Survey of Bioluminescent Coleoptera in the Atlantic Rain Forest of Serra da Paranapiacaba in São Paulo State (Brazil)

Levantamento de Coleópteros bioluminescentes na Mata Atlântica da Serra da Paranapiacaba

Abstracts

Brazil is the country with the largest number of bioluminescent beetle species in the world. However, estimates suggest that this number could much be higher, since many species remain to be discovered. In this work we made a survey of the species of bioluminescent beetles in Serra de Paranapiacaba - the largest remnant of Atlantic Forest in São Paulo State. The survey was done at Intervales State Park, Carlos Botelho State Park and municipality of Tapiraí and the following species were collected: Aspisoma lineatum, Aspisoma physonotum, Aspisoma fenestrata, Cratomorphus besckey, Cratomorphus distinctus, Photinus penai, Photinus sp₁, Photinus sp₉, Ethra aff. malledicta or axilaris, Ethra aff. adicta , Lucidotini incertae sedis, Cladodes flabellicornis, Cladodes demoulini, Amydetes lucernuta, Bicellonycha sp₈, Bicellonycha ornaticollis, Pyrogaster lunifer, Pyrogaster moestus, Pyrogaster sp₂, Pyrogaster sp₅, Pyrogaster sp₆, Photuris lugubris, Photuris sp₁, Photuris sp₇, Stenophrixotrix sp₁, Brasilocerus sp₁, Pseudophengodes sp₁, Hapsodrilus pyrotis, Hypsiophtalmus sp₁, Ptesimopsia luculenta, Pyroptesis cincticollis, Pyrearinus brevicolis, Pyrearinus candelarius, Pyrearinus micatus, Pyrophorus divergens. Our data show that Serra de Paranapiacaba is the second richest area in São Paulo state, especially in elaterids, with unique species typical of this area and species common to other investigated sites such as the Biological Station of Boracéia (in Salesópolis county) and the urbanized areas in the between Campinas - Sorocaba- São Paulo, originally covered to the Atlantic Rainforest.

Lampyridae; Phengodidae; Elateridae; Elateroidea; Staphylinidae; bioluminescence; Fireflies; Atlantic Rainforest

O Brasil é o país que possui o maior número de espécies de coleópteros bioluminescentes no mundo. Entretanto, estimativas sugerem que este número possa ser bem maior, dado que muitas espécies ainda não foram descritas. Neste trabalho foi realizado um levantamento das espécies de coleópteros bioluminescentes em três localidades na Serra da Paranapiacaba - a maior área remanescente contígua de Mata Atlântica no país, com associação ao respectivo habitat. No Parque Estadual Intervales, Parque Estadual Carlos Botelho e Tapiraí foram coletadas as seguintes espécies: (Lampyidae) Aspisoma lineatum, Aspisoma physonotum, Aspisoma fenestrata, Cratomorphus besckey, Cratomorphus distinctus, Photinus penai, Photinus sp₁, Photinus sp₉, Ethra aff. malledicta ou axilaris, Ethra aff. adicta, Lucidotini incertae sedis, Cladodes flabellicornis, Cladodes demoulini, Amydetes sp₁, Bicellonycha sp₈, Bicellonycha ornaticollis, Pyrogaster lunifer, Pyrogaster moestus, Pyrogaster sp₂, Pyrogaster sp₅, Pyrogaster sp₆, Photuris lugubris, Photuris sp₁, Photuris sp₇; (Phengodidae) Stenophrixotrix sp₁, Brasilocerus sp₂, Pseudophengodes sp₁; (Elateridae) Hapsodrilus pyrotis, Hypsiophtalmus sp₁, Ptesimopsia luculenta, Pyroptesis cincticollis, Pyrearinus brevicolis, Pyrearinus candelarius, Pyrearinus micatus, Pyrophorus divergens. Estes dados mostram que esta constitui a segunda área mais rica em espécies luminescentes do Estado de São Paulo, depois da Est. Biológica de Boracéia, especialmente em elaterídeos, com espécies únicas características destas localidades e espécies comuns è outras áreas investigadas, como as da Estação Biológica de Boracéia (Salesópolis, SP) e áreas urbanizadas no triângulo Campinas - Sorocaba - São Paulo, originalmente cobertas por Mata Atlântica.

Lampyridae; Phengodidae, Elateridae; Elateroidea; Staphylinidae; vaga-lumes; bioluminescência; mata atlântica

Introduction

Brazil has the largest diversity of luminescent beetles in the world, about 500 described species, corresponding about 23% of described species in the world (Costa 2000; Viviani et al. 2010). These species are distributed mainly in the three main families of Elateroidea: Lampyridae, Phengodidae and Elateridae. Furthermore, two species of luminescent Staphylinidae were reported (Costa et al. 1986; Rosa 2010). These species are distributed mainly in the following biomes: Atlantic Rainforest, Cerrado (Savannah), Pantanal (Marshies) and Amazon Rainforest. Among these ecosystems, the Atlantic Rainforest is one of the richest and also most threatened ones, currently with only around 7-8% of the original cover or 11.4-16% when considering the the small woods (Ribeiro et al. 2009).

Despite their biodiversity, taxonomic and systematic studies on bioluminescent beetles in Brazil are still scarce. Detailed modern taxonomical studies are found mainly for Elateridae by Costa e collaborators (Costa et al. 1988, Costa 1971a, 1971b; 1972, 1975). However, the families Lampyridae and Phengodidae lack recent reviews and their systematics remains troublesome. Some studies on the biology and ecology were performed for some species of Elateridae (Costa 1975, Costa et al. 1988) and Phengodidae (Costa et al. 1999, Viviani & Bechara 1997). In the family Lampyridae, bionomic studies were done for two semi-aquatic species of Aspisoma spp. (Costa et al. 1988, Viviani 1989), Aspisoma lineatum Gyllenhal, 1817 (Viviani et al., 2012), some lampyrid species of São Paulo State (Viviani 2001) and for Photuris fulvipes Blanchard 1837(Rosa 2007). Recently, the genus Amydetes was revised with the description of several new species (Da Silveira and Mermudes, 2014).

Considering the growing importance of fireflies and bioluminescent beetles as potential environmental indicators (Viviani et al. 2010), mainly for the impact of artificial night-lighting (Lloyd 2006; Viviani et al., 2010; Hagen et al., 2014), and their importance for scientific studies and in biotechnology as source of bioluminescent reagents (Viviani 2007), it is urgent to make biodiversity surveys to aid conservation and bioprospection programs.

The Atlantic Forest is, after the Amazon, the second largest forest type in South America. The last corridor of this forest is located mostly in São Paulo state, being composed of a series of protected areas with more than 17.300 km² of forests (Galleti Jr et al. 2008). Considering the lack of knowledge about bioluminescent beetles in Brazil, in the nineties, and especially from 2006 we made surveys in the Atlantic rain forest. We already made lists of species occurring near the urban areas of Campinas, Rio Claro, Sorocaba and Votorantim cities in São Paulo State, which were mostly covered by seasonal Atlantic rain forest in the past (Viviani 2001, Viviani et al. 2010) and more recently about the bioluminescent species occurring in the hotspot of Biological Station of Boracéia located in the second largest remnant of Atlantic rain forest of Serra do Mar, along the coastline (Viviani & Santos 2012).

In this study we present a first survey made from 2007-2012 in the largest and most preserved remnant of Atlantic rain forest in São Paulo State, which is located along of Serra da Paranapiacaba, including three main investigated sites: (I) Cachoeira do Chá in the municipality of Tapiraí at its Northern border; (II) Parque Carlos Botelho in the middle and (III) Parque Intervales in the Southern part.

Materials and Methods

Habitats description. The investigated sites are located in a largest continuous remnant of Atlantic rain forest along the scarp chain Serra da Paranapiacaba, which is a subdomain of the Serra do Mar scarp chain, located about 100 km from the coast between the hydrographic basins: Sorocaba-Médio Tietê, Paranapanema and Ribeira-Southern Shore. The main kind of forest is dense mountain ombrophyl forest (Fig.1) with the following phytophysionomic divisions: (I) mountain forest located over the mountains which include tall trees; (II) humid forest located between mountains over water bodies, which includes similar vegetation to the former habitat but with the presence of species better adapted to humid environments; (III) marshy areas at the borders of water bodies and streams which display typical grasses, and (IV) open fields surrounding the reservation or along thoroughfares opened in the forest. The following sites were investigated:

Figure 1.
(Upper panel) Map of São Paulo State showing the investigated sites; (Lower panels) main vegetation types on the study area: (a) Mountain Atlantic Rainforest in the top of hills in Carlos Botelho State Park, here the view of the Submontane Forest in the Ribeira Valey Basin is shown; (b) Humid forest at Carlos Botelho State Park. View of a marshy area along riparian woods in the Taquaral stream; (c) Open field with vegetal species more light tolerant, including grasses at the ground level and creepers at the edge of woods. This image represents a site with occurrence of Bicellonycha sp8. - a brazilian synchronous firefly - in Intervales State Park. Figura 1. (Painel superior) Mapa do Estado de São Paulo, com os pontos onde foram realizados os levantamentos; (Paineis inferiores) principais tipos de vegetação nas áreas de estudo. (a) Floresta Atlântica Montana no Parque Estadual Carlos Botelho, aqui é mostrado a vista para a Floresta Submontana no Vale do Ribeira; (b) Floresta pluvial no Parque Carlos Botelho. Vista de uma área brejosa de mata ripária no Rio Taquaral; (c) Área aberta com espécies vegetais mais tolerantes è luminosidade, como gramíneas e lianas. Esta imagem apresenta um local com ocorrência de Bicellonycha sp8 - um vagalume sincrônico brasileiro - no Parque Estadual Intervales.

Cachoeira do Chá, Tapiraí. (24°01′47.50"S - 47°3429.47"W, 595m asl). This is a trail located along a stream at the Northern border of the large remnant, borthered by the route SP-79 and located 15 km from Tapiraí municipality. In this site the mountains display an escarpment ending at Ribeira River Valley.

Carlos Botelho. (24°03′46.48"S - 47°59′30.31"W, 763m asl). This park features a rugged terrain in the upper level followed by escarpment ending at the Ribeira River Valley in the lower part (Fig.1). Located in the north, at higher elevations there is a watershed of Ribeira River Valley Basin and another (Taquaral river) of Paranapanema River Basin. The mountains have dense rain forest (Montana forest), whereas in the lower part of the drive at the southeast, the vegetation is composed mainly by lowland tropical rain forest (Submontana forest). The survey sites within this unit are located near the headquarters, in the Northern part of the Park in the municipality of São Miguel Arcanjo.

Fazenda Intervales. (24°16′12.46"S - 48°25′17.11"W, 826m asl). This site is located on the Serra da Paranapiacaba, and also on the Planalto de Guapiara, with very rough relief cut by headwater streams. The study site in this reserve was located between Guapiara and Ribeirão Grande municipalities. The present forest coverage is characterized as Montana forest as described above and the differential in this area is the geological and geomorphological formation, with many valleys and hills. The investigated areas were all located in the protected area, with the trails in the northwestern part of the Park.

Collecting techniques. The collecting and observations were originally done in 1991 and 1993 at Intervales, and then yearly from 2007-2012. In Intervales, collecting and observations were done in the period of October-April and eventually in July. In Carlos Botelho the observations and collecting were made weekly between July and December from 2007-2010. In Tapiraí, observations were done from 2009-2012, with sporadic visits in the same period from November through March.

Adult lampyrids and elaterids were located by their own light at night and collected in flight with entomological net, or in the bushes or in the litterfall, during the period from August to April. They were also collected on the foliage during the day, especially diurnal lampyrids. Luminescent click-beetles and even some lampyrids such as Cratomorphus distinctus Olivier 1895 firefly could also be attracted to green chemiluminescent light sticks. Adult male phengodids were collected, in rare occasions, on soil and on the grass. Firefly larvae were collected on the undergrowth, in litterfall and soil at night, by the location of their glows (Viviani 2001, Viviani et al. 2010). Phengodid larvae were also collected at night in the soil, in embankments, and eventually on the underbrush by location of their luminescence (Viviani & Bechara 1997).

Identification. Specimens of fireflies and their larvae were identified first by comparison with scientific collection of Professor Viviani at UFSCar, which was previously identified by comparison with the collections of Museu de Zoologia da Universidade de São Paulo (São Paulo, Brazil), Natural History Museum of Paris (France) and British Museum (London,UK). Some species were identified by Dra. S. P. Rosa during her visit at the Natural History Museum of Paris (France). Elateridae species were previously identified by Dra. C. Costa (MZUSP) and Dra. S. P. Rosa. Several lampyrid and phengodid species could not be identified, and were catalogued by the name of the genus followed by a specific number (separated in morph species), according to Viviani & Bechara (1997) and Viviani (2001). The specimens were deposited in the collection in UFSCar under curatorship of V. Viviani.

Results and Discussion

Taxonomic survey. Thirty one species of luminescent beetles were cataloged in the Serra da Paranapiacaba: 20 Lampyidae, 7 Elateridae, 4 Phengodidae and 1 Staphylinidae (Table 1).

Table 1
Species of bioluminescent beetles collected in the three different sites of Serra de Paranapiacaba. Tabela 1. Espécies de besouros bioluminescentes coletados nas três localidades de estudo na Serra de Paranapiacaba.

Taxon	Tapiraí	PECB	PE Intervales	Reference collection
Lampyridae
Cratomorphini
Aspisoma lineatum Gyllenhal		ok (4)		Col. MZUSP
Aspisoma physonotum Gorham, 1884			ok (2,3)	Col. MZUSP
Aspisoma sp2	ok (1,2)		ok (1,2)	Col. V. Viviani
Cratomorphus besckei Olivier 1895	ok (2)		ok (2)	Col. MZUSP
Cratomorphus giganteus Druty, 1782	ok (1)	ok (1)	ok (1)	Col. MZUSP
Photinini
Photinoides jenai McDormott, 1963	ok* (1,2,3,4)	ok (1,2,3,4)	ok (1,2,3,4)	Col. MZUSP
Photinus sp1		ok (1)	ok (1)	Col. V. Viviani
Photinus sp9	ok (2)			Col. V. Viviani
Lucidotini
Ethra aff. malledicta or axilaris		ok (1)		*
Ethra aff. adicta			ok (1)	*
Lamprocerini
Cladodes demoulini			ok (1)	Col. MZUSP
Cladodes flabellicornis		ok (1)		Col. V. Viviani
Amydetinae
Amydetes lucernuta			ok (4)	-
Photurinae
Bicellonycha sp.		ok (3,4)		-
Bicellonycha sp8	ok* (3,4)		ok (3,4)	Col. V. Viviani
Pyrogaster lunifer Eschscholtz, 1822		ok (1)		Col. V. Viviani
Pyrogaster moestus Germar 1824		ok (1,2)	ok (1,2)	Col. V. Viviani
Pyrogaster sp3			ok (1)	PORTO (2011)
Photuris lugubris Gorham 1881		ok (1,2)	ok (1,2)	Col. V. Viviani
Photuris sp7			ok (1)	Col. V. Viviani
Phengodidae
Mastinocerini
Stenophrixotrix sp1	ok (1)	ok (1)	ok (1)	Col. V. Viviani
Brasilocerus sp1	ok (1,2)			-
Phrixotrix hirtus Olivier, 1909	ok* (2)			Col. V. Viviani
Phengodini
Pseudophengodes sp1			ok (1,2)	-
Elateridae
Agrypininae/Pyrophorini
Hapsodrilus pyrotis Germar, 1841	ok (1)		ok (1)	Col. MZUSP
Pyroptesis cincticollis Germar 1841	ok (1,3)		ok (1,3)	Col. MZUSP
Ptesimopsia luculenta Germar, 1841			ok (1,2)	Col. MZUSP
Pyrearinus brevicollis Eschsdioltz 1829		ok (1)		Col. MZUSP
Pyrearinus candelarius Germar 1841			ok (1)	Col. V. Viviani
Pyrearinus micatus Costa 1978	ok (1,2)		ok (1,2)	Col. V. Viviani
Pyrophorus divergens Eschscholtz 1829	ok (1)	ok (1)	ok (1)	Col. V. Viviani

• Legenda: ok = ocorrência; ok* = ocorrência mas sem material fixado; (1) = floresta Montana; (2) = floresta úmida; (3) = áreas brejosas; (4) = áreas abertas. ** identificação pela chave.
• Legend: ok = collected; ok* = occurence recorded but no fixed material; (1) = montane forest; (2) = umid forest; (3) = marshy areas; (4) = open field. ** identificação pela chave.

The richness of each family of luminous beetles in each conservation unit is summarized in Figure 2.

Figure 2.
Number of bioluminescent Coleoptera species of different families in each site ofthe Serra da Paranapiacaba. Figura 2. Número de espécies de coleopteros bioluminescentes por família em cada local de coletas e observações na Serra da Paranapiacaba.

Lampyridae (Fig. 3). The number of observed lampyrid species was 10 in Carlos Botelho and 14 in the Intervales. In Tapiraí only five species were found (see Table 1). The smaller number found in Tapiraí could be due to the lower frequency and smaller scanned area of collecting made in this site. The following species were unique to Carlos Botelho: Aspisoma lineatum, Photinus sp₁, Ethra aff maledicta or axillaris, Lucidotini incertae sedis, Cladodes flabellicornis, Pyrogaster lunifer, Pyrogaster sp₅ and Pyrogaster sp₆. Seven species were found at Intervales but were not been found in Carlos Botelho and Tapiraí: Aspisoma physonotum, Aspisoma fenestrata, Amydetes lucernuta, Bicellonychia ornaticollis, Pyrogaster moestus, Photuris sp₁ and Photuris sp₇. There is a unique species which was collected only in Tapiraí - Photinus sp₉ - which displays a different bioluminescent signal from other Photinus species collected in Brazil, being characterized by a bimodal flash with bioluminescence spectrum peaking at 561 nm which is slightly blue-shifted in relation to other species of the genus found in the region (Viviani et al. 2010). Phengodidae. Only four species of phengodids were found in Paranapiacaba. In Carlos Botelho only the arboreal Stenophrixotrix sp₁ larvae were found. In Tapiraí Brasilocerus sp₂ and Phrixotrix hirtus were found and in Intervales Pseudophengodes sp₁. The arboreal Stenophrixotrix sp₁ was common to all three areas. When compared with Boraceia, the number of species and abundance of phengodids was lower. The following species were found in both places: Brasilocerus sp2, Phrixotrix hirtus and Stenophrixotrix.

Figure 3.
Some species of fireflies sampled on SA de Paranapiacaba: (a) Cratomorphus giganteus; (b) Cratomorphus besckei; (c) Aspisoma lineatum; (d) Aspisoma physonotum; (e) Aspisoma sp₂; (f) Photinoides jenai; (g) Photinus sp₁; (h) Photinus sp₉; (i) Aethra aff. malledicta or axilaris; (j); (k) Amydetes lucernuta; (l) Cladodes flabellicornis; (m) Cladodes demoulini; (n) Photuris lugubris; (o) Photuris sp₇; (p) Pyrogaster moestus; (q) Pyrogaster lunifer; (r) Pyrogastes sp₃ (ref. Diego S. Porto). Figura 3. Algumas especies de lampirídeos coletadas na SA de Paranapiacaba: (a) Cratomorphus giganteus; (b) Cratomorphus besckei; (c) Aspisoma lineatum; (d) Aspisoma physonotum; (e) Aspisoma sp₂; (f) Photinoides penai; (g) Photinus sp₁; (h) Photinus sp₉; (i) Aethra aff. malledicta ou axilaris; (j); (k) Amydetes lucernuta; (l) Cladodes flabellicornis; (m) Cladodes demoulini; (n) Photuris lugubris; (o) Photuris sp₇; (p) Pyrogaster moestus; (q) Pyrogaster lunifer; (r) Pyrogastes sp₃ (ref. Diego S. Porto).

Elateridae (Fig. 4). The studied area was especially rich in luminescent click-beetles, with eight species: Hapsodrilus pyrotis Germar 1841, Hypsiophtalmus sp₁, Ptesimopsia luculenta, Pyrearinus brevicollis, Pyrearinus candelarius, Pyrearinus micatus, Pyrearinus sp., Pyrophorus divergens. All these species were found at Intervales.

Figure 4.
Bioluminescent click-beetles of SA de Paranapiacaba: (a) Pyrophorus divergens; (b) Ptesimopsia luculenta; (c) Ptesimopsia cincticollis; (d) Pyrearinus candelarius; (e) Pyrearinus micatus, (f) Pyrearinus brevicollis; (g) Hapsodrilus pyrotis. Figura 4. Elaterideos bioluminescentes da SA de Paranapiacaba: (a) Pyrophorus divergens; (b) Ptesimopsia luculenta; (c) Ptesimopsia cincticollis; (d) Pyrearinus candelarius; (e) Pyrearinus micatus, (f) Pyrearinus brevicollis; (g) Hapsodrilus pyrotis.

In Tapiraí, four species of bioluminescent click-beetles were found: Pyrophorus divergens, Hypsiophtalmus sp₁, Pyrearinus micatus and Pyrearinus meatus: The species Hapsodrylus pyrotis, Ptesimopsia luculenta and Pyrearinus candelarius were not observed in Tapiraí and Carlos Botelho. Comparatively, Boraceia Biological Station showed only three species (Pyrophorus divergens, Pyrearinus micatus and Hapsodrilus pyrotis) and the municipalites of Campinas and Sorocaba four species (Hapsodrilus ignifer, Pyrearinus candelarius, Pyrearinus micatus and Pyrophorus divergens).

Considering that the surveys were conducted in the same morphoclimatic domain and continuous geographic Mountain Tropical Rain Forest, it was surprising that several of these species were not found more widely distributed among these areas. This could be due to two reasons: (1) there is more heterogeneity in the habitats than assumed, and the same kind of specific habitats were not sampled in the three localities, or (2) because the survey was not conducted simultaneously in the three areas. In this case, the seasonality of adults of a given species may vary between areas.

Comparison with other sites of Atlantic forest. When comparing this survey to Biol. St. of Boraceía, which is the second largest remnant of Atlantic Rainforest in São Paulo State, Paranapiacaba remnant showed a slightly smaller biodiversity. The following species were found in common in these two sites: Aspisoma physonotum Gorham 1884, Cratomorphus besckei Olivier 1895, Cratomorphus distinctus Olivier 1895, Photinus penai Germar 1824, Cladodes flabellicornis Motsch 1853, Pyrogaster moestus Germar 1824, Pyrogaster sp₂, Photuris lugubris Gorham 1881, Photuris sp₇, Haspodrilus pyrotis Germar 1841, Pyrophorus divergens Eschsholtz 1829 and Pyrearinus micatus Costa 1978. Paranapiacaba forests were especially rich in elaterids, whereas Boracéia was richer in fireflies. When compared with the fragmented remnants of the semi-seasonal forests and secondary growths of urbanized areas of Campinas, Sorocaba and Rio Claro municipalities, only the following species were found in common: Aspisoma lineatum, Aspisoma physonotum Gorham 1884, Cratomorphus distinctus Olivier 1895, Photinus sp₁, Photinus sp₂, Amydetes lucernuta, Bicellonycha ornaticollis Blanch, Pyrogaster moestus Germar 1824, Photuris sp₇, Pyrearinus micatus Costa 1978 and Pyrophorus divergens Eschsholtz 1829. Thus, the species: Aspisoma fenestrata, Ethra aff maledicta or axillaris, Bicellonycha sp₈, Pyrogaster lunifer Eschsholtz., Photuris sp₁, Brasilocerus sp₂, Hypsiophtalmus sp₁, Ptesimopsia luculenta, Pyrearinus brevicolis and Pyrearinus sp₁ were apparently unique to Paranapiacaba forests.

The frequency of species per family in places where they are visibly more abundant is shown on Fig. 5. There is an apparent trend for greater richness for all families in Mountain Forest. More refined analysis with quantification of abundance is however required, to confirm whether the diversity of habitats is significant or not. Marshy areas seem to harbor a lower number of species.

Figure 5.
Distribution per habitat of the main families of bioluminescent beetles in Serra da Paranapiacaba. Figura 5. Distribuição por habitats de familias de besouros bioluminescentes da Serra de Paranapiacaba.

Conclusions

The Serra de Paranapiacaba displays the second largest diversity of luminescent beetles in the Atlantic rain forest morphoclimatic domain of São Paulo state, with 34 species. Some species are common to sampled areas of semi-seasonal forest close to urbanized areas of inner São Paulo state and to the second largest and most preserved area of Biological Station of Boracéia. This area is especially rich in elaterid luminescent beetles, but displays poorer diversity of Phengodidae.

Acknowledgements

We are grateful to Dra. Simone Policena Rosa (MZUSP) for the identification of the specimens to Édipo F. V. da Silva and Pedro R. Busana (UFSCar) for assistance in the field activities and to Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Biota Project - FAPESP (Proc. 2011/19427-9) for financial support.

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Publication Dates

• Publication in this collection
  Mar 2016

History

• Received
  21 Apr 2015
• Reviewed
  16 Oct 2015
• Accepted
  28 Oct 2015