ORIGINAL PAPER

New records of marine Gastrotricha from São Sebastião Island(Brazil) and the description of a new species

André R. S. Garraffoni1 & Maikon Di Domenico2,3 & Rick Hochberg4

Received: 26 September 2015 /Revised: 20 March 2016 /Accepted: 23 March 2016 /Published online: 21 April 2016# Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2016

Abstract Until a decade ago, the marine gastrotrich fauna ofBrazil was virtually unknown, but, since then, there has been anincrease in the number of reports of new and known speciesfrom the states of São Paulo, Rio de Janeiro, and Bahia. Here,we report new records of five marine gastrotrichs from sublit-toral sediments around São Sebastião Island, Brazil, collectedby scuba. The genus Acanthodasys (Macrodasyida), represent-ed by two undescribed species, is reported for the first time inthe Southern Hemisphere; Crasiella fonseci is reported for thefirst time in São Sebastião Island and outside of its type locality,and Dendrodasys aff. rubomarinus and Aspidiophorustentaculatus (Chaetonotida) are reported for the first time inFome Beach. Dactylopodola todaroi sp. nov. is distinguishedfrom its congeners by the following combination of characters:red ocellar granules, four TbAwith two lateral tubes longer thanthe two medial tubes, five TbL (1+1+3), and seven TbP. Inaddition to new geographic records of Gastrotricha, we provideestimates of the diversity of this taxon on the Brazilian coast, asonly 3 % of the total coastline has been sampled so far. We

further examine our findings in light of a recent survey of theCaribbean ecoregions.

Keywords Macrodasyida . Dactylopodolidae .

Dactylopodola . Thaumastodermatidae . Planodasyidae .

Chaetonotidae . Atlantic Ocean

Introduction

Gastrotrichs are acoelomate microinvertebrates (most speci-mens are less than 1mm in length) commonly found inmarineand freshwater sediments (Balsamo and Todaro 2002; Todaroand Hummon 2008), and are some of the most abundant in-vertebrates in the marine meiofauna (Hochberg 1999).

The Gastrotricha comprises more than 820 species(Balsamo et al. 2015), traditionally divided into the two ordersMacrodasyida and Chaetonotida. The first order comprisesnine families and 33 genera of mostly marine species withstrap-shaped bodies, pharyngeal pores, and, usually, numer-ous adhesive tubes present in the anterior, lateral, and posteriorbody regions (Todaro and Hummon 2008; Balsamo et al.2015). Freshwater species are rare (Ruttner-Kolisko 1955;Kisielewski 1987; Garraffoni et al. 2010; Todaro et al. 2012;Araújo et al. 2013; Kånneby and Wicksten 2014). TheChaetonotida includes seven families and 30 genera of ma-rine, brackish, and freshwater species, with most species (ca.two-thirds) described from various freshwater habitats(Balsamo and Todaro 2002; Balsamo et al. 2008).Gastrotrichs have a cosmopolitan distribution, but most spe-cies are described from Europe and North America(Kisielewski 1991; Todaro and Rocha 2004, 2005; Balsamoet al. 2008), because most of the specialists in the group arehoused in institutions on these two continents. This taxon is a

Communicated by P. Martinez Arbizu

* André R. S. Garraffoniarsg@unicamp.br

1 Laboratório de Evolução de OrganismosMeiofaunais, Departamentode Biologia Animal, Instituto de Biologia, Universidade Estadual deCampinas, CP 6109, 13083-970 Campinas, SP, Brazil

2 Museu de Zoologia BProf. Dr. Adão José Cardoso^, Universidade deCampinas, Instituto de Biologia, CP 6109, 13083-863 Campinas, SP,Brazil

3 Present address: Centro de Estudos do Mar, Universidade Federal doParaná, CP 50.002, 83255-000 Pontal do Paraná, PR, Brazil

4 Department of Biology, University of Massachusetts Lowell,Lowell, MA 01854, USA

Mar Biodiv (2017) 47:451–459DOI 10.1007/s12526-016-0486-1

representative of the Bleast-known taxonomic groups^(Appeltans et al. 2012).

Until 11 years ago, the marine habitats of Brazil wererecognized as terra incognita for gastrotrich fauna (Todaroand Rocha 2004, 2005; but see Forneris 1985). However,the studies of Todaro and Rocha (2004, 2005) revealednew gastrotrich biodiversity along the northern coasts ofthe state of São Paulo and in the southern part of theState of Rio de Janeiro. In these initial studies, the authorsdescribed one new species (Macrodasys fornerise) and re-ported 42 additional species (most of these species appearto be undescribed). More recently, Todaro (2012, 2013)described two new species originally found in his earlierstudies: Pseudostomella dolichopoda (reported asPseudostomella sp. in Todaro and Rocha 2004) andPtychostomella lamelliphora (reported as Ptychostomellasp. in Todaro and Rocha 2004). Hochberg (2014) de-scribed a new species of Planodasyidae, Crasiella fonseci,collected on Tamanduá Island in Caraguatatuba (northerncoast of the state of São Paulo). Also, in 2014, Araujoet al. (2014) described a second new species ofPseudostomella from Brazil, P. squamalongispinosa, whichwas collected on the northeast coast of Brazil. Here, weprovide an assessment of gastrotrich diversity for the firsttime at Fome Beach, São Sebastião Island, from the depthrange of 8–10 m, and also describe a new species ofDactylopodola (Dactylopodolidae).

Materials and methods

Fieldwork was conducted (12/12/2014, 17/07/2015) at SãoSebastião Island (where the municipality of Ilhabela ishoused) in Fome Beach (23°44’27^S; 45°16’01^W;Fig. 1). We collected the top 15 cm of sediment at adepth of 8–10 m (total of 8 liters of sediments).Sediments were placed in plastic buckets and gastrotrichswere extracted at the Unicamp, in Campinas. Thegranulometric characteristics of the sediment at FomeBeach are: mean grain size (phi) 0.55, size class coarsesand, sorting (phi) 0.46, sorting class well sorted, skew-ness 1.25, and kurtosis 5.40 (Garraffoni et al. 2016).Extraction and analysis of gastrotrichs were performed fol-lowing the protocol reported by Hochberg and Atherton(2010). The specimens were sorted with a ZeissDM2000, mounted on glass slides, observed in vivo undera Zeiss Axioskop 2 plus equipped with differential inter-ference contrast (DIC). All examined specimens and pho-tographs are deposited in the Gastrotricha collection at theZoological Museum (ZUEC GCH) of the University ofCampinas, São Paulo, Brazil. Here, we provide a photo-graph of each taxon and a brief diagnosis of the majortaxonomic structures.

Results

Although the main concern of the present study was theassessment of gastrotrich fauna, we could also observe thatthe meiofauna in general at Fome Beach is highly diverseand abundant. In the sorted sediment, we were able toidentify the following taxa: Cephalochordata (amphioxus:abundant), Acoelomorpha (abundant), Halammohydra(Cnidaria: abundant), Proseriata (Platyhelminthes: abun-dant), Kalyptorhyncha (Platyhelminthes: abundant),Nematoda (several families, among them Desmocolidae,Draconometidae: very abundant), Polychaeta (Syllidae,Cirratulidae, Ctenodrilidae, Protodrilidae, Diurodrilidae),Rotifera (not abundant), Kinorhyncha (not abundant), andNemertea (abundant).

Among gastrotrichs, we were able to positively identifytwo known species: Aspidiophorus tentaculatus(Chaetonotida: Chaetonotidae) and Crasiella fonseci(Macrodasyida: Planodasyidae). We also identified a newspecies of Dactylopodola (below). Moreover, we providedetails of three species that appear to be new to sciencebut require further information before making formal de-scriptions: Acanthodasys sp. 1 and sp. 2 (Macrodasyida:Thaumastodermatidae) and a morphotype similar toDendrodasys rubomarinus, but here considered as D. aff.rubomarinus (Macrodasyida: Dactylopodolidae). We en-countered several specimens from seven genera(Chaetonotus , Aspidiophorus , Halichaetonotus ,Xenotricula, Paradasys, Ptychostomella, Thaumastoderma)that could not be confidently ascribed to known speciesbased on available data. Finally, as reported by Garraffoniet al. (2016), at least four distinct morphotypes ofTetranchyroderma are present at Fome Beach.

Order Chaetonotida Remane, 1925 [Rao and Clausen,1970]

Family Chaetonotidae Zelinka, 1889Genus Aspidiophorus Voigt, 1904Aspidiophorus tentaculatus Wilke, 1954(Fig. 2a, b)Description: Body length 240–260 μm (adult, n=3), elon-

gate body; pharynx 70–85 μm long with distinct posteriorbulb; PhJIn at U35-38; large number of dorsal pedunculatedscales (more than 100); presence of cephalion andhypostomium; a pair of cephalic tentacles posterior of thetactile cilia in the head (Fig. 2a, b).

Ecology: rareGeographical distribution: Cosmopolitan species (Atlantic

and Indian Oceans, Mediterranean and Red Seas [Hochberget al. 2014 and references within]; Brazil: Paraty: VermelhaBeach, Ubatuba: Prumirim Beach [Todaro and Rocha 2005]São Sebastião Island: Fome Beach).

Remarks: The morphometric data (total body length, phar-ynx and furca lengths) of the specimens from Fome Beach are

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larger than those reported by Hochberg et al. (2014).However, these morphometric differences are considered torepresent natural variation in the population and unlikely tosignal the presence of a new species.

Order Macrodasyida Remane, 1925 [Rao & Clausen,1970]

Planodasyidae Rao and Clausen, 1970Crasiella Clausen, 1968Crasiella fonseci Hochberg, 2014(Fig. 2c–e)Description: Body length 410–500 μm (adult, n=7). Strap-

shaped body, three-lobed head without piston pits; small ter-minal mouth; 130 μm pharynx length, pharyngeal pores atU30; cilia present along lateral and dorsolateral surfaces;many TbA and TbVL (not quantified); eight TbP arrangedon a pair of small caudal pedicles; epidermal glands are nu-merous in the pharyngeal region and along lateral body mar-gins of the trunk (Fig. 2a–c).

Ecology: Very abundant (probably the most abundantmeiofaunal taxon in Fome Beach)

Geographical distribution: Known only from Brazil(Caraguatatuba: Tamanduá Island [Hochberg, 2014]; SãoSebastião Island: Fome Beach).

Remarks: Our specimens are in agreement with the originaldescription (Hochberg 2014).

Family Thaumastodermatidae Remane, 1929Acanthodasys Remane, 1927Acanthodasys sp. 1(Fig. 3a–c)Description: Body length 620–850 μm (adult, n=2).

Body is mostly strap-shaped with a very small pair ofcaudal pedicles. TbA 6–7 per side, forming two distinctgroups, each one arranged forming two transversal rows.TbVL 20 per side. Scales are eye-shaped and spinedscales (uniancres) are present; uniancres present dorsallyand ventrally, though much larger on the dorsal side (dor-sal: 17 μm, ventral 5–8 μm); three TbP: two terminaltubes and one medial tube on each very small pedicle(Fig. 3a–c).

Ecology: Abundant

Fig. 1 Sampling locality alongthe coast of the state of São Paulo(Brazil) from Fome Beach, SãoSebastião Island

Mar Biodiv (2017) 47:451–459 453

Geographical distribution: Known only from Brazil (SãoSebastião Island: Fome Beach).

Acanthodasys Remane, 1927Acanthodasys sp. 2(Fig. 3d–f)Description: Body length 421–686 μm (adult, n=4). Strap-

shaped body with a distinct pair of caudal pedicles. TbA 6 perside and distributed along the mouth margin in a single row.TbVL 7 per side. Scales are eye-shaped with a central depres-sion and spined scales (uniancres) also present; scales coverentire body; large uniancres on the dorsal surface and smalluniancres on the ventral surface; TbP present as two terminaltubes and two medial tubes (Fig. 3d–f).

Ecology: RareGeographical distribution: Known only from Brazil (São

Sebastião Island: Fome Beach).Remarks: The genus Acanthodasys is reported from coastal

localities throughout Europe, the Mediterranean and BlackSeas, the Caribbean, and the coastal United States (reviewedin Atherton and Hochberg 2012). In the present study, wereport two undescribed species of Acanthodasys, which isthe first report of the genus in the Southern Hemisphere.These two potentially new species from Brazil are similar tothe four described species that possess a combination of spine-less scales and uniancres: A. aculeatus Remane, 1927,A. arcassonensis Kisielewski, 1987, A. caribbeanensisHochberg and Atherton, 2010, and A. paurocactus Athertonand Hochberg, 2012. Clausen and Båmstedt (2000) also reporttwo undescribed species that possess a similar armature. In our

study, Acanthodasys sp. 1 appears to differ from these previ-ous species by the number and size of the uniancres on bothdorsal and ventral sides; Acanthodasys sp. 2. differs from allother species by the presence of eye-shaped scales with acentral depression similar to A. caribbeanensis, but differingfrom this species in the size and distribution of the spines. Amore formal description of these species awaits the applica-tion of scanning electron microscopy (SEM) to verify the finestructure of the scales and spines and provide more accuratecounts of the adhesive tubes, which are often difficult to visu-alize in DIC micrographs.

Family Dactylopodolidae Strand, 1929Genus Dendrodasys Wilke, 1954Dendrodasys aff. rubomarinus Hummon, 2011(Fig. 4)Description: Body length 260–276 μm (adult, n=3). Head

that is slightly triangular, protruding anteriorly, with lobessomewhat rounded, and one pair of pestle organs. TbA oneper side; TbL and TbD absent. TbP arise from elongatecaudum: one TbP per side at the base of the caudum andtwo TbP arising terminally at bifurcation; testes paired and

Fig. 3 Acanthodasys sp. 1, DIC photos: a habitus, b dorsal view of thehead region, c ventral view of the pharyngeal region; Acanthodasys sp. 2,DIC photos: d habitus, e dorsal view of the head region, f ventral view ofthe trunk region. an, uniancre; TbL, lateral adhesive tubes; TbP, posterioradhesive tubes; sp, eye-shaped spineless scale. Scales bars: b, c, e, f:35 μm; a, d: 74 μm

Fig. 2 Aspidiophorus tentaculatus, DIC photos: a dorsal view of thehead region, b dorsal view of the posterior end; Crasiella fonseci, DICphotos: c habitus, d dorsal view of the head region, e dorsal view of theposterior end. ci, cilia; hy, hypostomium; ph, pharynx; pp, pharyngealpores; TbA, anterior adhesive tubes; TbP, posterior adhesive tubes; te,testes. Scales bars: a, b: 40 μm; c: 74 μm; d, e: 35 μm

454 Mar Biodiv (2017) 47:451–459

present laterally at the anterior third of the intestine. A possibleoocyte and caudal organ are present lateral to the intestine andon the left side of the trunk.

Ecology: RareGeographical distribution: Known only from Brazil (São

Sebastião Island: Praia Grande [Todaro and Rocha 2004],Fome Beach).

Remarks: There are six described species of Dendrodasys:D. affinis Wilke, 1954, D. gracil is Wilke, 1954,D. rubomarinus Hummon, 2011, D. duplus Lee, Chang andKim, 2014, D. ponticus Valkanov, 1957, and D. pacificusSchmidt, 1974. Two unidentified species are also recognized(Valbonesi and Luporini 1984; Todaro and Rocha 2004). Thesix described species can be distinguished based on the shapeof the anterior margin of the head (crescent or triangular), thepresence, number (one or two pairs), position (dorsal or ven-tral), and shape (knob-like, pyriform, lobiform) of the pestleorgans, the presence and shape (round or triangular) of thehead lobes, and the number of testes (uni- or bi-lateral). Ourspecimens most closely resemble Dendrodasys rubomarinusbecause both have somewhat rounded head lobes, a neck con-striction occurring at the pharyngeal pores, and bilateral testes,but the Brazilian specimens can be distinguished fromD. rubomarinus by the lower number of sensory hairs andpestle organs that are globular in shape and not knob-like.One of the unidentified species collected by Todaro andRocha (2004: Fig 3b) appears similar in morphology to thespecimens we collected. Due to the low number of speciesdescribed for the genus, and their limited spatial distribution,we consider our specimens to belong to the same speciescollected by Todaro and Rocha (2004) that still awaits formaldescription.

Family Dactylopodolidae Strand, 1929Genus Dactylopodola Strand, 1929Dactylopodola todaroi sp. nov.(Figs. 5 and 6)

Material examined

Three individuals, examined by light microscopy (DIC). Afourth specimen was fixed in 95 % ethanol for DNA analysis.

Fig. 4 Dendrodasys aff. rubomarinus, DIC photos: a habitus, b dorsalview of the head region, c dorsal view of the posterior end. co, caudalorgan; ep, epidermal gland; oo, oocyte; po, pestle organ; TbP, posterioradhesive tubes. Scales bars: 40 μm

Fig. 5 Dactylopodola todaroi sp. nov., schematic drawing of the ventralview. Scale bar: 65 μm. e, red ocellar granule; eg, egg; co: caudal organ;in: intestine; ph, pharynx; PhIJ, pharyngeo-intestinal junction; pp,pharyngeal pores; TbA, anterior adhesive tubes; TbL, lateral adhesivetubes, TbP posterior adhesive tubes; te, testes

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Type material

Holotype. Photographs of one specimen (adult), collectedfrom São Sebastião Island (municipality of Ilhabela),Fome Beach, State of São Paulo, Brazil (23°44’27^S;45°16’01^W) at 8–10 m depth, on 17/07/2015, availableat the Museum of Zoology, University of Campinas,Brazil, under accession number ZUEC GCH 26. The358-μm-long adult specimen shown in Fig. 6 is no longerextant (International Code of Zoological Nomenclature,Articles 73.1.1 and 73.1.4).

Paratypes. Photographs of two specimens (adult), col-lected from São Sebastião Island (municipality ofIlhabela), Fome Beach, State of São Paulo, Brazil(23°44’27^S; 45°16’01^W) at 8–10 m depth, on 17/07/2015, available at the Museum of Zoology,University of Campinas, Brazil, under accession num-bers ZUEC GCH 27 and ZUEC GCH 28.

Etymology

The specific epithet honors Dr. M. Antonio Todaro, who hasgreatly contributed to our understanding of marinegastrotrichs in Brazil and around the world.

Autapomorphies

Five TbL arranged as 1+1+3; length of the third and fifthTbL is shorter than the fourth; two neck constrictions thatdemarcate the head and trunk regions.

Diagnosis

Dactylopodolawith a body of 300–358 μm long (n=2). Bodydivided into head, neck, and trunk regions; trunk terminateswith a pair of caudal feet. Two neck constrictions demarcatethe head and trunk regions. Rounded head bears bilateral red

Fig. 6 Dactylopodola todaroi sp.nov., DIC photos: a habitus, bdorsal view of the head region, cdorsal view of the posterior end, dtrunk region showing intestinefilled with diatom and testes, edorsal view of the posterior end, fventral view of the head region, gtrunk region showing the egg andthe intestine filled with diatom.Scales bars: a–e: 50 μm; f, g:45 μm. The asterisks indicate theposition of the ocellar granules ina fixed specimen. e, red ocellargranule; ci, cilia; co, caudal organ;in, intestine; ph, pharynx; pp,pharyngeal pores; TbA, anterioradhesive tubes; TbL, lateraladhesive tubes, TbP, posterioradhesive tubes; te, testes

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ocellar granules (Figs. 5 and 6a, e, f). Terminal mouth leads toa pharynx with pharyngeal pores at U20. Sensory cilia dorsalon head. Four TbA originate from the body surface; two lat-eral tubes are long and two medial tubes are short. TbL ar-ranged as 1+1+3 on each side of the trunk. Caudal pediclesbear seven TbP per side, of unequal length. Locomotory ciliacover the ventral head region and are arranged in two columnsthat extend posteriorly on the trunk. Intestine straight and nar-row; anus at U86. Testes positioned lateral to the intestine andfilled with sperm; caudal organ present; frontal organ notobserved.

Body shape

The three measured animals have total lengths of 300 and358 μm, and maximum trunk widths of 68 and 73 μm, respec-tively. The head has a length (to the first neck constiction atU21) of 60–68 μm (n=2) and a maximal width of 42–60 μm(n = 2). The neck, which begins at U21 (constrictionwidth=33 μm), contains the anterior portion of the midgut.The second neck constriction at U30 has a width of 36 um.The trunk gradually widens from 35 μm to 61 μm and thennarrows posteriorly. At the transition to the caudal pedicles,the body is 27 μm wide (n=2) (Figs. 5 and 6a).

Adhesive tubes

Adhesive tubes arranged in anterior (TbA), lateral (TbL), andposterior series (TbP) (Figs. 5 and 6a). The four TbA arepresent at U11 and point laterally (Figs. 5 and 6a, b): thetwo longest tubes (12 μm each) are positioned at the lateralmargin of the head, while the two shorter tubes (4 μm each)are more medial in position. All TbA originate directly fromthe surface of the head. Five TbL per side are present along thelateral margins of the trunk (TbL 1–5) and distributed as 1+1+3; the two anterior-most tubes (TbL 1–2 at U33 and U46)have lengths of 12 and 18 μm, respectively. Around U69,there is a group of three adhesive tubes (TbL 3–5; Figs. 5and 6a). The anterior and posterior tubes (TbL 3 and TbL 5)are 8–9 μm long, while the medial is 13 μm long (TbL 4).Posteriorly, there is a pair of caudal pedicles. On the edge ofeach pedicle are seven adhesive tubes (TbP) (Figs. 5 and 6a,c): the most lateral tube is 6 μm long, the middle four tubes are8 μm long, and the medial tubes are 7 and 6 μm long.

Cilia

Cilia are distributed along the lateral edges of the headup to the region of the TbA. Locomotory cilia aresparsely distributed over the ventral head (from U9 toU27) and then split into two columns that extend downthe intestinal region (U27-U90).

Digestive tract

The terminal mouth is 8 μm wide. The pharynx isstraight, 54 μm long, and 17 μm wide. It ends atU21. Pharyngeal pores are present slightly anterior tothe pharyngeo-intestinal junction approximately at U20.The ventral anus is present at U86. Several diatoms arepresent in the midgut.

Reproductive organs

A pair of testes is situated lateral to the intestine and extendfrom approximately U38 to U60 (Figs. 5 and 6d). A caudalorgan, approximately 40 μm long, is position at U68-U73.The organ, which was difficult to observe, appeared to beventral in position and positioned between the cross-striatedlongitudinal muscles on either side of the body. The organ alsoappeared to be Bcharged^ with spermatozoa; no muscles wereevident around the organ. A frontal organ (seminal receptacle)was not observed. One mature egg dorsally in a more medialposition (Fig 6g).

Taxonomic remarks

Currently, the genus Dactylopodola includes ten marinespecies (reviewed in von und zu Gilsa et al. 2014;Todaro et al. 2015). Among them, only D. baltica(Remane, 1926), D. roscovita (Swedmark, 1967), andD. nadine Todaro et al. 2015 have anterior eye spots(red). Although Dactylopodola todaroi sp. nov. is quitesimilar to Dactylopodola baltica as described by Remane(1926), it differs in the number and sizes of TbA (4 TbAwith two different sizes vs. 2–3 TbA with the same size)and in the number of TbL (5 TbL [1+1+3] vs. 6 TbL[1 + 1+1+ 3]). The number and sizes of TbA and thenumber of TbL of Dactylopodola todaroi sp. nov. arethe same as those reported by Todaro and Rocha (2004)to D. baltica collected from Brazil, but both differ in thesize of TbL 3–5 (3 and 5 smaller than 4 vs. 3–5 progres-sively shorter) and the number of TbP (7 TbP vs. 4 TbP).Until new samples become available, we prefer to keepDactylopodola todaroi sp. nov. and D. baltica sensuTodaro and Rocha (2004) as different species. However,we did not dismiss the possibility that both taxa are con-specific. The new species can be distinguished from itstwo remaining Bred-eyed^ congeners by the followingcombination of characters: TbA (4 in D. todaroi n. sp.vs. 2 in D. roscovita and 3 in D. nadine); TbL (5 [1+1+3] in D. todaroi n. sp. vs. 9 [1+1+7] in D. roscovitaand 4 [1+1+2] in D. nadine) and TbP (7 in D. todaroin. sp. vs. 12 in D. roscovita and 4 in D. nadine).

Mar Biodiv (2017) 47:451–459 457

Discussion

Based on the findings of this study and previous records byseveral other researchers (Forneris 1985; Todaro and Rocha2004, 2005; Todaro 2012, 2013; Araujo et al. 2014; Hochberg2014; Garraffoni et al. 2016), we conclude that the Brazilianmarine gastrotrich fauna is composed of 22 formally describedspecies (four endemic and 18 reported in other parts of theworld) and another 42 undescribed species of bothMacrodasyida and Chaetonotida. It is important to highlightin the current study that we are only reporting a portion (6) ofour findings, while at least another 12 species from eight gen-era have been collected and will be described if more datahave been collected: Chaetonotida: Aspidiophorus sp.,Chaetonotus spp., Halichaetonotus sp., Xenotrichula sp.;Macrodasyida: Paradasys sp., Ptychostomella spp.,Tetranchyroderma spp., and Thaumastoderma sp. Thereason for this partial report is due to incomplete data onthese 12 species, which require additional collections andobservations for a more accurate assessment. So, with theaddition of these currently undescribed species, thereappears to be at least 18 species of Gastrotricha from FomeBeach, which makes it the beach with the highest gastrotrichdiversity on the north coast of São Paulo State. Comparingwith the estimative done by Todaro and Rocha (2004) on SaoSebastião Island, the highest species count per locality in theirstudy was Casthelianos Beach with 14 species, followed byGrande Beach with 11, Jabaquara Beach with four, Belugawith three, and Siriuba, Vila, and Velasquez with one each.The reasons for such high diversity require explanation andcan probably be ascertained after (1) additional meiofaunalgroups have been explored in more detail at the taxonomiclevel to determine if they too have unusually high diversityand (2) the physico-chemical parameters of the beach havebeen well characterized. Still, if we include the present biodi-versity findings from Fome Beach with findings from othersampling locations on the north coast of São Paulo State aswell as those of Rio de Janeiro State and Bahia State (Todaroand Rocha 2004, 2005; Araujo et al. 2014; Hochberg 2014;Garraffoni et al. 2016), we conservatively estimate that thereare at least 60 species in Brazil. To date, gastrotrich biodiver-sity has only been described from a total of 201 km ofBrazilian coastline, which makes up less than 3 % of the7490 km of coastline available for sampling. Thus, we expectour knowledge of gastrotrich biodiversity in Brazil to increasesubstantially as new sites are sampled and new specimens areexamined using both classical methods (DIC and SEM) andmolecular techniques. Considering the recent gain of gastro-trich descriptions from the Caribbean Sea (Hochberg 2008,2010; Hummon 2010; Kieneke et al. 2013a, b, 2015;Hochberg and Atherton 2010, 2011; Atherton and Hochberg2012; Todaro and Leasi 2013; von und zu Gilsa et al. 2014;Hochberg et al. 2014; Kånneby et al. 2014; Todaro et al. 2014),

we predict that increased sampling of Brazilian beaches, partic-ularly in the north, will extend the biogeographic range of sev-eral Caribbean species southward or reveal cases of endemismin the Tropical Northwestern Atlantic.

Acknowledgments This work was supported by grants from FundaçãodeAmparo à Pesquisa do Estado de São Paulo – FAPESP (2011/50317-5;2012/08581-0; 2014/23856-0), Conselho de Pesquisa e Desenvolvimento– CNPq (306558/2010-6; 478825/2013-7), and Fundo de Apoio aoEnsino, à Pesquisa e Extensão – FAEPEX (2031/2015). We would liketo thank Felipe Pi for the assistance with the scuba diving; Dr. AlexanderKieneke and an anonymous reviewer are also acknowledged for offeringsuggestions that greatly improved the paper.

References

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