Check List 9(3): 631–639, 2013 © 2013 Check List and Authors Chec List ISSN 1809-127X (available at www.checklist.org.br) Journal of species lists and distribution n

1* 2

istributio New distribution records of Dinophyta in Brazilian waters

D Paulo Francisco Granja Jardim and Luciana de Souza Cardoso

1 Universidade Federal do Paraná, Setor de Ciências Biológicas, Departamento de Botânica, Jardim das Américas. CEP 81531-990. Curitiba, PR,

raphic Brasil. g 2 Universidade Federal do Rio [email protected] do Sul, Instituto de Biociências, Av. Bento Gonçalves 9500, Prédio 43433. CEP 91501-970. Porto Alegre, RS, eo Brasil.

G * Corresponding author. Email: n o

Abstract:

otes New records of marine are reported for the municipality of Tramandaí (29°59’05” S; 50°08’01”

N W), including new records, not only for the state of Rio Grande do Sul, but also for Brazil. These records were based on sampling carried out in the region in 1976, by Kremer and Rosa (1983), as well as new samples taken in 2011, in the same location as the previous work. In total, we present 20 new records, 13 of them for Rio Grande do Sul and 7 for Brazil as a whole.

et al. Floristic studies of dinoflagellates in Brazil are relatively Water (STSW), which has temperatures higher than 14°C few, but amonget al. the several important contributions are and salinities between 33.5 and 36 (Moller 2008). those of Passavante (1979) for the state of Pernambuco; The discharge of the La Plata River and Patos Lagoon water Passavante (1982) for Ceará, Kremer and Rosa (1983) contributes large amounts of dissolved nutrients, as well for the region of Tramandaí-Imbé in Rio Grande do Sul, as lowering the salinity along the southern coast of Brazil, Cardoso (1995, 1997, 1998) for northern NeoceratiumSanta Catarina in primarily depending on the season and El Niño events. El Island and Arvoredo Island in Santa Catarina, Koening Niño events cause a high discharge of fresh wateret al. in the and Lira (2005) with species of the genus region due to high rainfall, generating a broad zone of Pernambuco, Haraguchi and Odebrecht (2010) and Islabão low-salinity and nutrient-rich water (Moller 2008; and Odebrecht (2011) for the area between Albardão- Seeliger and Odebrecht 2010).et al The water temperature and Chuí, Rio Grande do Sul, and the Cabo da Santa Marta in salinity were used to classify the kind of water during this Santa Catarina. The small number of studies of Dinophyta study, according to Moller . (2008). in Brazilian waters increases the chances of finding new The study area is located on the northern coast of RS occurrences for both states and the country, especially in the municipality of Tramandaí (29°59’05” S, 50°08’01” because the world biodiversity is estimatedet at al. 1555 species W), where two sampling points were selected: the beachm (Gómez 2005) in the marine environment. According area and near the Petrobras monobuoy. Aliquots from to the Brazilian Flora List (Odebrecht 2012), 352 qualitative samples (collected with a plankton net, 48 μ Dinophyta taxa have been recorded for Brazilian marine mesh) taken in the region in 1976 (Kremer and Rosa 1983) waters, 170 of them occurring in Rio Grande do Sul (RS). were inspected. These were donated by the herbarium This paper provides comments regarding the (Registration Number in Alarich Schultz’s herbarium - HAS and distribution of 19 taxa newly recorded in RS and / or 2419, 2421, 2430, 2436, 2443, 2442, 2478, 2484, 2490, Brazil, seeking to establish distribution patterns related to 2496, 2500 and 2504) of the Natural Sciences Museum the seasonality of different water bodies off this coast. (MCN) of the Zoobotanical Foundation of Rio Grande do The region is influenced by the convergence of the Sul (RS-FZB). New sampling in these same locations was Tropical Water (TW,et T> al. 20°C, S> 36) (Emilsson 1961) and conducted in summer (January, February and March) and the Subantarctic Shelf Water (SASW; T 4-15°C, S 33.70 winter (July, August and September) of 2011. Qualitative to 34.15; Sverdrup 1942; Thomsen 1962), and the (plankton net, 64 μm mesh) and quantitative (Van mixture of these water masses with the Subtropical Water, Dorn bottle) sampling followed established protocols also called the South Atlantic Central Water (SACW). The for these analyses (Sournia 1978). Some physical and South Atlantic Central Water (SACW) is a water mass chemical variables were measured simultaneously rich in nutrients flowing into the pycnocline region, etwith al. during the sampling: water temperature (with a mercury temperatures between 6 and 20° C and salinity between thermometer), salinity (measured later in the laboratory 34.6 and 36 (Braga and Niencheski 2006; Silveira in a hydrometer) and water transparency (Secchi disk). 2000). In addition to these water masses, the region Optical microscopy (Olympus IX30 and Zeiss West) receives a considerable influence of continental discharges and the sodium hypochlorite technique for observation of from the La Plata River and Patos Lagoon. The influence of thecal plates (Balech 1971) were used for the identification the Plata Plume Water (PPW; T >10°C, S <33.5) significantly of the species,et al. using basic references (Schiller 1933, 1937; affects an area extending from the Mar del Plata in Sournia 1978, 1984, Balech 1964b, 1971,Preperidinium 1978, 1988; Argentina to the coast of Santa Catarina, with seasonally Balech Protoperidium1984), and in some cases, specific references variable flow. The Plata Plume Water (PPW) mixed with for each taxon. Some species of the genus Tropical Water (TW) forms another water mass with Mangin and Bergh were identified through potential importance in the region: the Subtropical Shelf an epifluorescence microscope (Zeiss Axion Vert 135)631 Jardim and Cardoso | New records of Dinophyta in Brazil

using a fluocromo Calco fluor white specific for cellulose to this genus, requiring revisionsNeoceratium and nomenclatural hexacanthum with the aim of highlighting the plates cellulosic (Fritz and discussions about this taxon. Triemer 1985). Zeiss Camera and AxioCam ErC5s software When considered a variety of was utilized to the capture of the images. For quantitative (Gourret) F. Gómez, D. MoreiraN. and hexacanthum P. López-Garcia, which analysis, an inverted microscope (Zeiss Telaval 31) was may cause confusion in determining its geographical employed, using the Utermöhl (1958) method, where distribution. As a variety of it has been a fixed volume of 25 mL was counted completely.et al. The recorded only in France (Mediterranean Sea) (Balech 1988)N. new occurrences for RS and/or Brazil were verified by hexacanthumleading to the conclusion that its distribution is restricted consulting the Brazilian Flora List (Odebrecht 2012). and rare. Based on its morphological characteristics, C. The distribution of species as well as comments aestuarium has a regularly reticulate body (Kremer and Protoperidinium.regarding the ecology and taxonomy are included for each Rosa 1983) and a left N.horn hexacanthum twisted antapically, unlike taxon, in addition to tabulation (1’ and 2a) for the genus . The latter may be a morphotype adapted to At the end of the comments of each colder waters, unlike , which has a wide species there is a literature where illustrations for each distribution (in all regions) along the Brazilian coast,C. taxon are found. The taxa validation and the geographical aestuarium.and is most often associated with the Brazil Current distribution on a global scale wereet confirmed al. by consulting (Balech 1988). According to Fig. 1, it was evident that is the Algae Base (Guiry and Guiry 2012) and World Register Cladopyxis brachiolata Stein of Marine Species (Appeltans 2012). Photographs of some species were taken using a microscope (Leica DM750) with attached camera (DFC290HD) and the This species was found only in the winter of 2011 software Leica Application Suite Version 3.4.1. The species (Table 1), at a temperature of 13°C and salinity 34.63; illustrated in thePreperidinium photographs wereand Protoperidinium the most frequent and/ associated with SACW. Absent in the quantitative samples. or for which a satisfactoryProtoperidinium view of the plates was obtained Size: 50 µm (long); 44 µm (wide) (1 specimen). (for the genera ). Distribution: In Brazil, recorded in the Northeast (BA) The genus was prior identified by and Southeast (ES, RJ) regions. First occurrence for RS. It combination of morphology of the plates, one located in occurs in the Tropical Atlantic, Mediterranean Sea, Indian the ventral (1’) and a dorsal region (2a). The first may be Ocean, western Pacific and a coral reef in Australia (Wood Ortho, Meta or Para depending on if it has contact with four, 1963). five or six plates, respectively. The second may be Quadra, Comments: Balech (1964b) found this species in a Penta or Hexa following the same concept mentioned higher temperature (24°C) than in the present study, and above. Besides these standard tabular characteristics in similar salinities. Balech found mainly variations in such as size, shapeet al and the cingulate displacement were the size of the horns. The single specimen found in this important for accurate identification for the genus study accorded with the morphology described by Balech (Hoppenrath . 2009). (1964b), and differed from the report of Balech (1964a), Of the 20 new occurrences found in this study, 13 were where the horns were longer. Notably, the study of Balech Protoperidiniumnew for the state of Rio Grande do Sul, Gonyaulaxand 7 for Brazil (Table (1964b) was conducted in Argentinean waters, near the Neoceratium1). The largest number of new records was for members of site of the present study, while Balech (1964a) reported (55%), followed by (15%) and species from Pacific waters. aestuarium (10%). Schroder The Details of the morphology can be viewed in Balech (1988, plate diegensis 71, figs. 1-3). Kofoid (Figure 1A) This species was recorded-1 only in winter 2011 (Table 1), with a density of 20 ind.L . Occurred at low temperature This species occurred-1 only in quantitative samples in (13-14°C), and salinities from 29.29 (beach point), and August 2011 (20 ind.L ) (Table 1), at a temperature of 34.63 (monobuoy point), conditions associated with PPW 15°C and salinity 31.36; associated with PPW. and SACW, respectively. Size: 72 µm (long); 35 µm (wide) (1 specimen). Size: 200 – 250 µm (long); 70 – 80 µm (wide); 30 µm Distribution: In Brazil, recorded in the North (AP, (length of anterior body); 42 µm (length of posterior body); AP) and Northeast (RN) regions. First occurrence for RS. 160 – 210 µm (length of apical horn); length of horns 20 Species distributed from tropical waters (Gulf of Mexico) µm (right) and 25 µm et(left) al (2 specimens). to temperate waters (UK, North Sea) (Guiry and Guiry Distribution: First occurrence in Brazil. This species 2012). was found by Balech . (1984) in Argentinean waters, The details of the morphology and the tabulation can mentioned as uncommon and rarely recorded around the Gonyaulaxbe viewed in fragilis Balech (Schütt) (1988, plate Kofoid 74, fig. 10) world, only for the Mediterranean Sea and the Adriatic Sea Neoceratium(Balech 1988). et al Comments: This species belongs to the genus This species was only observed in summer 1976 (Table created by Gómez . (2009), due to its 1), at a temperature of 23°C and salinity 27.2; associated Neoceratiumessential characteristics for this taxon and living in the with PPW. marine environment. However, it wasn’t Neoceratium transferred to Size: 44µm (long); 35µm (wide) (1 specimen). et al, which leads to confusion in identifying needs. Distribution: In Brazil the species is recorded only in Based upon the characteristics defined in to the North (AP) and the South (SC). First occurrence for Gómez . (2009), this species should be transferred RS. It has a worldwide distribution from the tropics 632 to Jardim and Cardoso | New records of Dinophyta in Brazil

temperate zones (Guiry and Guiry 2012). 29.29 to 36.35; associated with PPW, SASW and SACW. Comments: According to Cardoso (1998), this species Size: 120 – 155 µm (long); 50 – 60 µm (wide) (4 prefers warm water. Cardoso (1998) foundet it al. at similar specimens). Neoceratium dens temperatures to the present study (21 to 27°C), but at Distribution: First occurrence for RS and Brazil. This higher salinities (32.91 to 34.73). Balech et al. (1984) species was referred to as (Ostenfeldet recorded this species in Argentinean waters, butGonyaulax did not al.and Schmidt) F. Gómez, D. Moreira and P. López-Garcia, hyalinamention the temperature and salinity. Balech (1984) asand N. was dens distinguished from thisN. by balechii Meave del Castillo speculated about the similarity of this species to (2003). Therefore, many previous records of taxa such Ostenfeld and Schmidt. The single specimen found in Brazil likely refer to . Its occurrence in this study had the dimensions within those recorded by in Brazil should be reviewed, because of the taxonomic Cardoso (1998), which suggests that this morphotype is change. Widely distributedet al. in the Mexican Pacific, also adapted to the southwest Atlantic. with records on the coasts of Ecuador, Peru and Chile The details of the morphology and the tabulation can (Meave del Castillo 2003). Gonyaulaxbe viewed in Cardoso spinifera (1998). (Claparède and Lachmann) Comments: TheNeoceratium specimens dens found in samples from Diesing winter 2011et al. were quite similarN. balechii to those recorded by Balech (1988)N. dens as . Recently, Meave Del Castillo (2003) described , differentiating This species occurred only in winter 1976 (Table 1), at it from . The former has the two antapical horns a temperature of 16°C and salinity from 27.1; associated small, with the left horn slightly more robust, whereaset the al with PPW. latter has the left anti-apical horn longer, reaching almost Size: 49 µm (long); 37 µm (wide) (1 specimen). to midlength of the apical horn (Meave del Castillo . Distribution: In Brazil it is found in all regions: North Neoceratium2003). setaceum (Jørgensen) F. Gómez, D. (PA), Northeast (RN), Southeast (RJ) and South (SC). Moreira and P. López-Garcia First occurrence for RS. According to Esqueda-Lara and Hernández-Becerril (2010) it is a tropical and subtropical species. This species occurred only in summer 1976 (Table Comments: It has also been found in Antarctica (Balech 1), in temperatures from 23 to 25.5°C and salinity 27.2; 1976), but is not considered an inhabitant of these waters, associated with PPW. where it was noted as invasive. Prefers cold waters, Size: 180 µm (long); 50 µm (wide) (1 specimens). occurring in low temperatures (2.15 to 15.55°C) and Distribution: In Brazil, its distribution extends to all salinities between 33 and 35 (Balech 1988). regions of the coastal zone: Northern (AP, AP), Northeast The details of the morphology and the tabulation can (RN, BA), Southeast (SE, SP, RJ) and South (PR, SC). First Lingulodiniumbe viewed in Balech polyedrum (1988, plate (F.Stein) 74, figs. J.D.Dodge 1-4) occurrence for RS. The world distribution includesNeoceratium both pentagonumtropical and temperate regions (Guiry and Guiry 2012). (Figure Comments: Species very similar to 1B-C) (Gourret) F. Gomez, D.Moreiraet al and P.Lopez- Occurred only in summer 1976 (Table 1), at a Garcia, differs from it by owning a typically longer apical temperature of 23°C and salinity 27.2; associated with horn of up to 408 et µm al (Gonçalves . 2006), a lower PPW. diameter cell and antapical left horn of greater length (20 Size: 40-55 µm (long); 35-45 µm (wide) (2 specimens). - 50 µm - Gonçalves . 2006). Distribution: In Brazil, recorded in North (AP), This species seems to be very uncommon in the Northeast (MA, RN, PE) and South (SC). First occurrence area, because Balech (1988), over decades of study, did for RS. Elsewhere, it is recorded in tropical and temperate not find it in the southwest Atlantic. According to Wood waters (Guiry and Guiry 2012). (1968) it is a tropical species, but closer inspection of the Comments: Species easily identifiable by its records indicates that it occurs from tropical to temperate characteristic polyhedral shape in lateral view. In apical environments. It seems to be more associated with view has almost circular outline (Cardoso 1998). the Brazil Current, because it frequently occurs off the Balech (1988) listed this species from the Brazil northern coast of theet country al. (Wood 1968). Current, always occurring rarely in the southwest Atlantic The details of the morphology of this species can be (between 38°30’S and 43°33’W) and in warm waters. Preperidiniumviewed in Gonçalves meunieri (2006).(Pavillard) Elbrächter Kofoid (1911) cited this species as very abundant in the summer plankton in Baja California, reaching high (Figure densities in the summer peak, and that intra-specific 1E-G) variation occurred mainly in the size and development of Occurred only in both winters (Table 1), at temperatures Neoceratiumthe reticulum. balechii var. longum (Meave, Okolodkov from 13 to 16.5°C and salinity from 19 to 36.35. Present-1 and Zamudio) F. Gómez, D. Moreira and P. López-Garcia in quantitative samples with low density (40 ind.L ); associated with SACW, SASW and PPW. Size: 30 – 40 µm (long); 45 – 60 µm (wide) (3 (Figure 1D) specimens). This species occurred widely-1 in winter 2011 (Table 1), Distribution: In Brazil, it has only been found in with a peak density of 220 ind.L in August. It is recorded the North Region (PA). First occurrence for RS. It has a in temperatures from 13 to 16.5°C and salinities from cosmopolitan distribution in the world (Balech 1976). 633 Jardim and Cardoso | New records of Dinophyta in Brazil

Protoperidinium Comments: Balech (1988) mentioned it as a highly divergenssummer. Diplopeltopsisthermotolerant minor because it has records This species can be confused with Meta from -2 to 24°C. This speciesPreperidinium has a heterotypic synonym: Quadra (Ehrenberg) Balech because theyP. divergens have similar more (Paulsen) Pavillard, and was shapes, and theP. brochii,tabulation of plates is identical (1’ ; transferred to the genus in 1993 by 2a ). Balech (1988) considered Elbrächter (Guiry and Guiry 2012). angular than which we also observed, and the Tiny pores were observed (Figure 1G)P. on meunieri the entire latter has a strongly sinuous contour (Cardoso 1997), cell surface as reported by Balech (1988). According to different from the former, which also has a more prominent this author, the cingular membrane of has apical horn. Protoperidiniumdenticulate edge, detail anomaloplaxum observed in specimens (Balech) found. Balech The details of the morphology and the tabulation can Protoperidiniumbe viewed in Balech conicoides (1988, plate (Paulsen) 41, figs. 4-7) Balech

(Figure 1H-I) (Figure Species found only in 1976 (Table 1), in temperatures 2A-B) from 14.5 to 23°C and salinities from 21.5 to 27.8; This species occurred only in winter in both years associated with PPW. (Table 1), in temperatures from 14.5 to 16.5°C and salinity Size: 45 – 50 µm (long); 47 – 52 µm(wide) (2 specimens). from 19 to 36.35; associated with SACW and PPW. Distribution: First occurrence for RS and Brazil. The Size: 40 – 50 µm (long); 35 – 42 µm (wide)Ortho (3 only previous record was by Balech (1988) in Argentinean specimens). Hexa waters between latitudes 37°39’S and 38°42’S, whichMeta is Tabulation: Ventral tabulation of epitheca type adjacent to this studyPenta region (30°S). and dorsal type Tabulation: Ventral tabulation of epitheca type Distribution: First occurrence in Brazil. It occurs mainly Protoperidiniumand dorsal type sphaeirodeum. in temperate regions (Guiry and Guiry 2012). Comments: The material was first identified as Comments: This species is usually found at lower P. anomaloplaxum (P. Dangeard) Balech, but temperatures than in the present study (5-11°C) (Balech afterfrom thoroughP. sphaeirodeum analysis of other specimens, it was apparent 1988). Occurs more frequently in temperate and polar that the 2a plate in (penta) is distinct regions (Appeltans et al. 2012;Protoperidinium Guiry and conicum Guiry, 2012) (quadra). Furthermore, plate 1’ has which confirms it as a cold-water species. Balech (1988) a different morphology, with the sinuous edge with the 6’’ compared this species with P. conicoides is moreGran (Balech 1988). Balech (1988) cited it as a species with a (Balech), however the latter species is more elongated very particular distribution, as it was always seen in the anteroposteriorly, and the edge of same latitudes. Its distribution does appearProtoperidinium to be restricted convex. lipopodiumto the southwest Atlantic, because it has not been found The details of the morphology and the tabulation can elsewhere. This species, together with Protoperidiniumbe viewed in Balech elegans (1988, (Cleve) plate 26, Balech figs. 7-11). (Balech) Balech, have no records elsewhere, suggesting that they could be endemics of the southwest Atlantic. Only one individual of this species was observed, in a The details of the morphology and the tabulation can sample from 1976 (Table 1), at a temperature of 23°C and Protoperidiniumbe viewed in Balech brochii (1988, (Kofoidplate 22, figs. and 11-13) Swezy) Balech salinity of 21.5; associated with PPW. Meta Size: 165 µmQuadra (long); 100 µm (wide) (1 specimen). Tabulation: Ventral tabulation of epitheca type (Figure 1J-L) and dorsal typeP. elegans . This species occurred in both years (Table 1), in Distribution: First occurrence for RS and Brazil. temperatures from 14.5 to 23°C and salinity from-1 19 to Elsewhere, occurs in tropical Protoperidinium and temperate 31.36. Present in quantitative samples (20 ind.L ) only in granderegions (Guiry and Guiry 2012). August 2011. It is associated with PPW. Meta Comments: Species very similar to Size: 70-92 µmQuadra. (long); 75 µm (wide) (2 specimens). (Kofoid) Balech differingP. grande by having a greater width Tabulation: Ventral tabulation of epitheca type between the antapical horns – and larger hypotheca P. elegans height and dorsal type (Balech 1988). Furthermore, presents minimum Distribution: In Brazil it occurs in the Southeast (SP) values​ ​for larger diameter (112 µm) greater than and the South (SC). First occurrence for RS. Outside (85 µm), checking this variable in Balech (1988). Brazil, the species is widely distributed from the tropics Species widely variable in size (Balech 1988). to temperate zones (Guiry and Guiry 2012). According According to Balech (1988), it is a thermophilic species to Esqueda-Lara and Hernández-Becerril (2010) it is a and appears to be intolerant, and is rarely found in the subtropical and temperate species. southwest Atlantic. Comments: Balech (1988) mentioned that this species The details of the morphology and the tabulation can almost always occurred at temperatures above 13°C; in Protoperidiniumbe viewed in. Balech globulus (1988, (Stein)plate 43, Balech figs. 1-4). that study, it was found in temperatures from 9 to 20°C and salinities from 33 to 36. Cardoso (1997) recorded this (Figure 2C-D) species from southern Brazil in ranges of temperature in This species occurred in both years, more frequently in (21-27°C) and salinity (32-35) slightly higher than in-1 the summer-1 (Table 1), however with a maximum density (300 present study, with a maximum density of 100 ind.L ind.L ) in winter. The limits of temperatures (13-25.5°C)634 Jardim and Cardoso | New records of Dinophyta in Brazil

and salinity (21.5 to 34.63) were wide; associated with Argentinean waters, which probably confirms the opinion SASW, TW, SACW and PPW. Meta of Cardoso (1997), making the species an indicator for the Size: 30 µm (long);Hexa 25-30 µm (wide) (4 specimens) region. Tabulation: Ventral tabulation of epitheca type The minimum wide was below than reported by Balech and dorsal type . (1988). The other measures were within the limits found Distribution: In Brazil, it is distributed in the North by this author. (PA), Northeast (MA) and South (SC). First occurrence for The details of the morphology and the tabulation can RS. It is widely distributed in warm waters (Wood 1964). Protoperidiniumbe viewed in Balech pacificum (1988, plate (Kofoid 31, figs. and 1-4). Michener) F. J. Comments: Species easily recognized by its globular R. Taylor and Balech ex Balech shape and small dimensions. It was observed difficulties in the visualization of the plates even with Fritz and (Figure 2G) Triemer’s techniques (Fritz and Trimer 1985). ) in This species occurred only in summer 1976 (Table 1), Seems to be a tolerant species, because in southern-1 in temperatures from 23 to 25.5°C and salinities from 27.2 Brazil it occurred in high densities (100 cells. L to 33; associated with PPW. summer (Cardoso 1997), and in high abundance in winter Size: 50 – 65 µm (long); 55 – 60 µm (wide)Meta (2 in this study. specimens). Quadra The details of the morphology and the tabulation can Tabulation: Ventral tabulation of epitheca type Protoperidiniumbe viewed in Cardoso granii (1997). (Ostenfield) Balech and dorsal type . Distribution: First occurrence for RS and Brazil. Elsewhere, it is known from tropical and temperate This species occurred in both years analyzed (Table regions (Balech 1988; Okolodkov 2008). Balech (1988) 1), over a range of temperatures from 13 to 25.5°C and found specimens of this species always near the Malvinas salinities 29.29 to 34.63; associated with SASW, SACW and Current, with few records above 14.5 °C. PPW. Meta Comments: The wide was larger than found by Balech Size: 58 µm (long);Penta 52 µm (wide) (1 specimen). (1988). Tabulation: Ventral tabulation of epitheca type The details of the morphology and the tabulation can and dorsal type . Protoperidiniumbe viewed in Balech rampii (1988, (Balech) plate 40, Balech figs. 13-18). Distribution: In Brazil, it has been recorded in the Northeast (PE, BA), Southeast (ES, RJ) and South (SC). First occurrence for RS. A widely distributed species, from This species was recorded only in summer 1976 (Table tropical to polar regions (Guiry and Guiry 2012). 1), at a temperature of 23°C and salinity of 21.5; associated Comments: Balech (1988) mentioned a possible with PPW. Para Protoperidiniumpreference for cold-temperate mite waters, based on its Size: 55 µm (long);Quadra. 50 µm (wide) (1 specimen). global distribution. Balech cited its resemblance to Tabulation: Ventral tabulation of epitheca type (Pavillard) Balech, which differs and dorsal type especially in having less-developed horns, non-excavated Distribution: First occurrence for RS and Brazil. No cingulum and the equatorial region of the cell is less world records were found in the literature, except by rounded (Balech 1988). BalechProtoperidinium (1988) for Argentinean pacificum waters. The details of the morphology and the tabulation can Comments: BalechMeta (1988) in commentedP. pacificum on and its Parasimilarity in P. Protoperidiniumbe viewed in Balech lipopodium (1988, plate (Balech) 40, figs. 9-12). Balech torampii , differentiating this with respect to plate 1’ ( (Figure ) andP. pacificumthe size of the horns. These distinctive features 2E-F) were observed in the specimensP. analyzed.rampii The antapical This species was recorded only in 1976 (Table 1), in horns of are slightly longer, and have a less temperatures from 16 to 25°C and salinities from 27.2 to sinuous contour compared with . 32.1; associated with PPW. According to Balech (1988), it is widely distributed in Size: 45 – 50 µm (long); 43 – 52 µm (wide)Meta (2 the coldP. anomaloplaxum waters of Argentina, and P. lipopodium,rarely above 14°C, but was specimens). Quadra. recorded in this study at a higher temperature. Together Tabulation: Ventral tabulation of epitheca type with no other record and dorsal type exists in the literature, except in the southwest Atlantic, Distribution: In Brazil, it has been recorded only in the making it a likely indicator of the region. South region, in SC by Cardoso (1997). First occurrence for The details of the morphology and the tabulation can RS. It has not been recorded elsewhere (Guiry and Guiry Protoperidiniumbe viewed in Balech steinii (1988, (Jorgensen) plate 47, figs. Balech 5-8). 2012), except by Balech (1988). Comments: Species with characteristic shape (piriformis) with the presence of two antapical spines This species was found in both years analyzed, more curved to the right, with the left larger than the right often in summer-1 (Table 1), but with maximum density in (Cardoso 1997). winter (40 ind.L ). It was recorded in temperatures from Cardoso (1997) suggested that this species is restricted 15 to 25°C and salinities from 27.1 to 32.1; associated with to the southwest Atlantic, because the only record prior TW and PPW. to her study was by Balech (1988). Currently, there is Size: 60 – 70 µm (long); 30 – 45 µm (wide) ( 3 no record of this species other than southern Brazil and specimens). 635 Jardim and Cardoso | New records of Dinophyta in Brazil

Meta P. ovum Penta Tabulation: Ventral tabulation of epitheca type were details that distinguished the two species. P. steinii, and dorsal type . is more ovoid, and the epitheca and the hypotheca are Distribution: In Brazil, it has been recorded in all the same size (Balech 1988), which differs from P. steinii regions: Northern (AP, AP), Northeast (BA), Southeast whichMeta hasand aPenta higher) and epitheca P. ovum than Para hypotheca and Hexa (Cardoso) are (SE, SP, RJ) and South (SC). First occurrence for RS. It is 1997). Furthermore, the 1’ and 2a plates of distributed in tropical and temperate regions (Guiry and ( ( Guiry Protoperidinium2012). ovum different. Comments: At first sight, the species was confused The details of the morphology and the tabulation can with (Schiller) Balech, but there be viewed in Cardoso (1997). A B C

D E F

G H I

J K L

Figure 1. Ceratium aestuarium Lingulodinium polyedrum Neoceratium balechii longum Preperidinium meunieri P. meunieri P. meunieri Protoperidinium anomaloplaxum P. anomaloplaxum P.A) brochii P. ;brochii B-C) ; D) var. ; E) , overview; F) , apical view; G) , apical view (optical microscope); H) , overview; I) , view of the plate 1’; J) , overview. K) , view of the plate 1’; L) , view of the plate 2a. 636 Jardim and Cardoso | New records of Dinophyta in Brazil

Protoperidinium symmetricum (Halim) Balech Protoperidinium abei (Figure between 20 and 26°C, associated with the Brazil Current. 2H-J) It is very similar to P. abei (Paulsen) This species was observed only in summer 2011 Balech,P. symmetricumhowever can be easily distinguished mainly by the (Table 1) in temperatures from 23 to 23.5°C and salinities morphology (Balech 1988). is longer and narrower from 29.61 to 33.88; associated with TW. Absent in the than ., and in the latter the antapical spine quantitative samples. Ortho is easily viewed. Size: 75 µm (long);Hexa 76 µm (wide) (1 specimen). It was difficult in identifying the species found in Tabulation: Ventral tabulation of epitheca type 1976. Many cells were damaged due to prolonged action and dorsal type . of formaldehyde along of 35 years, and the calcofluor Distribution: The only previous record was from analyses were impossible to run because that. Besides that, Southeast Brazil (RJ). First occurrence for RS. some was rare and it difficult to obtain photographs of Comments: According to Balech (1988), this is a them. Therefore, we intend to confirm some new tropical species found in Brazilian waters in temperatures occurrences in future work, with records made by A B C

D E F

G H I

J

Figure 2. P. conicoides P. conicoides P. globulus P. globulus P. lipopodium P. pacificum P. symmetricum, A)P. symmetricum, overview; B) P. symmetricum, view of the plate 1’; C) , overview; D) , view of the plate 2a; E-F) ; G) , overview; H) overview. I) , view of the plate 1’; J) , view of the plate 2a.. 637 Jardim and Cardoso | New records of Dinophyta in Brazil

photographs. Protoperidinium of dinoflagellates, other microscopical techniques should Most species found here for the first time in Rio Grande be used, such as scanning electron and fluorescence do Sul and /or Brazil belong to the genus . microscopy, because of the great difficulty of identifying This disproportion is associated with the greater dinoflagellates with optical microscopy alone. difficulty in identifying Gonyaulaxspecies from this genus, which are This relatively large number of first occurrences of distinguished by the epithecal plates, added to a lack of species illustrates the need for more taxonomic and studies on this genus. was second in number ecological research on dinoflagellates in Brazilian waters, of first records, for the same reason, requiring analysis in order to better understand an important group that is of the cellulosic plates. For more comprehensive studies being neglected in Brazil for lack of specialists. Table 1.

New records for Rio GrandeJan/76 do Sul andFeb/76 Brazil (*).Mar/76 (B= beach,Jul/76 M = Aug/76monobuoy)Sep/76 Jan/11 Feb/11 Mar/11 Jul/11 Aug/11 Sep/11 Ceratium aestuarium*

Cladopyxis brachiolata B,M Gonyaulax diegensis B Gonyaulax fragilis M Gonyaulax spinifera B Lingulodinium polyedrum B Neoceratium balechii var. longum* B Neoceratium setaceum B,M M B,M Preperidinium meunieri B Protoperidinium anomaloplaxum* B,M B,M B B,M Protoperidinium brochii B M B,M Protoperidinium conicoides* B B B,M B,M Protoperidinium elegans* B,M M M Protoperidinium globulus B Protoperidinium granii B M B M M M B Protoperidinium lipopodium M B, M Protoperidinium pacificum* B B M Protoperidinium rampii* M B Protoperidinium steinii B Protoperidinium symmetricum B B B B,M B B M M B,M

Acknowledgments: e do Talude da Região Sudeste-Sul do Brasil

We thank Petrobras for the cooperation and Ceratium . São Paulo: Editora da support provided during the sampling, M.Sc. Zulanira Meyer Rosa and the Universidade de São Paulo (EDUSP). Zoobotanical Foundation for donating the samples from 1976, and to the Cardoso, L.S.Biociências 1995. O gênero SCHRANK (Dinophyta, Ceratiaceae) chemist Cacinele M. Rocha for providing the materials for sampling from na ilha do Arvoredo e na PraiaProtoperidinium de Ponta das canas, Santa Catarina, the Water, Sediment and Fish Biology Laboratory (Ceclimar /UFRGS). Brasil. 3 (1): 3-41 The MS was reviewed and corrected by PhD. Janet W. Reid from JWR Cardoso, L.S. 1997.Biociências O gênero BERGH (Dinophyta, Associates. Peridiniaceae) na Ilha do Arvoredo e Praia de Ponta das Canas, Santa Catarina, Brasil. 5(1): 3-33. Biociências Appeltans, W., P. Bouchet, G.A. Boxshall, C. De Broyer, N.J. De Broyer, N.J. De Cardoso, L.S. 1998. Dinoflagelados da Ilha do Arvoredo e da Praia de WorldVoogd, RegisterD.P., Gordon, of Marine B.W. Hoeksema,Species T. Horton, M. Kennedy, J. Mees, BoletimPonta das do Canas, Instituto Santa de OceanografiaCatarina, Brasil. 6(1): 3-54. G.C.B. Poore, G. Read, S. Stöhr, T.C Walter and M.J. Costello. (ed). 2012. Emilsson, I. 1961. The shelf and coastal Waters off SouthernDinoflagelados Brazil. Cladopyxis. Electronic Database accessibleRevista del at microplantónicos marinos del Pacífico 11(2): central 101-112. de México (Isla Isabel, Museohttp://www.marinespecies.org. Argentino de Ciencias Captured Naturales on Bernandino20 September Rivadavia 2012. , Esqueda-Lara,Nayarit y costas K. and de Jalisco D.U. Hernández-Becerril.y Colima) 2010. Balech,Hidrobiologia E. 1964a. El género “ ”(Dinoflagellata). . México: Instituto de Ciencias del 1(4): 27-39. Boletim del Instituto de Biologia Mar y Limnologia. 206 p. Balech,Marina E. 1964b. El plancton de Mar del Plata durante el período 1961- Fritz, L. andJournal R.E. of Triemer. Phycology 1985. A rapid simple technique utilizing 1962 (Buenos Aires, Argentina). calcofluor white M2R for the visualization of dinoflagellate thecal 4: 3-60 Revista del Museo Argentino de Ciencias Naturales plates. Acta botanica Croatica 21: 662-664. Balech,Bernandino E. 1971. Rivadavia Microplancton, Hidrobiologia de la campaña oceanográfica Gómez, F. 2005. A list of free-living dinoflagellateNeoceratium species in the world’s productividad III. Publicación oceans. 64(1): 129-212. Ceratium Instituto Antártico Argentin 3(1): 1-204. Gómez, F., D. MoreiraProtist and P. López-García. 2009. gen. nov., Balech, E. 1976. Clave ilustrada de dinoflagelados antárticos. a New Genus for All Marine Species Currently Assigned to Revista delo 11:Museo 1-99. Argentino de Ciencias Naturales, (). 161: 35-54. Balech,Hidrobiologia E. 1978. Microplancton de la campaña oceanografica Gonçalves,Dinoflagelados C.P., C.M.M. e tintinídeosPimenta, D.R. da Tenenbaum, região central E.T. da Gomes, Zona J. Econômica Eduardo, produtividadeLos IV. dinoflagelados del Atlantico Sudoccidental ExclusivaM.C.Q. Mendes, Brasileira M. Menezes, – Guia de M.M.F. identificação Hatherly and S.C. Viana. 2006. 5(7): 137-219. Balech, E. 1988. . Madrid: AlgaeBase . Rio de Janeiro: Museu Ministério de agricultura pesca y alimentación. 310 p. Revista de Nacional. 288 p. Balech,Investigaciones E., R. Akselman, del Desarollo H.R. Benavides, Pesquero R.M. Negri. 1984. Suplemento Guiry, M.D. and G.M. Guiry. 2012. . World-wide electronic a “Los Dinoflagelados del Atlantico Sudoccidental’’. publication, National University of Ireland, Galway. Electronic 4: 5-20 Database accessible at http://www.algaebase.org. Captured on 20 Braga, E.S. and L.F.H. Niencheski.In 2006. Composição das massas de água September 2012. Biota e seus potenciaisO produtivos Ambiente Oceanográfico na área entre da o CaboPlataforma de São Continental Tomé (RJ) Haraguchi,Neotropica L. and C. Odebrecht. 2010. Dinophysiales (Dinophyceae) e o Chuí (RS); p. 161-218 C.L.D.B. Rossi-Wongtschowski and L.S. no extremo Sul do Brasil (inverno de 2005, verão de 2007). Madureira (ed.). 10(3). 638 Jardim and Cardoso | New records of Dinophyta in Brazil

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