Fragmentation effects on beta diversity of fragmented and conserved landscapes: insights about homogenization and differentiation processes

Araújo, Felipe de Carvalho; Mendes, Carolina Njaime; Souza, Cleber Rodrigo de; Miranda, Kaline Fernandes; Morelli, Mariana Caroline Moreira; Coelho, Polyanne Aparecida; Figueiredo, Maria Auxiliadora Pereira; Carvalho, Warley Augusto Caldas; Castro, Gislene Carvalho de; Fontes, Marco Aurélio Leite; Santos, Rubens Manoel dos

doi:10.1590/0102-33062020abb0261

Felipe de Carvalho Araújo ^** Corresponding author: carvalhoaraujo_f@yahoo.com.br

Biology Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0003-0210-8763

Carolina Njaime Mendes

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0002-2377-7600

Cleber Rodrigo de Souza

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0002-4122-2748

Kaline Fernandes Miranda

Biology Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0002-2898-4270

Mariana Caroline Moreira Morelli

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0001-8061-7016

Polyanne Aparecida Coelho

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0002-4397-7298

Maria Auxiliadora Pereira Figueiredo

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0001-5205-1502

Warley Augusto Caldas Carvalho

Federal University of São João del Rei, 36301-160, São João del Rei, MG, Brazil

http://orcid.org/0000-0002-1524-1531

Gislene Carvalho de Castro

Federal University of São João del Rei, 36301-160, São João del Rei, MG, Brazil

http://orcid.org/0000-0003-0951-5986

Marco Aurélio Leite Fontes

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0001-8467-4667

Rubens Manoel dos Santos

Forest Science Department, Federal University of Lavras, 37200-000, Lavras, MG, Brazil

http://orcid.org/0000-0002-4075-462X

* Corresponding author: carvalhoaraujo_f@yahoo.com.br

Figures (5)
Tables (2)

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Figure 1
Map of municipalities and localities where sampling was conducted over Espinhaço Range. PESRM represents the protected landscapes and Sabará and Brumadinho fragmented landscapes.

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Figure 2
Non-metric multidimensional scaling of the three landscapes with a total of five vegetation types, three in Brumadinho, four in Sabará and five at PESRM. The distance among plots indicates the similarity among them regarding the beta diversity.

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Figure 3
Boxplot of beta diversity (turnover and nestedness) in the three studied landscapes. B = Brumadinho; S = Sabará; R = PESRM. Asterisks represent significance in the analysis of similarity (ANOSIM). Each box encompasses the 25th to 75th percentiles; the median is indicated by the horizontal line within the box and the other horizontal lines outside the box indicate the 10th and 90th percentiles.

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Figure 4
Boxplot of the beta diversity components (turnover and nestedness) in the three relationships: B/S = fragmented landscapes (Brumadinho and Sabará); R/S = protected x fragmented landscape (PESRM and Sabará); R/B = protected x fragmented landscape (PESRM and Brumadinho). B = Brumadinho; S = Sabará; R = PESRM. Asterisks represent significance in the analysis of similarity (ANOSIM). Each box encompasses the 25th to 75th percentiles; the median is indicated by the horizontal line within the box and the other horizontal lines outside the box indicate the 10th and 90th percentiles.

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Figure 5
A conceptual framework showing consequences of reducing habitats using ecotone vegetation as an example. Abrupt transitions increase distance from propagules from species due habitat loss. While gradual transition allows higher species flow, abrupt transition enhances beta diversity leading to a potential differentiation process.

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Table 1
Distribution of plots in the sampled landscapes in the Espinhaço Range. Floristic relationships and similarities between vegetation types are presented in the supplementary material, Figure S3.

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Table 2
Characterization of vegetation types sampled at all landscapes surveyed.

Landscape	Vegetation types	Landscape characterization	All plots/plots per vegetation type (20 x 20 m)	Total richness
PESRM	- Semi-deciduous seasonal forest; - Ironstone; - Brazilian savannah; - Ecotone forest/ironstone and - Ecotone forest/Brazilian savannah.	Intact landscape without anthropogenic disturbance or destruction	25/5	186
Brumadinho	- Semi-deciduous seasonal forest; - Ironstone; - Brazilian savannah and - Anthropogenic vegetation called ‘anthropogenic pastures’ - open area of secondary succession after disturbance.	Landscape fragmented by mining, with consequent absence of ecotones and presence of grass	40/10	131
Sabará	- Semi-deciduous seasonal forest; - Ironstone; - Brazilian savannah; - Ecotone and - Anthropogenic vegetation called ‘anthropogenic pastures’ - open area of secondary succession after disturbance.	Landscape fragmented by mining, presence of ecotones with Eucalyptus spp. plantation, presence of grass	50/10	146

Vegetation type	Description
Semideciduous seasonal forest	Composed mainly by trees, this forest environment is characterized by the loss of 30 to 60 % of its leaves in the dry season, conditioned by the double climatic seasonality (*Veloso et al. 1991Veloso HP, Rangel Filho ALR, Lima JCA. 1991. Classificação da vegetação brasileira, adaptada a um sistema universal. Rio de Janeiro, Instituto Brasileiro de Geografia e Estatística.; Oliveira-Filho 2009*Oliveira-Filho AT. 2009. Classificação das fitofisionomias da América do Sul cisandina tropical e subtropical: proposta de um novo sistema-prático e flexível-ou uma injeção a mais de caos? Rodriguésia 60: 237-258.).
Ironstone	It is included in the definition of rupestrian grassland sensu latu (*Silveira et al.* 2015Silveira FAO, Negreiros D, Barbosa NPU, Buisson E, Carmo FF. 2015. Ecology and evolution of plant diversity in the endangered campo rupestre: a neglected conservation priority. Plant Soil. doi: 10.1007/s11104-015-2637-8 https://doi.org/10.1007/s11104-015-2637-... ). Occurs on ferruginous fields, mostly above 1000 m altitude (Vincent & Meguro 2008Vincent RDC, Meguro M. 2008. Influence of soil properties on the abundance of plant species in ferruginous rocky soils vegetation, southeastern Brazil. Brazilian Journal of Botany 31: 377-388.). It contains a high richness of herbaceous species and high endemism on the layer of compacted ironstone; the occurrence of trees and shrubs is restricted to depressions, where the soil layer is deeper (Fernandes 2016**Fernandes GW. 2016. The Megadiverse Rupestrian Grassland. In: Fernandes GW (ed). Ecology and conservation of mountaintop grasslands in Brazil. Springer. 3-12).
Brazilian savannah	Characterized by low-altitude, fertile soils with high aluminum content. The vegetation comprises trees and shrubs, with dense understory and soil covered by grasses (*Schaefer et al. 2016Schaefer CEGR, Corrêa GR, Cândido HG, et al. 2016. The Physical Environment of Rupestrian Grasslands (Campos Rupestres) in Brazil: Geological, Geomorphological and Pedological Characteristics, and Interplays. In: Fernandes GW (ed). Ecology and conservation of mountaintop grasslands in Brazil. Springer. 15-49.). Plant diversity is mediated by the presence of quartzitic outcrops, fire (Fernandes 2016Fernandes GW. 2016. The Megadiverse Rupestrian Grassland. In: Fernandes GW (ed). Ecology and conservation of mountaintop grasslands in Brazil. Springer. 3-12), depth of the water table and by soil characteristics (Ribeiro & Walter 2008*Ribeiro JF, Walter BMT. 2008. As principais fitofisionomias do bioma cerrado. In: Sano SM, Almeida SP, Ribeiro JF (eds). Cerrado: Ecologia e flora. Brasília: Embrapa Informação Tecnológica. 151-212.).
Ecotone	“…a transition between two or more vegetation type; it is a junction zone or tension belt which may have considerable linear extent but is narrower than the adjoining community areas themselves.” (Odum 1953Odum EP. 1953. Fundamentals of ecology. Philadelphia, W.B. Saunders Company.). Additionally, ecotones from PESRM have demonstrated be floristically and phylogenetically distinct vegetation type with own particularities (*Souza et al. 2020*Souza CR, Paula GGP, Mendes CN, et al. 2020. Local-scale tree community ecotones are distinct vegetation types instead of mixed ones: a case study from the Cerrado-Atlantic forest ecotonal region in Brazil. Australian Journal of Botany. doi: 10.1071/BT19108 https://doi.org/10.1071/BT19108... ).
Anthropogenic vegetation: ‘anthropogenic pastures’	Open area of secondary succession after disturbance covered by grasses.

[1] * Corresponding author: carvalhoaraujo_f@yahoo.com.br

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Fragmentation effects on beta diversity of fragmented and conserved landscapes: insights about homogenization and differentiation processes

ABSTRACT