<p><strong>Two new species of <em>Aaptos</em> (Demospongiae, Hadromerida) from Brazil (western Atlantic)</strong></p>

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Zootaxa 3750 (4): 357–366 www.mapress.com /zootaxa / Copyright © 2013 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3750.4.4 http://zoobank.org/urn:lsid:zoobank.org:pub:78D7A4C5-69D8-4503-A1A4-83BD5DBF1B5E

Two new species of Aaptos (Demospongiae, Hadromerida) from Brazil (western Atlantic) MARIANA DE S. CARVALHO1,3, SUZANE M. DA SILVA2 & ULISSES PINHEIRO2 1

Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Invertebrados, 20940-040, Rio de Janeiro, Rio de Janeiro State, Brazil. E-mail*: [email protected] 2 Universidade Federal de Pernambuco, Centro de Ciências Biológicas, Departamento de Zoologia, Av. Nelson Chaves, s/n, C. Universitária, Recife, 50373-970, Pernambuco State, Brazil. Email: [email protected] 3 Corresponding author

Abstract Twenty-one species of Aaptos Gray, 1867 are known world-wide, of which only three were reported from Brazil. Two new species of this genus are here described from the Brazilian coast (Potiguar Basin, Northeastern Brazil): A. hajdui sp. nov. and A. potiguarensis sp. nov. Both possess only one category of strongyloxeas and one of styles, although both with wide size variation, suggesting that the diagnosis of the genus should be revised. Previous Brazilian records of A. aaptos have their status re-evaluated here, and only three species of the genus can be considered valid in Brazil: A. glutinans, A. hajdui sp. nov. and A. potiguarensis sp. nov. Key words: Porifera, Suberitidae, Taxonomy, Rio Grande do Norte State, Potiguar Basin

Introduction Aaptos Gray, 1867 is a cosmopolitan genus with 21 species known world-wide, although many other may be hidden among the widespread records of A. aaptos (Schmidt, 1864; van Soest 2002, van Soest et al. 2013). The genus poses special challenges to the Porifera taxonomists due to a shortage of conspicuous anatomical characters. Species may have strongyloxeas in three overlapping size categories frequently hard to tell apart and the intermediate and smaller spicules are occasionally oxeas, styles or tylostyles. The distinction between the species is very difficult as the descriptions currently provide little basis for species definition, because spicule dimensions, general morphology and skeletal details vary little between species in the literature (Kelly-Borges & Bergquist 1994). Only three species of Aaptos are known from Brasil: A. aaptos from Espírito Santo State (Solé-Cava et al. 1981), Santa Catarina State (Mothes & Lerner 1994, as A. aff. aaptos), and Atol das Rocas (Moraes 2011); A. bergmanni de Laubenfels, 1950 from Bahia State (Hechtel 1976) and Pernambuco State (Boury-Esnault 1973 as A. aaptos); and A. glutinans Moraes, 2011 from Atol das Rocas (Moraes 2011). This is the first time that the genus is found at the continental shelf of Rio Grande do Norte State. There are other several records of unidentified Aaptos spp. from Brazil, viz. Bahia, Rio Grande do Norte (Atol das Rocas) Espírito Santo, São Paulo, and Santa Catarina States (see Muricy et al., 2011). In this paper, we describe two new species of Aaptos from Potiguar Basin, Rio Grande do Norte State, Brazil and provide a brief review of bibliographic records of other species of the genus from Brazil.

Material and methods The Potiguar Basin is located on the north continental shelf of Rio Grande do Norte and Ceará States (Northeastern Brazil; Fig. 1). The studied specimens were collected by trawling on board of the R.V. ‘Astro Garoupa’ in two Accepted by J.Hooper: 2 Dec. 2013; published: 20 Dec. 2013

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campaigns (BPOT 2–3) by the Project of Environmental Characterization and Monitoring of Potiguar Basin (Muricy et al. 2008). Samples were fixed and preserved in 70% ethanol. Dissociated spicule slides and thick section mounts were made through usual procedures for Demospongiae (Hajdu et al. 2011). Spicules measurements are presented as minimum–mean–maximum and length/width. They were measured all mixed together and size categories of strongyloxeas lengths were determined with Sturges’ algorithm (Sturges 1926) to define spicule categories. A total of 100 spicules of each spicule type were measured for each specimen. The skeletal and spicular features were observed in light microscope. The holotype and paratypes of A. hajdui. sp. nov. and A. potiguarensis sp. nov. are deposited in the Porifera collection of Universidade Federal de Pernambuco (UFPEPOR) and schyzotypes are deposited in the Porifera collection of Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ).

FIGURE 1. Map showing the sampling sites in Potiguar Basin, Rio Grande do Norte state (Northeastern Brazil).

Systematics Class DEMOSPONGIAE Sollas, 1885 Order HADROMERIDA Topsent, 1894 Family SUBERITIDAE Schmidt, 1870 Genus Aaptos Gray, 1867 Diagnosis. Lobate or spherical sponges with a radial skeleton, often consisting of confluent globular or lobate units. Surface smooth or tuberculate-papillate, usually rough to the touch. Some species show a distinct colour

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change when taken out of the water. Fibrillar collagen is often present in the subectosomal region and may be diagnostic in the degree of density, the depth, and pattern of disposition. Skeleton strictly radiate, with tracts and single spicules issuing from the centre of the lobe or globular body. Primary tracts or swathes of megascleres diverge or fan towards the surface where they intersperse with a dense palisade of smaller spicules. Choanosomal megascleres are strongyloxeas in a large size range that may or may not form distinct size categories, spicules in the dermal palisade usually form a distinct size category and may be oxeas, styles or subtylostyles (emended from Van Soest 2002). Type species. Ancorina adriatica Gray, 1867 (by original designation) = Aaptos aaptos (Schmidt, 1864; sensu Van Soest 2002).

Aaptos hajdui sp. nov. (Figure 2, 3A–B; Table 1) Type material. Holotype: UFPEPOR 62—Campaign BPOT 02, station 04 (Potiguar Basin, Rio Grande do Norte State, 04°37’31.7”S–36°46’0.7”W), 70–101 m depth, coll. R/V ‘Astro Garoupa’, 14/V/2003. Schyzoholotype: MNRJ 16684. Paratypes: UFPEPOR 181—Campaign BPOT 03, station A3 (Potiguar Basin, Rio Grande do Norte State, 04°49´58.0”S–36°12´12.0”W), 50 m depth, coll. R/V ‘Astro Garoupa’, 13/XI/2003. Schyzoparatypes: MNRJ 16688. Diagnosis. Aaptos with an ectosomal skeleton with subectosomal cavities, composed of styles forming a palisade at the surface and bouquets formed by strongyloxeas branching below the surface. Spicules are one category of strongyloxeas with telescopic or mucronate tips (occasionally rounded ends), 485–1475 µm in length and 6–30 µm thick; and one category of styles, 242–582 µm in length and 2.4–9.6 µm thick. Description. Specimens are globular or sub-spherical (up to 9.7 cm in diameter, holotype; Fig. 2A), with some incrustations of calcareous substrate. Oscules rounded, up to 1 mm in diameter, although only few are visible. Surface optically smooth, but microhispid. Consistency firm, almost incompressible. Color of the holotype is unknown and the paratype (UFPEPOR 181) is orange alive, turning dark gray, almost black after fixation in ethanol, with a light interior. Skeleton. Ectosomal skeleton with subectosomal cavities and composed of styles forming a discontinuous palisade at the surface, in which the ends of the strongyloxeas are also present. Choanosomal skeleton with large tracts composed of strongyloxeas with a radial arrangement branching below the surface, forming bouquets (Fig. 2B,C). Collagen is present in a low density mainly within the ectosome throughout. Spicules. Megascleres in two categories (Tab. 1). Choanosomal strongyloxeas (with variations to styles; Figs. 2D,E; 3)—smooth, straight or slightly curved, usually with telescopic or mucronate ends, with a wide size range: 485–1475 µm in length and 6–30 µm thick. Ectosomal styles (Fig. 2F)—smooth, straight, occasionally slightly curved: 242–582 µm in length and 2.4–9.6 µm thick. The distinction between strongyloxeas and styles is very subtle. Here, we consider strongyloxeas as fusiform or slightly fusiform and styles are isodiametric. TABLE 1. Spicule measurements of Aaptos hajdui sp. nov. and Aaptos potiguarensis sp. nov. Smallest length–mean length–largest length / Smallest width–mean width–largest width, in micrometers. Specimens

Strongyloxeas

Styles

UFPEPOR 62—holotype

649–1076.7–1475 / 10–19.7–30

252–342.7–524 / 2.4–3.4–7.2

UFPEPOR 181— paratype

485–896.9–1445 / 6–13.2–20

242–378.8–582 / 2.4–6.0–9.6

UFPEPOR 45—holotype

301–793.7–1562 / 8–20.6–43

220–350–590 / 3.7–7.4–11

UFPEPOR 46—paratype

301–779.1–1484 / 8–23.5–41

300–405–520 / 2.5–5.5–7.5

Aaptos hajdui sp. nov.

Aaptos potiguarensis sp. nov.

Etymology. The species is named after Prof. Dr. Eduardo Hajdu in recognition for his great contribution to our understanding of the systematic of sponges. NEW AAPTOS FROM BRAZIL (WEST ATLANTIC)

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FIGURE 2. Aaptos hajdui sp. nov. A. Holotype (UFPEPOR 62). B. Choanosomal skeleton. C. Detail of the ectosomal skeleton, showing the bouquets near the surface and the ectosomal cavities. D. Strongyloxeas. E. Telescopic and mucronate tips of the strongyloxeas. F. Styles. Scale bars: A = 5 cm. B, C = 500 µm. D = 100 µm. E, F = 50 µm.

Distribution. Provisionally endemic from Potiguar Basin (Rio Grande do Norte State, north-eastern Brazil). Ecology. Bryozoans and polychaetes were found associated to both specimens. This species occurs from 50 to 101 m depth, on rocky substrate. Remarks. Aaptos hajdui sp. nov. approaches A. aaptos very closely. Overall morphology and spicule dimensions are similar. Both species differ though in some traits such as the presence of larger styles in the new species (up to 582 µm in length) and the occurrence of styles to subtylostyles in A. aaptos (sensu Schmidt, 1864; Kelly-Borges & Bergquist, 1994 and van Soest 2002). In addition, Kelly-Borges & Bergquist (1994) have examined the holotype and verified the presence of flexuous styles and subtylostyles, not observed in the new species. Aaptos hajdui sp. nov. is considerably different from the other Brazilian species. Its smaller and thicker strongyloxeas and the presence of only one category of styles differentiate the new species from A. glutinans,

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which has strongyloxeas always larger than 925 µm and two categories of styles (Tab. 2). Among the records of Aaptos from Brazil, only A. aff. aaptos sensu Mothes & Lerner (1994) is close to the new species in spiculation (Tab. 1, 2), except for the fact that it shows variations of styles to subtylostyles and tylostyles, a character not found in A. hajdui sp. nov. Aaptos hajdui sp. nov. is also different from the three species known from the Caribbean. In A. bergmanni and A. duchassaingi strongyloxeas are always smaller than 950 µm (see Table 2). There are no measurements reported of oxeas and styles of A. pernucleata (Carter, 1870), but the presence of oxeas and the black color easily differentiate it from the new species. Other species of the genus (A. alphiensis Samaai & Gibbons, 2005; A. ciliata (Wilson, 1925); A. confertus Kelly-Borges & Bergquist, 1994; A. globosum Kelly-Borges & Bergquist, 1994; A. horrida (Carter, 1886); A. kanuux Lehnert, Hocevar & Stone, 2008; A. laxosuberites (Sollas, 1902); A papillata (Keller, 1880); A. robustus (Plotkin & Janussen, 2008); A. rosacea Kelly-Borges & Bergquist, 1994; A. tentum Kelly-Borges & Bergquist, 1994; and A. vannamei de Laubenfels, 1935) possess other kinds of megascleres in their spicule set, such as oxeas, tylostyles and/or subtylostyles, thus also differing from the new species. TABLE 2. Comparative table of spicular measurements of the species of Aaptos of the western Atlantic. Measurements are expressed in µm, as minimum–mean–maximum length / width value whenever available. n.r., not reported. Data are from original descriptions, unless stated otherwise. Legend: St, style; S, subtylostyle; T, tylostyle; O, oxeas. Species

Strongyloxeas

Ectosomal spicules

Distribution

A. aaptos Schmidt, 1864

I. 1053–1911 x 12–31 II. 490–955 x 10–23

St–S: 135–230 x 1–5

Mediterranean Sea, NW Atlantic

A. aaptos sensu van Soest & Stentoft (1988)

1000–1400 x 10–30 (styles, occasionally oxeotes)

St: 180–260 x 2–4 (occasionally sinuous)

Barbados

A. aaptos sensu Kelly-Borges & Bergquist (1994)

I. 1053–1502–1911 x 12–29–31 II. 490–705–955 x 10–16–23

St: 364–388–509 x 5–7–8 S: 270–318–354 x < 1

Mediterranean Sea, Algiers

A. aaptos sensu Soest (2002; lectotype)

I. 1500–1900 x 12–45 II. 750–1000 x 10–25

St –S: 135–230 x 1–5

Mediterranean Sea

A. aaptos sensu Moraes (2011) I. 1000–1725 x 5–17 II. 220–490 x 4–10

St: 182–303 x 2–4

Atol das Rocas, Brazil

A. aaptos sensu Solé-Cava et al. 1064–1322–1522 x 17–34.2–47 (1981)

St: 250–319–425 x 3–6– 11.5

Guarapari, Espírito Santo State, Brazil

A. aff. aaptos sensu Mothes & Lerner (1994)

427–1633 x 6.9–41.4

St –T: 171–541.5 x 2.3– 13.8

Santa Catarina State, Brazil

A. bergmanni de Laubenfels, 1950

Up to 950 x 15

St: 150 x 2.5

Bermudas

A. bergmanni sensu BouryEsnault (1973) as A. aaptos

n.r.

St: 300–760 x 4.5–15

Pernambuco State, Brazil

A. durissima (Carter, 1882)

No measurements reported

A. duchassaingi (Topsent, 1889)

Up to 800 x 25 (styles)

n.r.

Caribbean and Mexico

A. glutinans Moraes, 2011

925–1375 x 7–14

St I: 172–296 x 2–7 St II: 240–570 x 3–12

Atol das Rocas, Brazil

A. pernucleata (Carter, 1870)

O: no measurements reported

St: no measurements reported

Caribbean

St: 549–569 (rare)

Bahia, Brazil

A. spp. sensu Hajdu et al. (2011) I. 1046–1320 x 23–30

South Africa

II. 208–950 x 5–25

NEW AAPTOS FROM BRAZIL (WEST ATLANTIC)

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FIGURE 3. Distribution frequency of the size-classes of the strongyloxeas of Aaptos hajdui sp. nov. (A. UFPE POR 62; B. UFPE POR 181) and A. potiguarensis sp. nov. (C. UFPE POR 45; D. UFPE POR 46). (n=100). x-axis size in microns; y-axis frequency in number of spicules per size class.

Aaptos potiguarensis sp. nov. (Figure 3C–D, 4; Table 1) Type material. Holotype: UFPEPOR 45—Campaign BPOT 02, station 33 (Potiguar Basin, Rio Grande do Norte State, 05°00´03.9”S–36°25´12.1”W), 7–10.2 m depth, coll. R/V ‘Astro Garoupa’, 28/V/2003. Schyzoholotype: MNRJ 16685. Paratype: UFPEPOR 46—Campaign BPOT 02, station 33 (Potiguar Basin, Rio Grande do Norte State, 05°00´03.9”S–36°25´12.1”W), 7–10.2 m depth, coll. R/V ‘Astro Garoupa’, 28/V/2003. Schyzoparatype: MNRJ 16686. Diagnosis. Aaptos with an ectosomal skeleton without subectosomal cavities, where small styles form a palisade at the surface. Spicules are one category of strongyloxeas with rounded tips (301–1562 µm in length and 8–43 µm thick) and one category of styles (220–590 µm in length and 2.5–11 µm thick). Description. Specimens are massive, up to 18.8 cm in length and 10.9 cm width (holotype; Fig. 4A). The base has incrustations of calcareous substrate. Oscules rounded, up to 3 mm in diameter, dispersed on the surface. Surface is irregular, uneven. Consistency is firm, almost incompressible. Color is unknown in the holotype, and light brown alive and beige in spirit, in the paratype. Skeleton. Ectosomal skeleton with strongyloxeas and styles crossing the surface. Small styles form a palisade at the surface, in which the ends of the strongyloxeas are also present. Choanosomal skeleton is very dense, composed mainly of tracts of strongyloxeas with a radial arrangement, protruding beyond the surface of the sponge (Fig. 4B). Little collagen is seen.

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FIGURE 4. Aaptos potiguarensis sp. nov. A. Holotype (UFPEPOR 45). B. Choanosomal skeleton. C. Strongyloxeas. D. Styles. Scale bars: A = 5 cm. B = 500 µm. C, D = 100 µm.

Spicules. Megascleres in two categories (Tab. 1, Fig. 4C–D). Choanosomal strongyloxeas (Fig. 4C)—smooth, highly fusiform, usually straight or slightly curved, with rounded tips: 301–1562 µm in length and 8–43 µm thick. Ectosomal styles (Fig. 4D)—smooth, straight, occasionally curved: 220–590 µm in length and 2.5–11 µm thick. The distinction between strongyloxeas and styles is very subtle and, as in Aaptos hajdui sp. nov., we consider strongyloxeas as fusiform or slightly fusiform and styles are isodiametric. Etymology. The specific epithet potiguarensis is refers to its type locality, from the Potiguar Basin. Distribution. Provisionally endemic from Potiguar Basin (Rio Grande do Norte State, north-eastern Brazil). Ecology. Bryozoans and polychaetes were found associated to both specimens. This species occurs in shallow waters, from 7 to 10 m depth on rocky substrate. Remarks. Nine species of the genus Aaptos have tylostyles and/or subtylostyles as ectosomal spicules, thus differing from A. potiguarensis sp. nov.: A. aaptos (holotype and lectotype descriptions sensu Van Soest 2002 and Kelly-Borges & Bergquist 1994); A. alphiensis; A. globosum; A. kanuux; A. laxosuberites; A papillata; A. robustus; A. rosacea; and A. tentum. Other species such as A. confertus and A. horrida possess oxeas as dermal or additional spicules, and A. pernucleata have true oxeas as main megascleres (as observed in figures of Carter, 1870). Aaptos aaptos possesses two categories of strongyloxeas, in a similar size range of the strongyloxeas of A. potiguarensis sp. nov., although some other differences can be observed. Besides having styles to substylostyles as dermal spicules, A. aaptos (holotype and lectotype, see Table 2) has its category smaller than in A. potiguarensis sp. nov. Aaptos hajdui sp. nov. is different from A. potiguarensis sp. nov. due to the telescopic/mucronate ends of the strongyloxeas in the former, in addition to the differences in the skeleton, as the presence of subectosomal cavities and the megascleres of the choanosomal skeleton forming bouquets near the surface. Besides, A. hajdui sp. nov. is NEW AAPTOS FROM BRAZIL (WEST ATLANTIC)

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globular and almost black after fixation, while A. potiguarensis sp. nov. is massive, turning beige after fixation in ethanol.

Discussion The genus Aaptos has 21 species known worldwide (van Soest et al. 2013), with six of them with records from the Western Atlantic (Tab. 2): A. aaptos; A. bergmanni; A. duchassaingi (Topsent, 1889); A. durissima (Carter, 1882); A. glutinans; and A. pernucleata. However, Aaptos aaptos has a worldwide distribution, which should be carefully re-examined. The possession of three categories of megascleres that overlap in size was put forward by Kelly-Borges & Bergquist (1994) as a key diagnostic characteristic in a genus that has a dearth of defining characters. We have found that, although there is always a large range in the primary choanosomal strongyloxeas (typically from 500 to 1000 µm), there are several species in which size categories are difficult to discern within this range, even by statistical analysis (Fig 3). Examples of such species include: A. bergmanni, A. ciliata, A. duchassaingi, A. horrida, A. niger, A. nuda, A. pernucleata, A. suberitoides, and A. vannamei (see Table 2). Size frequency distributions of spicule length in specimens of A. hajdui sp. nov. and A. potiguaris sp. nov. (Fig. 3) show little tendency to form clearly different size categories amongst the strongyloxeas, In order to accommodate these difficulties, we have emended the diagnosis of Aaptos accordingly. Kelly-Borges & Bergquist (1994) also emphasized a ‘cortical region’ defined by the presence of collagen in various conformations and densities, and differences in the arrangement of the primary skeletal tracts between species. Figure 2 shows the strongly divergent split tracts of A. hajdui sp. nov., and Figure 4 shows the linear swathes of spicules in A. potiguaris sp. nov., that do not diverge at the surface. Kelly-Borges & Bergquist (1994) and van Soest (2002) very clearly articulate key diagnostic characters that help to differentiate in this difficult genus. We propose that the main characters for the distinction of the species of Aaptos should consider also the external morphology, color, morphology and size of the megascleres, and details of the skeleton, for example, presence of ectosomal cavities, ectosomal palisade or bouquets. Some differences between the previous records of A. aaptos from Brazil and the lectotype described by van Soest (2002) and Kelly-Borges & Bergquist (1994) were noticed. Solé-Cava et al. (1981) did not find the smaller category of strongyloxeas (490–1000 µm) in the specimens of Guarapari, Espírito Santo State (Tab. 2). Furthermore, they exhibit styles larger than the lectotype (Tab. 2). The larger strongyloxeas of this record differentiate it from both new species. Mothes & Lerner (1994) found only one category of megascleres ranging from 427 to 1633 µm in length in specimens of A. aff. aaptos from Bombinhas, Santa Catarina State, which differs from the type due to the presence of a second category of megascleres including tylostyles, up to 541 µm in length (Tab. 2). The new Brazilian species described here do not have tylostyles as well. Although the specimen identified as A. aaptos described by Moraes (2011) from Atol das Rocas has two categories of strongyloxeas and one of styles, the second category of strongyloxeas is smaller than in the type (220–490 µm in the Brazilian specimen, while always larger than 490 µm in the types; see Table 2). The two categories of strongyloxeas are clearly separated in the specimens described by Moraes (2011), which differentiates it from both new species, in addition to the smaller styles in the Atol das Rocas’ species (Tab. 2). Hajdu et al. (2011) described Aaptos spp. from Bahia with two categories of strongyloxeas and one of styles (Tab. 2). Aaptos spp. sensu Hajdu et al. (2011) has no choanosomal spicule tracts, ectosomal cavities and ectosomal bouquets and in these characters it is considerably different from A. hajdui sp. nov, and closer to A. potiguarensis sp. nov. However, the styles in the Bahia’s specimens are rare and differ from the new species in length and thickness. A. potiguarensis sp. nov. has styles smaller and thinner than in Aaptos spp. (Tab. 1, 2). Besides, Aaptos spp. is globular, yellow, and dark brown after fixation, differing from the new species. Thus, we suggest that all the Brazilian material should be revised. Aaptos aaptos was also recorded to Brazil by Boury-Esnault (1973) based on the material collected in Pernambuco State by the Calypso Expedition. This record was synonymized with A. bergmanni by Hechtel (1976), who at the same time recorded this species to Bahia State. Solé-Cava et al. (1981) confirmed this synonymy arguing that the Boury-Esnault's (1973) specimens had only styles. However, in the original description, A.

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bergmanni has strongyloxeas up to 950 µm and styles with 150 µm, and the figures available in Boury-Esnault (1973) show that they are actually a smaller category of strongyloxeas, not styles. Furthermore, the spicular dimensions of the specimens studied by Boury-Esnault (1973) do not approach those of any species of Aaptos, neither with A. bergmanni nor with A. aaptos, which possesses strongyloxeas and true styles. Thus, we consider invalid all records of A. bergmanni from Brazil (from Bahia and Pernambuco State; sensu Hechtel, 1976), until the revision of these specimens. They probably belong to another species, yet undescribed in the Brazilian coast, and are better referred to as Aaptos sp. According to Van Soest et al. (2013), A. durissima was described from the Caribbean. However, despite the name of the species has been proposed by Carter (1882, p. 357 as Trachya durissima), the description of the species was published by Carter (1876, p. 393) from South Africa’s material. Anyway, according to his description, A. durissima presents strongyloxeas with tylostyle-like modifications, differing from the styles of both new species. Based on the above discussion, only three species of Aaptos can be considered valid in Brazil: A. glutinans, A. hajdui sp. nov. and A. potiguarensis sp. nov., while the records of A. aaptos and A. bergmanni are not accepted here and should be better examined. A comprehensive revision of all Brazilian Aaptos is in progress.

Acknowledgements CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior), CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico), FAPERJ (Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro), FACEPE (Fundação de Amparo à Ciência e Tecnologia do estado de Pernambuco) and CENPES– PETROBRAS are deeply thanked for the provision of grants and fellowships. The authors are thankful to Guilherme Muricy (Museu Nacional, UFRJ) for critically reading the manuscript and for help with the identification.

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