Research Article |
Corresponding author: Celine S. S. Lopes ( celinelopes@outlook.com ) Academic editor: James Reimer
© 2019 Celine S. S. Lopes, Hellen Ceriello, André C. Morandini, Sérgio N. Stampar.
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:
Lopes CSS, Ceriello H, Morandini AC, Stampar SN (2019) Revision of the genus Ceriantheomorphe (Cnidaria, Anthozoa, Ceriantharia) with description of a new species from the Gulf of Mexico and northwestern Atlantic. ZooKeys 874: 127-148. https://doi.org/10.3897/zookeys.874.35835
|
The present study presents a revision of the genus Ceriantheomorphe Carlgren, 1931, including redescriptions of the two presently recognized species, Ceriantheomorphe ambonensis (Kwietniewski, 1898) and Ceriantheomorphe brasiliensis (Mello-Leitão, 1919), comb. nov., and a description of the new species Ceriantheomorphe adelita sp. nov.
Biogeography, cnidarian taxonomy, North America, Pacific Ocean, South America
Ceriantharia is a subclass within the cnidarian class Anthozoa, consisting of species commonly known as tube anemones. This taxon has several taxonomic inconsistencies (
The genus Ceriantheomorphe was described by
However, assigning Cerianthus ambonensis as “Ceriantheomorphe ambonensis” would have been premature because the simple description made by
More than two decades after the description of Ceriantheomorphe,
Despite this taxonomic confusion, Ceriantheomorphe brasiliensis had been listed as an endangered species in Brazil for over 10 years (
This study aims to present a taxonomic review of the genus Ceriantheomorphe including a redescription of the holotype of C. brasiliensis, a redescription of C. ambonensis, and the description of a new species from the North Atlantic.
Twenty specimens of Ceriantheomorphe were sampled by SCUBA: sixteen of C. brasiliensis from the South Atlantic, three from the North Atlantic, and one, C. ambonensis, from the Pacific Ocean (Table
List of Ceriantheomorphe specimens in this study. Abbreviations: ES = Espírito Santo State; RJ = Rio de Janeiro State; SP = São Paulo State; SC = Santa Catarina State;
Species | Country | Locality | Coordinates | Museum code |
---|---|---|---|---|
C. brasiliensis | Brazil | Guanabara Bay-RJ | 22°49'6''S, 43°8'45''W | MNRJ 200 |
Arraial do Cabo-RJ | 23°0'4''S, 42°0'29''W |
|
||
Araçá Beach-SP | 23°48'58''S, 45°24'24''W |
|
||
Araçá Beach-SP | 23°48'58''S, 45°24'24''W |
|
||
Cagarras Islands-RJ | 23°1'55''S, 43°11'58''W |
|
||
Canasvieiras-SC | 27°25'31''S, 48°27'0.2''W |
|
||
Camburi Beach-ES | 20°16'39''S, 40°16'29''W |
|
||
Camburi Beach-ES | 20°16'39''S, 40°16'29''W |
|
||
Rio de Janeiro-RJ | – |
|
||
Urca-RJ | – |
|
||
Zimbro Beach-SP | 23°49'27''S, 45°25'4''W |
|
||
Sabacu Island-RJ | 23°0'43''S, 44°22'7''W | MNRJ 2766 | ||
Uruguay | José Ignacio-Maldonado | 35°00'S, 54°24'2''W |
|
|
La Paloma-Rocha | 34°42'3''S, 54°0.5'W |
|
||
La Paloma-Rocha | 34°42'3''S, 54°0.5'W |
|
||
Punta del Diabo | 34°04'S, 53°29'W |
|
||
C. adelita sp. nov. | Mexico | Punta de Almagre-Tamaulipas | – |
|
United States of America | Pass A’Loutre-Louisiana | – |
|
|
Port Aransas, Corpus Christi-Texas | – |
|
||
C. ambonensis | Indonesia | Jakarta Bay-Jakarta | – |
|
The morphology of all specimens was studied through internal anatomy and cnidome studies, both based on criteria adopted by
All protomesenteries/directive mesenteries (P) were measured. Five quartets of mesenteries were measured for each specimen. We also divided the metamesenteries (type M and type m) value and betamesenteries (type B and type b) value to calculate the ratio between these mesentery types. We calculated the proportion occupied by protomesenteries in the gastrovascular cavity using the following equation:
F (length of protomesentery) × 100 / E (length of gastrovascular cavity)
The cnidome study was based on the sampling of 30 cnidae capsules for each cnida type from each body region (superior tip of marginal and labial tentacles, actinopharynx region, column, metamesenteries and betamesenteries). Each cnida was classified according to their shape based on different authors (
Subclass Ceriantharia Perrier, 1893
Suborder Spirularia den Hartog, 1977
Family Cerianthidae Milne-Edwards & Haime, 1851
Cerianthidae with fertile mesenteries, except for directives. Two pairs of mesenteries connected to the siphonoglyph. Mesenteries grouped in quartets following M, B, m, b order (after
Ceriantheomorphe brasiliensis (Mello-Leitão, 1919).
Ceriantheomorphe brasiliensis (Mello-Leitão, 1919) new comb., Ceriantheomorphe ambonensis (Kwietniewski, 1898), Ceriantheomorphe adelita sp. nov.
Southwestern Atlantic (Brazil and Uruguay), Gulf of Mexico (United States of America and Mexico), Central West Pacific (Java Sea, Indonesia).
Cerianthus brasiliensis Mello-Leitão, 1919: 38–39.
Ceriantheomorphe brasiliensis
sensu
Ceriantheomorphe brasiliensis
(not) –
Holotype: MNRJ 200 • adult individual (16.5 cm long), Guanabara Bay, Rio de Janeiro, Brazil (22°49'6"S, 43°8'45"W), Mello-Leitão leg. (Fig.
Dissected specimens of Ceriantheomorphe brasiliensis from southwestern Atlantic. A Individual
Large cerianthid, 8.5–24 cm long and 1.5–13.8 cm wide. 132–392 marginal tentacles arranged in (1)1123.1123 and 108–384 labial tentacles arranged in (1)1122.1122 or (1)1123.1123. Pharynx occupies about 8–27% of total body length. Five pairs of protomesenteries, of which two pairs connected to the siphonoglyph, (directives and P2). Gastrovascular cavity takes up to 33–72% of total body length. All fertile mesenteries, except for directives. Number of mesenteries about 170–642. Directives mesenteries longer than protomesenteries P3, P5 and metamesenteries m, except by m of the 2nd and 3rd cycles. Protomesenteries (P2) longer than all mesenteries, extending up to the aboral pore (Fig.
Measurements of 30 cnida capsules for each cnida type in 6 distinct body regions of Ceriantheomorphe brasiliensis (N = 16). Information inside parentheses indicates cnidae length and width, respectively, and information outside parentheses indicates average of cnidae size.
Body part/cnida type | Ceriantheomorphe brasiliensis |
---|---|
Marginal tentacles | |
Microbasic b-mastigophore type I | 65.56 (50.50–80.63) × 13.13 (7.57–18.69) |
Microbasic b-mastigophore type II | 38.23 (27.96–48.5) × 4.99 (3.13–6.86) |
Microbasic b-mastigophore type III | 31.16 (18.36–43.97) × 3.96 (1.97–5.95) |
Microbasic b-mastigophore type IV | 16.55 (10.61–22.49) × 4.10 (2.2–6.01) |
Microbasic b-mastigophore type V | 27.87 (18.01–37.73) × 7.02 (1.6–5.42) |
Labial tentacles | |
Microbasic b-mastigophore type I | 48.75 (36.89–60.61) × 9.11 (5.41–12.82) |
Microbasic b-mastigophore type II | 34.93 (25.2–44.66) × 5.12 (3.65–6.6) |
Microbasic b-mastigophore type III | 28.27 (17.20–39.35) × 4.03 (1.71–6.35) |
Microbasic b-mastigophore type IV | 24.11 (17.25–30.97) × 2.73 (1.64–3.83) |
Microbasic b-mastigophore type V | 26.10 (15.03–37.18) × 3.29 (1.79–4.79) |
Pharynx | |
Atrichous type I | 38.33 (26.15–50.52) × 5.95 (2.68–9.22) |
Microbasic b-mastigophore type I | 52.64 (35.56–69.73) × 8.38 (5.43–11.33) |
Microbasic b-mastigophore type II | 44.39 (32.10–56.68) × 6.09 (3.28–8.91) |
Microbasic b-mastigophore type III | 34.97 (21.86–48.09) × 3.35 (2.13–4.57) |
Microbasic b-mastigophore type V | 27.62 (23.37–31.88) × 2.81 (2.19–3.43) |
Column | |
Ptychocyst type I | 71.99 (56.21–87.77) × 24.41 (13.75–35.08) |
Ptychocyst type II | 77.14 (50.15–94.14) × 14.12 (8.86–19.38) |
Atrichous type I | 48.85 (30.09–67.61) × 11.09 (4.41–17.78) |
Microbasic b-mastigophore type I | 41.33 (26.47–56.2) × 6.25 (3.96–8.54) |
Microbasic b-mastigophore type IV | 28.14 (23.83–32.45) × 3.12 (2.48–3.76) |
Microbasic b-mastigophore type V | 29.95 (22.51–37.4) × 3.03 (2.08–3.98) |
Holotrichous | 50.95 (30.04–71.86) × 14.88 (7.3–22.46) |
Mesenteries M | |
Microbasic b-mastigophore type I | 51.58 (35.0–68.17) × 10.09 (6.41–13.77) |
Microbasic b-mastigophore type IV | 22.25 (10.93–33.58) × 5.76 (2.25–9.28) |
Microbasic b-mastigophore type III | 20.03 (13.3–26.77) × 4.90 (2.91–6.9) |
Mesenteries b | |
Microbasic b-mastigophore type I | 54.65 (39.57–69.74) × 10.44 (7.16–13.73) |
Microbasic b-mastigophore type II | 33.69 (24.83–42.56) × 5.01 (3.32–6.7) |
Microbasic b-mastigophore type III | 19.97 (12.1–27.85) × 4.17 (1.95–6.4) |
Microbasic b-mastigophore type IV | 19.59 (8.62–30.56) × 4.06 (2.24–5.89) |
Southwestern Atlantic-Brazil (from the State of Espírito Santo (20.5°S) to Rio Grande do Sul (33.7°S) State) and Uruguay (34°S). This species was only observed in shallow waters (1–40 m depth).
(MNRJ 200) (Fig.
Cerianthus ambonensis
Kwietniewski, 1898: 426;
Cerianthus sulcatus McMurrich, 1910: 28–30.
Ceriantheomorphe ambonensis
–
(
Small cerianthid, 3.8 cm long and 2.1 cm wide. 48 marginal tentacles and 72 labial tentacles, both disposed in three cycles. Directive marginal and labial tentacles absent. Marginal tentacles arrangement: (0)1123.1121.1213.1213... Labial tentacles arrangement: (0)112.1121.1121.1121… Pharynx occupies about 18% of total body length. Hyposulcus and hemisulci absent. Gastrovascular cavity occupies about 55% of total body length. Three pairs of protomesenteries, all connected to the siphonoglyph (directive mesenteries, P2 and P3). About 96 mesenteries arranged in M,B,m,b (Fig.
Measurements of 30 cnida capsules for each cnida type in 6 distinct body regions of Ceriantheomorphe ambonensis (N = 1). Information inside parentheses indicates cnidae length and width, respectively, and information outside parentheses indicates average of cnidae size.
Body part/cnida type | Ceriantheomorphe ambonensis |
---|---|
Marginal tentacles | |
Microbasic b-mastigophore type II | 36.02 (23.16–48.89) × 6.18 (4.89–7.47) |
Microbasic b-mastigophore type IV | 19.54 (14.42–24.66) × 6.18 (4.89–7.47) |
Microbasic b-mastigophore type V | 18.90 (16.21–21.60) × 2.56 (2.22–2.90) |
Labial tentacles | |
Microbasic b-mastigophore type I | 46.84 (42.40–51.28) × 8.05 (6.46–9.65) |
Microbasic b-mastigophore type II | 30.31 (26.15–34.47) × 4.58 (3.30–5.87) |
Microbasic b-mastigophore type III | 27.68 (24.16–31.20) × 3.54 (2.83–4.25) |
Microbasic b-mastigophore type V | 23.52 (18.13–28.92) × 2.82 (2.05–3.59) |
Pharynx | |
Atrichous | 40.36 (33.48–47.25) × 5.99 (4.81–7.17) |
Microbasic b-mastigophore type I | 50.45 (44.63–56.28) × 7.49 (5.92–9.07) |
Microbasic b-mastigophore type II | 36.49 (32.28–40.70) × 5.17 (3.58–6.76) |
Microbasic b-mastigophore type III | 29.92 (24.42–35.42) × 3.59 (2.48–4.71) |
Column | |
Ptychocyst | 61.96 (53.31–70.62) × 21.63 (17.22–26.05) |
Atrichous | 48.50 (41.69–55.32) × 11.38 (8.74–14.03) |
Microbasic b-mastigophore type I | 41.45 (34.51–48.39) × 9.64 (8.74–10.54) |
Holotrichous | 55.10 (47.45–62.76) × 14.97 (11.27–18.68) |
Mesenteries M | |
Microbasic b-mastigophore type I | 49.11 (43.91–54.31) × 9.24 (6.92–11.57) |
Microbasic b-mastigophore type IV | 19.03 (16.70–21.37) × 4.99 (3.38–6.61) |
Mesenteries b | |
Microbasic b-mastigophore type IV | 22.34 (16.34–28.34) × 5.93 (4.10–7.76) |
Indonesia, shallow waters.
Small individual, with 3.8 cm long and 2.1 cm wide. 48 marginal tentacles and 72 labial tentacles, both disposed in three cycles. Marginal tentacles arrangement: (0)1123.112…, labial tentacles arrangement: (0)112.112.112… Small pharynx, occupies 18% of total body length. Hyposulcus and hemisulci absent. Well-marked siphonoglyph with three pairs of mesenteries connected to it (one pair of directive mesenteries and two pairs of protomesenteries). Long protomesenteries (P2) extending to the terminal pore and longer than other mesenteries. Directive mesenteries shorter than all mesenteries. Protomesenteries (P3) shorter than metamesenteries (M and m) and longer than betamesenteries (B and b). 96 mesenteries arranged in M,B,m,b (Fig.
Ceriantheomorphe brasiliensis
Carlgren, 1931 (in part): 2–6;
Holotype:
Large cerianthid, 19 cm long and 5.0–7.3 cm wide. 192–352 marginal tentacles (2.4–3.0 cm long in preserved animal) and 144 to 336 labial tentacles (0.5–2.0 cm long in preserved animal), both disposed in four cycles. Marginal tentacles arrangement: (0)1123.1122.1122.1123.1122…, labial tentacles arrangement: (0)1123. 1122… Siphonoglyph well-marked by two protuberant tissues. Three pairs of protomesenteries (directive mesenteries, P2 and P3), all connected to the siphonoglyph. Well distinct hyposulcus and hemisulci absent. Protomesenteries (P3) longer than metamesenteries (m). Ratio from 2.7–5.2% between metamesenteries (M × m) and 3% between betamesenteries (B × b). Directive mesenteries, P2 and P3, extend up to 30.5%, 92.5% and 56.4% of total gastrovascular cavity length, respectively. Cnidome (Fig.
Measurements of 30 cnida capsules for each cnida type in 6 distinct body regions of Ceriantheomorphe adelita sp. nov. (N = 3). Information inside parentheses indicates cnidae length and width, respectively, and information outside parentheses indicates average of cnidae size.
Body part/cnida type | Ceriantheomorphe adelita sp. nov. |
---|---|
Marginal tentacles | |
Microbasic b-mastigophores II | 39.19 (34.20–44.18) × 5.04 (4.07–6.01) |
Microbasic b-mastigophores V | 25.12 (20.16–30.09) × 3.04 (2.09–3.99) |
Labial tentacles | |
Microbasic b-mastigophores I | 48.72 (39.22–58.22) × 6.71 (5.24–8.19) |
Microbasic b-mastigophores II | 36.32 (28.18–44.46) × 4.77 (3.55–6.00) |
Pharynx | |
Atrichous | 41.66 (32.23–51.09) × 5.74 (4.13–7.35) |
Microbasic b-mastigophores I | 51.43 (40.10–62.77) × 7.7 (6.25–9.15) |
Microbasic b-mastigophores II | 44.2 (35.29–53.11) × 5.13 (3.97–6.29) |
Microbasic b-mastigophores III | 36.75 (27.57–45.93) × 3.36 (2.53–4.20) |
Column | |
Atrichous | 48.12 (38.88–57.37) × 9.28 (7.38–11.19) |
Ptychocysts type I | 55.42 (50.08–60.77) × 13.92 (9.49–18.35) |
Microbasic b-mastigophores I | 41.74 (47.80–35.68) × 6,0 (4.16–7.84) |
Ptychocysts type II | 64.3 (58.8–69.8) × 17.1 (15.0–19.2) |
Mesenteries M | |
Microbasic b-mastigophores IV | 18.77 (23.27–14.27) × 4.24 (2.5–5.99) |
Mesenteries b | |
Microbasic b-mastigophores II | 38.76 (34.01–43.51) × 4.36 (3.53–5.20) |
Microbasic b-mastigophores III | 19.94 (15.46–24.42) × 4.79 (3.98–5.60) |
Microbasic b-mastigophores IV | 23.37 (19.45–27.29) × 3.12 (2.15–4.10) |
The specific name “adelita” is an allusion to an important group of women that fought during the Mexican Revolution. Occasionally, they adopted the identities of men to join in combat against the enemy.
Gulf of Mexico (Northern Mexico) to North Atlantic (North Carolina, United States of America), shallow waters.
Both Ceriantheomorphe brasiliensis and C. adelita sp. nov. have labial and marginal tentacles disposed in four cycles, whereas C. ambonensis has its tentacles arranged in three cycles. All three species have distinct labial and marginal tentacles arrangements (Table
Comparison of morphological characters between species of the genus Ceriantheomorphe.
Characters | Ceriantheomorphe brasiliensis | Ceriantheomorphe adelita sp. nov. | Ceriantheomorphe ambonensis |
---|---|---|---|
Number of marginal tentacles | 132–392 | 192–352 | 48**-150* |
Number of labial tentacles | 108–384 | 144–336 | 72**-150* |
Tentacular cycles | 4 | 4 | 3* |
Arrangement of marginal tentacles | (1)1123... | (?)1122... | (0)112...** |
Arrangement of labial tentacles | (1)1122… | (?)1122… | (0)112…** |
Proportion between pharynx in relation to body length | 8–27% | 21% | 18%** |
Siphonoglyph | Two pairs of mesenteries connected | Three pairs of mesenteries connected | Three pairs of mesenteries connected** |
Proportion of gastrovascular cavity in relation to body length | 33–72% | 56% | 55%** |
Ratio between mesenteries | 1.2–3.1% (B × b); 1.1–3.1%(m × M) | 3% (B × b); 2.7–5.2% (m × M) | 4% (B × b); 2.2–3.5% (M × m)** |
P1 (directive mesenteries) | Longer than P3, P5, betamesenteries (B and b) and metamesenteries (m), except for m of the 2nd and 3rd cycles. Shorter than P2, P4 and metamesenteries (M). | Longer than betamesenteries (B and b) and metamesenteries (m). Shorter than P2, P3 and metamesenteries (M). | Shorter than mesenteries.** |
P2 | Longer than mesenteries | Longer than mesenteries | Longer than mesenteries |
P3 | Longer than P5, betamesenteries (b) and betamesenteries (B), except for B of the 1st and 2nd cycles. Shorter than directive mesenteries, P2, P4 and metamesenteries (M and m). | Longer than directive mesenteries, betamesenteries (B and b) and metamesenteries (m). Shorter than P2 and metamesenteries (M). | Longer than directive mesenteries and betamesenteries (B and b). Shorter than P2 and metamesenteries (M and m). |
P4 | Longer than directive mesenteries, P3, P5, betamesenteries (B and b) and metamesenteries (m), except for m of the 2nd cycle. Shorter P2 and metamesenteries (M). | Absent | Absent |
P5 | Longer than betamesenteries (b) and betamesenteries (B), except for B from 1st to 4th cycles. Shorter than directive mesenteries, P2, P3, P4 and metamesenteries (M and m). | Absent | Absent |
Proportion of directive mesenteries in the gastrovascular cavity | 36.6% | 30.5% | 2.3%** |
Proportion of protomesenteries P2 in the gastrovascular cavity | 88.8% | 92.5% | 85.7% |
Proportion of protomesenteries P3 in the gastrovascular cavity | 12.2% | 56.4% | 14.2% |
Proportion of protomesenteries P4 in the gastrovascular cavity | 38.8% | Absent | Absent |
Proportion of protomesenteries P5 in the gastrovascular cavity | 11.1% | Absent | Absent |
As a result of the disjunct distribution of specimens identified as Ceriantheomorphe brasiliensis (Mexico+US/Brazil+Uruguay) and the incomplete description of Cerianthus ambonensis made by
Cnidome of Ceriantheomorphe brasiliensis. A Microbasic b-mastigophore type I B microba-sic b-mastigophore type II C Microbasic b-mastigophore type III D microbasic b-mastigophore type IV F microbasic b-mastigophore type VI G microbasic b-mastigophore type V H ptychocyst type I I atrichous type I J holotrich K ptychocyst type II. Scale bars: 15 µm.
Based on morphological characters and biogeographic perspectives (Table
Cnidome of Ceriantheomorphe ambonensis. A Microbasic b-mastigophore type I B microbasic b-mastigophore type II C microbasic b-mastigophore type III D microbasic b-mastigophore type IV E microbasic b-mastigophore type V F microbasic b-mastigophore type VI G atrichous H holotrich I ptychocyst. Scale bars: 15 µm.
Cnidome of Ceriantheomorphe adelita sp. nov. holotype. A Atrichous B ptychocyst type I C holotrich D ptychocyst type II E microbasic b-mastigophore type I F microbasic b-mastigophore type II G microbasic b-mastigophore type III H microbasic b-mastigophore type IV I microbasic b-mastigophore type V. Scale bars: 15 µm.
Some authors have discussed the taxonomic value of mesenteriel organization regarding the assignment and identification of species (
Our results also demonstrated that the use of ratios (division of the values) between metamesenteries (M × m) and betamesenteries (B × b) for each quartet can be useful to distinguish species of Ceriantheomorphe. In specimens of C. brasiliensis, the ratio between metamesenteries (M × m) and betamesenteries (B × b) ranged from 1.1 to 3.1% and from 1.2 to 3.1%, respectively. In comparison, the ratios observed in C. adelita sp. nov. are from 2.7 to 5.2% between metamesenteries and 3% between betamesenteries, while in C. ambonensis they range from 2.2 to 3.5% between metamesenteries and 4% between betamesenteries.
Similar to the ratio between metamesenteries (M × m) and betamesenteries (B × b), the proportion of protomesenteries found in the gastrovascular cavity was also useful to distinguish Ceriantheomorphe species in our study. While protomesenteries (P3) in C. adelita sp. nov. extend over half of the entire gastrovascular cavity length (56.4%), the ones in C. brasiliensis and C. ambonensis are much shorter (12.2% and 14.2%, respectively). Furthermore, we found differences between species while comparing protomesenteries length (Table
Characters observed |
|
This study |
---|---|---|
Specimen size | 8.5 cm | 3.8 cm |
Number of marginal tentacles | About 150 | 24 |
Number of labial tentacles | About 150 | 36 |
Arrangement of both tentacles | 3 cycles | 3 cycles |
Pharynx region | About 2.5 cm | 0.7 cm long and 2.0 cm wide |
Hyposulcus and hemisulci | No information | Absent |
Gastrovascular cavity | Noinformation | 2.1 cm long and 2.0 cm wide |
Siphonoglyph | No information | 0.7 cm long and 0.3 cm wide / 3 pairs of mesenteries connected. |
Mesenteries | Numerous | 96 |
Arrangement of mesenteries | No information | M,B,m,b |
Cnidome | No information | Spyrocists, microbasic b-mastigophores (six types), atrichous (one type), ptychocyst and holotrichous. |
Currently, the genus Ceriantheomorphe has a wide geographic distribution; one species is restricted to the warm temperate northwest Atlantic (Gulf of Mexico and United States of America), another to the warm temperate southwestern Atlantic (southeast and South of Brazil and Uruguay) and C. ambonensis is recorded from tropical Central Indo-Pacific, Sunda Shelf (Indonesia) (
Disjunctive distribution patterns are exhibited by some marine invertebrates, even those having a free-swimming phase that would benefit wide dispersal, for instance, the bivalve Macoma balthica Linnaeus, 1758 (
This work was partly funded by FAPESP (grants 2015/24408-4, 2015/21007-9, 2016/04962-0 and 2016/50389-0), CNPq (grants 304961/2016-7) and Acordo CAPES/CNPq – PROTAX II (grants 88882.156878/2016-01). We would like to thank Dr. Priscila A. Grohmann from IB-