean foothills, in forested areas and in pasture and croplands. Annual proportion burned varies with land tenure, being highest in National Parks. This study is the first complete regional map of fire disturbance in a South American savanna. This detailed regional data provides a unique opportunity for increasing the accuracy of global carbon emission calculations.     

Floristic composition,plant species abundance,and soil properties of montane savannas in the Gran Sabana,Venezuela

Floristic composition, species abundance, and soil properties were studied in slope, flat and disturbed savannas in the northern part of the Gran Sabana, Venezuela. All savannas presented shallow soils (<30 cm depth) with high content of sand and low content of clay. In general, the soils were poor in nutrients and strongly acidified. The major difference between the soils was the content of the stony fraction, which was significantly higher (P<0.05) in the slope savannas than in the flat savannas. A total of 57 dicot, 42 monocot, and 7 fern species were recorded in all studied savannas. In the flat and slope savannas predominated the monocot species, while in the disturbed savanna predominated the dicots. The families with the largest number of species were Poaceae (19.8%), Cyperaceae (13.2%), Asteraceae (10.4%) and Melastomataceae (8.5%). The number of species in the flat savannas was higher than that of the slope savannas. The lowest plant species richness was associated to slope savannas and their high content of stony fraction of the soils. The highest floristic similarity was found between slope savannas, and the lowest between disturbed savanna and slope savannas. The most abundant life forms in the studied savannas were perennial (42.4%) and annual (24.5%) herbs, followed by suffruticoses (16.0%) and shrubs (12.3%), and the less frequent was lianas (4.7%). The disturbed savanna showed the higher richness and diversity index. Trachypogon plumosum (Poaceae) was the most abundant species in all studied savanna.     

Distribution and vegetation dynamics of humid savannas in Africa and Asia

Abstract. A review is presented on the literature about the distribution of savannas in humid climates in Africa and Asia and their vegetation dynamics. Sections are devoted to African lowland and montane savannas (the latter divided into southern, eastern, western and northern African), Madagascar, Indian subcontinent, SE Asia and New Guinea. It is concluded that the extension of savannas under humid climatic conditions and the relation to the distribution of forests is a function of cultivation, grazing by domestic and wild animals, present and previous climate, geomorphology and soil characteristics. Once established, savannas are often maintained by fires, both natural and man-made. Montane savannas are generally brought about by man's clearing, cultivation and burning. Fire is a stochastic variable; it creates an ecotone sensu stricto (an environmentally stochastic stress zone) at the forest/savanna border. On the other hand, if geomorphology and soil are the determinants, the transition between forest and savanna would have the character of an ecocline (a gradient zone) with fundamentally different conditions. In humid African lowland climates forests expand into savannas if the latter are not maintained by man. Whether forests also expand in less humid climates is disputed. In montane areas forest expansion may be delayed on degraded soils and when diaspores are lacking.     

Nuclear ribosomal internal transcribed spacer 1 (ITS1) variation in the Anastrepha fraterculus cryptic species complex (Diptera,Tephritidae) of the Andean region

The nuclear ribosomal internal transcribed spacer 1 (ITS1) was sequenced for Anastrepha fraterculus (Wiedemann, 1830) originating from 85 collections from the northern and central Andean countries of South America including Argentina (Tucumán), Bolivia, Perú, Ecuador, Colombia, and Venezuela. The ITS1 regions of additional specimens (17 collections) from Central America (México, Guatemala, Costa Rica, and Panamá), Brazil, Caribbean Colombia, and coastal Venezuela were sequenced and together with published sequences (Paraguay) provided context for interpretation. A total of six ITS1 sequence variants were recognized in the Andean region comprising four groups. Type I predominates in the southernmost range of Anastrepha fraterculus. Type II predominates in its northernmost range. In the central and northern Andes, the geographic distributions overlap and interdigitate with a strong elevational effect. A discussion of relationships between observed ITS1 types and morphometric types is included.     

Submillennium-scale migrations of the rainforest-savanna boundary in Colombia: C wiggle-matching and pollen analysis of core Las Margaritas

We present the 11?150-cal-yr-long pollen record Laguna Las Margaritas (3°23′N, 73°26′W; 290 m altitude), located at a site sensitive to climatic change near the transition from the Amazonian rainforest to the savanna of the Llanos Orientales in Colombia. In the 10-m-long core nine AMS ¹⁴C bulk samples show ages from 9760±60 to 854±36 BP and provide initial time control. Thirty-one additional AMS ¹⁴C samples of selected macrofossils provide time control from 6250 to 4050 cal BP; for this interval precise time control was obtained by ¹⁴C wiggle-match dating. From 11?150 to 9100 cal BP, grass savanna dominated the landscape while gallery forest along the drainage system was poorly developed. Water availability was lower than today and the length of the dry season longer. From 9100 to 7330 cal BP gallery forest expanded pointing to wetter conditions. From 7330 to 5430 cal BP savanna was increasingly replaced by forest, but with alternating abundance of both vegetation types. From 5430 to 2500 cal BP forest and wooded savanna dominated the western Llanos Orientales suggesting high precipitation rates. Expansion of Mauritia palm forest at 2500 cal BP reflects increasing water availability and stagnant water environments. Increasing savanna between 2500 and 1000 cal BP may represent a combined natural and anthropogenic signal. Development from open savanna to forest during the Middle Holocene is synchronous with a decreasing caloric seasonality, and a southward migration of the Intertropical Convergence Zone, suggesting that the large-scale climatic and vegetational change in the Colombian savannas is precession-forced. High-frequent migration of the savanna-rainforest boundary started around 6000 cal BP and continued at least to 3000 cal BP. Precipitation regimes in northwest and northeast South America seem opposite: dry conditions in the Colombian savanna area seem to reflect a La Niña setting of the climate system; wet conditions reflect an El Niño setting. Δ¹⁴C fluctuations reflect changes of solar activity and we tested the hypothesis that Δ¹⁴C fluctuations correspond to climatic and vegetational change. For the interval 6250 to 4050 cal BP, we applied ¹⁴C wiggle-match dating (WMD), i.e. matching a series of radiocarbon ages in ¹⁴C BP from the sediment, with the dendrochronology-based calibration curve, and we also wiggle-matched the pollen record vs. the Δ¹⁴C record. In this first attempt in a neotropical ecosystem, we could not find unambiguous support that changes in solar activity did trigger climatic and vegetational change in the savannas of the Llanos Orientales.     

The effect of environmental heterogeneity on clonal behaviour of Prunella vulgaris L.

Melinis minutiflora Beauv. (Poaceae) is an African grass that is invading mid-elevation Trachypogon savannas in Venezuela. The objective of this study was to investigate the influence of soil fertility, competition and soil disturbance in facilitating Melinis' invasion and growth in these savanna sites. We manipulated soil fertility by adding nitrogen (+N), phosphorus and potassium (+PK), or nitrogen, phosphorus, and potassium (+NPK). We simultaneously manipulated the competitive environment by clipping background vegetation. In a separate experiment, we mechanically disrupted the soil to simulate disturbance. We hypothesized that germination and growth were bottlenecks to early establishment in undisturbed savanna, but that disturbance would alleviate those bottlenecks. We measured Melinis seed germination and subsequent establishment by adding seeds to all plots. We examined Melinis growth by measuring biomass of Melinis seedling transplants, 11 months after they were placed into treatment plots. Germination and establishment of Melinis from seed was extremely low. Of the 80,000 seeds applied in the experiment, only 28 plants survived the first growing season. Mortality of Melinis seedling transplants was lowest in PK fertilized plots, but in the absence of PK mortality increased with N additions and clipping. By contrast, fertilization of the savanna with NPK greatly increased Melinis seedling biomass and this effect was greatly enhanced when competition was reduced (e.g. clipping). Melinis transplant growth responded strongly to soil disturbance- a response not fully explained by removal of competitors (clipping) or changes in soil nutrients and moisture. We suspect that disruption of the soil structure allowed for greater root proliferation and subsequent plant growth. We believe that native savanna is relatively resistant to Melinis invasion, since Melinis seedlings persisted in intact savanna but exhibited little or no growth during the first year. The significant enhancement of Melinis seedling growth with clipping and nutrient additions suggests that low soil nutrients and the presence of native savanna species are important factors in the ability of native savanna to resist Melinis establishment. However, the potential for Melinis growth increases enormously with soil disturbance.     

Plant introduction,naturalization, and invasion in French Guiana (South America)

Continental tropical ecosystems are generally viewed as less vulnerable to biological invasions than island ones. Their apparent resistance to invasive alien species is often attributed to their higher native biota diversity and complexity. However, with the increase of human activities and disturbances and the accelerate rate of introductions of plant species, these apparently resilient continental ecosystems are now experiencing alien plant naturalization and invasion events. In order to illustrate this emergent phenomenon, we compiled a list of all known introduced and naturalized plant species in French Guiana (Guiana Shield, South America). A total of 490 alien plants were recorded, about 34% of which are currently naturalized, mainly species belonging to the Acanthaceae and Fabaceae (Faboideae) in the Eudicotyledons, and Poaceae (grasses) and Arecaceae (palms) in the Monocotyledons. The coastal dry and wet savannas appears to be vulnerable to plant invasion (with 165 naturalized species, about 34% of the alien flora), especially by Acacia mangium (Mimosaceae) and Melaleuca quinquenervia (Myrtaceae) which are forming localized but dense monotypic stands. Both tree species, intentionnally introduced for reforestation, rehabilitation, and as garden ornamentals and have the potential to spread with increasing human disturbances The number and abundance of naturalized alien plants in the relatively undisturbed tropical lowland rainforests and savannas remains still very low. Therefore, surveillance, early detection, and eradication of potential plant invaders are crucial; moreover collaboration with neighbouring countries of the Guiana Shield is essential to prevent the introduction of potentially invasive species which are still not present in French Guiana.     

Changes in plasma estradiol, progesterone and LH level in the menstrual cycle of the adult female Japanese monkey during the mating season

Adult 15 female Japanese monkeys showing regular menstrual cycles were subjected to the daily blood sampling for the measurement of estradiol (E2), progesterone (P) and biological LH in the mating season. Monkeys were maintained under controlled conditions in a standardized environment. Of the 35 cycles observed, 18 (51.4%) were estimated as anovulatory cycles and 17 (48.6%) were ovulatory cycles. The anovulatory cycles were classified into three types according to the peak level of E2 (Type I: E2 less than 50 pg/ml 3 cycles, Type II: E2 less than 170 pg/ml 7 cycles, Type III: E2 greater than 170 pg/ml 8 cycles). The ovulatory cycles were classified into two Types according to the peak level of P (Type IV: P less than 5.0 ng/ml 5 cycles, Tyep V: P greater than 5.0 ng/ml 12 cycles). The menstrual cycle was 27.5 +/- 7.8 days. The differences between mid cycle LH surge and P level in Type IV and in Type V were statistically significant. It was revealed that female Japanese monkeys kept under controlled condition in the mating season showed high incidence of various types of anovulatory cycles and that the ovulatory cycles with low P elevation in the mid luteal phase showed low LH and P secretions on the mid cycle date.     

Inhibition of plastid division by ampicillin in the pteridophyte Selaginella nipponica Fr. et Sav

We investigated the effect of the beta-lactam antibiotic, ampicillin, on plastid division in the pteridophyte Selaginella nipponica. Guard cells of plantlets treated with 1 mM ampicillin only often had one plastid, whereas guard cells of untreated plantlets had two to four plastids. We generated a S. nipponica cell culture system and used it to investigate the effects of ampicillin. Treatment with 1 mM ampicillin had no effect on cell division in culture. We classified cultured cells into four types based on the number of plastids they contained: one (Type I), two (Type II), three or four (Type III) and more than five (Type IV). After 3 d in culture, the percentage of each cell type (I-IV) was 29.5, 46.7, 20.9, and 1.9%, respectively. Subsequently, the percentage of Types III and IV increased gradually, reaching 61.9 and 11.4%, respectively, after 15 d in culture in the absence of ampicillin. When 1 mM ampicillin was added, there was a minimal increase in the number of Type III and IV cells, with high percentages of Type I and II cells (32.4 and 45.7%, respectively) after 15 d. These results suggest that ampicillin inhibits plastid division in S. nipponica.     

Strategic procurement of fish by the pumé: A South American “fishing culture”

Information is presented on fishing effort, efficiency, techniques, and catch composition for Pumé men, women, and children along with a conceptual model of fishing as a food procurement strategy. The Pumé are a native lowland South American group living in the topical savanna region of southwestern Venezuela characterized by seasonal flooding. Results are discussed in relation to the Pumé environmental and social situation, and briefly compared with results from other lowland South American groups.     

A whole range of fine structural criteria for immunohistochemically identified LH cells in rats

Summary Pituitaries from normal, young and adult male rats were fixed either in sublimate-formalin or in glutaraldehyde-osmium. In adjacent Paraplast sections, almost all the gonadotrophs were immunostained with both LH and FSH antisera. The rat LH and FSH antisera used were shown to be highly specific by the absorption test and by double antibody radioimmunoassay. Thin and thick adjacent Epon sections were prepared for EM and immunohistochemical examination. Cells stained with the rat LH antiserum were identified by LM, and then observed in detail by EM. On the basis of these observations we suggest that the LH cells are arranged in a sequence of basophils, i.e., Types II/III, III, III/IV and IV: Type II/III basophils are elongate with a cytoplasmic process and less vesiculated. They have morphological features of Type II (classical thyrotrophs) and also of Type III basophils. Type III basophils are oval in shape and moderately vesiculated. Both Types II/III and III basophils can be divided into two classes of cell characterized mainly by the existence of only small secretory granules (150–220 nm in diameter) (Type A) or by the coexistence of small and large (350–500 nm) (Type B). Type III/IV basophils are cells intermediate between types III and IV basophils, and moderately vesiculated with an abundance of secretory granules (150–300 nm in diameter). Type IV basophils are large, spherical or oval cells whose RER cisternae are conspicuously dilated; they contain less numerous secretory granules (150–300 nm in diameter). It is concluded that LH cells are not a single cell type, but include a wide range of subtypes.     

Phylogeny of Chaetanthera (Asteraceae: Mutisieae) reveals both ancient and recent origins of the high elevation lineages

Penalized likelihood analysis of previously published chloroplast DNA (cpDNA) ndhF sequences suggests that the central-southern Andean genus Chaetanthera diverged ca. 16.5 million years (my) ago, well before the uplift of the Andes to their present heights. Penalized likelihood analysis based on new nuclear ribosomal DNA (rDNA) internal transcribed spacer (ITS) sequences indicates that the most relictual lineages occupy high elevation Andean habitats that did not exist until some 10my later. This result is contrary to the expectation that younger habitats should be occupied by phylogenetically younger lineages. The results are interpreted with respect to the development of aridity in lowland habitats during the Miocene and Pliocene, which presumably extinguished the lowland relatives of the high elevation taxa or, in effect, forced them upwards in search of adequate moisture. As the more northerly lineages were being displaced upward, others diversified in the mediterranean-type climate area of central Chile, giving rise to additional high elevation taxa again, at an early date, as well as lowland taxa. Some species of Chaetanthera from lowland central Chile appear as the phylogenetically youngest taxa, suggesting secondary adaptation to lowland aridity. At the same time, at least two high elevation species, Chaetanthera peruviana and Chaetanthera perpusilla, appear to have been derived recently from a lower elevation ancestor, while some middle to low elevation taxa seem to have evolved recently out of a high elevation complex. The results suggest that the younger high elevation habitats have served as both "cradle" and "museum" for Chaetanthera lineages.     

Late Holocene history of savanna gallery forest from Carimagua area, Colombia

The pollen record of a 65cm long core Laguna Carimagua-Bosque (4 degrees 04'N, 70 degrees 13'W) shows the late Holocene environmental history from a lake located within the gallery forest of the savannas of the Llanos Orientales of Colombia. Nine AMS radiocarbon dates of the organic deposits show that the core represents the period from ca. 1300(14)CyrBP to the present. The lake evolved from an active drainage system.During the period from ca. 1300 to 875(14)CyrBP (zone CMB-Ia), Mauritia-dominated swamp and gallery forest was present, dominated by Cecropia, and later also Acalypha and Alchornea. From 875 to 700(14)CyrBP (zone CMB-Ib), the lake was completely surrounded by gallery forest. Mauritiella and Cecropia occurred around the lake. Cecropia pioneer forest reached its greatest abundance and became gradually replaced by a more species-rich gallery forest, including Acalypha, Alchornea, Euterpe/Geonoma, Moraceae/Urticaceae, Piperaceae, and Virola. From 700 to 125(14)CyrBP (zone CMB-II), Cecropia lost its dominant role, and Mauritiella palms became more frequent. The main vegetation categories were swamp forest, gallery forest, understory elements, savanna shrubs and trees, and grass savanna. From 125(14)CyrBP to recent (zone CMB-III), the plant diversity in the gallery forest became highest, Mauritiella became very abundant, and among the savanna elements, woody Didymopanax increased.Comparison of four pollen records from savanna sites shows that pollen of savanna vegetation is markedly underrepresented in lake sediments when the lake lies within the gallery forest. As most of the drainage system of a savanna is hidden by gallery forest, we also expect a significant underrepresentation of the savanna ecosystem in river-transported pollen assemblages.     

Seasonal diet composition in the Crested bobwhite in savannas of central-eastern Venezuela

The diet of Crested Bobwhite (Colinus cristatus, Phasianidae) in a non-flooded savanna located in central-eastern Venezuela was studied. Food items obtained from crops were sorted, counted, and identified mostly to species level. Birds are omnivorous and their diets comprised mostly seeds, some invertebrates, and green vegetation; besides, they consumed snail shells during the breeding season. Seeds dominated in the diet during most of the year but the relative use of seeds, invertebrates, and green vegetation changed seasonally. Some management guidelines in order to increase the abundance of this important game bird in neotropical savannas are proposed.     

Nutrient Limitation to Primary Productivity in a Secondary Savanna in Venezuela1

We examined nutrient limitation to primary productivity in a secondary savanna in the interior branch of the Coastal Range of Venezuela, which was converted from forest to savanna more than 100 years ago. We manipulated soil nutrients by adding nitrogen (+N), phosphorus and potassium (+PK), and nitrogen, phosphorus, and potassium (+NPK) to intact savanna. Eleven months after fertilization, we measured aboveground biomass and belowground biomass as live fine roots in the top 20 cm of soil, and species and functional group composition in response to nutrient additions. Aboveground biomass was highest in the NPK treatment ([mean g/m2]; control = 402, +N = 718, +PK = 490, +NPK = 949). Aboveground production, however, appeared to be limited primarily by N. Aboveground biomass increased 78 percent when N was added alone but did not significantly respond to PK additions when compared to controls. In contrast to aboveground biomass, belowground biomass increased with PK additions but showed no significant increase with N (depth 0–20 cm; [mean g/m2]; control = 685, +N = 443, +PK = 827, +NPK = 832). There was also a 36 percent increase in root length with PK additions when compared to controls. Whole savanna shoot:root ratios were similar for control and +PK (0.6), while those for +N or +NPK fertilization were significantly higher (1.7 and 1.2, respectively). Total biomass response (above + belowground) to nutrient additions showed a strong N and PK co‐limitation ([mean g/m2]; control = 1073, +N = 1111, +PK = 1258, +NPK = 1713). Aboveground biomass of all monocots increased with N additions, whereas dicots showed no response to nutrient additions. Trachypogon spp. (T. plumosus+T. vestitus) and Axonopus canescens, the two dominant grasses, made up more than 89 percent of the total aboveground biomass in these sites. Trachypogon spp. responded to NPK, whereas A. canescens, sedges, and the remaining monocots only responded to N. Even though nutrient additions resulted in higher aboveground biomass in N and NPK fertilized plots, this had little effect on plant community composition. With the exception of sedges, which responded positively to N additions and increased from 4 to 8 percent of die plant community, no changes were observed in plant community composition after 11 months.