Desmacella Schmidt, 1870 from Brazil: Description of two new species and a review of records (Desmacellida: Demospongiae: Porifera)

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http://dx.doi.org/10.11646/zootaxa.4034.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:5C08231F-2280-4FF8-A042-F94F04466AE3

Desmacella Schmidt, 1870 from Brazil: Description of two new species and a review of records (Desmacellida: Demospongiae: Porifera) THAYNÃ CAVALCANTI1, 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 Three species of Desmacella Schmidt, 1870 are described from the Brazilian coast: Desmacella microsigmata sp. nov., Desmacella tylovariabilis sp. nov. and Desmacella annexa Schmidt, 1870. The new species were compared with Desmacella species from the Atlantic Ocean, which differ by the size of their spicules and by the presence of microspined sigmas. Desmacella annexa was found to have microspined toxiform microxeas and sigmas. Desmacella now contains 31 species distributed worldwide. Key words: Porifera, taxonomy, marine biodiversity, deep waters, Atlantic Ocean

Introduction The genus Desmacella has 29 species worldwide, and includes sponges with an hispid surface, reticulate, plumoreticulate and vaguely halichondrioid choanosomal skeleton, megascleres tylostyles or styles and microscleres sigmas and raphides (Hajdu & van Soest 2002). The Atlantic Ocean has 17 known species of Desmacella (van Soest et al. 2015). Nine species occur in the Western Atlantic: D. annexa Schmidt, 1870, D. jania Verrill, 1907, D. meliorata Wiedenmayer, 1977, D. polysigmata van Soest, 1984, D. pumilio Schmidt, 1870, D. suberitoides (Burton, 1932), D. vagabunda Schmidt, 1870, D. vestibularis (Wilson, 1904), and D. vicina Schmidt, 1870. Up until the present there have been one description of Desmacella species from Brazil, as Desmacella aff. pumilio (Cosme & Hajdu 2010), but only Desmacella annexa has been mentioned in the literature and identified to species level (Hajdu et al. 2004, from Southeast Brazil). Furthermore, specimens of the genus have been recorded six times between the Bahia and Rio Grande do Sul States (see Muricy et al. 2011), although not identified to species level. Here, we review records of Desmacella from Brazil, and describe two new species.

Material and methods The specimen of Desmacella microsigmata sp. nov. was collected from trawl site by PETROBRAS (Petróleo do Brasil S/A) in 2011, as part of the Project ‘Campanha de Monitoramento Ambiental do Projeto de Caracterização Ambiental do Talude Continental na Bacia Potiguar/ Rio Grande do Norte/ BR’ (BPot). It was preserved in 80% ethanol and deposited in the Coleção de Porifera of Universidade Federal de Pernambuco (UFPEPOR). The specimen of Desmacella tylovariabilis sp. nov. was collected from dragging by PETROBRAS in 2003, as part of the Project ‘Caracterização Ambiental de Águas Profundas da Bacia de Campos’ (CENPES). The specimens of Desmacella annexa was collected from trawl site by REVIZEE Project, in 1998 (Hajdu et al. 2004). The specimens of Desmacella tylovariabilis sp. nov. and Desmacella annexa are deposited at Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ) and it was preserved in 80% ethanol. Dissociated spicules mounts, SEM preparations and skeletal sections were made using classical procedures for Demospongiae (Hajdu et al. 2011).

364 Accepted by J. Hooper: 12 Oct. 2015; published: 29 Oct. 2015

Images of specimens, sections (Leica DM 500 microscope) and SEM (Quanta 200 FEG) preparations were obtained digitally. Taxonomic comparisons were made with data tabulated for Atlantic species of Desmacella catalogued in the World Porifera Database (van Soest et al. 2015).

FIGURE 1. Location of the collection sites of Desmacella species from Brazil. (1) Desmacella microsigmata sp. nov. (Rio Grande do Norte State); (2) Desmacella tylovariabilis sp. nov. (Rio de Janeiro State); (3) Desmacella annexa (São Paulo State). NEW DESMACELLA FROM BRAZIL

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Systematics Class Demospongiae Sollas, 1885 Subclass Heteroscleromorpha Cárdenas, Perez & Boury-Esnault, 2012 Order Desmacellida Morrow & Cárdenas, 2015 Family Desmacellidae Ridley & Dendy, 1886 Genus Desmacella Schmidt, 1870 Synonymy (see Hajdu & van Soest 2002). Definition. Desmacellidae with tylostyles, occasionally styles, with reticulate, plumo-reticulate and vaguely halichondrioid choanosomal skeleton, microscleres sigmas, and raphides; the latter may be absent (Hajdu & van Soest 2002).

Desmacella microsigmata sp. nov. (Figures 1, 2, Table 1) Type locality: Brazil, Rio Grande do Norte State, Potiguar Basin. Type specimens: Holotype. UFPEPOR 1759, Bacia Potiguar (04° 36.6848’ S; 036° 46.6926’ W), Rio Grande do Norte State, Brazil, depth 157 m, trawl, coll. Petrobrás, (21/V/2011). Diagnosis. Desmacella microsigmata sp. nov. is an encrusting sponge, characterized by the presence of tylostyles (286.3 x 3.9 µm, length x width) with elliptical tyles situated at varied positions at their bases, and small sigmas with microspined ends (14.6 µm, chord length). External morphology (Fig. 2A). Encrusting sponge, 1 x 1.5 cm (length x width) (Fig. 2A). Hispid surface, consistency is fragile, oscules were not seen, colour in situ is unknown, brownish-purple in ethanol. The specimens were collected in the same dredge as Aiolochroia crassa (Hyatt, 1875) and stored in the same container. It is possible that the A. crassa pigments stained the Desmacella specimens (Cavalcanti et al. 2014). Skeleton (Fig. 2B). The ectosomal skeleton consists in terminations of the ascending choanosomal tracts. Choanosomal skeleton is composed of poor spongin fibres and tylostyles forming bouquets of spicules projecting through the surface. Sigmas are randomly distributed. Spicules (Fig. 2C–F). Tylostyles (177–286.3–425 x 2–3.9–7 µm, length x width): long, thin, smooth, straight to slightly curved, with elliptical tyles situated at varied positions at their bases (Fig. 2C, F); Sigmas (12–14.6–19 µm, chord length): small, thin, abundant and with microspined ends (Fig. 2D, E). Ecology. The specimens of Desmacella microsigmata sp. nov. was found associated with the bryozoan Canda alsia Winston, Vieira & Woollacott, 2014. Distribution (Fig. 1). Brazil: Northeastern Region: Rio Grande do Norte State: off Potiguar Basin. Etymology. The species name refers to the small size of the sigma. Remarks. Desmacella microsigmata sp. nov. belongs to the genus by the possession of tylostyles and sigmas, and a skeleton consisting of plumose bundles of tylostyles. Desmacella microsigmata sp. nov. differs from other Atlantic species by its elliptical tyles with variously located along the basal part of the shaft of the megasclere and dimensions of the spicules. The new species differs from D. annexa, D. digitata, D. polysigmata, D. pumilio and D. vicina by having just one category of sigmas. Desmacella grimaldii, D. informis, D. inornata, D. suberitoides, D. topsenti, D. vagabunda and, D. vestibularis are distinguished by the possession of larger tylostyles and sigmas bigger than those of D. microsigmata sp. nov. (see Table 1). The new species differs from D. infundibuliformis, D. jania and D. meliorata, by the encrusting growth form of the new species and its tiny sigmas (see Table 1). Desmacella peachi sensu Ferrer-Hernandez (1914) is distinguished from D. microsigmata sp. nov. by the presence of raphides, sinuous tylostyles and lacking sigmas. Finally, in spite of D. corrugata’s description (Bowerbank, 1866, from Azores) not having included the dimensions of its spicules, it can still be differentiated from D. microsigmata sp. nov. due to the former’s tylostyles lacking variously located subterminal tyles.

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Barbados / 100

São Paulo State (Brazil) / 153

Azores, Celtic Seas, United Kingdom / Not recorded

Sahelian Upwelling, Senegal / 25–30

Azores / 927

D. annexa Schmidt, 18701,2

D. annexa sensu van Soest & Stentoft 19883

D. annexa (present study)

D. corrugata (Bowerbank, 1866)1,4

D. digitata (Lévi, 1960)1,5

D. grimaldii (Topsent, 1890)1,6

Arctic Ocean, Azores / 228.6

Aegean Sea, Alboran Sea, Azores, Western Mediterranean / 100–270

Bermuda, Caribbean Sea, Mexico / not recorded

Bahamas, Caribbean Sea / not recorded

D. infundibuliformis (Vosmaer, 1885)8,9

D. inornata (Bowerbank, 1866)1,4

D. jania Verrill, 19071,10

D. meliorata Wiedenmayer, 19771,11

Ireland / 457–1024

Florida / 350

Desmacella tylovariabilis sp. nov.

D. informis (Stephens, 1916)

Rio de Janeiro State (Brazil) / 1130

Desmacella microsigmata sp. nov.

1,7

Region Found / Depth (m)

Potiguar Basin (Brazil) / 157

Species

Lobular / Crimson red

Massive and irregularly lobulate / White (dried)

Amorphous / Dirty gray (dried)

Leaf-like/ Not recorded

Thinly encrusting / Brown

Globular / Not recorded

Finger-like / Light cream

Not recorded / Blood-red

Fragment, encrusting / Cream (fixed)

Thin film / Not recorded

Encrusting / Not recorded

Massive / Beige (fixed)

Encrusting / Dark brown (fixed)

Shape / Color

210–230 / 3.5–4.5

220–250 (styles to tylostyles)

190–1000 / 6–18

250 / 500

180–1300 / 8–27

390–1900 / 8–30

180–270 / 1–2

present, size not given

286–392.5–521 / 3–8.1–14

280–700 / 2.5–8

present, size not given

315–616.0–1050/ 6–11.0–16

177–286.3–425 / 2–3.9–7

Tylostyles

37 / 2

37–40

20–45

25

26–45

28–45

22–26

present, size not given

19–29.8–38

28–42

14–over 100

25–34.2–48

12–14.6–19

Sigmas I

-

-

-

-

-

-

Raphides

-

-

-

-

-

-

-

-

54–76.7–90 (toxiform)

53–115 / 0.5–2 (toxiform)

thin oxeas (size not given)

-

-

......continued on the next page

14–18

-

9–11.6–14

11–15

-

-

-

Sigmas II

TABLE 1. Comparative micrometric data on the spicules, shape, colour and overview of distribution of the living species of Desmacella Schmidt, 1870 from Atlantic Ocean. Values are in micrometres (µm), expressed as follows: minimum–maximum or minimum–mean–maximum length/width. References are numbered in parentheses and listed after the table.

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Not recorded

Florida, Caribbean Sea, Greater Antilles, Gulf of Mexico / 98.7

Tristan Gough / 80–140

Azores / Not recorded

Florida / 30–44

Galapagos Islands, Namibia, Eastern Philippines / 16

Florida /Not recorded

D. pumilio Schmidt, 18701,13,14,16

D. suberitoides (Burton, 1932)1,15

D. topsenti (Burton, 1930)1,16

D. vagabunda Schmidt, 18701,14

D. vestibularis (Wilson, 1904) 1,17

Thinly encrusting / Not recorded

Encrusting / Not recorded

Encrusting / Black

Club-shaped, thinly to massively encrusting / Greyish brown

Sprawling body / Beigesandy

600 / 12

240–630 / 8–16

600

250–730 / 5–10

1000 / 18 (choanosomal); 600 / 12 (ectosomal)

320–1400 / 9–17

36

12–36

14–over 100

43

28

30–46

30–37.3–42

513– 575.4–635 / 10–15.2– 19 (styles to strongyles)

Sigmas I -

Tylostyles present, long and sinuous, size not given

12

-

-

-

-

12–27

10–11.6–15

-

Sigmas II

-

-

-

-

-

-

-

present, thin and, curved size not given

Raphides

References: (1) Lehnert et al. (2005); (2) Topsent (1936); (3) van Soest & Stentoft (1988); (4) Bowerbank (1866); (5) Lévi, (1960); (6) Topsent (1890); (7) Stephens (1916); (8) Vosmaer (1885); (9) Fristedt (1887); (10) Verril (1907); (11) Wiedenmayer (1977); (12) Ferrer-Hernandez (1914); (13) Van Soest (1984); (14) Schmidt (1870); (15) Burton (1932); (16) Burton (1930); (17) Wilson (1904).

D. vicina Schmidt, 18701,14

Encrusting / Yellowish brown

Belize, Caribbean Sea / 100

D. polysigmata van Soest, 19841,13

Not recorded

D. peachi sensu Ferrer-Hernandez, 19141,12

Shape / Color

Region Found / Depth (m)

Spain / not recorded

Species

TABLE 1. (Continued)



FIGURE 2. Desmacella microsigmata sp. nov. (UFPEPOR 1759) in SEM. A, preserved specimen; B, thick section showing the ectosome and choanosome; C, size variation of tylostyles; D, sigma; E, details of microspines on the tip of a sigma; F, variation of tylostyle bases. Scale bars: A = 1 cm; B = 200 µm; C = 100 µm; D = 5 µm; E = 2 µm; F = 10 µm.

Desmacella tylovariabilis sp. nov. (Figures 1, 3, Table 1) Type locality: Brazil, Rio de Janeiro State, Campos Basin.

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FIGURE 3. Desmacella tylovariabilis sp. nov. (MNRJ 7339) in SEM. A, preserved specimen; B, thick section showing the ectosome and choanosome; C, size variation of tylostyles; D, variation of tylostyle bases; E, sigmas; F, details of microspines on the tip of a sigma. Scale bars: A = 1 cm; B = 200 µm; C = 400 µm; D–E = 10 µm; F = 2 µm.

Type specimens: Holotype. MNRJ 7339, Campos Basin (22° 58’ S; 42° 1’W), Rio de Janeiro State, Brazil, 1130 m depth, dragging, coll. CENPES/UFRJ, (07/II/2003). Diagnosis. Desmacella tylovariabilis sp. nov. is the only Desmacella in the Tropical Western Atlantic with the combination of large tylostyles (616.0 x 11 µm, length x width) bearing rounded tyles in varied positions, and sigmas with microspined ends (34.2 µm, chord length). External morphology (Fig. 3A). Massive sponge, 6 x 4.5 cm (length x width) (Fig. 3A). Hispid surface, without distinct oscules, fragile consistency and colour beige (fixed in ethanol).

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Skeleton (Fig. 3B). Ectosomal skeleton with plumose bundles of tylostyles. Choanosomal skeleton composed of a vaguely halichondrioid skeleton; poorly developed spongin fibres. Sigmas appear to be distributed randomly. Spicules (Fig. 3C–F). Tylostyles (315–616.0–1050 x 6–11.0–16 µm, length x width): long, thin and smooth, straight to slightly curved, rounded tyles situated at varied positions at their bases (Fig. 3C, D); Sigmas (25–34.2– 48 µm, chord length): thin, abundant and microspined ends (Fig. 3E, F). Distribution (Fig. 1). Brazil: Southeastern Region: Rio de Janeiro State: Campos Basin. Etymology. The specific epithet tylovariabilis refers to the variation observed on the tyles of the tylostyles, tylo = from the Greek tyle, meaning knob or lump, and varius = from the Latin, meaning different or various. Remarks. Desmacella tylovariabilis sp. nov. is assigned to the genus because of its tylostyles and sigma spicules, and vaguely halichondrioid skeleton formed by large tylostyles. There are four species similar to D. tylovariabilis sp. nov.: D. grimaldii, D. informis, D. inornata and D. topsenti. They occur in the North Atlantic Ocean, and have similarities regarding spicule dimensions (see Table 1). However, the above species differ from D. tylovariabilis sp. nov. which has rounded tyles situated at varied positions at their bases. The new species differs from D. annexa, D. digitata, D. polysigmata, D. pumilio and D. vicina, by the presence of only one category of sigmas. Desmacella tylovariabilis sp. nov. is distinguished from D. infundibuliformis, D. jania, D. meliorata, D. vagabunda and D. vestibularis, by the new species’ larger tylostyles and sigmas (see Table 1). Desmacella corrugata differs from the new species by the different morphology of its megascleres. Additionally, the new species differs from D. suberitoides as that species presents two categories of tylostyles. Finally, D. tylovariabilis sp. nov. has massive growth form and larger spicules than those of D. microsigmata sp. nov. (Table 1).

Desmacella annexa Schmidt, 1870 (Figures 1, 4, Table 1) For full synonymy see Muricy et al. 2011. Specimens examined. MNRJ 2860 (Hajdu et al. 2004), São Paulo State (25º 36.988’ S; 45º 13.571’ W), Brazil, 153 m depth, REVIZEE South, trawl, station 6686 (28) (13/I/1998), det. E. Hajdu et al. External morphology (Fig. 4A). Fragments, thinly encrusting 2 x 1 cm (length x width). Oscules were not found, vaguely hispid surface, consistency is fragile and colour cream in ethanol. Skeleton (Fig. 4B). There is no special ectosomal skeleton. Choanosomal skeleton formed by large tylostyles making massive plumose bundles; well developed spongin fibres. Abundant sigmas (I and II) appear to be randomly distributed. Toxiform microxeas were not recognized. Spicules (Fig. 4C–I). Tylostyles (286–392.5–521 x 3–8.1–14 µm, length x width): long, thin, erect and smooth (Fig. 4C–D); Sigmas I (19–29.8–38 µm, chord length) and Sigmas II: (9–11.6–14 µm, chord length) thin, abundant and terminally microspined (Fig. 4F, G, I); Toxiform microxeas (54–76.7–90 µm, length) long, thin, curved and microspined (Fig. 4E, H). Geographical distribution. Florida (Schmidt 1870), South and West Iceland (Stephens 1921), Adriatic Sea (Pansini 1987), Mediterranean (Vacelet 1969), Naples (Pulitzer-Finali 1978), Spain (Bibiloni 1981), Azores (Boury-Esnault & Lopes 1985), Barbados (van Soest & Stentoft 1988), Alboran Sea (Boury-Esnault et al. 1994; Sitj & Maldonado 2014) and Aegean Sea (Voultsiadou 2005). Brazil: São Paulo State: off Guarujá and off Ilhabela (Hajdu et al. 2004; Hajdu & Lopes 2007; Muricy et al. 2011). Remarks. Van Soest & Stentoft (1988) described the typical morphology of D. annexa. The species’ spicules consist of tylostyles, two categories of sigmas and toxiform microxeas. Here, we provide for the first time illustrations using scanning electron microscopy of D. annexa, the microspination of its sigmas and toxiform microxeas. The presence of microspination on microscleres is also observed in other species of Desmacella such as D. austini Lehnert et al. (2005, from the Pacific coast of Canada) (Lehnert et al. 2005) and Desmacella aff. pumilio (from Campos Basin, Brazil) (Cosme & Hajdu 2010). Desmacella austini has microspines on the tips of its two categories of sigmas, while Desmacella aff. pumilio has microspines on the tips of one category of sigma. We believe that this new feature probably occur in other populations of D. annexa, however this hypothesis can be confirmed only with SEM images. Records biogeographically akin to the Floridian type locality, such as the Brazilian ones (see Hajdu et al. 2004), are likely conspecific. Nevertheless, the other more disjunct records need to be reviewed with great care.

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FIGURE 4. Desmacella annexa (MNRJ 2860) in SEM. A, preserved specimen; B, thick section showing the ectosome and choanosome; C, size variation of tylostyles; D, tylostyle base; E, size variation of toxiform microxeas; F, sigma I and sigma II; G, details of sigma I microspines; H, details of toxiform microxea microspines; I, details of sigma II microspines. Scale bars: A = 1 cm; B = 200 µm; C = 100 µm; D = 10 µm; E = 20 µm; F = 10 µm; G–H = 2 µm; I = 1 µm.

Discussion Desmacella has a strong similarity to Biemna Gray, 1867, which has a similar, but denser skeleton (see Hajdu & van Soest 2002). Desmacella and Biemna share also the possession of a massive shape, sigmas and raphides. However, both genera differ in many features: Desmacella has tylostyles or styles of one or more sizes whereas typical Biemna has (subtylo-)styles of a single size (Hajdu & van Soest 2002). Furthermore, Desmacella might have vaguely halichondrioid choanosomal skeletons versus plumose or plumoreticulate choanosomal skeleton in Biemna. Additionally, commata and microxeas (microspined) are found in Biemna but not recorded in Desmacella (except for toxiform microxea in Desmacella annexa).

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Some species of Desmacella have been known for almost a century, and as such lack precise or accurate information about their morphology, and as such difficult to differentiate from more recently described species. For example, four species from the North Atlantic, viz. Desmacella grimaldii, D. informis, D. inornata and D. topsenti, share the presence of tylostyles and sigmas with very similar dimensions, and possibly they belong to a single species complex. By comparison, some species such as Desmacella annexa, discussed above, have widespread, disjunct distributions, in which populations may represent a cryptic species complex of two or more taxa. These anomalies reinforce the necessity to reexamine the poorly known or poorly described species using standardized technologies (such as SEM), to revise Desmacella.

Acknowledgments We thank Dr Eduardo Hajdu (Museu Nacional, Universidade Federal do Rio Janeiro) for the loan of specimens. The authors are grateful to PETROBRAS for donating the type material and providing collection data. Authors thank Dr Janaina Melo, Diego Oliveira, and Josineide Correia for SEM facilities at CETENE (Centro de Tecnologias Estratégicas do Nordeste), Adélia Alliz, MSc Joana Sandes, MSc Ana Carolina Almeida, Dr Paulo Santos (UFPE), Dr André Esteves (UFPE) and to Dr Leandro Vieira (UFPE) for technical support. G.S. thanks CAPES for providing a doctoral scholarship. We further thank CNPq (Edital PROTAX: 562320/2010-5 and Universal 454908/2014-8 to UP), and FACEPE (APQ-0839-2.04/08 to UP), for providing grants and/or fellowships.

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