Vegetational change in the Balearic Islands (Spain) during the Holocene

New information about the vegetational history of the Balearic Islands during part of the Holocene derives from the results of a pollen study of a 20‐metre‐deep core from Cala Galdana on the SW coast of Minorca.

The lower part of the pollen diagram at the beginning of the Atlantic Period records significant quantities of pollen of Corylus and also local taxa indicating swamp vegetation. In the middle part, Corylus is greatly reduced, AP percentages decrease appreciably and Olea and Ericaceae reach important values, while Plantago Poaceae and Cerealia increase significantly. In the upper part, a rise in Chenopodiaceae occurs, during which a slight increase of the arboreal taxa takes place with greater dryness. Most recently, vegetation changes reflect human activity.     

Effect of vegetation data collection strategies on estimates of relevant source area of pollen (RSAP) and relative pollen productivity estimates (relative PPE) for non-arboreal taxa

Thirteen surface moss samples were collected for pollen analysis from an area of heathland in western Norway. Vegetation composition at different distances around the sampling locations was measured using three different survey methods; rooted frequency within a sub-divided 1 m × 1 m quadrat, visual estimates of cover within a 1 m × 1 m quadrat and a modified form of the ‘circle-walking method’. Extended R-value analysis was used to explore the pollen–vegetation relationships for five main taxa, Calluna vulgaris, Vaccinium-type, Cyperaceae, Poaceae and Potentilla-type. The estimates of relevant source area of pollen obtained were similar regardless of the vegetation survey method. Values obtained were always under 4 m. However, estimates of relative pollen productivity and the background pollen component (proportion of pollen coming from vegetation growing beyond the relevant source area of pollen) differ markedly depending on the method of vegetation survey chosen. This has important implications for the quantitative reconstruction of past vegetation cover.     

Coastal dune vegetation and pollen representation in south Buenos Aires Province, Argentina

Aim To establish the relationship between coastal dune vegetation and its pollen representation as an aid to interpret Holocene vegetation dynamics and environmental changes from pollen assemblages. Location The study area is situated on the temperate Atlantic coast of south Buenos Aires Province, Argentina (c. 39° S and 61°20′ W). Methods The vegetation of the active dune area adjacent to the beach was described on the basis of its floristic composition from 25 plots. Classification of the vegetation into distinct zones was carried out by cluster analysis. Surface samples were collected from each vegetation stand and analysed for their pollen composition. Pollen percentage data were analysed using principal components analysis in order to investigate the degree to which the different vegetation units can be distinguished by their pollen spectra. Pollen–vegetation relationships for selected taxa were explored using simple scatter plots and indices of association, under‐ and over‐representation. Indices of floristic diversity and palynological richness were used to assess the representation of the vegetation in the pollen spectra. Results and conclusions Five vegetation zones are defined on the basis of species composition and their quantitative variation: back shore, mobile dunes, slacks, semi‐fixed and fixed dunes. Pollen assemblages from back shore, mobile dunes and slacks are clearly differentiated from semi‐fixed and fixed dunes. Pollen assemblages differ considerably from the associated vegetation composition. Major discrepancies are caused by large differences in pollen and vegetation proportion of Hyalis argentea and Discaria americana. There is a considerable proportion of non‐local pollen in every spectrum. Pollen representation in the coastal dunes at Monte Hermoso is influenced by differences in pollen production, dispersal and preservation of individual taxa as well as by the spatial distribution of the vegetation, the topography of the dune system and the wind pattern. The pollen–vegetation relationship established in this study has important implication for understanding and interpreting fossil pollen records from coastal dune environments.     

Modern vegetation–pollen–climate relationships for the Pampa grasslands of Argentina

Aim To analyse the relationships between potential natural vegetation, pollen and climate in order to improve the interpretation of fossil pollen records and provide the background for future quantitative palaeoclimatic reconstructions. Location Pampa grasslands of Argentina, between 33–41° S and 56–67° W. Methods Modern pollen data were obtained from a pollen data base developed by the Grupo de Investigación de Paleoecología y Palinología, Universidad Nacional de Mar del Plata, Argentina (143 surface samples and 17 pollen types). Analysis of pollen and climate data involved multivariate statistics (cluster analysis and principal components analysis), scatter diagrams, Pearson’s correlation and isopoll mapping. Results Vegetation patterns at regional scales (grasslands and xerophytic woodlands) and local scales (edaphic communities) were identified by cluster analysis of pollen surface samples. The main climatic variables that appear to constrain the vegetation distribution and abundance of taxa are mean annual precipitation, annual effective precipitation and summer temperature. Individual pollen types such as Chenopodiaceae, Apiaceae, Cyperaceae, Prosopis, Schinus, Condalia microphylla and other xerophytic taxa are good indicators of moisture regime. Many pollen types are significantly correlated with summer temperature. The modern vegetation–pollen–climate relationships vary in a broadly predictable manner, supporting the contention that fossil pollen assemblages can be related to particular climatic characteristics. Main conclusions An expanded suite of modern analogues facilitated new insights into vegetation–pollen–climate relationships at the regional scale in Pampa grasslands. Relationships between individual pollen types and climate are appraised at a regional scale and new modern analogues are presented. The results provide the basis for improved vegetation and climate reconstruction from fossil records of the study area.     

Vegetation patterns,pollen deposition and distribution of non-pollen palynomorphs in an ice-wedge polygon near Kytalyk (NE Siberia), with some remarks on Arctic pollen morphology

In ice-wedge polygon mires, small-scaled microrelief of ridges enclosing small depressions results in a short-distance vegetation mosaic. The correct recognition of these landscape elements in palaeoecological studies of peat sections in order to reconstruct their patterns and dynamics requires insight in the short-distance relationship between vegetation and pollen deposition. This paper presents an analysis of pollen surface samples in a high-resolution (1 m) transect across an ice-wedge polygon near Kytalyk (NE Siberia), including a discussion on the morphology of some critical pollen types and non-pollen palynomorphs (NPPs). We found a strong correlation between vegetation and surface elevation and a fair correspondence between pollen deposition and vegetation. Distribution of NPPs reflects surface elevation well, with algal spores dominating deep spots and testate amoebae prevailing on higher spots. Peak pollen/spore values unrelated to high species coverages (e.g. of Salix, Betula, Sphagnum, Poaceae) indicate that single plants within a population may cause the bulk of the pollen production. The absence of pollen of taxa with an important presence in the vegetation (e.g. Utricularia) must be attributable to low pollen productivity. Distributional patterns point at pollen transport by water in the polygon troughs/depressions. Our study shows that Arctic pollen records mainly reflect short-distance vegetation patterns. Palaeosequences consequently allow accurate reconstruction of local microtopography and its dynamics, but should not be over-interpreted in terms of changing (over)regional vegetation patterns and associated drivers.     

Variability in glacial and Holocene marine pollen records offshore from west southern Africa

The distribution of pollen in marine sediments is used to record vegetation changes over the past 30,000 years on the adjacent continent. A transect of marine pollen sequences from the mouth of the river Congo (∼5°S) to Walvis Bay and Lüderitz (∼25°S) shows vegetation changes in Congo, Angola and Namibia from the last glacial period into the Holocene. The comparison of pollen records from different latitudes provides information about the latitudinal shift of open forest and savannahs (Poaceae pollen), the extension of lowland forest (rain forest pollen) and Afromontane forest (Podocarpus pollen), and the position of the desert fringe (pollen of Caryophyllaceae, Chenopodiaceae and Amaranthaceae). High Cyperaceae pollen percentages in sediments from the last glacial period off the mouth of the river Congo suggest the presence of open swamps rather than savannah vegetation in the Congo Basin. Pollen from Restionaceae in combination with Stoebe-type pollen (probably from Elytropappus) indicates a possible northwards extension of winter rain vegetation during the last glacial period. The record of Rhizophora (mangrove) pollen is linked to erosion of the continental shelf and sea-level rise. Pollen influx is highest off river mouths (10–2000 grains year⁻¹ cm⁻²), close to the coast (300–6000 grains year⁻¹ cm⁻²), but is an order of magnitude lower at sites situated far from the continent (<10 grains year⁻¹ cm⁻²).     

Quantifying the relationship between airborne pollen and vegetation in the urban environment

The goal of this study was to quantitatively assess the relationship linking vegetation and airborne pollen. For this, we established six sampling stations in the city of Thessaloniki, Greece. Once every week for 2 years, we recorded airborne pollen in them, at breast height, by use of a portable volumetric sampler. We also made a detailed analysis of the vegetation in each station by counting all existing individuals of the woody species contributing pollen to the air, in five zones of increasing size, from 4 to 40 ha. We found the local vegetation to be the driver of the spatial variation of pollen in the air of the city. Even at very neighbouring stations, only 500 m apart, considerable differences in vegetation composition were expressed in the pollen spectrum. We modelled the pollen concentration of each pollen taxon as a function of the abundance of the woody species corresponding to that taxon by use of a Generalized Linear Model. The relationship was significant for the five most abundantly represented taxa in the pollen spectrum of the city. It is estimated that every additional individual of Cupressaceae, Pinaceae, Platanus, Ulmus and Olea increases pollen in the air by approximately 0.7, 0.2, 2, 6 and 5%, respectively. Whether the relationships detected for the above pollen taxa hold outside the domain for which we have data, as well as under different environmental conditions and/or with different assemblages of species representing them are issues to be explored in the future.     

Pugillo Di Licheni Dell'Abissinia E Dell'Eritrea

Abstract

The long pollen sequences from four volcanic lakes of the Lazio région are presented in order to outline the history of flora and vegetation on the Tyrrhenian side of Central Italy during the last hundred thousand years. The four long pollen records correlate satisfactorily with each other and reflect climatic changes at least at regional scale. Particular attention is paid to the development of arboreal vegetation during the last glacial. The local name “Lazio Complex” is given to the seven main fluctuations of trees of the Pleniglacial (from about 60,000 to about 30,000 years B.P.). The history of Zelkova, Abies, Picea, Fagus, Corylus, Ulmus, Carpinus betulus, Tilia in central Italy is outlined.     

Differentiating vegetation types from eastern South American ecosystems based on modern and subfossil pollen samples: evaluating modern analogues

In south and southeast Brazil land use caused profound changes in natural vegetation and consequently the value of the pollen composition in surface samples as modern analogues. In order to test the capability of modern pollen to represent the natural vegetation, three different time slices of pollen assemblages from 27 sites spread over southern and south-eastern Brazil and the Misiones Province in Argentina were collated. Pollen samples from the pre-colonization period, selected from the moment just before abrupt changes evidenced on pollen diagrams caused by the colonization process throughout the last 500 years, were assumed to represent the natural vegetation conditions once the climate remained stable within this period. Thus we used pre-colonization assemblages to compare with modern samples to explore to what extent surface pollen may be biased in representing the natural vegetation types. Furthermore, to compare man made vegetation change to climate driven vegetation change we also compared to these 20 out of 27 samples dated to 3,000 years bp. Guided by ordination and cluster analysis, but using abundance thresholds of indicator taxa we classified the pollen spectra of pre-colonization time into seven groups consistent with the main vegetation types in the area. Ordination analyses capture the differentiation between grassland and forested vegetation and between tropical and subtropical vegetation types. Comparing the pre-colonization with other time slices we observed that based on Poaceae abundance, 70 and 85 % respectively of sites from 3,000 bp and modern assemblages maintained their classification. Based on finer classification criteria these values decreased to 40 and 52 % respectively. Square chord dissimilarity indicates that colonization impact altered the pollen composition as strongly as 3,000 years of climate induced vegetation change. The surface samples still represent important environmental gradients; however, their use as modern analogue requires careful treatment and eventual exclusion of highly impacted sites.     

First pollen record for Ancyloscelis apiformes (Apidae: Emphorini) in Central Amazon

Abstract

The first pollen study for the solitary bee Ancyloscelis apiformes (Emphorini) in a nest aggregation in the Central Amazon was made in October 2012 at the Sucupira apiary in east Manaus, Amazonas, at 03° 00? 05″ S, 59° 51? 05″ W. A total of 15 samples/nests (N1–N15) were analysed, including intact pollen loads (stored inside the nest) and post-emergence residues, which were acetolysed and mounted in glycerine gelatin. From each sample, 500 pollen grains were counted to determine the pollen frequencies and diversity (H′) and evenness (J′) indices. The most representative pollen types were Bonamia ferruginea (N1 = 95.6% and N2 = 96.00%) and the Ipomoea (N3 = 96.40%), both of which belong to the Convolvulaceae. In addition, Byrsonima crispa (N10 = 93.40%) and Malpighia glabra (N7 = 90.20% and N9 = 97.60%), representatives of the Malpighiaceae, were classified as temporary specialisation events with frequencies above 90%. In this study, A. apiformes, which is considered an oligolectic bee species, presented a polylectic tendency since it collected large percentages of pollen types from plants in distinct families that were not phylogenetically related.     

Tallgrass prairie pollen assemblages in mid-continental North America

The mid-continent of North America has experienced dramatic and abrupt climate change during the Holocene, but the response of grassland vegetation to past climate change has been difficult to quantify. To improve interpretation of tallgrass prairie vegetation from pollen assemblages, we acquired and analysed a surface sample set collected from 25 small ponds (less than 10 ha surface area) in the largest contiguous remnant of tallgrass prairie in the USA. We compared these tallgrass prairie assemblages to 476 modern pollen samples classified as “prairie” in the North American Surface Sample database. We then compared the surface pollen assemblages with fossil pollen assemblages from sediment cores at two sites in Kansas—Cheyenne Bottoms and Muscotah Marsh—using the modern analog technique. Pollen assemblages in the Flint Hills surface samples were very similar to each other, with an average squared chord distance of 0.19. They were different than other modern grassland pollen assemblages mainly due to higher percentages of pollen from six woody taxa: Carya, Cornus, Juniperus, Juglans, Maclura, and Platanus. Arboreal pollen percentages ranged from 17 to 62 % and did not correlate with woody cover among sites. Cheyenne Bottoms was open grassland for the past 25,000 years, but it did not have many tallgrass prairie analogs. Muscotah Marsh did not have many grassland analogs over the past 30,000 years, possibly due to its position on the prairie-forest border or its surrounding wetland vegetation.     

A comparison of sedimentary DNA and pollen from lake sediments in recording vegetation composition at the Siberian treeline

Reliable information on past and present vegetation is important to project future changes, especially for rapidly transitioning areas such as the boreal treeline. To study past vegetation, pollen analysis is common, while current vegetation is usually assessed by field surveys. Application of detailed sedimentary DNA (sedDNA) records has the potential to enhance our understanding of vegetation changes, but studies systematically investigating the power of this proxy are rare to date. This study compares sedDNA metabarcoding and pollen records from surface sediments of 31 lakes along a north–south gradient of increasing forest cover in northern Siberia (Taymyr peninsula) with data from field surveys in the surroundings of the lakes. sedDNA metabarcoding recorded 114 plant taxa, about half of them to species level, while pollen analyses identified 43 taxa, both exceeding the 31 taxa found by vegetation field surveys. Increasing Larix percentages from north to south were consistently recorded by all three methods and principal component analyses based on percentage data of vegetation surveys and DNA sequences separated tundra from forested sites. Comparisons of the ordinations using procrustes and protest analyses show a significant fit among all compared pairs of records. Despite similarities of sedDNA and pollen records, certain idiosyncrasies, such as high percentages of Alnus and Betula in all pollen and high percentages of Salix in all sedDNA spectra, are observable. Our results from the tundra to single‐tree tundra transition zone show that sedDNA analyses perform better than pollen in recording site‐specific richness (i.e., presence/absence of taxa in the vicinity of the lake) and perform as well as pollen in tracing vegetation composition.     

The southern limit of the Mediterranean vegetation in the Sahara during the Holocene

Holocene records from the southern Sahara in Niger allow a reconstruction of the vegetation history and inform us about the former extension of the Mediterranean. Both pollen and charcoal analyses evidenced the direct contact of Sudanian and Saharan savannas during the middle Holocene at about 19°N, whereas at 20°N the transition from the Saharan savanna to the desert was found. In southwestern Libya (26°N) a combination of a Saharan desert vegetation and a semi‐desert Artemisia shrub on the plateaus demonstrated the contact with Mediterranean influenced formations. Regular ash and charcoal layers in middle‐Holocene sediments of the northern Niger prove an early interference of man with the vegetation development. One has to imagine that, in combination with the cattle‐keeping and the later metal production, man could have changed the former northern Sudanian vegetation into the present Sahelian savanna system from the middle Holocene on.     

Myrmecochorous dispersal distances: a world survey

Abstract. Myrmecochorous dispersal distances are reviewed; the seed dispersal curve generated by ants shows a characteristic peak at short distances and a long tail, a shape suited to small densities of safe sites. Mean global distance is of 0.96 m (n= 2524) with a range of 0.01–77 m. Data have been broken down by geography (Northern hemisphere v. Southern hemisphere), taxonomy (ant subfamilies) and ecology (vegetation: sclerophyllous v. mesophyllous). Although a statistical difference exists between dispersal curves from the Northern hemisphere and the Southern hemisphere, this may be an artefact of lack of data from mesophyllous myrmecochores from this hemisphere. The four ant subfamilies do show also numerical differences but could not be subjected to statistical analysis. A difference between the shape of dispersal curve for sclerophyllous myrmecochores and mesophyllous myrmecochores has also been detected. We hypothesize that this difference is related to the myrmecological communities from both types of vegetation: dispersing ants from sclerophyllous vegetation would have smaller nest densities and/or bigger foraging areas than dispersing ants from mesic environments.     

Cell division study in ‘pollen mother cells’ and ‘pollen grains’ of in vivo and ex vitro plants in Drimiopsis botryoides

The various normal and abnormal stages of meiosis and pollen mitosis of Drimiopsis botryoides are described, and a comparison between naturally propagated in vivo and tissue culture derived ex vitro plants in respect to their cytological behaviour presented. We also describe the floral morphology and investigate the relationship between the floral developmental stages and the progression of microgametogenesis. In total, 33 bivalents are observed in diakinesis, which indicate the diploid number 2n = 66 and this number is cross-checked by a haploid set of n = 33 chromosomes in pollen mitosis. Only 6.8% and 4.9% meiotic abnormalities were recorded on in vivo and ex vitro plants, respectively, which led to the formation of non-viable pollen. Finally, the microspores have to develop into tri-cellular male gametophyte. Only 0.2% pollen grains are found with a micro-nucleus. Though the higher pollen viability was recorded on both in vivo (89.3 ± 4.1%) and ex vitro (92.1 ± 4.6%) plants, but surprisingly the pollen germination rate is extremely low with 13.6 ± 1.74% and 21.3 ± 1.55%, respectively. The present study obviously enriches the cytological database of D. botryoides and may help future research on androgenesis and genetic improvement.     

Holocene forest development along the Setesdal valley, southern Norway, reconstructed from macrofossil and pollen evidence

The Setesdal valley in South Norway runs north to south for 200 km, from alpine vegetation at 1200 m, passing the tree-line at around 1000 m, through Boreal forests, to Nemoral forest at sea level. The Holocene vegetation history and its altitudinal differentiation were reconstructed using pollen percentages and influx and plant macrofossil concentration records from four lakes along an altitudinal transect. During the early Holocene (c. 10500–8000 cal b.p.) Betula pubescens, Pinus sylvestris, Alnus, and Corylus expanded in the lowlands. Only Pinus and B. pubescens reached 1000 m asl (Lille Kjelavatn). Only B. pubescens reached Holebudalen (1144 m asl) at about the same time as it arrived in the lowlands. Between c. 8000–3000 cal b.p. mixed deciduous forest developed around Dalane (40 m asl) and to a lesser extent around Grostjørna (180 m asl), birch woodland with pine surrounded Lille Kjelavatn and birch woodland occurred at Holebudalen. From c. 3000 cal b.p. to present, the vegetation at Dalane hardly changed except for slight human impact and the immigration of Picea abies. At Grostjørna Pinus expanded. At Lille Kjelavatn Pinus disappeared and Betula became sparse as at the tree-line today. Betula retreated from Holebudalen thus leaving it above the tree-line in low-alpine vegetation. The strengths and weaknesses of pollen and plant macrofossil data were assessed for forest reconstructions. Where local pollen production is low, as near the tree-line, percentages of long-distance tree pollen can be misleadingly high. Pollen influxes of Betula and Pinus were much smaller near their altitudinal limits than at lower altitudes, although their macrofossils were equally abundant. The limited dispersal capacity of macrofossils documents the local presence of species and the character of the local vegetation, although macrofossils of some tree taxa are rarely found. Pollen and plant macrofossil evidence complement each other to provide a more complete reconstruction of Holocene tree-limits and tree-lines and hence climate changes, than either form of evidence alone.     

A 250 year comparison of historical, macrofossil and pollen records of aquatic plants in a shallow lake

1. Sedimentary remains of aquatic plants, both vegetative (turions, leaves, spines) and reproductive (fruits, seeds, pollen), may provide a record of temporal changes in the submerged vegetation of lakes. An independent assessment of the degree to which these remains reflect past floristic change is, however, rarely possible. 2. By exploiting an extensive series of historical plant records for a small shallow lake we compare plant macrofossil (three cores) and pollen (one core) profiles with the documented sequence of submerged vegetation change since c. 1750 AD. The data set is based on 146 site visits with 658 observations including 42 taxa classified as aquatic, spanning 250 years. 3. Approximately 40% of the historically recorded aquatic taxa were represented by macro‐remains. In general macrofossils underestimated past species diversity, with pondweeds (three of eight historically recorded Potamogeton species were found) particularly poorly represented. Nonetheless, several taxa not reported from historical surveys (e.g. Myriophyllum alterniflorum and Characeae) were present in the sediment record. 4. The pollen record revealed taxa which left no macro‐remains (e.g. Littorella uniflora), and the macrofossil record provided improved taxonomic resolution for some taxa (e.g. Potamogeton) and a more reliable record of persistence, appearance and loss of others (e.g. Myriophyllum spp. and Nymphaeaceae). 5. Detrended correspondence analysis indicated that changes in the community composition evidenced by the palaeolimnological and historical records were synchronous and of a similar magnitude. Both records pointed to a major change at around 1800, with the historical record suggesting a more abrupt change than the sedimentary data. There was good agreement on a subsequent change c. 1930. 6. The palaeolimnological data did not provide a complete inventory of historically recorded species. Nevertheless, these results suggest that combined macrofossil and pollen records provide a reliable indication of temporal change in the dominant components of the submerged and floating‐leaved aquatic vegetation of shallow lakes. As such palaeolimnology may provide a useful tool for establishing community dynamics and successions of plants over decadal to centennial timescales.     

An extinct species of Basella: pollen evidence from sediments (~80 ka) in Kerala,India

Abstract

The genus Basella belongs to family Basellaceae and is comprised of five species. Out of these, Basella alba is pantropical in distribution, B. paniculata is endemic to south-eastern Africa and the other three species viz. B. excavata, B. leandriana and B. madagascariensis are endemic to Madagascar. The palynological study of an organic layer buried ~80 ka in Chaganacherry, Kerala revealed a rich diversity of rainforest pollen and estuarine/marine palynomorphs indicating warm and humid climate coupled with sea level high stand. The Basella pollen recorded from this sediment shows intermediate characters of two extant Basella species. While it resembles B. paniculata in being pantocolpate (20), it shows affinity with B. alba in sexine characters. The recovered pollen was therefore, ascertained to be a new species, named here as B. keralensis sp. nov. The age correlation of the studied sediment was carried out by fingerprinting (Rare Earth Elements) of Youngest Toba Tuff (YTT) (glass shards) which spread in the atmosphere through a volcanic eruption in Sumatra ~74 ka and also with marine palynomorph markers correlating the interglacial Marine Isotopic Stage 5a (~80 ka). It is inferred that earlier, the genus Basella comprised of six species out of which Basella keralensis sp. nov. became extinct and until now neither the fossil record nor the extant form of this is known in records from any part of the World.     

Composition,biomass and structure of mangroves within the Zambezi River Delta

We used a stratified random sampling design to inventory the mangrove vegetation within the Zambezi River Delta, Mozambique, to provide a basis for estimating biomass pools. We used canopy height, derived from remote sensing data, to stratify the inventory area, and then applied a spatial decision support system to objectively allocate sample plots among five strata. Height and diameter were measured on overstory trees, saplings and standing dead trees in nested plots, and biomass was calculated using allometric equations. Each of the eight mangrove species occurring in Mozambique exist within the Delta. They are distributed in heterogeneous mixtures within each of the five canopy height classes, not reflecting obvious zonation. Overstory trees averaged approximately 2000 trees ha^?1, and average basal area ranged from 14 to 41 m² ha^?1 among height classes. The composition of the saplings tended to mirror the overstory, and the diameter frequency distributions suggest all-aged stands. Above-ground biomass ranged from 111 to 483 Mg ha^?1 with 95 % confidence interval generally within 15 % of the height class mean. Despite over 3000 trees ha^?1 in the small-tree component, 92 % of the vegetation biomass is in the overstory live trees. The objective inventory design proved effective in estimating forest biomass within the 30,267 ha mangrove forest.