STRUCTURE OF 15 HECTARES PERMANENT PLOTS OF TERRA FIRME DENSE FOREST IN CENTRAL AMAZON

ESTRUTURA DO COMPONENTE ARBÓREO EM 15 HA DE PARCELAS PERMANENTES DE FLORESTA DENSA DE TERRA FIRME NA AMAZÔNIA CENTRAL

ABSTRACT

Terra Firme dense forest occurs in 65% of the Amazonian region and is characterized by the high diversity of plant species and high occurrence of rare plant species. The objective of this work was to analyze the horizontal structure of 15 hectares of plots, randomly chosen from a set of 400 ha of permanent plots situated at the Experimental Site of Embrapa, Manaus, Amazonas State, Brazil. All individuals with DBH ≥ 10 cm were identified at the level of species in 2005. The VI values (Values of importance) were separated in classes of range with one (01) standard deviation and the individuals distributed according to DBH classes of 10 cm in amplitude. In total, 8771 individuals were identified, distributed into 264 species and 53 families. A larger number of individuals is concentrated in a few number of species, such as Protium hebetatum D.C. Daly, Eschweilera coriacea (DC.) S.A. Mori and Licania oblongifolia Standl, which sum up 21% of the total number of individuals and 12% of the VI. There is a high occurrence of rare species (36%). The families Sapotaceae, Lecythidaceae and Burseraceae together accounted for 39% of the total number of individuals. The diametric structure in an inverse-J shape shows that 80% of the individuals are concentrated in the DBH classes ranging from 10 to 30 cm. The results highlight the high occurrence of rare species and those with low density, and with distribution restricted to some plots, particularly those belonging to the lowest classes of VI, which deserve special attention in actions for biodiversity conservation and forest management.

Keywords:
Phytosociology; Tree species; Amazon

RESUMO

A floresta densa de terra firme ocorre em 65% da região Amazônica e é caracterizada por possuir alta diversidade de espécies vegetais e elevada ocorrência de espécies raras. Este trabalho analisou a estrutura horizontal de 15 ha de parcelas aleatoriamente selecionadas de um conjunto de 400 ha de parcelas permanentes, localizadas no Campo Experimental da Embrapa em Manaus, AM.Todos os indivíduos com DAP ≥ 10 cm foram identificados em nível de espécie em 2005. Os valores de VI (Valor de Importância) foram separados em classes de amplitude de um (01) desvio padrão e os indivíduos distribuídos segundo classes de DAP de amplitude de 10 cm. Ao todo foram identificados 8771 indivíduos, distribuídos em 264 espécies e 53 famílias. Um maior número de indivíduos concentra-se em poucas espécies, tais como Protium hebetatum D.C.Daly, Eschweilera coriacea (DC.) S.A. Mori e Licania oblongifolia Standl. que somaram21% do número total de indivíduos e 12% do VI. Há elevada ocorrência de espécies raras (36%). As famílias Sapotaceae, Lecythidaceae e Burseraceae somaram 39% do número total de indivíduos. A estrutura diamétrica em forma de J-invertido mostra que80% dos indivíduos estão concentrados nas classes de 10-30 cm de DAP. Os resultados chamam a atenção para a elevada ocorrência de espécies raras e daquelas com baixa densidade, e com distribuição restrita a algumas parcelas, principalmente aquelas pertencentes às menores classes de VI, as quais devem merecer atenção especial nas ações de conservação da biodiversidade e manejo florestal.

Palavras-chave:
Fitossociologia; Espécies arbóreas; Amazônia

1. INTRODUCTION

Brazil, a country with approximately one-third of the remaining rain forests in the world, is one of the most important repositories of global biodiversity (PAS 2008). In the Amazon, many areas are formed by mosaic of habitats, with different sets of plant species occurring in adjacent areas on different substrates. It can be seen, in general, , that the distribution of tropical tree species can occur due to habitat preferences and/or variations in their evolutionary history (PITMAN et al., 2001; NOVAES FILHO et al., 2007).

The dynamics and structure of rain forests vary across the Amazon basin in an east-west gradient, following a pattern that matches variations in soil fertility (QUESADA et al., 2009). To describe these patterns of variation, using large data sets, such as those in the present study, is extremely important to enhance the understanding of the mechanisms that determine the distribution of species in nature (CORONADO et al., 2009).

About 65% of the Amazon region is covered by a forest type named terra firme dense forest. This plant typology is the most representative of the Amazon rainforest, mainly characterized by high richness and diversity of species with few individuals of each species and generally showing high dissimilarity floristic between adjacent plots (FERREIRA; PRANCE, 1998; LIMA FILHO et al., 2001; CONDÉ; TONINI, 2013).

Oliveira and Amaral (2004) reported the occurrence of 239 species in a study on one (01) hectare of terra firma dense forest in Manaus, evidencing the high diversity in this typology. Climatic and soil factors such as more humid climates and less seasonal and soils relatively fertile in nutrients, have been cited as possible responsible for a greater diversity of plants in the Western Amazon (GENTRY, 1988, QUESADA et al., 2009). Other authors such as Phillips et al. (1994) report the higher species richness to the natural dynamics of mortality of trees, where forests with high rates of mortality and recruitment would be more diversified.

Most studies in the Amazon uses, on average, one sample area of one (01) hectare for studies of floristic composition and structure. This study contributes to this subject, using a large data set. Thus, the objective of this study was to describe the horizontal structure of 15 hectares of terra firma dense forest in central Amazonia, Manaus, state of Amazonas, in order to contribute to enhance the knowledge on the organization of species in the community, therefore, helping in forest conservation actions and forest management in the Amazon.

2. MATERIAL AND METHODS

2.1. Study area

The study area is located in a terra firme dense forest in Central Amazonia, in the Experimental Field of Embrapa Amazônia Ocidental, Manaus, in the state of Amazonas, Brazil, at the coordinates 2° 32' 12.75" S; 60°0'14.62" W. The area comprises the project developed in the scope of Rede de Monitoramento da Dinâmica de Florestas da Amazônia Brasileira-REDEFLOR (Monitoring Net of Brazilian Amazon Forest Dinamics).

The weather in the area is "Am" type according to Köppen classification, with annual rainfall of 1,355 to 2,839 mm. Average annual temperature is 27.6 °C, with air relative humidity ranging from 84 to 90% over the year. The most humid months are from December to May, and the driest ones are from July to October, when those months receive less than 100 mm of rain (RADAM, 1978).

The average altitude ranges from 60 to 160 m (REGIS, 1993).The predominant soils in the area are Yellow Latosol with texture ranging from more clayey on the plateaus to sandy in the lower parts. They are soils predominantly covered by vegetation of terra firme dense forest, with emergent canopy (IBGE, 1999), consisting of trees ranging from medium to large sizes, reaching up to 55 m of height (REGIS, 1993).

2.2. Data collection and analysis

The plots were delimited at random in the Experimental Field, representing 15 plots of 100 x 100 m. All individuals with diameter at breast height (DBH) e" 10 cm were labeled and botanically identified at the species level in 2005. The plant material was identified by means of comparisons with dried plants available in the herbarium of the Instituto Nacional de Pesquisas da Amazônia - INPA (National Institute of Amazonian Research) as well as by consulting the literature on native species of the Amazon (RIBEIRO et al., 1999). The botanical names were conferred at the Missouri Botanical Garden Web page (MOBOT, 2012), APGII system.

Calculation of the plant structural parameters, such as basal area (BA), absolute and relative dominance (ADo, RDo), abundance (N), absolute and relative density (AD, RD) and absolute and relative frequency (AF, RF) for the composition of the Importance Value Index (IVI) were obtained according to Mueller-Dumbois and Ellenberg (1974), calculated on Mata Nativa 2 software (CIENTEC, 2006). The estimate of the ecological importance of families in the evaluated community was calculated by the Family Importance Index (FII), calculated by the sum of the diversity (number of species of the family/total number of species), relative density and dominance (MORI; BOOM, 1983) using the Microsoft Excel for Windows. Classes of IV (%) were defined using for the upper limit of the first class, the average value minus 0.5 standard deviations (M-0.5* standard deviation) and thereafter, they were added with 1 standard deviation. The diameter classes were defined with an amplitude of 10 cm.

3. RESULTS

Out of the fifteen sampled hectares, 8,771 individuals were identified, belonging to 264 species and 53 families. The number of trees and basal area per plot ranged from 497 to 688 and from 23.4 to 32.7 m2, respectively. The ten most important species in the study area, on the Importance Value (IV) basis, Figure 1, represent 24% of total IV, which were Protium hebetatum D.C. Daly, Eschweilera coriacea (DC.) S.A. Mori, Licania oblongifolia Standl., Pouteria minima T.D.Penn. and Ocotea cernua (Nees) Mez, the ones that presented the highest IV values. Only the first three species have 20.7% of the total number of identified individuals. The complete list of species, with their respective IV values is presented in the appendix.

Figure 1
Value index (VI%) for the 20 especies with the highest values at the Experimental site of Embrapa Amazônia Ocidental, Manaus, Amazonas State, Brazil.
Figura 1
Composição do Valor de Importância-VI(%) para as 20 espécies com os maiores valores no Campo Experimental da Embrapa Amazônia Ocidental, Manaus, AM, Brasil.

The most important families, according to the Family Importance Value (FIV), in a decreasing order were Sapotaceae, Lecythidaceae, Burseraceae, Fabaceae-Mimosoideae and Chrysobalanaceae (Figure 2), in which the three first families grouped approximately 39% of the total number of individuals. A great occurrence of rare species was found (1 individual/plot-1ha), whose values ranged from 30 to 42% of the total number of species recorded in the sampled plots. The families Fabaceae-Mimosoideae, Fabaceae-Faboideae and Fabaceae-Caesalpinoideae, in a decreasing order, are the subfamilies that showed the largest number of species contributing to this value.

Figure 2
Familiar importance value (VIF%) for the 20 families with the highest values at the Experimental site of Embrapa Amazônia Ocidental, Manaus, Amazonas State, Brazil.
Figura 2
Composição do Valor de Importância Familiar - VIF (%) para as 20 famílias com os maiores valores, no Campo Experimental da Embrapa Amazônia Ocidental, Manaus, AM, Brasil.

A great variability is found in the occurrence of species distributed in IV classes (%) within the 15 sampled plots (Figure 3), especially in the two smallest classes that group values lower than 0.66%. Those species correspond to approximately 83% of the species identified in the study area.

Figure 3
Distribution of the number species within plots by class of VI (%) for the 15 plots of 100 x 100m, situated at the Experimental site of Embrapa Amazônia Ocidental, Manaus State, Brazil.
Figura 3
Distribuição do número de espécies nas parcelas segundo as classes de VI (%) para 15 parcelas de 100 x 100m localizadas no Campo Experimental da Embrapa Amazônia Ocidental, Manaus-AM, Brasil.

Plots 15, 150 and 273 have a larger number of exclusive species within the smallest class (<0.096%) and as the IV values increase, a more even distribution of species in the 15 plots is found. Of the 85 species grouped in the smallest class of IV (<0.096%), 41 species appear with exclusive occurrence. These species have low density, with 87% of them occurring with only one (01) individual and 13% with two to six. Duckesia warty (Ducke) Cuatrec. (Humiriaceae), presents six individuals that were registered only in the plot 106, which is on the low topographic position, containing higher content of moisture and with sandy texture. Considering the plots 15, 273 and 150, these sum 39% of the number of species, with the lowest value of IV and with occurrence restricted to these plots. The fact of working with a large set of data, allows better observation of these variations, giving greater reliability to the observed results (Table 1).

Table 1
Horizontal structure of 15 ha of permanente plots in a terra firme dense forest in Central Amazônia
Tabela 1
Estrutura horizontal de 15 ha de parcelas permanentes de uma floresta densa de terra firme na Amazônia Central

Family	Scientific name	N	DR	FR	DoR	VI	VI (%)
Burseraceae	Protium hebetatum Daly	1037	11.82	0.87	5.42	18.10	6.03
Lecythidaceae	Eschweilera coriacea (DC.) S.A. Mori	471	5.37	0.87	3.72	9.95	3.32
Chrysobalanaceae	Licania oblongifolia Standl.	310	3.53	0.87	3.28	7.68	2.56
Sapotaceae	Pouteria minima T.D.Penn.	293	3.34	0.87	2.85	7.06	2.35
Lauraceae	Ocotea cernua(Nees) Mez s.l.	258	2.94	0.87	2.30	6.10	2.03
Malvaceae	Scleronema micranthum Ducke	197	2.24	0.87	2.63	5.74	1.91
Chrysobalanaceae	Licania apelata (E.Mey.) Fritsch	172	1.96	0.87	2.06	4.89	1.63
Lecythidaceae	Eschweilera collina Eyma	176	2.01	0.87	1.98	4.85	1.62
Lecythidaceae	Couratari alta Kunth.	125	1.42	0.87	1.97	4.26	1.42
Sapotaceae	Chrysophyllum sanguinolentum (Pierre) Baehni	146	1.66	0.69	1.67	4.03	1.34
Moraceae	Naucleopsis caloneura (Huber) Ducke	170	1.94	0.87	1.02	3.83	1.28
Moraceae	Brosimum rubescens Taub.	106	1.21	0.87	1.73	3.81	1.27
Elaeocarpaceae	Sloanea excelsa Ducke	92	1.05	0.87	1.63	3.54	1.18
Urticaceae	Pourouma myrmecophyla Ducke	144	1.64	0.81	1.08	3.53	1.18
Fabaceae-Mimosoideae	Inga obidensis Ducke	132	1.50	0.87	1.10	3.47	1.16
Myrtaceae	Psidium araca Raddi	152	1.73	0.87	0.87	3.47	1.16
Myristicaceae	Iryanthera coriacea Ducke	146	1.66	0.87	0.75	3.28	1.09
Lecythidaceae	Couratari stellata A.C.Sm.	74	0.84	0.87	1.42	3.13	1.04
Myristicaceae	Virola micheliiHeckel	128	1.46	0.81	0.82	3.09	1.03
Chrysobalanaceae	Licania micrantha Miq.	86	0.98	0.87	1.21	3.06	1.02
Combretaceae	Buchenavia grandis Ducke	31	0.35	0.75	1.92	3.03	1.01
Humiriaceae	Vantanea macrocarpa Ducke	93	1.06	0.87	1.05	2.97	0.99
Sapotaceae	Pouteria laurifolia (Gomes) Radlk	128	1.46	0.69	0.77	2.93	0.98
Sapotaceae	Micropholis trunciflora Ducke	89	1.01	0.81	1.05	2.87	0.96
Vochysiaceae	Qualea albiflora Warm	49	0.56	0.75	1.53	2.84	0.95
Fabaceae- Caesalpinioideae	Sclerolobium helanocarpus Ducke	75	0.85	0.75	1.16	2.77	0.92
Myristicaceae	Osteophloeum platyspermum (A.DC.) Warb.	54	0.62	0.69	1.38	2.69	0.90
Sapotaceae	Glycoxylon pedicellatum (Ducke) Ducke	90	1.03	0.87	0.77	2.66	0.89
Sapotaceae	Pouteria venosa (Mart.) Baehni ssp. amazonica T.D.Penn.	74	0.84	0.75	1.05	2.64	0.88
Goupiaceae	Goupia glabra Aubl.	32	0.36	0.58	1.69	2.64	0.88
Burseraceae	Protium pilosissimum Engl.	126	1.44	0.52	0.59	2.55	0.85
Burseraceae	Trattinnickia burserifolia Mart.	60	0.68	0.81	1.03	2.52	0.84
Fabaceae-Faboideae	Bocoa viridiflora (Ducke) R.S.Cowan	85	0.97	0.75	0.75	2.47	0.82
Apocynaceae	Aspidosperma marcgravianum Woodson	36	0.41	0.81	1.19	2.41	0.80
Fabaceae- Caesalpinioideae	Eperua duckeana R.S.Cowan	72	0.82	0.46	1.04	2.32	0.77
Olacaceae	Minquartia guianensis Aubl.	75	0.85	0.64	0.80	2.29	0.76
Burseraceae	Protium heptaphyllum (Aubl.) ssp. ulei (Swart) Daly	82	0.93	0.81	0.49	2.24	0.75
Annonaceae	Unonopsis duckei (R.&P.)Macba.	83	0.95	0.81	0.37	2.13	0.71
Fabaceae- Caesalpinioideae	Eperua glabriflora (Ducke) R.S.Cowan	50	0.57	0.87	0.68	2.11	0.70
Sapotaceae	Micropholis guyanensis (A.DC.) Pierre ssp.duckeana (Baehni)	60	0.68	0.69	0.73	2.11	0.70
Vochysiaceae	Erisma bicolor Ducke	37	0.42	0.81	0.84	2.07	0.69
Caryocaraceae	Caryocar pallidum A.C. Smith	12	0.14	0.52	1.40	2.06	0.69
Nyctaginaceae	Neea oppositifolia Ruiz & Pav.	69	0.79	0.81	0.45	2.05	0.68
Chrysobalanaceae	Licania heteromorpha Benth.	59	0.67	0.69	0.62	1.99	0.66
Sapotaceae	Manilkara bidentata (A.DC.) A. Chev.	39	0.44	0.64	0.89	1.97	0.66
Fabaceae-Mimosoideae	Zigia racemosa (Ducke) Barneby & J.W. Grimes	44	0.50	0.75	0.70	1.95	0.65
Moraceae	Clarisia racemosa Ruiz & Pav.	29	0.33	0.69	0.91	1.93	0.64
Lecythidaceae	Lecythis barnebyi Mori	31	0.35	0.64	0.88	1.87	0.62
Melastomataceae	Mouriri callocarpa Ducke	37	0.42	0.87	0.57	1.86	0.62
Sapotaceae	Pouteria caimito (Ruiz & Pav.) Radlk.	41	0.47	0.75	0.59	1.81	0.60
Chrysobalanaceae	Licania laevigata Prance	26	0.30	0.58	0.93	1.81	0.60
Moraceae	Brosimum lactescens (S.Moore) C.C.Berg.	24	0.27	0.46	1.06	1.79	0.60
Fabaceae- Caesalpinioideae	Tachigali cf. myrmecophila Ducke	42	0.48	0.87	0.44	1.79	0.60
Lecythidaceae	Eschweilera atropetiolata S.A.Mori	47	0.54	0.75	0.50	1.78	0.59
Fabaceae-Mimosoideae	Enterolobium schomburgkii Benth.	33	0.38	0.69	0.71	1.78	0.59
Annonaceae	Bocageopsis multiflora (Mart.) R.E.Fr.	45	0.51	0.81	0.45	1.77	0.59
Fabaceae-Faboideae	Swartzia recurva Poepp.	47	0.54	0.64	0.54	1.71	0.57
Myristicaceae	Virola calophylla Warb.	57	0.65	0.69	0.28	1.62	0.54
Annonaceae	Guatterria olivacea R.R.Fr.	45	0.51	0.81	0.27	1.59	0.53
Fabaceae-Faboideae	Andira trifoliata S.A.Mori	50	0.57	0.46	0.56	1.59	0.53
Sapotaceae	Ecclinusa guianensis Eyma	30	0.34	0.46	0.77	1.57	0.52
Fabaceae-Faboideae	Swartzia schomburgkii Benth. Var. guyanensis R.S. Cowan	20	0.23	0.52	0.76	1.51	0.50
Euphorbiaceae	Hevea guianensis Aubl.	47	0.54	0.40	0.56	1.50	0.50
Lecythidaceae	Lecythis graciena S.A.Mori	34	0.39	0.64	0.46	1.49	0.50
Fabaceae-Mimosoideae	Dinizia excelsa Ducke	3	0.03	0.17	1.23	1.44	0.48
Malvaceae	Theobroma sylvestre Mart.	53	0.60	0.69	0.14	1.44	0.48
Opiliaceae	Agonandra brasiliensis Miers	20	0.23	0.58	0.62	1.43	0.48
Apocynaceae	Geissospermum argenteum Woodson	23	0.26	0.64	0.48	1.38	0.46
Fabaceae-Faboideae	Andira parviflora Ducke	14	0.16	0.46	0.75	1.37	0.46
Meliaceae	Trichilia micropetala T.D.Penn.	40	0.46	0.69	0.21	1.36	0.45
Fabaceae-Mimosoideae	Inga alba (Sw) Willd.	44	0.50	0.58	0.28	1.35	0.45
Dichapetalaceae	Tapura amazonica Poepp. & Endl.	45	0.51	0.58	0.26	1.35	0.45
Fabaceae-Mimosoideae	Piptadenia suavolens Miq.	18	0.21	0.52	0.60	1.33	0.44
Clusiaceae	Distomovita brasiliensis D’ Arcy	37	0.42	0.69	0.20	1.31	0.44
Sapindaceaea	Talisia cf. cupularis Radlk.	41	0.47	0.64	0.20	1.30	0.43
Moraceae	Helicostylis scabra (Macbr.)	32	0.36	0.69	0.24	1.30	0.43
Lauraceae	Licaria guianensis Aubl.	33	0.38	0.64	0.28	1.29	0.43
Simaroubaceae	Simarouba amara Aubl.	21	0.24	0.58	0.44	1.26	0.42
Lecythidaceae	Gustavia elliptica S.A.Mori	37	0.42	0.69	0.13	1.25	0.42
Lauraceae	Aniba hostmaniana (Ness) Mez.	37	0.42	0.64	0.17	1.22	0.41
Humiriaceae	Endopleura uchi (Huber) Cuatrec.	21	0.24	0.58	0.41	1.22	0.41
Moraceae	Helianthostylis sprucei Baill.	44	0.50	0.58	0.14	1.22	0.41
Anacardiaceae	Anacardium parvifolium Ducke	16	0.18	0.46	0.55	1.20	0.40
Fabaceae-Mimosoideae	Inga gracilifolia Ducke	34	0.39	0.64	0.16	1.19	0.40
Sapotaceae	Pouteria reticulata (Engl.) Eyma	13	0.15	0.40	0.59	1.15	0.38
Olacaceae	Pytchopetalum olacoides Benth.	29	0.33	0.52	0.27	1.12	0.37
Vochysiaceae	Ruizterania cassiquiarensis (Spruce ex Warm.)	18	0.21	0.52	0.37	1.10	0.37
Malvaceae	Lueheopsis rosea (Ducke) Burret	22	0.25	0.52	0.32	1.09	0.36
Fabaceae-Mimosoideae	Stryphnodendron sp.	26	0.30	0.58	0.21	1.08	0.36
Fabaceae-Mimosoideae	Parkia pendula (Willd.) Walp.	4	0.05	0.23	0.80	1.08	0.36
Lecythidaceae	Lecythis usidata Camb.	18	0.21	0.64	0.24	1.08	0.36
Fabaceae- Caesalpinioideae	Macrolobium limbatum Spruce ex Benth	33	0.38	0.46	0.22	1.06	0.35
Lauraceae	Sextonia rubra (Mez) Van der Werff	13	0.15	0.46	0.45	1.06	0.35
Lauraceae	Mezilaurus itauba (Meissn.) Taubert ex Mez	20	0.23	0.58	0.24	1.05	0.35
Vochysiaceae	Vochysia vismiaefolia Spruce ex Warm.	13	0.15	0.40	0.49	1.04	0.35
Malvaceae	Sterculia pruriens (Aubl.) K.Schum.	25	0.28	0.52	0.22	1.03	0.34
Fabaceae-Faboideae	Swartzia corrugata Benth.	26	0.30	0.52	0.19	1.01	0.34
Sapotaceae	Pouteria oblanceolata Pires	17	0.19	0.46	0.30	0.95	0.32
Euphorbiaceae	Conceveiba guianensis Aubl.	24	0.27	0.46	0.21	0.95	0.32
Lauraceae	Licaria sp.	20	0.23	0.58	0.13	0.93	0.31
Fabaceae-Mimosoideae	Inga stipularis DC.	19	0.22	0.46	0.24	0.92	0.31
Moraceae	Brosimum utile (H.B.K.) Pittier ssp. ovatifolium (Ducke) C.C.Berg.	15	0.17	0.40	0.33	0.90	0.30
Violaceae	Rinorea racemosa (Mart.) Kuntze	26	0.30	0.52	0.06	0.88	0.29
Annonaceae	Xylopia calophylla R.E. Fr.	21	0.24	0.52	0.10	0.86	0.29
Fabaceae-Mimosoideae	Parkia multijuga Benth.	13	0.15	0.40	0.30	0.85	0.28
Bignoniaceae	Jacaranda copaia (Aubl.) D. Don.	22	0.25	0.46	0.13	0.84	0.28
Simaroubaceae	Simaba polyphylla (Cavalcante) W.Thomas	21	0.24	0.52	0.08	0.84	0.28
Rubiaceae	Duroia fusifera Hook. F. ex K. Schum.	18	0.21	0.52	0.11	0.84	0.28
Chrysobalanaceae	Couepia longipendula Pilg.	19	0.22	0.35	0.25	0.81	0.27
Melastomataceae	Tococa guianensis Aubl.	17	0.19	0.52	0.09	0.80	0.27
Fabaceae-Faboideae	Dipteryx magnifica Ducke	12	0.14	0.40	0.25	0.79	0.26
Fabaceae-Mimosoideae	Abarema jupunba (Willd.) Britton &Killip	21	0.24	0.29	0.26	0.79	0.26
Fabaceae-Faboideae	Diplotropis rodriguesii Lima	18	0.21	0.35	0.22	0.77	0.26
Fabaceae-Faboideae	Paramachaerium ormosioides Ducke	13	0.15	0.52	0.09	0.76	0.25
Malvaceae	Apeiba echinata Gaertner	10	0.11	0.52	0.12	0.75	0.25
Vochysiaceae	Qualea paraensis Ducke	9	0.10	0.23	0.41	0.74	0.25
Fabaceae-Faboideae	Swartzia reticulata Ducke	13	0.15	0.40	0.18	0.74	0.25
Icacinaceae	Emmotum aff. nitens Benth. Ex Miers	15	0.17	0.46	0.10	0.73	0.24
Burseraceae	Protium rubrum Cuatrec.	27	0.31	0.29	0.14	0.73	0.24
Clusiaceae	Platonia insignis Mart.	11	0.13	0.40	0.20	0.73	0.24
Solanaceae	Duckeodendron cestoides Kuhlm.	6	0.07	0.29	0.37	0.72	0.24
Fabaceae-Mimosoideae	Inga suberosa T.D.Penn.	12	0.14	0.52	0.05	0.71	0.24
Euphorbiaceae	Micranda siphonoides Benth.	13	0.15	0.46	0.05	0.67	0.22
Malvaceae	Theobroma subincanum Mart.	15	0.17	0.40	0.08	0.66	0.22
Apocynaceae	Ambelania duckei Markgr.	12	0.14	0.46	0.05	0.65	0.22
Malvaceae	Pseudobombax munguba (Mart. & Zucc) Dugand	17	0.19	0.35	0.11	0.65	0.22
Melastomataceae	Miconia reglelii Cogn.	9	0.10	0.46	0.06	0.62	0.21
Meliaceae	Carapa guianensis Aubl.	23	0.26	0.06	0.29	0.61	0.20
Fabaceae- Caesalpinioideae	Peltogyne paniculata Benth.	8	0.09	0.40	0.11	0.61	0.20
Erythroxylaceae	Erythroxylum amplum Bth.	15	0.17	0.35	0.08	0.60	0.20
Meliaceae	Trichilia septentrionales C.DC.	10	0.11	0.40	0.08	0.59	0.20
Vochysiaceae	Qualea acuminata Aubl.	8	0.09	0.29	0.20	0.58	0.19
Sapotaceae	Manilkara amazonica (Huber) Stand.	11	0.13	0.29	0.16	0.57	0.19
Bixaceae	Bixa orellana L.	15	0.17	0.35	0.05	0.57	0.19
Lauraceae	Licania sp.	11	0.13	0.29	0.15	0.57	0.19
Fabaceae-Mimosoideae	Parkia nitida Miq.	9	0.10	0.35	0.11	0.56	0.19
Fabaceae-Mimosoideae	Inga cordatoalata Ducke	14	0.16	0.35	0.05	0.56	0.19
Humiriaceae	Vantanea guiannensis (Aubl.) Ducke	10	0.11	0.35	0.09	0.56	0.19
Urticaceae	Pouroma guianensis Aubl.	12	0.14	0.35	0.07	0.55	0.18
Lauraceae	Ocotea cymbarum H.B.K.	11	0.13	0.29	0.12	0.53	0.18
Humiriaceae	Vantanea micrantha Ducke	11	0.13	0.29	0.11	0.53	0.18
Bignoniaceae	Tabebuia serratifolia (Vahl) Nichols	6	0.07	0.35	0.11	0.52	0.17
Annonaceae	Pseudoxandra coriacea R.E.Fr.	9	0.10	0.35	0.07	0.52	0.17
Anacardiaceae	Astronium lecointei Ducke	7	0.08	0.29	0.14	0.51	0.17
Melastomataceae	Mouriri angulicosta Morley	7	0.08	0.29	0.10	0.47	0.16
Myrtaceae	Myrcia Paivae O. Berg	8	0.09	0.35	0.03	0.47	0.16
Annonaceae	Annona ambotay Aubl.	12	0.14	0.23	0.10	0.47	0.16
Humiriaceae	Sacoglotis matogrossensis Aubl.	9	0.10	0.29	0.07	0.46	0.15
Rubiaceae	Chimarrhis sp.	7	0.08	0.35	0.03	0.46	0.15
Peraceae	Pogonophora schomburgkiana Miers ex Benth.	7	0.08	0.29	0.08	0.45	0.15
Fabaceae-Faboideae	Dipteryx odorata (Aubl.) Willd.	6	0.07	0.29	0.08	0.44	0.15
Moraceae	Brosimum parinarioides Ducke	6	0.07	0.12	0.24	0.43	0.14
Melastomataceae	Miconia elaeagnoides Cogn.	7	0.08	0.29	0.05	0.42	0.14
Fabaceae-Mimosoideae	Pithecellobium elegans Ducke	6	0.07	0.12	0.24	0.42	0.14
Fabaceae-Faboideae	Swartzia ulei Harms	12	0.14	0.17	0.10	0.41	0.14
Sapotaceae	Pouteria ambelaniifolia (Sandwith) . T.D.Penn	5	0.06	0.23	0.12	0.41	0.14
Sapotaceae	Chrysophyllum sparsiflorum Klotzsch ex Miq.	5	0.06	0.23	0.11	0.39	0.13
Lauraceae	Licaria canella (Meissn.) Kosterm.	7	0.08	0.23	0.08	0.39	0.13
Euphorbiaceae	Pausandra macropetala Ducke	12	0.14	0.17	0.07	0.38	0.13
Euphorbiaceae	Mabea subsessilis Pax & K. Hoffm.	10	0.11	0.23	0.03	0.38	0.13
Lecythidaceae	Cariniana sp.	3	0.03	0.17	0.17	0.37	0.12
Apocynaceae	Couma utiles Mart.	5	0.06	0.23	0.08	0.37	0.12
Burseraceae	Protium divaricatum Engl.	4	0.05	0.17	0.12	0.34	0.11
Lauraceae	Ocotea sp.	8	0.09	0.17	0.07	0.33	0.11
Salicaceae	Casearia grandiflora Cambess	6	0.07	0.23	0.03	0.33	0.11
Monimiaceae	Siparuna sp.	6	0.07	0.23	0.02	0.32	0.11
Quiinaceae	Touroulia guianensis Aubl.	6	0.07	0.23	0.02	0.32	0.11
Lauraceae	Aniba megaphylla Mez	5	0.06	0.23	0.03	0.32	0.11
Fabaceae- Caesalpinioideae	Hymenea parvifolia Huber	4	0.05	0.23	0.04	0.31	0.10
Combretaceae	Terminalia dichotoma G.Meyer	4	0.05	0.23	0.04	0.31	0.10
Euphorbiaceae	Croton lanjouwensis Jabl.	5	0.06	0.17	0.08	0.31	0.10
Fabaceae-Faboideae	Dipteryx polyphylla Huber	4	0.05	0.17	0.09	0.31	0.10
Apocynaceae	Aspidosperma album (Vahl.) R. Bem.	4	0.05	0.17	0.09	0.31	0.10
Lecythidaceae	Lecythis prancei S.A. Mori	4	0.05	0.23	0.03	0.31	0.10
Rubiaceae	Palicourea corymbifera Mull. Arg.	9	0.10	0.17	0.03	0.31	0.10
Moraceae	Sorocea guilleminiana Gaudich.	9	0.10	0.17	0.03	0.31	0.10
Fabaceae-Faboideae	Dipteryx sp.	4	0.05	0.06	0.20	0.30	0.10
Sapotaceae	Poteuria platyphylla (A.C.Sm.) Baehni	3	0.03	0.17	0.09	0.30	0.10
Sapotaceae	Micropholis sp.	3	0.03	0.17	0.08	0.29	0.10
Lecythidaceae	Lecythis poiteaui Berg.	6	0.07	0.12	0.10	0.29	0.10
Clusiaceae	Symphonia globulifera L.	7	0.08	0.12	0.09	0.28	0.09
Sapotaceae	Pouteria petiolata T.D. Penn.	4	0.05	0.17	0.06	0.28	0.09
Euphorbiaceae	Conceveiba martiana Baill	5	0.06	0.12	0.10	0.27	0.09
Urticaceae	Cecropia sciadophylla Mart.	5	0.06	0.17	0.04	0.27	0.09
Fabaceae-Faboideae	Platymiscium duckei Huber	4	0.05	0.17	0.05	0.27	0.09
Euphorbiaceae	Mabea sp.	6	0.07	0.17	0.01	0.26	0.09
Chrysobalanaceae	Couepia bracteosa Benth.	4	0.05	0.12	0.09	0.26	0.09
Clusiaceae	Vismia japurensis Reichardt	4	0.05	0.17	0.04	0.26	0.09
Violaceae	Rinorea guianensis Aubl.	5	0.06	0.17	0.02	0.25	0.08
Apocynaceae	Couma macrocarpa Barb.	4	0.05	0.17	0.03	0.25	0.08
Moraceae	Brosimum acutifolium Huber ssp. Interjectum C.C.Berg	3	0.03	0.12	0.10	0.25	0.08
Sapotaceae	Pradosia cochlearia(Lecomte) T.D.Penn	3	0.03	0.17	0.04	0.25	0.08
Monimiaceae	Siparuna guianensis Aubl.	4	0.05	0.17	0.03	0.24	0.08
Sapotaceae	Pouteria peruviensis (Aubrév.) Bernardi	3	0.03	0.12	0.09	0.24	0.08
Ulmaceae	Ampelocera edentula Rusby	4	0.05	0.17	0.02	0.24	0.08
Apocynaceae	Hymatanthus sucuba (spruce)Woodson	3	0.03	0.17	0.03	0.24	0.08
Annonaceae	Guatteria sp.	4	0.05	0.17	0.01	0.23	0.08
Euphorbiaceae	Glycydendron amazonicum Ducke	3	0.03	0.17	0.02	0.23	0.08
Fabaceae-Faboideae	Swartzia tomentifera Harms	6	0.07	0.12	0.04	0.23	0.08
Annonaceae	Xylopia amazonica R.E. Fr.	3	0.03	0.17	0.01	0.21	0.07
Fabaceae-Faboideae	Ormosia grossa Rudd	2	0.02	0.12	0.08	0.21	0.07
Putranjivaceae	Drypetes variabilis Uittien	6	0.07	0.12	0.02	0.21	0.07
Anisophylleacea	Anisophyllea manausensis Pires & W.A.Rodrigues	3	0.03	0.12	0.05	0.20	0.07
Caryocaraceae	Caryocar villosum (Aubl.) Pers.	2	0.02	0.12	0.06	0.20	0.07
Rubiaceae	Warszewiczia schwackei K.Schum.	3	0.03	0.12	0.05	0.20	0.07
Humiriaceae	Duckesia verrucosa (Ducke) Cuatr.	6	0.07	0.06	0.06	0.18	0.06
Peraceae	Pera heteranthera (Schrank) I.M. Johnst.	2	0.02	0.12	0.04	0.18	0.06
Lecythidaceae	Corythophora rimosa W.A.Rodrigues	2	0.02	0.12	0.04	0.18	0.06
Linaceae	Roucheria punctata Ducke	3	0.03	0.12	0.03	0.18	0.06
Sapotaceae	Pouteria guianensis Aubl.	4	0.05	0.06	0.07	0.17	0.06
Lauraceae	Aniba canellila (H.B.K.)Mez	3	0.03	0.12	0.02	0.17	0.06
Icacinaceae	Emmotum acuminatum (Benth.) Miers	2	0.02	0.12	0.03	0.17	0.06
Anacardiaceae	Anacardium spruceanum Engl.	1	0.01	0.06	0.10	0.17	0.06
Lauraceae	Aniba rosaeodora Ducke	3	0.03	0.12	0.01	0.16	0.05
Rubiaceae	Chimarrhis duckeana del Prete	3	0.03	0.12	0.01	0.16	0.05
Rubiaceae	Duroia saccifera (Mart.) Hook. F. ex K. Schum.	3	0.03	0.12	0.01	0.16	0.05
Apocynaceae	Couma sp.	2	0.02	0.12	0.02	0.16	0.05
Moraceae	Brosimum potabile Ducke	2	0.02	0.12	0.01	0.15	0.05
Burseraceae	Protium subserratum Engler	2	0.02	0.12	0.01	0.15	0.05
Annonaceae	Xylopia brasiliensis Spreng.	2	0.02	0.12	0.01	0.15	0.05
Myrtaceae	Myrcia magna Legrand	2	0.02	0.12	0.01	0.15	0.05
Myristicaceae	Iryanthera juruensis Warb.	2	0.02	0.12	0.01	0.15	0.05
Fabaceae-Faboideae	Swartzia cuspidata Spruce ex Benth.	2	0.02	0.12	0.01	0.15	0.05
Myristicaceae	Virola caducifolia W.A. Rodrigues	2	0.02	0.12	0.01	0.15	0.05
Fabaceae-Mimosoideae	Abarema sp.	2	0.02	0.12	0.00	0.14	0.05
Chrysobalanaceae	Licania adolphoduckei Prance	2	0.02	0.12	0.00	0.14	0.05
Fabaceae-Faboideae	Dipteryx punctata (Blake) Amshoff	1	0.01	0.06	0.07	0.13	0.04
Fabaceae-Mimosoideae	Stryphnodendron pulcherrimum (Willd.) Hochr.	1	0.01	0.06	0.05	0.12	0.04
Urticaceae	Cecropia concolor Willd.	1	0.01	0.06	0.05	0.12	0.04
Lecythidaceae	Couratari guianensis Aubl.	2	0.02	0.06	0.03	0.11	0.04
Fabaceae-Mimosoideae	Parkia decussata Ducke	1	0.01	0.06	0.04	0.11	0.04
Sapindaceaea	Matayba sp.	3	0.03	0.06	0.01	0.11	0.04
Fabaceae-Faboideae	Hymenolobium sericeum Ducke	1	0.01	0.06	0.03	0.10	0.03
Fabaceae- Caesalpinioideae	Dialium guianense Steud.	1	0.01	0.06	0.03	0.09	0.03
Clusiaceae	Vismia sp.	1	0.01	0.06	0.02	0.09	0.03
Euphorbiaceae	Alchornea discolor Klotzsch	2	0.02	0.06	0.01	0.09	0.03
Urticaceae	Cecropia purpurascens C.C. Berg	2	0.02	0.06	0.01	0.09	0.03
Annonaceae	Guatteria poeppigiana	2	0.02	0.06	0.01	0.09	0.03
Sapotaceae	Pouteria eugenifolia (Pierre)Baehni	1	0.01	0.06	0.02	0.09	0.03
Malpighiaceae	Byrsonima crispa Juss.	1	0.01	0.06	0.01	0.08	0.03
Fabaceae-Mimosoideae	Pithecellobium racemosum Ducke	1	0.01	0.06	0.01	0.08	0.03
Malpighiaceae Anderson	Byrsonima duckeana W.R.	1	0.01	0.06	0.01	0.08	0.03
Euphorbiaceae	Croton cajucara Benth.	1	0.01	0.06	0.01	0.08	0.03
Celastraceae	Maytenus guyanensis Klotzsch	1	0.01	0.06	0.01	0.08	0.03
Melastomataceae	Miconia sp.	1	0.01	0.06	0.01	0.08	0.03
Araliaceae Frondin	Schefflera morototoni (Aubl.)	1	0.01	0.06	0.01	0.08	0.03
Malvaceae	Scleronema praecox Ducke	1	0.01	0.06	0.01	0.08	0.03
Fabaceae-Mimosoideae	Stryphnodendron guianensis (Aubl.)Benth.	1	0.01	0.06	0.01	0.08	0.03
Fabaceae-Faboideae	Swartzia tessmannii Harms	1	0.01	0.06	0.01	0.08	0.03
Rubiaceae	Albertia edulis A. Rich.	1	0.01	0.06	0.00	0.07	0.02
Fabaceae-Faboideae	Andira micrantha Ducke	1	0.01	0.06	0.00	0.07	0.02
Boraginaceae	Cordia sp.	1	0.01	0.06	0.00	0.07	0.02
Chrysobalanaceae	Couepia canomensis (Mart.) Benth. ex Hook.f.	1	0.01	0.06	0.00	0.07	0.02
Lecythidaceae	Eschweilera rhododendrifolia (Knuth) A.C.Sm.	1	0.01	0.06	0.00	0.07	0.02
Myrtaceae	Eugenia diplocampta Diels	1	0.01	0.06	0.00	0.07	0.02
Rubiaceae	Ferdinandusa elliptica Pohl.	1	0.01	0.06	0.00	0.07	0.02
Meliaceae	Guarea sp.	1	0.01	0.06	0.00	0.07	0.02
Quiinaceae	Lacunaria jenmani (Oliv.)Ducke	1	0.01	0.06	0.00	0.07	0.02
Fabaceae- Caesalpinioideae	Macrolobium angustifolium(Benth.) R.S.Cowan	1	0.01	0.06	0.00	0.07	0.02
Simaroubaceae	Simaba cedron Planch.	1	0.01	0.06	0.00	0.07	0.02
Monimiaceae	Siparuna amazonica Mart.	1	0.01	0.06	0.00	0.07	0.02
Lamiaceae	Vitex cimosa Bert. ex Spreng	1	0.01	0.06	0.00	0.07	0.02
Fabaceae-Mimosoideae	Zigia juruana (Harms) L.Rico	1	0.01	0.06	0.00	0.07	0.02
Fabaceae- Caesalpinioideae	Zollernia paraensis	1	0.01	0.06	0.00	0.07	0.02

The distribution of individuals in diameter classes in 10 cm of amplitude (Figure 4) shows a J-inverse distribution, characteristic of natural forests. The total amplitude of diameters ranged from 10 to 210 cm, where it is observed the occurrence of individuals in almost all diameter classes, except for classes 140-180 cm and 190-200 cm.

Figure 4
Total number of individuals by classes of diameter of 10 cm range, at the Experimental site of Embrapa Amazônia Ocidental, Manaus, Amazonas State, Brazil.
Figura 4
Número total de indivíduos por classes de diâmetro de amplitude 10 cm, no Campo Experimental da Embrapa Amazônia Ocidental, Manaus, AM, Brasil.

Most individuals (61%) is distributed in the smallest diameter class (10-20 cm). The first two classes (10-30 cm) concentrate 80% of the total number of individuals in the 15 hectares sampled. Buchenavia grandis Ducke (Combretaceae) and Andira parviflora Ducke (Fabaceae-Faboideae) occurred in diametric class 180-190 cm; Caryocar pallidum A.C. Sm. (Caryocaraceae) and Dinizia excelsa Ducke (Fabaceae-Mimosoideae), in the largest class (200-210 cm).

4. DISCUSSION

In general, the evaluated terra firme forest has floristic composition and structure that are characteristic of this plant type, with most individuals concentrated in few species and few families concentrating the highest percentage of species richness. These results are corroborated by other studies carried out in the same plant typology (PRANCE; RODRIGUES, 1976; MILLIKEN, 1998; LIMA FILHO et al., 2004; OLIVEIRA; AMARAL, 2004; OLIVEIRA et al., 2008).

The Burseraceae family, despite the less richness in species compared to the Fabaceae, showed higher FIV mainly due to higher number of individuals observed for that family. This family (Burseraceae) appears with the third highest FIV and is the one with the species with the highest IV in the area (Protium hebetatum). This shows the great variability in the tropical forests related to the abundance of individuals and diversity in families.

Gama et al. (2005) and Oliveira et al. (2008) reported in their studies that Protium hebetatum and Eschweilera coriacea are the species most commonly recorded in terra firme forests in the Amazônia. Those species have wide geographical distribution and the highest values of abundance. This was also observed in this study, where the density and dominance, in general, contributed to the greater importance of the species in the area.

The greater relative density found for Protium hebetatum has decisively contributed to a higher value of dominance and consequent higher value of IV. The ten species, with the highest IV values showed similar frequency values, with wide distribution in the sampled plots.

The variability in the distribution of the number of species per IV classes, within the plots, especially in the smaller classes can be attributed to the influence of the rare species. Plots 15 and 150 were those that had the highest occurrence of these species in the total set of 15 plots, also being the ones with the highest percentage of exclusive species, occurring with low abundance.

Several studies have reported the occurrence of a significant number of rare species in the Amazonian terra firme forests. These values are similar to those found in this study, on average 34% of the total number of species (FERRERIA; PRANCE, 1998; OLIVEIRA et al., 2008). Gentry (1988) and Quesada et al. (2009), among others, reported the "preference" for habitats, probably due to variations in soil characteristics, among other factors, as those responsible for the great diversity and dynamics in tropical forests. The high diversity is favored by the occurrence of rare species, whose mechanisms are poorly understood and discussed (Silva, 2010).

In the context of the IV classes, our results show that some species with low abundance have restricted occurrence in the plots and deserve special attention along with rare species (1 individual/plot), in order to avoid local extinctions due to management actions. Except for the first lower IV class, it can be considered in general that all plots have species of all IV classes, suggesting that based on this classification, the horizontal structure observed is well represented in the set of plots.

The shape of the distribution curve of the diameters in J-inverted in this study is typical of rain forests with a high proportion of trees with DBH <30cm (FERREIRA; PRANCE, 1998; LIMA FILHO et al., 2001; HAUGAASEN et al., 2006; GUIMARÃES et al., 2009). The diametric structure found in this work suggests that the area had not suffered major disturbances and the natural dynamics of mortality and recruitment of new individuals, due to the occurrence of small natural clearings may be responsible for the observed distribution (Oliveira et al., 2008). The occurrence of some individuals in classes of diameter greater than 80 cm also suggests that the forest has not been affected by large disturbances, in which it is observed, for example, individuals of Dinizia excelsa with DBH of approximately 200 cm

5. CONCLUSIONS

The use of a large set of data from the 15 permanent plots, allowed us to observe more broadly the variations in the composition and distribution of the species throughout the study area. This was evidenced by the high occurrence of rare species, of species with low density and distribution restricted to some plots. This fact leads us to reflect on how to properly define and discuss the ecological importance of these species, considering that the indexes usually used may not represent their function in the plant community. Studies on the relationship between the species with their occurrence environment can assist in this regard.

Generally, the species and most abundant families, observed in this study are those considered as typical of terra firme dense forest in the Amazon.

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Publication Dates

• Publication in this collection
  Jul-Aug 2016

History

• Received
  24 Oct 2013
• Accepted
  01 Dec 2015