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Research Article
The Cis bilamellatus species-group (Coleoptera, Ciidae) in southern Africa: Cis mooihoekite sp. n. and new distributional records
expand article infoIgor Souza-Gonçalves‡, Cristiano Lopes-Andrade‡
‡ Universidade Federal de Viçosa, Viçosa, Brazil

Corresponding author: Igor Souza-Gonçalves ( igao_bio@yahoo.com.br )

Academic editor: Yasen Mutafchiev
© 2018 Igor Souza-Gonçalves, Cristiano Lopes-Andrade.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Souza-Gonçalves I, Lopes-Andrade C (2018) The Cis bilamellatus species-group (Coleoptera, Ciidae) in southern Africa: Cis mooihoekite sp. n. and new distributional records. African Invertebrates 59(1): 25-35. https://doi.org/10.3897/afrinvertebr.59.22269
ZooBank: urn:lsid:zoobank.org:pub:C9DB8E9C-335F-4962-AD2F-66F5FFC07812
Open Access

Abstract

Cis mooihoekite sp. n. is described based on specimens collected at two localities in the province of Mpumalanga, South Africa. The new species is included in the Cis bilamellatus species-group, which comprises species with a single plate on both anterocephalic edge and anterior pronotal edge in males, females with pronotum usually widest near the posterior end and gradually narrowing anteriorly and both sexes with dual elytral vestiture. Cis mooihoekite sp. n. can be distinguished from the other South African species in the group by the pronotum devoid of a median impunctate line, pronotal plate angularly emarginate forming two small and triangular horns with acute apex and anterocephalic edge with very acute corners. New geographical records and information on the host fungi of Cis pickeri Lopes-Andrade, Matushkina, Buder & Klass, 2009 are provided.

Keywords

Minute tree-fungus beetles, Ciinae , Ciini , Ethiopian, Western Cape Province

Introduction

Cis Latreille is the most diverse genus of Ciidae with about 400 described species and a worldwide distribution (Lopes-Andrade et al. 2009, Oliveira et al. 2013, Lawrence 2016). It includes more than half of all described Ciidae species and some of them are organised into artificial species-groups (Lawrence 1971, Lopes-Andrade 2008, Oliveira et al. 2013, Lawrence 2016).

The sub-Saharan Ciidae are represented by 72 described species (Lopes-Andrade et al. 2009, Orsetti and Lopes-Andrade 2016, Souza-Gonçalves and Lopes-Andrade 2017), of which 49 belong to Cis. In southern Africa, comprising Botswana, Lesotho, Namibia, Republic of South Africa, Swaziland and the southern tip of Mozambique, there are 16 described species of Cis: C. afer Fähraeus, 1871, C. aster Souza-Gonçalves & Lopes-Andrade, 2017, C. bimucronatus Motschoulsky, 1851, C. caffer Fähraeus, 1871, C. capensis Mellié, 1849, C. delagoensis Pic, 1916, C. makebae Souza-Gonçalves & Lopes-Andrade, 2017, C. mandelai Souza-Gonçalves & Lopes-Andrade, 2017, C. masekelai Souza-Gonçalves & Lopes-Andrade, 2017, C. muriceus Mellié, 1849, C. neserorum Souza-Gonçalves & Lopes-Andrade, 2017, C. stalsi Souza-Gonçalves & Lopes-Andrade, 2017, C. pickeri Lopes-Andrade, Matushkina, Buder & Klass, 2009, C. regius Orsetti & Lopes-Andrade, 2016, C. testaceus Fähraeus, 1871 and C. urbanae Souza-Gonçalves & Lopes-Andrade, 2017.

Cis pickeri is the only species belonging to the Cis bilamellatus species-group described from the Ethiopian region (biogeographic regions sensuMorrone 2015). The bilamellatus-group also includes nine more species described from other biogeographic regions: C. australis Blackburn, 1888, C. bilamellatus Wood, 1884, C. biscutatus Fauvel, 1904, C. clarki Blair, 1940, C. laminicollis Blair, 1940, C. lineatosetosus Pool, 1917, C. setiferus Blackburn, 1888 and C. walkeri Blair, 1940, from the Australian region; and the Palaearctic C. onyosi Viñolas & Muñoz-Batet, 2015, described from a single locality in Spain and recently reported from France (Rose 2017, Viñolas and Muñoz-Batet 2015). There are several undescribed species from the Neotropical and Oriental regions that could be placed in this group (Araujo et al. 2015, Lawrence 2016, Lopes-Andrade pers. obs.). Cis bilamellatus was introduced into southeast England during the nineteenth century and has subsequently expanded its distribution to Wales, southern Scotland, Ireland, the Channel Islands and north-west France (Orledge et al. 2010). Recently, it has also been recorded from Chile (Lopes-Andrade and Lüer 2014), making it one of the most widespread and successful invasive ciid species in temperate and subtropical areas.

The aim of this paper is to describe a new species of Cis belonging to the bilamellatus-group and provide new geographical records of this group in southern Africa.

Materials and methods

Museum abbreviations are as follows:

ANICAustralian National Insect Collection, CSIRO Entomology (Canberra, Australian Capital Territory, Australia)

CELC Coleção Entomológica do Laboratório de Sistemática e Biologia de Coleoptera da Universidade Federal de Viçosa (Viçosa, Minas Gerais, Brazil)

CMNCanadian Museum of Nature (Ottawa, Ontario, Canada)

SANCSouth African National Collection of Insects (Pretoria, Gauteng, South Africa)

The new species described here and additional material of C. pickeri were collected and documented by the staff of the SANC, mostly during ten years of a project on parasitoids of Ciidae in South Africa (Neser 2012). The original coding of the morphospecies (Neser 2012) has been revised and some of them have been described (Orsetti and Lopes-Andrade 2016, Souza-Gonçalves and Lopes-Andrade 2017).

Terms for external morphology and male terminalia of ciids follow Lawrence et al. (2011), Lawrence (2016) and Lopes-Andrade and Lawrence (2005, 2011), but see also Oliveira et al. (2013) for an explanation on the use of “tegmen”. The following abbreviations are used for measurements (in mm) and ratios: BW (basal width of scutellar shield), CL (length of antennal club measured from base of the eighth to apex of the tenth antennomere), EL (elytral length along the midline), EW (greatest width of both elytra), FL (length of antennal funicle measured from base of the third to apex of the seventh antennomere), GD (greatest depth of body measured in lateral view), GW (greatest diameter of eye), PL (pronotal length along midline), PW (greatest pronotal width), SL (length of scutellar shield), TL (total length counted as EL+PL, i.e. excluding head). The GD/EW and TL/EW ratios indicate the degree of body convexity and elongation, respectively.

Transcription of labels, dissection, photography and measurement of specimens follow the methods provided by Araujo and Lopes-Andrade (2016). Specimens of the new species were measured from all localities. The number of measured specimens depended on the number of available specimens and the observed variation in size. Differences are given in “Variation”, together with standard measurements (mean and standard deviation) and ratios. Discussion about the morphology of female terminalia of the bilamellatus-group was not possible due to low availability of specimens for dissection or to lack of data in literature. Data on host fungi extracted from labels are summarised in the sections “Host fungi”, together with corresponding number of records in each fungus species and indicating breeding records. The criteria, provided in Orledge and Reynolds (2005) for determining breeding records, have been followed. The distribution map (Fig. 13) was created in the freeware QGIS 2.14.2-Essen.

Results

Cis mooihoekite Souza-Gonçalves & Lopes-Andrade, sp. n.

http://zoobank.org/03353CBF-C605-417C-957D-E3B1CC8DF17E
Figs 1–8, 9–12

Type locality

“Mooihoek Farm” (near Wakkerstroom), coordinates 27°13'S, 30°32'E (Pixley Ka Seme Local Municipality, Gert Sibande District, Mpumalanga Province).

Etymology

The species name refers to the mooihoekite, a copper iron sulphide mineral, which was discovered in 1972 in South Africa and bears a sheen that resembles the dorsal colour of this species.

Diagnosis

The new species differs from C. pickeri, as described by Lopes-Andrade et al. (2009), in the following features: TL less than 1.30 mm; pronotum devoid of a median impunctate line; male with anterocephalic edge with acute corners (Figs 1, 4 and 7) and pronotal plate angularly emarginate forming two small and triangular horns with acute apex (Fig. 7); male abdominal sex patch about one-quarter of the length of the first ventrite at midline (Figs 3 and 4).

Figures 1–8. 

Cis mooihoekite sp. n. Male holotype (Figs 1–7): 1 Dorsal view, note acute corners of male anterocephalic edge (arrows) 2 Lateral view 3 Part of first abdominal ventrite, showing sex patch (arrow) 4 Ventral view, note prosternal carina (yellow arrow), abdominal sex patch (white arrow) and acute corners of anterocephalic edge (black arrows) 5 Oblique view of head, pronotum and part of anterior portion of elytra with scutellar shield 6 Frontal view of head and pronotum, note frontal lamina of head (arrow) 7 Dorsal view of anterior portion of elytra with scutellar shield, pronotum and part of head; note small triangular horns of pronotal plate (large arrows) and acute corners of anterocephalic edge (small arrows) 8 Female paratype. Scale bars: 0.5 mm (1–2, 4, 8); 0.1 mm (3); 0.2 mm (5–7).

Male holotype (Figs 1–7). Adult fully pigmented and in good condition, except for lacking the left antenna and five tarsi. Measurements in mm: TL 1.19, PL 0.43, PW 0.51, EL 0.77, EW 0.53, GD 0.39. Ratios: PL/PW 0.84, EL/EW 1.46, EL/PL 1.81, GD/EW 0.74, TL/EW 2.27. Body elongate, convex, dorsum and venter reddish-brown; antennae yellowish-brown with darker club, palpi and tarsi yellowish-brown; dorsal vestiture distinctly dual, consisting of suberect and decumbent pale yellowish bristles easily discernible at high magnifications (90×); ventral vestiture of decumbent pale yellowish setae easily discernible at high magnifications (90×). Head (Figs 4–5) with anteriormost portion visible from above; dorsum with few shallow and sparse punctures, subglabrous, with microreticulate and dull sheen interspaces; anterocephalic edge with acute corners (Figs 1, 4 and 7), mesally raised and produced forming a short narrow lamina (Fig. 6) with an emarginate apex. Antennae with ten antennomeres, lengths as follows (in mm, right antennae measured): 0.05, 0.04, 0.03, 0.02, 0.01, 0.01, 0.01, 0.03, 0.02, 0.04 (FL 0.08 mm, CL 0.09 mm, CL/FL 1.13). Eyes coarsely facetted; with about 50 ommatidia; GW 0.11 mm. Gula 0.48 times as wide as long. Pronotum (Fig. 7) with coarse, deep and single punctation; punctures distributed irregularly, separated from each other by two puncture-widths or less on disc and one puncture-width or less close to the sides; interspaces, microreticulate with a dull sheen; vestiture single, consisting of suberect (0.02–0.03 mm) pale yellowish bristles; anterior edge projected forwards and upwards as a short, broad plate angularly emarginate forming two very close and small triangular horns with acute apex, each bearing a tuft of bristles (Fig. 7); lateral edges finely crenulate, not explanate and not visible when seen from above, except for posteriormost portion. Scutellar shield subtriangular, bearing a few punctures and decumbent bristles; BW 0.07 mm and SL 0.05 mm. Elytra with dense and dual punctation; large punctures coarse, confused, deep, about twice as large as small punctures; irregular interspaces, somewhat rugose and with a dull sheen; vestiture subseriate and distinctly dual, consisting of long (0.03–0.04 mm) suberect and small (0.01–0.02 mm) decumbent pale yellowish bristles, both arising from minute punctures. Metathoracic wings developed, apparently functional. Hypomera with sparse, shallow punctation; each puncture bearing a fine decumbent seta; interspaces, microreticulate and dull sheen. Prosternum biconcave, tumid and bearing a longitudinal carina at midline (Fig. 4); interspaces, microreticulate. Prosternal process parallel-sided, as long as prosternum at midline; apex rounded. Protibiae with maximum width about one-quarter of its length; apical edge devoid of spines; outer apical angle projected in a small tooth. Meso- and metatibiae without spines on apical edge. Metaventrite with coarse, shallow punctures; interspaces, microreticulate and dull sheen; discrimen about one-quarter the length of metaventrite at midline. Abdominal ventrites with moderately dense, coarse and shallow punctures, separated from each other by one puncture-width or less and each bearing a pale yellowish seta; interspaces, microreticulate and dull sheen; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.14, 0.06, 0.06, 0.06, 0.07; first abdominal ventrite bearing a margined, circular, fully exposed sex patch at middle, with a transverse diameter of 0.03 mm (Figs 3 and 4).

Male paratypes. Terminalia (Figs 9–12) with sternite VIII (Fig. 9) with posterior margin subrounded, bearing short setae at middle and long setae at corners; anterior portion membranous. Tegmen (Fig. 11) about 2.6x as long as wide, widest near apex; sides more or less parallel-sided, but somewhat sinuate near apex; apex with a small emargination on both sides (visible in the dorsal view; Fig. 11); base triangular. Basal piece (Fig. 10) subrounded, about 1.7x as wide as long. Penis (Fig. 12) about as long as tegmen, about 4.7x as long as wide; subcylindrical, parallel-sided along basal three-fifths and then converging to apex, which bears a slight emargination; base rounded. Measurements in mm (n = 4): TL 1.20–1.25 (1.21 ± 0.03), PL 0.43–0.45 (0.44 ± 0.01), PW 0.45–0.48 (0.47 ± 0.01), EL 0.75–0.80 (0.77 ± 0.02), EW 0.50–0.53 (0.51 ± 0.02), GD 0.38–0.43 (0.39 ± 0.02). Ratios (n = 4): PL/PW 0.89–1.00 (0.89 ± 0.04), EL/EW 1.43–1.55 (1.50 ± 0.05), EL/PL 1.67–1.82 (1.73 ± 0.08), GD/EW 0.75–0.81 (0.77 ± 0.03), TL/EW 2.29–2.40 (2.37 ± 0.05).

Figures 9–12. 

Cis mooihoekite sp. n., sclerites of aedeagus of a paratype from the type locality; the anterior ends are placed to the right and the posterior ends are placed to the left. 9 Sternite VIII 10 Basal piece (top) and gross outline of basal piece (bottom) 11 Tegmen (top) and gross outline of tegmen excluding membranes (bottom), note emarginations near apex (arrows) 12 Penis. Scale bar: 0.1 mm.

Female paratypes (Fig. 8). Anterior edge of head truncate, not raised nor produced. Pronotum widest near the posterior end and gradually narrowing anteriorly; anterior edge of pronotum rounded, not developed. Protibia with outer apical angle not toothed but expanded and with several socketed spines along the apical edge. Otherwise like males, but without an abdominal sex patch. Measurements in mm (n = 3): TL 1.13–1.25 (1.18 ± 0.06), PL 0.38–0.45 (0.42 ± 0.04), PW 0.43–0.53 (0.47 ± 0.05), EL 0.75–0.80 (0.77 ± 0.03), EW 0.50–0.60 (0.55 ± 0.05), GD 0.40–0.43 (0.42 ± 0.01). Ratios (n = 3): PL/PW 0.86–0.94 (0.89 ± 0.04), EL/EW 1.33–1.50 (1.40 ± 0.09), EL/PL 1.76–2.00 (1.85 ± 0.13) GD/EW 0.73–0.85 (0.76 ± 0.08), TL/EW 2.08–2.25 (2.16 ± 0.09).

Type series

Holotype: ♂ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm nr Wakkerstroom, 27°13'S, 30°32'E, 15.vii.2008, O&S Neser \ Ex bracket fungus Thelephora sp., BF#125 \ NATIONAL COLL. OF INSECTS Pretoria, S. Afr. \ Cis mooihoekite Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratype: ♂ (CELC, dissected) “SOUTH AFRICA: MPU, Mooihoek Farm nr Wakkerstroom, 27°13'S, 30°32'E, 15.vii.2008, O&S Neser \ Ex bracket fungus Coriolus versicolor, BF#140 \ NATIONAL COLL. OF INSECTS Pretoria, S. Afr.”; ♂ (ANIC) and ♀ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm nr Wakkerstroom, 27°13'S, 30°32'E, 15.vii.2008, O&S Neser \ Ex bracket fungus, BF#129 \ Ex bracket fungus Coriolus versicolor, BF#129 \ NATIONAL COLL. OF INSECTS Pretoria, S. Afr.”; 2 ♂♂ (1 CELC; 1 CMN) and 2 ♀♀ (1 CELC; 1 CMN) “RSA: Transvaal, 11 Km SE PilgrimsRest [sic], 1400m, 11–31.xii.85, S & J Peck, relict native forest edge FIT-malaise”. All paratypes additionally labelled “Cis mooihoekite Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi

Thelephora sp. (Thelephoraceae), one record; and Trametes versicolor (L.) Lloyd (Polyporaceae), two records.

Distribution

Cis mooihoekite sp. n. was collected in two localities: one in the northern Mpumalanga, dominated by Grassland (Mooihoek Farm); and the other in southern Mpumalanga, dominated by Savannah (Transvaal). Both localities are 1,000 m a.s.l. and have urban areas (Fig. 13).

Figure 13. 

Known distribution of southern African species of the bilamellatus-group: Cis mooihoekite sp. n. (black circle) and Cis pickeri Lopes-Andrade, Matushkina, Buder & Klass (black triangle).

Comments

This species corresponds to Cis sp. I in Neser (2012) and was collected together with C. neserorum, the morphospecies Cis sp. C, Cis sp. Q (see explanation on morphospecies’ coding in the section on Materials and Methods), Xylographus madagascariensis Mellié, 1849, the invasive species Ceracis tabellifer (Mellié, 1849) and the parasitoids Astichus micans Neser, 2012 (Hymenoptera: Eulophidae: Entiinae) and A. silvani Neser, 2012.

Cis pickeri Lopes-Andrade, Matushkina, Buder & Klass, 2009

Additional material

10 ♂♂ (4 CELC; 6 SANC) and 7 ♀♀ (2 CELC; 5 SANC) “SOUTH AFRICA, W.Cape, Palmiet River R. nr. Kleinmond, 34°20'S, 18°58'E, 6.iv.2008, S. Neser \ Ex bracket fungus, BF#71 \ NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 3 ♂♂ (1 CELC; 2 SANC) “SOUTH AFRICA, WCAPE, Prince Alfred'S, Pass N of Knysna, 33°58'S, 23°09'E, 5.xi.2009, S & OC Neser \ Ex bracket fungus #213 \ Ex bracket fungus Coriolus versicolor, BF#213 \ NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 4 ♂♂ (2 CELC; 2 SANC) and 5 ♀♀ (2 CELC; 3 SANC) “SOUTH AFRICA WCape, Garden of Eden Indig. Forest, nr. Knysna, 34°02'S, 23°12'E, 1.iii.1991, AJ Hendricks \ Emerged from log of Nuxia floribundaBUDDLEJACEAE, UA677B \ NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 5 ♀♀ (2 CELC; 3 SANC) “SOUTH AFRICA WCape, Heatherlands, George, 33°57'S, 22°57'E, 11.iv.1990, AJ Urban \ Emerged from Dais cotinifoliaTHYMELAEACEAE, with fruiting body of fungus ?Lenzites, UA601 \ NATIONAL COLL. OF INSECTS Pretoria, South Africa”; ♀ (SANC) “SOUTH AFRICA WCape, Woodifield Farm nr. George, 33°58'S, 22°32'E, 9.iv.1990, AJ Urban & AJ Hendricks \ Adults collected on dry fruiting bodies of Russula capensis BASIDIOMYCETE [sic] on Pinus radiata, UA598 \ NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All additionally labelled “Cis pickeri Lopes-Andrade et al., 2009; I. Souza-Gonçalves det.”.

Host fungi

Laetiporus sp. (Fomitopsidaceae), one record; Russula capensis A. Pearson (Russulaceae), one record; and Trametes versicolor (L.) Lloyd (Polyporaceae), one breeding record.

Distribution

Cis pickeri was described based on specimens from two field collections: one during the Swedish South Africa Expedition in the 1950s (at Viljoenspas) and the other to the Western Cape Province in 2003 (at Ceres) (Lopes-Andrade et al. 2009). The field collections made by the staff of the SANC during a project on ciid parasitoids (Neser 2012) resulted in five additional localities for this species: Palmiet River Nature Reserve, Prince Alfred'S, Pass, Garden of Eden, Heatherlands and Woodifield Farm. All localities in which C. pickeri occurs are areas of Fynbos vegetation (Fig. 13).

Comments

This additional material was collected together with C. neserorum, the morphospecies Cis sp. C, Cis sp. D, Cis sp. H, Cis sp. J, Cis sp. Q, Cis sp. S, Cis sp. Y (see explanation on morphospecies’ coding in the section on Materials and Methods) and the invasive species Cer. tabellifer.

Discussion

Cis mooihoekite sp. n. is here included in the bilamellatus-group, which is characterised by the following features (based on Lawrence 2016): (i) presence of a single plate on both anterocephalic edge and anterior pronotal edge in males; (ii) females with pronotum usually widest near the posterior end and gradually narrowing anteriorly; (iii) dual elytral vestiture (sometimes difficult to determine); and (iv) outer apical angle of protibia toothed in males but expanded with several socketed spines in females. Information on the morphology of sclerites of male terminalia is available for species of the bilamellatus-group, except for C. biscutatus and C. lineatosetosus (Lopes-Andrade et al. 2009, Lawrence 2016). The aedeagus is characterised by tegmen widest at middle or widest near apex, usually bearing an emargination on both sides of the apex and penis usually subcylindrical and without struts, as long as or slightly longer than tegmen.

Amongst African species, Cis mooihoekite sp. n. not only differs from C. pickeri in the features mentioned in the section on “Diagnosis”, but also in its geographic distribution. Cis mooihoekite sp. n. occurs only in areas of Grassland or Savannah at Mpumalanga Province, far from the coast, while C. pickeri occurs only in areas of Fynbos vegetation at Western Cape Province, close to the coast (Fig. 13).

Amongst non-African species of the bilamellatus-group, Cis mooihoekite sp. n. mostly resembles C. onyosi from Spain and C. clarki and C. setiferus from Australia. It differs from C. onyosi in the anterocephalic edge with corners more acute and further away from the median anterocephalic plate (Fig. 7) and the comparatively more trapezoidal pronotum with straight base (Viñolas and Muñoz-Batet 2015) (Fig. 7); it differs from C. setiferus in bearing smaller and margined abdominal sex patch (Figs 3 and 4) and anterocephalic edge with corners more acute (Lawrence 2016) (Figs 1, 4 and 7); and from C. clarki in the smaller body length and male abdominal sex patch, larger and less seriate elytral punctures, comparatively shorter pronotal plates and anterocephalic edge with acute corners (Lawrence 2016).

Except for C. onyosi, all described species of the bilamellatus-group occur in the Southern Hemisphere, mostly in subtropical and temperate areas (Lopes-Andrade et al. 2009, Lawrence 2016). The occurrence of C. onyosi in Europe, recently described from Spain and reported from France (Rose 2017, Viñolas and Muñoz-Batet 2015), may well be a case of biological invasion. In such case, C. onyosi would be the second invasive species of the bilamellatus-group to reach the Northern Hemisphere, after C. bilamellatus (Orledge et al. 2010).

Acknowledgments

We would like to especially thank John F. Lawrence, who first recognised the new species in the material belonging to CMN and Ottilie Neser, Riaan Stals and Ros Urban for the great opportunity to work on hundreds of ciids belonging to SANC. We also thank Glenda Orledge, John F. Lawrence, Vivian E. Sandoval Gómez, one anonymous reviewer and the editor Yasen Mutafchiev for valuable corrections and suggestions to the text. I.S.G. thanks the Programa de Pós-Graduação em Ecologia (Universidade Federal de Viçosa, Brazil) for the academic support. Financial support was provided by Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG; Edital 01/2016 - Demanda Universal, APQ-02675-16), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; research grant to C.L.A. n° 307116/2015-8) and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES; doctorate degree grant to I.S.G., PROTAX – Edital n° 001/2015).

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