Morphological Description of Cyrtopleura Costata (Bivalvia: Pholadidae) from Southern Brazil
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ARTICLE Morphological description of Cyrtopleura costata (Bivalvia: Pholadidae) from southern Brazil Nicole Stakowian¹ & Luiz Ricardo L. Simone² ¹ Universidade Federal do Paraná (UFPR), Setor de Ciências Biológicas, Departamento de Zoologia (DZOO), Programa de Pós-Graduação em Zoologia. Curitiba, PR, Brasil. ORCID: http://orcid.org/0000-0002-3031-783X. E-mail: [email protected] ² Universidade de São Paulo (USP), Museu de Zoologia (MZUSP). São Paulo, SP, Brasil. ORCID: http://orcid.org/0000-0002-1397-9823. E-mail: [email protected] Abstract. The aim of the study is to describe in detail, for the first time, the internal and external anatomy of Cyrtopleura costata, which displays ellipsoid and elongated valves with beige periostracum, the anterior adductor muscle unites the valves in the pre- umbonal region, with abduction capacity in its dorsal half, sparing the ligament. Two accessory valves are identified: the mesoplax (calcified) located in the umbonal region; and the protoplax (corneus) above the anterior adductor muscle. Internally there is a pair of well-developed apophysis that supports the labial palps and the pedal muscles, and support part of the gills. The posterior half of mantle ventral edge is fused and richly muscular, working as auxiliary adductor muscle. The siphons are completely united with each other, the incurrent being larger than the excurrent. The foot is small (about ⅛ the size of the animal). The kidneys extend laterally on the dorsal surface, solid, presenting a brown/reddish color. The style sac is well developed and entirely detached from the adjacent intestine. The intestine has numerous loops and curves within the visceral mass. The fecal pellets are coin-shaped. The present study certainly may be used as comparative scenario for specimens from other regions of the species range. Keywords. Taxonomy; Bivalve; Morphology; Myida; Pholadoidea. INTRODUCTION Pholadidae Lamarck, 1809 is a family of bur- rowing bivalves present into different hard and The bivalves are the second largest class of the soft ground substrates, such as clay, limestone, phylum Mollusca in diversity and, accordingly, of rocks and mud (Turner, 1954). Several species high economic and ecological importance (Bieler have shown economic importance as an import- & Mikkelsen, 2006; Bieler et al., 2013; Gomes-dos- ant source of protein (Ribas, 2014) in Europe (Arias Santos et al., 2019). Bivalves constitute an ideal & Richter, 2012) and in the Philippines (Bin Ramli group to answer biological questions around & Yusop, 2016). The group has a set of features morphological adaptations to dig, drill, swim and that reflects the punch/burrowing habit, such attach on several substrates (Bieler, 2006; Bieler as accessory valves, apophyses, the presence of et al., 2013). spines along the valves surface, hinge ligament Although the available knowledge about bi- reduction (or loss) and anterior adductor muscle valves is relatively extensive and dates back to entered dorsally, which allows the valves to move several centuries, there are still many gaps to on a dorsoventral axis. This arrangement is exclu- be filled. Moreover, many species are still to be sive of the pholadid bivalves (Ansell & Nair, 1969; described and identified, as well as the current Ito, 2005; Monari, 2009; Jeon et al., 2012). known species must be better defined (Bieler Cyrtopleura costata (Linnaeus, 1758) is a bur- et al., 2013). The magnitude of megadiverse rowing pholadid, inhabitant of the infauna of mud- groups, such as Mollusca, hampers large-scale dy regions of the intertidal from the east coast of research, and in classical morphological stud- Canada to southern Brazil (Rios, 2009; Velásquez, ies, it is usual a functional single organ anatomy 2017; Cullain et al., 2018). The Cyrtopleura galleries characterization. In this framework, morpholog- have a narrow opening and an internal chamber ical generalizations for other groups and speci- that contributes to increased complexity of the mens are common. In addition, there are scant ecosystem, acting as ecosystem engineers (Pinn attention to soft parts anatomy, with studies et al., 2005; Haider et al., 2018). Despite being one focusing only on description of the shell, al- of the most conspicuous species of Pholadidae, though the taxonomic importance of the inter- with easily recognizable large individuals (Turner, nal organs. 1954), little is known about its anatomy, except Pap. Avulsos Zool., 2021; v.61: e20216108 ISSN On-Line: 1807-0205 http://doi.org/10.11606/1807-0205/2021.61.08 ISSN Printed: 0031-1049 http://www.revistas.usp.br/paz ISNI: 0000-0004-0384-1825 http://www.scielo.br/paz Edited by: Marcelo Veronesi Fukuda Received: 04/04/2020 Accepted: 30/11/2020 Published: 29/01/2021 Pap. Avulsos Zool., 2021; v.61: e20216108 Stakowian, N. & Simone, L.R.L.: Morphology of Cyrtopleura costata 2/7 for the macroscopic descriptions and brief note in basic RESULTS anatomy (Dall, 1889; Turner, 1954) and the ciliary focused study of Kellogg (1915). Cyrtopleura costata (Linnaeus, 1758) Studies of descriptive morphology can provide new (Figs. 1‑2) characters for phylogenetic studies, contributing towards the description of new species and for the survey of local Pholas costatus: Linnaeus, 1758: 669; Lamarck, 1818: 445; biodiversity, besides serving as the basis for further re- 1835: 45; Tryon, 1862: 201; Schepman, 1887: 156, 164. searches. Studies describing soft anatomy are scattered Capulus shreevei: Conrad, 1869: 105, pl. 13, fig. 3. for Pholadidae, with few examples (Dall, 1889; Turner, Scobina costata: Bayle, 1880: 242. 1954; Purchon, 1955; Morton, 1973; Wong, 1982; Turner Pholas (Barnea) costatus: Linnaeus, 1759 (sic): Grant & & Santhakumaran, 1989). Thus, this study aimed to de- Gale, 1931: 431. scribe the shell and soft parts morphology of C. costata Cyrtopleura (Scobinopholas) costata: Turner, 1954: 35, specimens from a population located in southern Brazil, pls. 17-18; Altena, 1968: 156, 176; 1971: 78, pl. 6, Paranaguá Bay, Paraná. The intention is, in the future, to figs. 10-11; Pacaud, 1998: 64-65, figs. 21-22. compare with other populations from the extraordinary Cyrtopleura costata: Perry & Schwengel, 1956: 94, pl. 19, wide geographic range of the species. fig. 124; Moore, 1961: 49; Turgeon, 1968: 71, fig. 43; Hoagland, 1983: 44; Spencer & Campbell, 1987: 54, pl. 9, figs. 19-20. MATERIAL AND METHODS Material examined: EUA. Florida, Pierce, Bathtub Beach, The specimens were collected in the Paranaguá MZSP 46850, 1 shell. BRAZIL. Bahia, Ilhéus, Olivença, Bay (State of Paraná, Southern Brazil) (25°43 53,17 S, MZSP 101530, 1 shell; Rio de Janeiro, Ibicuí, Almirante ′ ″ 48°44 63,05 W), then shortly relaxed in menthol for 30 Aratanha, MZSP 144243, 1 shell; São Paulo, Iguape, MZSP ′ ″ minutes, and fixed in formalin 10% for at least 24 hours, 20106, 1 shell; Paraná, Baía de Paranaguá, Comunidade with posterior conservation in ethanol 70%. Dissection de São Miguel, MZSP 143461, 8 specimens. was performed using standard techniques under stereo- microscope (Simone et al., 2015), with animals immersed Distribution: From eastern Canada, USA, México, Cuba, in alcohol. Internal structures were observed by remov- and South America, from Venezuela to southern Brazil ing the integument from the visceral mass and scrap- (Rios, 2009; Velásquez, 2017; Cullain et al., 2018). ing the gonadal tissue. Transverse sections of the labial palp were submitted to histological procedures (adapt- Habitat: sandy to silt-muddy bottom 40 cm depth. ed from Borzone et al., 2001). The drawings were done with the aid of a camera lucida. Additional conchological material from other sites was examined and termed just Description “shell”. The collected and dissected bivalves are treated as “specimens”. The material examined is in the results. Shell (Fig. 1A‑E, G, H): Opaque beige to white, thin, Abbreviations in the figures: aa, auxiliary adduc- with radial ornamentation extending from umbo region tor muscle; am, anterior adductor muscle; an, anus; ao, to ventral edge; more protruded (perceptible) in ante- aorta; ap, apophyses; au, auricle; bv, “blood” vessel; cc, rior and posterior regions (Figs. 1A, C, E). Shape ellip- cerebrovisceral connective; cg, cerebropleural ganglia; soid and elongated with anterior and posterior margin cv, ctenidial (efferent) vessel; dg, digestive gland; es, rounded (Figs. 1A, B). Posterior region narrowing grad- esophagus; esp, excurrent siphon; fg, food grooves; fvm, ually, extending posteriorly to base of siphonal process flap of visceral mass; fm, fusion between left and right (Figs. 1C, D). Anterior edge of valves reflected, anterior to mantle lobes; ft, foot; go, gonad; gp, gonóporo; ic, inner umbo (Fig. 1H); insertion of anterior adductor muscle up- ciliary connection of gill; id, inner demibranch; idm, in- wards, external to valves (Figs. 1D: am). Sockets formed sertion of outer demibranch in mantle; in, intestine; ip, below and posteriorly to reflection (Fig. 1H: sc), where inner hemipalp; is, incurrent siphon; it, inner tentacle; mantle flaps (Fig. 2I) are inserted. Periostracum brown, ki, kidney; mb, mantle border; me, mantle; mi, mantle translucent, well-developed near edges (Fig. 1D: pr). border inner fold; mm, mantle border middle fold; mo, Pallial line apparent, pallial sinus wide and shallow (~ ⅓ mantle border outer fold; mt, metaplax; np, nefróporo; valve length) (Fig. 1B). Apophysis with same thickness of oc, outer tissue connection of gill; od, outer demibranch; valves, long (~ half valve width) and