Two new cleistocheliferous species of Clathria of sciophilous habitats from Northeastern Brazil (Poecilosclerida: Demospongiae: Porifera)

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Article

ISSN 1175-5326 (print edition)

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ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3900.1.6 http://zoobank.org/urn:lsid:zoobank.org:pub:502D2292-E295-4CE8-9D89-72396685BD1F

Two new cleistocheliferous species of Clathria of sciophilous habitats from Northeastern Brazil (Poecilosclerida: Demospongiae: Porifera) GEORGE GARCIA SANTOS1 & ULISSES PINHEIRO1,2 1

Universidade Federal de Pernambuco, Centro de Ciências Biológicas, Departamento de Zoologia, Av. Nelson Chaves, s/n Cidade Universitária CEP 50373-970, Recife, PE, Brazil 2 Corresponding author. E-mail: [email protected]

Abstract We describe two new species of Clathria (Microciona) Bowerbank, 1862: C. (M.) crassitoxa sp. nov. and C. (M.) trairae sp. nov. from Paraíba State (Northeastern Brazil). All are sciophilous species and are represented by small fragments removed from the substrate. Both new species have cleistochelae, and are compared with their cleistocheliferous congeners, differing from all of them by the possession of different combinations of other megascleres and microscleres. Key words: sponges, Clathria, new species, cleistochelae, Paraíba State, taxonomy, Brazil

Introduction According van Soest (2009), sciophilous (shaded) marine habitats are characterized by mosaics of small and thinly encrusting faunal inhabitants, notably sponges, bryozoans and colonial tunicates. Sizes of these organisms usually are measured in millimeters rather than centimeters, making sampling and identification often problematic. In sponges in particular, the taxonomy of thinly encrusting specimens is challenging owing to the difficulties in sampling their thin tissue (Zea et al. 2014). Thinly encrusting species of Clathria Schmidt, 1862 are generally difficult to recognize in the field and from their simple spiculation (Hooper 1996). Clathria (Microciona) Bowerbank, 1862 have 99 valid described species, however, only two of them are recorded from the Brazilian Coast: C. (M.) calla de Laubenfels, 1934 and C. (M.) campecheae Hooper, 1996. Neither of these two Brazilian Clathria species have cleistochelae (viz. isochelae with central teeth (alae) nearly in contact or touching - see Boury-Esnault & Rützler 1997). In total, only nine species of cleistocheliferous Clathria are known worldwide, viz. Clathria (Microciona) bicleistochelifera van Soest et al., 2013 (from Cape Verde), C. (M.) cancapseptima van Soest et al., 2013 (from Cape Verde), C. (M.) cleistochela Topsent, 1925 (from Italy), C. (M.) echinata (Alcolado, 1984) from Cuba, C. (M.) elliptichela (Alander, 1942) from Celtic Seas, C. (M.) toxirecta (Sarà & Siribelli, 1960) from Italy, C. (Clathria) hjorti (Arnesen, 1932) from Mauritania, C. (Thalysias) sulfocleistochela Zea et al., 2014 (from Colombia), and C. (C.) tortuosa Uriz, 1988 (from Namibia). In this paper, we describe two new cleistocheliferous species of Clathria (Microciona), collected from sciophilous habitats in the coast of the Paraiba State (Northeastern Brazil Ecoregion).

Material and methods The specimens were collected during a faunistic survey conducted in the area of Baía da Traição city (6º41’19”S, 34º55’60”W), Paraíba State, Brazil (Figure 1). This is an important area for tourism and as such a more comprehensive knowledge of marine life in the region is essential. The specimens studied here were preserved in 92% ethanol. The sponges were deposited in the Coleção de Porifera of Universidade Federal de Pernambuco (UFPEPOR). Dissociated spicule mounts, scanning electronic microscopy (SEM) preparations and skeletal sections were made using classical procedures for Demospongiae (Hajdu et al. 2011). Images of specimens, Accepted by J. Hooper: 25 Nov. 2014; published: 19 Dec. 2014

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sections and SEM preparations were obtained digitally. Higher systematics of the species follows the Systema Porifera (Hooper 2002; Hooper & van Soest 2002). Taxonomic comparisons were made with valid species of Clathria with cleistochelae listed in the World Porifera Database (van Soest et al. 2014).

FIGURE 1. Location of the study area, Baía da Traição city, Brazil.

Results and discussion Systematics Class Demospongiae Order Poecilosclerida Topsent, 1928 Family Microcionidae Carter, 1875 Genus Clathria Schmidt, 1862 Subgenus Clathria (Microciona) Bowerbank, 1862 Synonymy. For synonymy see Hooper (2002). Definition. Clathria with persistently encrusting growth form, with hymedesmioid skeletal architecture consisting of a basal layer of spongin, typically with ascending, plumose, non-anastomosing spongin fibre nodes, and megascleres embedded and erect on basal layer (Hooper 2002). Type species. Microciona atrasanguinea Bowerbank, 1862 (by subsequent designation; Bowerbank, 1864: 188).

Clathria (Microciona) crassitoxa sp. nov. (Figures 2–3, Table 1) Type specimens: Holotype. UFPEPOR 1474, Baía da Traição (6º41’9.88”S, 34º55’55.62”W), Paraíba State,

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Brazil, 1 m depth, coll. G. G. Santos, 13 December 2012. Paratypes: UFPEPOR 1581, 1598, 1610, Baía da Traição (6º41’9.88”S, 34º55’55.62”W), Paraíba State, Brazil, 1 m depth, coll. G. G. Santos, 07 November 2013. Diagnosis. Thinly encrusting Clathria (Microciona) from the Atlantic with two size categories of cleistochelae and one category of oxhorn toxas, two categories of choanosomal principal styles and one category each of echinating acanthostyles and subectosomal styles. External morphology (Fig. 2A–E). A thinly encrusting sponge (< 1 mm thick) on stones and polychaete worm tubes (Fig. 2A–C). Consistency soft and fragile, easily torn. Surface is hispid and texture is velvety (Fig. 2D–E). No oscules were seen. Live-colour orange (Fig. 2A, C) to yellowish-beige (Fig. 2B), turning beige after preservation in ethanol.

FIGURE 2. Clathria (Microciona) crassitoxa sp. nov. A–C, photographs of fresh specimens taken out of water; D–E, surface details; F, cross section of skeleton showing microcionid skeletal structure; G, overview of peripheral skeleton showing cleistochelae and bouquets of subtylostyles. Scale bars: A, B = 1 cm; C = 1.5 cm; D = 3 mm; E = 1 mm; F = 100 µm; G = 200 µm.

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FIGURE 3. Spicule composition of Clathria (Microciona) crassitoxa sp. nov. (UFPEPOR 1474, holotype) in SEM. A, Choanosomal principal style (I); B–C, Choanosomal principal styles (II); D, auxiliary subtylostyle; E, echinating acanthostyle; F, detail of the head of A; G–I, details of respectively C–E; J, cleistochela I; K, cleistochelae II; L, oxhorn toxa. Scale bars: A = 100 µm; B–C = 40 µm; D = 50 µm; E = 15 µm; F = 10 µm; G = 8 µm; H = 3 µm; I = 4 µm; J = 10 µm; K = 8 µm; L = 2.5 µm.

Skeleton (Fig. 2F–G). Typical Clathria (Microciona) hymedesmioid – microcionid skeletal arrangement, with a basal layer of spongin from which short erect fibres arise echinated by megascleres (including echinating acanthostyles). Principal choanosomal megascleres and echinating acanthostyles stand erect on the substrate (thin sections of the skeleton), held erect by the basal layer of spongin, but also coring or echinating the erect fibre nodes slightly above the basal spongin layer (Fig. 2F). In thicker sections of the skeleton subectosomal auxiliary megascleres are arranged in disorganized bouquets, protruding through the surface (Fig. 2G). There is no second category of auxiliary spicule in the ectosomal region. Microscleres of variable abundance are scattered throughout the skeleton. Spicules (Fig. 3A–L). Megascleres: Choanosomal principal style with a large size range. Larger (I) principal style (328–570.0–715 / 16–22.0–26 µm) relatively long, robust, with an evenly rounded smooth head, straight to slightly curved shaft (Fig. 3A, F). Smaller (II) principal style (132–178.2–219 / 6.4–11.1–16.1 µm) robust, straight or slightly curved shaft, only occasionally rugose and most are entirely smooth, some with a subterminally constricted subtylote base (Fig. 3B, C, G). Subectosomal auxiliary subtylostyle (177–301.6–438 / 2.5–3.9–6.4 μm) straight, thin, long smooth shaft, with a slightly swollen microspined base (Fig. 3D, H). Echinating acanthostyle (62–68.8–80 / 6–7.1–8 μm) spined all over or only partially spined on the base and mid-section of the shaft, with spines sharp, stout, frequently bent towards the base of the spine (Fig. 3E–I). Microscleres: Cleistochelae (I) (22–26.0–29 μm long): with alae almost touching, with space between the alae filled in with a blade-like extension of the shaft (Fig. 3J). Cleistochelae (II) (15–16.1–16.8 μm long) shorter but similar to the previous (Fig. 3K).

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Regular palmate isochelae absent. Toxa (9–9.6–10 / 1 µm) small, oxhorn shape that is slightly bent with points faintly recurved, tapering relatively evenly from the central thickest part towards their points (Fig. 3L). Ecology. This sciophilous species is often found in reefs environments, encrusting undersides of dead corals and on tubes of the polychaete worm Sabellaria bella (Grube, 1870). Distribution (Fig. 1). Brazil: Northeastern Region: Paraíba State: Baía da Traição. Depth. 1 m. Etymology. Latin crassi, which means thick, in reference to the robust oxhorn toxas in this species. Remarks. Over 340 species of Clathria are listed in the World Porifera Database (van Soest et al. 2014). However, only eleven of these species have cleistochelae, including the two new species described here (see Table 1). All these species have important points of difference compared to our two new species. With the exception of C. (M.) bicleistochelifera (from Cape Verde), none of the other previously known nine species have two categories of cleistochelae. However C. (M.) bicleistochelifera differs from the new species by having a blood-red to red colouration alive, compared to Clathria (M.) crassitoxa sp. nov. that is distinctively yellow alive. This new species also has two size classes of principal styles, one class of echinating acanthostyles, one morphology of oxhorn toxas, but lacks normal palmate isochelae (Table 1). The possession of cleistochelae also differentiates Clathria (M.) crassitoxa sp. nov. from the only two previously known species of Clathria (Microciona) recorded from Brazil (Clathria (M.) calla de Laubenfels, 1934 and Clathria (M.) campecheae Hooper, 1996).

Clathria (Microciona) trairae sp. nov. (Figures 4–5, Table 1) Type specimens: Holotype. UFPEPOR 1478, Baía da Traição (6º41’8.88”S, 34º55’55.62”W), Paraíba State, Brazil, 1 m depth, coll. G. G. Santos, 07 November 2012. Diagnosis. Clathria (Microciona) trairae sp. nov. is the only species of Clathria (Microciona) in the Atlantic with two classes each of cleistochelae, toxas and principal styles, and one class of unmodified palmate isochelae, auxiliary subtylostyles and echinating acanthostyles. External morphology (Fig. 4A–E). Thinly to massively incrusting, up to 1 cm thick, with tubular elevations; surface is hispid from protruding spicules. Consistency spongy, compressible and fragile. Colour bright red alive, turning beige after preservation in ethanol. No oscules were seen. Skeleton (Fig. 4D, E). Microcionid to plumose skeletal architecture. The ectosome is a thin cellular pinacoderm (Figure E), supported by brushes of larger principal megascleres (Fig. 4D–E). In thicker sections of the skeleton there are intermediate tracts/ brushes of principal megascleres. Auxiliary megascleres are scattered in the subectosomal and ectosomal region, crowning the spongin fiber endings and protruding beyond the surface (Fig. 4D). In the choanosome the fiber columns are sparingly cored by spicules (principal and auxiliary megascleres) in cross section (Fig. 4D). In the basal plate, principal, auxiliary megascleres and echinating acanthostyles stand erect on the substrate (Fig. 4E). Microscleres are diffusely scattered throughout the skeleton. Spicules (Fig. 5A–K). Megascleres: Choanosomal principal style (I) (397–552.0–702 / 16–23.0–29 µm) long, thin, smooth, with straight to slightly curved shaft (Fig. 5A’, D); Choanosomal principal style (II) (196–260.0–325 / 13–15.4–16 μm) smooth, short, stout, usually straight (Fig. 5A”). Subectosomal auxiliary subtylostyle (190–299.0–493 / 3–4.9–7 μm) straight, thin, long, smooth shaft, with slightly swollen microspined base (Fig. 5B, E); Echinating acanthostyle (175–217.9–292 / 7–7.6–8 μm) slightly curved, slightly fusiform, tapering gradually to a sharp apex, base slightly swollen, roundish, irregularly acanthose, with variably sharp spines (resembling blunt warty thorns), and completely smooth shaft (Fig. 5C, F). Microscleres: Cleistochelae (I) (39–43.8–47 μm long) with alae almost touching, and with the space between the alae filled-in with a blade-like extension of the shaft (Fig. 5I); Cleistochelae (II) (16–19.0–24 μm long) shorter and with curved front alae (Fig. 5J); Oxeote toxa (I) (241–289.8–344 / 8–10.2–13 μm) long, stout, with a gradual shallow v-shaped curve (Fig. 5G); Oxhorn toxa (II) (6.4–20.9–45 / 1–1.5 μm) small, thin, shallow-curved centrally and slightly recurved points (Fig. 5K). Unmodified palmate isochelae (21–23.8–29 μm long) mostly with nearly straight, slender shaft, slightly curved alae (Fig. 5H). Ecology. Only one specimen of this species was found at Baía da Traição. This species was found in a sciophilous habitat. Distribution (Fig. 1). Brazil: Northeastern Region: Paraíba State: Baía da Traição.

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Habana (Cuba) / 13

Yucatan, Campeche / 2–120

Colombia (Caribbean) / 4.5–35

Naples (Itay), Azores / 10 Cape Verde Islands / 2–15

Cape Verde Islands / 16– 70 Morocco / 39

Namibia / 290

Clathria (M.) echinata (Alcolado, 1984)1

Clathria (M.) echinata (Alcolado, 1984)6

Clathria (Thalysias) sulfocleistochela Zea et al., 20147

Clathria (M.) cleistochela (Topsent, 1925)2 Clathria (M.) bicleistochelifera van Soest et al., 20132

Clathria (M.) cancapseptima van Soest et al., 20132 Clathria (Clathria) hjorti (Arnesen, 1932)2

Clathria (C.) tortuosa Uriz, 19884

166–450 / 2–7.8 (spiny or smooth heads)

210–750 / 2–5

190–299.0–493 / 3–4.9–7

177–301.6–438 / 2.5–3.9–6.4

Subectosomal Subtylostyles

Encrusting / not recorded

383–880 / 7–17

400–850 / 5–7

I - 136–336.8–726 / 16–21.1–36 II - 72–123.8–204 / 6–10.2–22 (spined or warty heads) -

ramose / bright red

Encrusting to ramose / cream

186–369.7–564 / 7–12.6–19

Encrusting / yellow to red

182–252 / 5–7.7

-

285–610 / 3–7

122–238.0–324 / 2–3.1–7

243–363.5–516 / 2–4.7–8

-

I - 1800 / 22 II - 140 / 8

I - 560–1010 / 36–46 II - 210–439 / 15–25

-

17.5–21

15–16 (as elliptical chelae)

15–19

14–16.6–21

16–21.3–27

I - 22–27.1–31 II - 13–16.9–20

-

75–113.0–168 / 4–6.9–11

13.7–16.6

7.5–13.2–16.1

15.8–36

14–25 (as “aspidoquelas”)

I - 39–43.8–47 II - 16–19.0–24

I - 22–26.0–29 II - 15–16.1–16.8

Cleistochelae

76–283.5 / 5–10.5

59–109.1–146 / 4.6–5.2–6.7

-

-

175–217.9–292 / 7–7.6–8

62–68.8–80 / 6–7.1–8

Echinating Acanthostyles

28–35 / 0.7

I - 45–300 / 0.5–4 II - 6–12 / 1–2

I - 80–120 / 3–4 II - 20–46 / 1–1.5

18–68.9–135

30–108.6–294

I - 60.2–100 / 0.7–2.5 II - 27.2–38.5 / 0.5–1.0 -

I - 146.1–642 / 2–5 (oxeote) II - 8.8–13.4 (oxhorn) -

40–1084 / 0.5–5

I - 241–289.8–344 / 8–10.2–13 (oxeote) II - 6.4–20.9–45 / 1–1.5 (oxhorn)

9–9.6–10 / 1 (oxhorn)

Toxas

-

14–16

-

-

13–15.8–19

16–19.3–22

-

11–13.5–16

16–20.2

16–26

21–23.8–29

-

Palmate Isochelae

References: (1) Alcolado (1984); (2) van Soest et al. (2013); (3) Alander (1942); (4) Uriz (1988); (5) Sarà & Siribelli (1960); (6) Gómez (2014, redescription); (7) Zea et al. (2014).

Clathria (M.) toxirecta (Sarà Napoli (Italy) / & Siribelli, 1960)5 40

I - 187–620 / 10.4–26 II - not recorded

I - 397–552.0–702 / 16– 23.0–29 II - 196–260.0–325 / 13– 15.4–16 (spiny or smooth heads) I - 270–920 / 9–31 II - 210–545 / 5–11

I - 328–570.0–715 / 16– 22.0–26 II - 132–178.2–219 / 6.4– 11.1–16.1

Principal Styles

very thin 199–258.4–333 / I - 199–220.9–261 encrusting / dark 4.7–8.1–9.5 / 2.9–3.8–4.8 orange to II - 123–153.9–185 / cinnamon 1.4–2.4–3.8 Encrusting / 85–560 / 5–16 202–371 / bright red 1.7–3.6 Encrusting / 183–370.2–492 / 6–8.2–11 204–283.9–363 / Blood-red or 1–1.9–2.5 bright red

subspherical / bright red or orange erect, tubular or massive-vase / orange or red

Encrusting with tubular elevations / bright red

Encrusting / yellow-beige to orange

Shape / colour

Celtic Seas / 400 Encrusting / not recorded

Paraíba State (Brazil) / 0.5

Clathria (Microciona) trairae sp. nov.

Clathria (M.) elliptichela (Alander, 1942)3

Paraíba State (Brazil) / 1

Location / depth (m)

Clathria (Microciona) crassitoxa sp. nov.

Species

-

-

-

238–288.6–345 / 2.5–3.8–5 (“quasitylotes”) -

-

-

-

-

20 (raphides)

-

-

Other spicules

TABLE 1. Comparative micrometric data on the spicules, shape, depth and distributions of the living species of cleistocheliferous Clathria Schmidt, 1862. Values are in micrometres (μm), expressed as follows: minimum–maximum or minimum–mean–maximum length/width. References are numbered in parentheses and listed at the foot of the table.

FIGURE 4. Clathria (Microciona) trairae sp. nov. (UFPEPOR 1478, holotype). A, specimen in situ at Baía da Traição; B, photograph of a fresh specimen taken out of water, showing surface details; C, tubular elevations with sand grains and spicules; D, thick section showing the ectosome and choanosome; E, detail of a thin section showing the thin ectosome with some spicules echinating the spongin fibres, microscleres and grains of sand. Scale bars: A–B = 1 cm; C = 1 mm; D = 700 µm; E = 150 µm.

Depth. 0.5 m. Etymology. The specific epithet trairae refers to the type locality of Baía da Traição, Brazil, which in Portuguese means ‘betrayer’. Remarks. Although all three species (Clathria (M.) bicleistochelifera, C. (M.) crassitoxa sp. nov. and C. (M.) trairae sp. nov.) share the possession of two size categories of cleistochelae, the latter differs from both the other species by having two morphologies of toxas, modified echinating (acantho)styles with only the vestige of spines on the base, and unmodified palmate isochelae (see Table 1 and Figure 5). Clathria (M.) trairae sp. nov. shares the presence of two categories of toxas with three species of cleistocheliferous Clathria, viz. Clathria (M.) cleistochela, C. (M.) elliptichela and C. (C.) tortuosa. However those other species do not have acanthostyles nor two categories of both cleistochelae and principal styles as found in the new species (see Table 1). The closest

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species to C. (M.) trairae sp. nov. is C. (M.) echinata, sharing a live red color and the presence of styles I, toxas I, and palmate isochelae. However, C. (M.) echinata has only one category of subtylostyles and have toxas and isochelae smaller than the new species (see Table 1). Furthermore, C. (M.) echinata shows a wide morphological variation in its growth form, ranging from tubular or vase-shaped, upright massive, subspherical and encrusting morphotypes (Gomez 2014). Finally, the presence of cleistochelae also differentiates Clathria (M.) trairae sp. nov. from the two previously known species of Clathria (Microciona) recorded from Brazil (Clathria (M.) calla de Laubenfels, 1934 and Clathria (M.) campecheae Hooper, 1996).

FIGURE 5. Spicule composition of Clathria (Microciona) trairae sp. nov. (UFPEPOR 1478, holotype) in SEM. A’, Choanosomal principal style (styles I); A”, Choanosomal auxiliary style (styles II); B, Auxiliary subtylostyle; C, Echinating acanthostyle; D–F, details of respectively A–C; G, oxeote toxa I; H, palmate isochelae; I, back and side views of cleistochelae I; J, cleistochelae II; K, oxhorn toxa II. Scale bars: A = 100 µm; B = 50 µm; C = 60 µm; D = 15 µm; E–F = 3.5 µm; G = 40 µm; H = 10 µm; I = 15 µm; J–K = 10 µm.

Discussion Topsent (1925) proposed the term “cleistochelae” for palmate isochelae in which the free alae from opposite sides (almost) touch and the shaft has a plate-like ridge (almost completely) filling the space between the alae. The cleistochelate condition of the palmate isochelae is shared with some other genera of Poecilosclerida, viz. Mycale Gray, 1867 (as “naviculichelae”, Hajdu 1999), Iophon Gray, 1867, Abyssocladia Lévi, 1964, and Antho Gray, 1867. However, the two new species described here display the typical morphology of Clathria (Microciona): an encrusting growth form, a reduced basal (hymedesmioid) spongin skeleton with erect fibre nodes cored by plumose tracts of principal and echinating spicules standing erect on the substrate (=‘microcionid’ architecture of Lévi

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1960), and smooth toxas (Hooper 2002). In Clathria, according van Soest et al. (2013), the similarity in cleistochelae micromorphology between all the species is striking and indicates the existence of a possibly related group of species in the Atlantic region transgressing the subgenus boundaries, e.g. Clathria (Clathria) Schmidt, 1862; Clathria (Microciona) and Clathria (Thalysias) Duchassaing & Michelotti, 1864. Genera such as Haliclona Grant, 1836, Clathria, and Mycale, are common in reef sciophilous habitats (Kobluk & van Soest 1989). For example, van Soest (2009) has shown that sponges inhabiting the undersides of coral rubble and crevices are not just juveniles of more massive species, but comprise a distinct assemblage of sponges. These have been largely over-looked in the past because they are usually cryptic and small, making sampling and identification difficult. These hidden habitats also complicate the collection of specimens, which by necessity has to be by indirect means, and only scuba diving and snorkeling permit an efficient study of these animals. In Brazil, the few studies that been carried out on the sciophilous sponge faunas (see Cedro et al. 2013; Santos et al. 2014a, b) resulted in the discovery of several species new to science. The two new species described here may be considered ‘moderately sciophilous’, since they occur at very shallow depths in very turbid waters. These sponges are restricted to marine environments with low water movement conditions, which might favor the high fragility of the ‘microcionid’ skeletal structure.

Acknowledgments G.S. thanks CAPES for providing a doctoral scholarship. Authors are thankful to Dra Janaina Melo, Dyego Oliveira, and Josineide Correia for SEM facilities at CETENE (Centro de Tecnologias Estratégicas do Nordeste), Dr. Carlos Perez (UFPE), M.Sc. Felipe Campos, Diego Aquino, M.Sc. Luis Aquino, M.Sc. Danielle Lopes, Adélia Alliz, Dr. José Barbosa (UFPB), Dr. Leandro Vieira (UFPE), Dra Adriane Pereira Wandeness (UFPB) and to Dr. André Esteves (UFPE) for technical support. We further thank CNPq (Edital PROTAX: 562320/2010-5), and FACEPE, for providing grants and/or fellowships.

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