Volume 53(9):115‑127, 2013 Anatomy of Phyllodina persica (Bivalvia: Tellinidae), and its first occurrence in southeastern Brazilian waters Rodrigo Cesar Marques1,2 Luiz Ricardo L. Simone2 Abstract This study presents a detailed anatomy of a rare Western Atlantic tellin, Phyllodina persica, under a comparative scenario. Some characters are shared with other tellinids such as the large hemipalps compared to gills; gills with outer demibranch with a single lamella absent from the pericardial region; the type‑V stomach associated with the style sac conjoined with the proxi‑ mal intestine, and distal intestine presenting a dorsal and ventral group of loops, separated by the transverse muscle. The stomach presents a laterally enlarged typhlosole, although shallow, without flange in the margins. This feature is not found in other tellinid species. Another noteworthy feature in the stomach is the aperture of both caeca, which are larger than the left pouch aperture, and as wide as the style sac aperture. Furthermore, there is an interesting small process in the anterior hinge, and a pair of oblique protractor muscles placed posteriorly to the anterior foot retractor muscle, being a new type of intrinsic muscle described in bivalves. In addition to anatomy, this study presents the southernmost record of P. persica, expanding its distribution to the southeastern region of Brazil. Key-Words: Tellinidae; Anatomy; Morphology; Phyllodina; Western Atlantic. Introduction Phyllodina Dall 1900 is a tellinid taxon of rare occurrence, characterized by an elongated oval outline, oblique and detached pallial sinus connected to the pallial line by a linear scar and a foliated concentric sculpture (Boss, 1969). It has traditionally been considered as a subgenus of Tellina Linné, 1758. Due to this distinctive characteristics, Phyllodina possibly constitutes a natural group (Boss, 1966, 1969). Recently this taxon, like of most subgenera belonging to Tellina, have been shifted to the genus status (e.g., Turgeon et al., 2009). Four species of Phyllodina are present in the American continents: Phyllodina fluctig‑ era (Dall, 1908) and P. pristiphora (Dall, 1900) from Mexican to Costa Rica Pacific coasts (Keen, 1971), P. persica (Dall & Simpson, 1901) and P. squamifera (Deshayes, 1855) in the Western Atlantic, ranging from North Carolina, USA (Boss, 1969), through Caribbean, and Brazilian Northeastern Coast (Tenório, 1984; Cauquoin, 1967). P. persica, the focus of the present paper, is characterized by a more ellipsoid outline and a sculpture less foliated than P. squamifera and P. pristophora, resembling P. fluctigera of the Eastern Pacific, a possible closely allied species (Boss, 1969). P. persica presents a very rare record, characterized so 1. Programa de Pós-Graduação em Zoologia, Instituto de Biociências da Universidade de São Paulo. E‑mail: marquesrc@yahoo.com.br 2. Museu de Zoologia, Universidade de São Paulo. Caixa Postal 42.494, 04218‑970, São Paulo, SP, Brasil. E‑mail: lrsimone@usp.br 116 Marques, R.C. & Simone, L.R.L.: Anatomy of Phyllodina persica far by few occurrences from Cuba through the Lesser Antilles, and the Colombian Caribean. The present reported occurrence in northern Rio de Janeiro State expands the geographic distribution of the species. All the registers are in depths below 25 m up to 400 m. All of these past registers are constituted only by valves, without soft parts preserved. Recently, in a marine research study carried out by Petrobras S/A, called Habitats Project, a well-preserved specimen was dredged from northern Rio de Janeiro State. Due to paucity of the Phylodina records, as well as the first occurrence of a sample with soft parts, the assessment of the anatomy of P. persica improves considerably the systematics of the taxon, this being a concern of the present paper in a comparative scenario. muscle; pr, foot protractor muscle; ps, septal siphonal muscle; pt, posterior lateral tooth; r, dorsal stomach ridge; r1, minor dorsal stomach ridge; ra, anterior foot retractor muscle; rc, right caecum; rd, radial siphonal muscle re, rectum; rg, ring around style sac aperture; rm, esophageal rim; rp, posterior foot retractor muscle; rs1, superior branch of siphonal muscle; rs2, inferior branch of siphonal muscle; sa, selection area; si, blood sinus; sm, siphonal muscle; sn, siphonal nerve; sp, intestine-style sac septum; ss, style sac; st, stomach; tp, projections around distal siphon tip; tm, middle transverse muscle; tt, siphonal tentacle; ty, typhlosole; vc, ventral channel; ve, ventricle; vg, visceral ganglion; vi, visceral sac; vs, vestibule. Material and Methods Phyllodina persica Dall and Simpson 1901 (Figs. 1‑40) The specimen is fixed in Alcohol 70% and dissected by standard techniques, immersed in alcohol. Pictures were taken with a digital camera mounted on stereomicroscope and later edited in Corel PhotoPaint®. Drawings were done with the aid of a camera lucida and later edited in Corel Draw®. Abbreviations in figures and text: aa, anterior adductor muscle; ab, aortic bulb; am, anterior transverse muscle above stomach; ap, dorsal-posterior appendix; bm, branchial muscle; bn, branchial nerve; an, anterior nerve; at, anterior lateral tooth; au, auricle; bt, bifid cardinal tooth; ci, circular siphonal muscle; cc, cerebral commissure; cg, cerebral ganglion; cm, cruciform muscle; cp, cerebro-pedal connective; cs, cruciform to siphonal muscle; ct, ctenidial axis; cv, cerebro-visceral connective; dd, diverticulum; de, outer demibranch; dh, dorsal hood; di, inner demibranch; es, excurrent siphon; fl, flap over style sac aperture; fn, pedal nerve; ft, foot; gd, gonoduct; gf, gastric flap; go, gonad; gp, gonopore; gs, gastric shield; he, outer hemipalp; hi, inner hemipalp; ig, intestinal food groove im, intestinal transverse muscle; is, incurrent siphon; ki, kidney; lg, ligament; la, lateral extension of typhlosole; lc, left caecum; le, external mantle lamella; li, internal mantle lamella; lm, middle mantle lamella; lo, longitudinal siphonal muscle; lp, left pouch; lt, laminate cardinal tooth; mp, pallial muscle; mo, mouth; ne, nephropore; ny, nymph; oe, esophagus; om, oblique protactor muscle; pa, posterior adductor muscle; pg, pedal ganglion; pl, palp; po, posterior pallial muscle; pp, papilla; pn, posterior nerve; pm, posterior transverse Description Tellina (Arcopagia; Phyllodina) persica: Dall & Simpson 1901: 479 (pl. 55, fig. 1). Tellina (Arcopagia) persica: McLean, 1951: 94. Tellina (Eurytellina) persica: Warmke & Abbott, 1961: 195 (pl. 29, fig. g). Tellina persica: Cauquoin, 1967: 227. Tellina (Phyllodina) persica: Boss, 1969: 255 (pl. 136, fig. 2); Rios, 1975: 241 (pl. 77, fig. 1157); 1994: 271 (pl. 93, fig. 1328); Abbott, 1974 497 (fig. 5646); Díaz & Puyana, 1994: 88 (fig. 219). Type locality: Mayaguez, Puerto Rico. Shell (Figs. 1‑3, 6‑11) General outline elliptic, elongated. Length ~18 mm. Length/height ratio ~1.65. Width/length ratio ~0.4. Umbo positioned on shell half as mall, pointed upturned. Anterior margin rounded. Ventral margin slightly rounded. dorso-posterior margin straight. Posterior margin bounded by angles well developed, rounded. Poorly rostrated. Walls relatively thick. Outer surface opaque white, suffused with yellow close to umbo; inner surface white suffused with red. Ornamentation with commarginal ribs, elevated, laminated, separated by broad sulcus; taller and more irregular in posterior and anterior surface than median; sulcus around six times wider than ribs (Fig. 6). Umbonal cavity deep. Cardinal teeth on infraumbonal region, comprising two teeth in both valves, disposed divergently (Figs. 10‑11); right cardinal posterior tooth bifid (Fig. 7: bt); right cardinal anterior Papéis Avulsos de Zoologia, 53(9), 2013 tooth laminate (Fig. 7); left cardinal posterior tooth laminate (Fig. 8: lt); left cardinal anterior tooth bifid (Fig. 8: bt). Cardinal laminate teeth not reaching cardinal plate margin (Fig. 9). Anterior and posterior lateral teeth present in both valves, sockets present in right valve only; anterior lateral teeth distal, ranging 117 from posterior third to posterior half of dorso-anterior margin, with taller elevation in half, turned dorsally upward (Fig. 10: at); posterior lateral teeth distal, located above posterior adductor scar level, in third posterior of dorso-posterior margin, equivalent to 30% of anterior lateral teeth length (Fig. 10: pt). Figures 1‑11: Shell and anatomic aspects of Phyllodina persica (MZSP 96780): 1, left valve, external view; 2, right valve, internal view; 3, dorsal view; 4, right view, valves and part of mantle removed; 5, detail of right mantle edge, internal view (gray arrow = thin muscles; white arrow = thick muscles); 6, detail of commarginal ornamentation on posterior surface of left valve; 7, right cardinal hinge tooth, inferior view; 8, left cardinal hinge tooth, inferior view; 9, right cardinal tooth and process anterior hinge (white arrow); 10, right hinge; 11, left hinge. Scales bar = 1.5 mm. 118 Marques, R.C. & Simone, L.R.L.: Anatomy of Phyllodina persica Small process on third posterior between cardinal teeth and anterior lateral teeth (Fig. 9). Ligament seated on nymph, brown, ranging from posterior side of umbo to half of dorso-posterior margin (Figs. 10‑11). Nymph wall shorter than lateral teeth, parallel to margin. Muscle scars well visible; both adductor muscle scars of equivalent size, each scar occupying ~10% of valve inner surface. Anterior adductor muscle scars elliptical, dorso-ventrally elongated. Posterior adductor muscle scars rounded. Pallial line parallel and at 1/10 of shell height distant from margin. Pallial sinus antero-posteriorly elongated, detached from pallial line, extending to half of valve; ventral margin straight, raised 45° from pallial line; posterior margins slightly pointed, dorsal margin slightly curved (Fig. 2). Cruciform muscle scar inconspicuous. Muscle system Anterior adductor muscle insertion area about 7% of shell surface area, dorsoventrally elongated, 3 times dorso-ventrally longer than antero-posterior length, divided by protractor muscle insertion into two regions (Figs. 4, 11); ventral region equivalent to 2/3 of anterior adductor insertion area; dorsal region equivalent to 1/3 of anterior adductor insertion area. Posterior adductor muscle rounded in cross section, located more ventrally than anterior adductor muscle; area equivalent to 5% of shell surface (Figs. 4, 11). Pair of anterior pedal retractor muscles covering anterior edge of foot (Fig. 25), bifurcating just below mouth (Fig. 20), extending to insertion area, just above anterior adductor muscle; insertion area wider anteriorly than posteriorly, ~1/3 of anterior adductor insertion area. Pair of posterior pedal retractor muscles running with each other from posterior edge of foot, bifurcating in superior third of shell height (Fig. 25); bifurcation angle ~50° (Fig. 28); insertion area ~40% of posterior adductor insertion area. Pair of pedal protractor muscles originating from posterior region, close to pericardial region, extending to anterior adductor insertion area; distal muscle bundles thin, becoming thicker proximally (Fig. 25); insertion area triangular, separating dorsal and ventral regions of adductor anterior, tapering anteriorly, extending to posterior third of anterior adductor muscle (Fig. 12). Umbonal muscles unidentifiable. Branchial muscle thin, ranging from below pericardial chamber region to infraumbonal cavity, just parallel to ctenidial axis (Fig. 36). Pallial muscles consisting of individual muscle bundles laterally arranged, ranging from base of three mantle folds to insertion area ~10% of shell height (Figs. 5, 19); two types of muscle bundles, thin bundles, equally spaced (Fig. 5: gray arrow) and thick bundles, about three times thickness of thin bundles, ventrally branched, spaced at 4 thin bundles (Fig. 5: white arrow). Siphonal muscle ranging from proximal aperture of siphons (Fig. 29) to lateral median region of shell, spreading in numerous fibers (Fig. 12); insertion area fan-like, antero-posteriorly elongated (Fig. 30); origin divided in two bundles of same thickness (Fig. 29: rs1), connected to proximal excurrent aperture, and rs2, below rs1, connected to proximal incurrent aperture. Siphonal septal muscle originating from each side of posterior adductor insertion area, inserting in proximal opening of excurrent siphon, parallel to siphonal septum (Figs. 4, 12: ps); length ~1/6 of shell size; area of origin spreading in ventral-anterior side of adductor muscle insertion. Siphonal muscle ranging from posterior branch of cruciform muscle to bottom of inhalant siphon (Figs. 4, 12: cs), consisting of few fibers, inconspicuous, of same length and thinner than siphonal septal muscle. Posterior pallial muscles (Fig. 12: po) on posterior side, originating below posterior adductor muscle, inserting in ventral-posterior margin, length ~15% of shell height; origin ~1/10 of posterior adductor muscle insertion area. Pair of oblique protractor muscles (Fig. 25: om) ranging from dorsal margin, above and between insertion area of anterior retractor muscles, descending obliquely, backward, each pair inserting laterally on anterior quarter of protractor muscles, bifurcating; thickness ~2/3 of esophagus diameter. Median transverse muscles present in visceral sac (Fig. 25), surrounding style sac (Fig. 32); insertion area same size of esophageal diameter. Anterior transverse muscle thinner than median transverse muscle, trespassing digestive diverticula, above right and left caeca (Fig. 32: am). Intestinal transverse muscles composed of flattened fibers (Fig. 32: im), separating intestinal loops in two groups. Posterior transverse muscles compound of indivisible set of muscle fibers aligned conjoining with connective tissue; set of muscle fibers, as wall, separating visceral cavity from pericardial region (Figs. 25, 32: pm); ventral portion thicker than dorsal portion; average thickness ~2/3 of median transverse muscle. Cruciform muscles total length equivalent to posterior adductor muscle (Fig. 31); anterior and posterior branches same size, located posterior to horizontal level of posterior adductor muscle; cruciform organ not grooved, closed; insertion area of posterior branch of cruciform muscles rounded, area ~¼ of posterior area of retractor insertion; anterior cruciform muscles insertion area subquadrate, same size as posterior cruciform insertion area (Fig. 12). Papéis Avulsos de Zoologia, 53(9), 2013 Pallial cavity Palps somewhat triangular; each hemipalp about 15% of shell surface area, anterior-posterior 119 length ~1.5 times longer than dorsal-ventral length; length of intersection of hemipalps to 35% of shell length; inner hemipalp ~1.2 times longer than outer hemipalp; folds area occupying 90% of internal inner Figures 12‑24: Phyllodina persica general anatomy: 12, whole right view, mantel partially removed; 13, loop pattern of intestine; 14, distal portion of right gill, semi-schematic transverse section; 15, middle portion of right gill, semi-schematic transverse section; 16, right hemipalps deflected, dashed line representing right gill removed; 17, incurrent siphon tip, partially sectioned; 18, excurrent siphon tip, partially sectioned; 19, transverse section of right mantle edge; 20, frontal view of anterior foot retractor muscle and mouth, with hemipalps partially sectioned (dotted lines representing the remainder of hemipalps); 21, transversal section of excurrent siphon semi-schematic; 22, right cerebral ganglion, lateral view; 23, visceral ganglia; 24, cerebral ganglia, frontal view. Scales bar = 1.5 mm. 120 Marques, R.C. & Simone, L.R.L.: Anatomy of Phyllodina persica hemipalp surface and 75% of internal outer surface areas; palp smooth area occupying dorsal edge and posterior area; palp folds originating from hemipalp intersection spreading toward palp dorsal edge (Fig. 16); folds becoming relatively wider distally. Each fold width ~1/30 of palp length on average (Figs. 16, 20). Gill ~80% times inner hemipalp area (Fig. 20); anterior-posterior axis of gill three times longer than dorso-ventral axis; outer demibranch with one lamella (Fig. 14), present only in posterior quarter of gill, lacking in pericardial region (Figs. 15, 26); inner demibranch with two lamella; inner lamella same size as outer lamella (Figs. 14, 15). Food groove running along ventral edge of inner demibranch (Fig. 15). Association gills-palp of Type III of Stasek; region of under-lapped gill extend to 2/5 of hemipalp intersection (Fig. 16). Mantle edge four times thicker than remaining mantle; thickened mantle edge occupying ~1/10 of shell height in middle region and up to 1/8 in anterior region (Fig. 4). Mantle edge threefold; depth of sulcus between middle and internal fold equivalent to 25% of height of thickened mantle edge; sulcus between external and middle fold equivalent to 1/7 of sulcus depth between middle and internal fold, middle fold with series of small, narrow, same sized, uniformly distributed papillae; papillae height equivalent to 6% of height of thickened mantle edge (Fig. 19); each papilla cylindrical, with tip rounded, separated from others by small space, almost touching each other. Ventral channel (vc) on both internal sides of mantle (Figs. 4, 12), ranging from anterior side and same level of anterior branch of cruciform muscles (Fig. 31) up to middle region (Fig. 4), distant from margin equivalent to 1/8 of shell height; ventral channel wall blunt and curved posteriorly, shortening forward. Foot flattened, ~40% of shell volume when retracted; distal tip pointed (Figs. 4, 25). Incurrent and excurrent siphons totally separate from each other, except for their base. Retracted siphons length equivalent to 1/3 of shell length. Siphons outer surface lacking pigment; inner surface of incurrent siphon smooth (Fig. 17); inner surface of excurrent siphon corrugated (Fig. 18). Incurrent siphonal tip with six little elevations (tp), separated by space equivalent to twice its width (Fig. 17); excurrent tip with six tentacles (tt); each tentacle three times mantle papillae (Fig. 18). Siphonal septum, thin, translucent, membrane-like, ranging from siphon base, between incurrent and excurrent proximal aperture, up to gill distal tip; gill connected to siphonal membrane by cilia. Cruciform membrane connecting dorsal region of cruciform muscles to base of incurrent siphon, thin, transparent. Proximal incurrent siphon diameter equivalent to 3/4 that of adductor posterior muscle, becoming gradually narrower distally, diameter equivalent to 1/6 that of adductor posterior muscle. Proximal excurrent siphon diameter ~1.2 times larger than excurrent siphon, becoming gradually narrower distally, with same diameter as distal incurrent siphon (Fig. 29); incurrent siphonal wall thickness ~22% of incurrent siphon total diameter; excurrent siphonal wall thickness ~20% of excurrent siphon total diameter. Radial, longitudinal and circular muscles lying in both siphonal walls; 28 radial muscles in siphon wall, spaced by interval 3 times their width. Three sets of longitudinal and three sets of circular siphonal wall muscles intercalate each one (Fig. 21); outer circular siphonal muscles ~30% of siphonal wall thickness on both siphons; outer longitudinal siphonal muscle ~30% on both siphons; middle and inner circular siphonal muscles thinner, ~10% of outer circular siphonal muscle; middle longitudinal muscle equivalent to 15% of siphonal wall thickness; inner longitudinal with same thickness of middle longitudinal muscle layer. Six longitudinal nerves present in inner longitudinal muscle layer, equally spaced (Fig. 21). Circulatory and excretory systems Pericardial structures occupying 25% of total visceral volume (Fig. 25). Ventricle thick, surrounding rectum, occupying 1/6 of pericardial chamber. Pair of auricles connected to middle third of gill. Auricles inserted in middle-anterior region of ventricle. Aortic bulb translucent, surrounding rectum, as intumescence (Figs. 26, 36), equivalent to 1/10 of ventricle size, located in third posterior of pericardial chamber length (Fig. 36). Kidney yellowish, solid, with fusiform lobes on both sides of pericardial chamber, just behind outer demibranch (Figs. 25, 36). Each lobe equivalent to 1/8 of pericardial chamber; anterior-posterior length two times dorso-ventral length. Nephropores as slits, 1/10 of kidney antero-posterior length, located just below ctenidial axis (Fig. 36). Digestive system Palps as described above: Mouth conic, with relatively thick lips, lacking folds (Fig. 20). Esophagus somewhat cylindrical, length ~75% of stomach length; thickness of ~25% of style sac thickness (Fig. 32). Inner surface folded only in region connected with stomach (esophageal rim – Fig. 37); remaining inner surface smooth. Stomach type V of Purchon, with style sac and intestine conjoined. Stomach oval, somewhat flattened ventrally, weakly positioned to Papéis Avulsos de Zoologia, 53(9), 2013 121 Figure 25: Phyllodina persica; topology of visceral structures, pericardial region, main muscle system and central nervous system. Asterisk indicating insertion area of posterior pallial muscles. Scales bar = 1.5 mm. Figures 26‑31: Phyllodina persica anatomy. 26, Pericardial region sectioned and posterior loops of intestine visible by transparency, with right lobe of kidney withdrawn (asterisk); 27, detail of anterior region with foot protractor muscle withdrawn, right view; 28, posterior foot retractor muscles, posterior view (dotted line representing the rectum); 29, detail of base of siphons showing branches of right siphonal muscle; 30, right siphonal muscle; 31, cruciform muscle and ventral channel, dorsal view. Scales bar = 1.5 mm. 122 Marques, R.C. & Simone, L.R.L.: Anatomy of Phyllodina persica right, few wider ventrally than dorsally; length 1.2 times of height. Ducts to digestive diverticula running from right cecum, spreading ventrally and anteriorly; digestive diverticula from left cecum spreading ventrally (Fig. 32). Digestive diverticula occupying middle and dorsal portions of visceral sac. Dorsal hood Figures 32‑39: Phyllodina persica anatomy.32, digestive system and associated muscles, right view; 33, section of intestine, median region; 34, section of style sac, distal portion; 35, section of style sac, proximal portion; 36, pericardial region and adjacent structures; 37, stomach sectioned on right side, dorsolateral view; 38, detail of dorsal hood aperture; 39, detail of right and left caeca and left pouch, with gastric fold deflected. Papéis Avulsos de Zoologia, 53(9), 2013 externally extruding and turned to left; internal surface area ~20% of stomach internal surface. Dorsal appendix (ap) small, balloon-like, smooth internally. Inner gastric surface smooth, except for sorting area between dorsal ridges (r, r1); sorting area weakly corrugated. Ridges (r, r1) equally sized, ranging from dorsal hood (Fig. 39), passing over dorsal surface of stomach, diverging from each another, up to right side (Fig. 37). Vestibule positioned in front of esophageal aperture, smooth, weakly protruded posteriorly, occupying ~15% of inner stomach surface. Gastric flap above left pouch, thick, flattened, with dorsal surface ~10% of stomach floor surface; divided into two lobes; anterior lobe equivalent to 90% of gastric flap; posterior lobe as small projection protruding laterally, equivalent to 10% of gastric flap volume. Major typhlosole as single structure, not differentiated, spreading over anterior floor of stomach, extending to both caeca (Figs. 37); left side of single typhlosole protruding (Fig. 37: la) laterally as a ridge in median region, gradually becoming faded; small flap emerging laterally, above anterior side of style sac aperture (Fig. 37: fl); intestinal groove of typhlosole present just in intestine and style sac (Figs. 33‑35), becoming shallow toward proximal portion of style sac and absent in stomach cavity. Both caeca placed between style sac aperture and vestibule, below esophagus level. Left and right caeca same size (Fig. 39); caeca aperture ~80% of style sac aperture. Left pouch protruding downward; floor plicated, with groove running parallel to left pouch axis; aperture antero-posterior elongated, with same width of base as gastric flap. Style sac aperture outflanked by elevated ring (Fig. 37: rg), somewhat thick, with tips rounded, fusing anteriorly with typhlosole. Style not visible. Gastric shield chitinous, transparent, spreading over left-dorsal, left and left-ventral internal surface of stomach; small portion spread into proximal style sac, other portion entering into left pouch and dorsal hood (Figs. 37‑39). Style sac straight, parallel to dorsal ventral axis (Figs. 25, 32); dorso-ventral length equivalent to 28% of shell height. Width equivalent to 40% of stomach length. Intestine becoming free from style sac terminally, performing several loops backward (Fig. 32); loops divided in two groups, dorsal and ventral (Fig. 13), separated by intestinal transverse muscles (im); 11 loops in ventral group; 8 loops in dorsal group; intestine width ~20% of style sac width, except for 4 first loops in dorsal group, where width equivalent to half of style sac width (Figs. 26, 32). Rectum passing through ventricle and aortic bulb in pericardium chamber (Fig. 36), running backward, between pair of posterior retractor muscles (Fig. 28) and around posterior adductor 123 muscle, turning down, ending in anus; anteriorly pointed; width equal to proximal portion of intestine. Anus simple located on ventral surface of posterior adductor muscle, at base of excurrent siphon. Genital system Gonads and ducts filling middle and ventral portion of visceral sac. Pair of genital ducts receiving branches from several gonadal acina along their length in posterior region of visceral sac. Genital pore simple, oval, located above and close to nephropore (Fig. 36). Nervous system Pair of cerebral ganglia oval (Fig. 27), posterior region slightly wider than anterior region and slightly dorso-ventrally flattened (Figs. 22, 24), located just to both protractor muscle insertions, almost mouth level (Fig. 20), equivalent to same size of esophagus diameter; cerebral commissure long, ~1.2 times wider than cerebrovisceral connective, almost as long as adductor muscle width, passing above mouth (Fig. 24). Pallial anterior nerve wide, ventral-anterior, obliquely directed, anteriorly branching in several nerves (Figs. 22, 27). Pair of pedal ganglia fused, composing a single structure, elliptical; equivalent to 75% of pair of visceral ganglia volume; located at same level of distal portion of style sac (Fig. 25). Pair of pedal nerves spreading downward from pedal ganglia. Pair of visceral ganglia partially fused, located between posterior retractor muscles and posterior adductor muscle, above level of latter muscle (Fig. 36); volume of each visceral ganglion approximately twice that of cerebral ganglion. Two lobes in each visceral ganglion; posterior lobe, rounded, equivalent to 40% of each ganglion volume; anterior lobe, rounded, slightly antero-posterior elongated, equivalent to 60% of each ganglion volume (Fig. 23). Cerebrovisceral connective about twice width of cerebropedal connective, running parallel to protractor muscle (Fig. 25). Pair of branchial nerve thin, emerging from ventral side of visceral ganglia; pair of posterior adductor nerve emerging from dorsal side of posterior lobe of visceral ganglia towards dorsal portion of posterior adductor muscle, just below posterior retractor muscle. Pair of siphonal nerves emerging from thickening of ventralposterior portion of visceral ganglia; thickness about twice posterior adductor nerve width (Fig. 23); no pallial or siphonal ganglia visible. Distribution: Cuba to Rio de Janeiro State (Brazil). 124 Marques, R.C. & Simone, L.R.L.: Anatomy of Phyllodina persica Habitat: sandy to silt-muddy bottom 25‑400 m depth. Material examined: BRAZIL. Rio de Janeiro; 5 km off Paraíba do Sul river mounth, ~21°36’S, 40°55’W, MZSP 96780, 1 specimen (Petrobras Habitats, sta. 27 RPS/FOZ‑34; 2009) Discussion One of the unique features distinguishing Phyl‑ lodina from other tellinids is the strong foliated concentric ornamentation. However, P. persica presents a less developed foliation, as well as an additional distinction of the oblique scales, as yet stated (Dall, 1900; Boss, 1969). Many of the internal anatomical features of P. persica are similar to others tellinids, such as a dorso-ventral elongated anterior adductor muscle, a long pair of siphons associated to a broad siphonal muscle extending, at least, to the half shell, a large palp with same size (or larger) than the gills, and a broad gastric shield. These general features have been described as representatives of Macominae and Tellininae (Yonge, 1949; Owen, 1953; Franc; 1960; Bright & Ellis, 1989; Gilbert, 1977; Beukema & Meehan, 1985; Ciocco & Barón, 1998; Barón & Ciocco, 1997; Simone & Wilkinson, 2008) that appear to be a set of stable characters within the Tellinidae, which may be considered synapomorphies of this family (Simone & Wilkinson, 2008). P. persica presents two noteworthy characters that seem to be constant through subfamily Tellininae: an outer demibranch with a single lamellae and a pericardial region with no recovering outer demibranch. Macominae, represented by genus Macoma, presents an outer demibranch with both lamellae throughout the gill (Caddy, 1969; Kennedy et al., 1989; Thiriot-Quiévreux & Wolowicz, 1996; Arruda & Domaneschi, 2005; although in Simone & Wilkinson, 2008, the assumed macominae representative, Macomona, has a “Tellininae” gill type). In P. persica the presence of outer demibranchs is just in the third distal region of the gill, whereas in others representatives of Tellininae, the outer demibranch seems to be located at half end of the gill (Simone & Wilkinson, 2008). The hinge in P. persica is like the other representatives of subfamily Tellininae. The lateral anterior hinge teeth are far from the cardinal hinge, a feature shared with other tellininines taxa such as Tellina s.s., Tellinella and Laciolina (Boss, 1969). However, a small process is observed between the cardinal teeth and anterior lateral teeth, present in both valves, a feature that has not been indicated by any previous study. This process could be interpreted as a muscular insertion area, maybe related to the branchial muscle insertion area. It is possible that this process occurs in other tellinids species. But, as this process is small and inconspicuous, it is possible that this structure has not been noted in previous studies. The adductor muscle system in P. persica is essentially similar to other representatives of Tellina in a wide sense, mainly by the anterior adductor being dorso-ventrally more elongated than the posterior adductor, as in Serratina capsoides (Lamarck, 1818), Elpidollina sp. (Simone & Wilkinson, 2008) and Tellina petitiana d’Orbigny, 1846 (Ciocco & Barón, 1997). Moerella nitens (Deshayes, 1846), Cadella se‑ men Hanley, 1844, Pinguitellina pinguis (Hanley, 1844) and Tellinides timorensis (Lamarck, 1818) (Simone & Wilkinson, 2008). All of these present anterior adductor muscle less elongated than P. persica. This character is common in others tellinids, such as Macoma balthica (Linnaeus, 1758) (Caddy, 1969) and M. biota (Arruda & Domaneschi, 2005), species that presents the most dorso-ventrally elongated anterior adductor muscles. The foot protractor muscle in P. persica presents an insertion area penetrating in the superior third of the anterior adductor muscle insertion area, not completely separating this muscle, as in S. capsoides, Moerella nitens, Pinguitellina pinguis, Tellinides timorensis (Simone & Wilkinson, 2008) and Tellina petitiana (Barón & Ciocco, 1997). Despite the same pattern of protractor muscles in taxon previously cited, P. persica presents a smaller anteriorposterior elongated protractor muscle insertion area, with more amorphous outline and with low anterior adductor penetration, very similar to the protractor insertion area shape of T. petitiana (Barón & Ciocco, 1997). This protractor shape and anterior adductor penetration degree is similar to other tellinoid families, such as Donacidae (Purchon, 1963; Narchi, 1978; Ansell, 1981; Passos & Domaneschi, 2004; Simone & Dougherty, 2004), Psamobiidae (Pohlo, 1972), Solecurtidae (Bloomer, 1905), and Semelidae (Domaneschi, 1995; Simone & Guimarães, 2008). Thus this character could be a basal one, as suggested by Simone & Wilkinson (2008).” The pericardial cavity organs have the same spatial arrangement as other tellinoids, with the pair of auricles and ventricle positioned anteriorly (Figs. 27, 36). There is a posterior translucent aortic bulb. The aortic bulb has not been noticed in others tellinids, except for Scro‑ bicularia plana (Purchon, 1955) and Macoma nasuta (Smith & Davis, 1965). However, the fact that this structure was not observed in other tellinids is due to their thin and fragile composition, turning this structure often inconspicuous, which could hinder Papéis Avulsos de Zoologia, 53(9), 2013 the observation of the aortic bulb. The kidney in this P. persica specimen presents lobes smaller than other tellinid representatives. There is a new kind of oblique muscle (Fig. 25: om), not described for other tellinids. This is a pair of muscles that connect to the lateral side of foot protractor fibers, running up and forward, passing over the esophagus, and inserting among anterior retractor muscle bundles, above the anterior portion of the esophagus, termed here as oblique protractor muscle. This muscle is very similar in thickness compared to other visceral sac transverse muscles. This pair of muscles can be assigned to the oblique protractor or a different kind of transverse muscle, such as the oblique protractor muscle, which seems to be “transverse” in dorsal view. However, as the oblique protractor insertion is just among the anterior retractor muscle insertion area, it is hypothesized that this muscle is more similar to the anterior foot retractor muscle, which can be derived from some retractor muscle bundles that turn backward during ontogeny. Despite the similar features, the oblique protractor muscles apparently have a different function from the retractor muscles. As the area of origin of the oblique protractor muscle is from both sides of the foot protractor muscle, it may be inferred that the contraction of the oblique protractor muscle acts more like a protractor muscle than a retractor muscle. This oblique protractor muscle could contract at the same time as the foot protractor muscle. This muscle could be homologous to the “middle-anterior accessory foot retractor muscle” figured in Semele sinensis (Simone & Guimarães, 2008, fig. 15). Nevertheless, these authors did not describe or mention this structure in their study. The stomach of P. persica (Figs. 37‑39) is very similar to the other tellinid type‑V stomachs, with the proximal intestine and style sac conjoined, such that the intestine becomes separated from the style sac subterminally (Fig. 35). However there are some notable differences between the P. persica stomach and other tellinids. The posterodorsal appendix in P. persica is simple, not extended as in Elpidollina sp. (Simone & Wilkinson, 2008) or multiplicated as in Serratina capsoides (Simone & Wilkinson, 2008) and some representatives of Donax (Simone & Dougherty, 2004). The typhlosole presents a conspicuous flange in all known tellinidoids (Purchon, 1987). However, in P. persica the major typhlosole is broad, flattened and spread on the anterior stomach floor and is not flanged. This suggests that the flow towards caeca is unsheltered by the major typhlosole margin, unlike that which occurs with other tellinoidean species. 125 Although this feature appears to be unique among Tellinoidea, this feature is also found in type‑V stomachs as in Cardiidae, Glaucomyidae, Ciprinidae, and Petricolidae (Purchon, 1987). The caeca aperture is also different from other tellinids; it is a quite wide in P. persica, with almost 8/10 of style sac aperture, and larger than the left pouch aperture. This condition seems to be exclusive in P. persica, whereas in polysiringian bivalve stomachs (which include most forms of tellinoidean), all embayment related to diverticula – left pouch, left caecum and right caecum – presents the same dimensions. In addition, the digestive structures of P. persica present a different character on the tip of the siphons aperture – tentacles occurring only on excurrent siphon (Figs. 17‑18). The tentacles in the siphonal aperture of some bivalve groups is correlated to sensory organs, related to control and protection of food capture (Ansell et al., 1999). Thus it is expected that there is an increased complexity in the development of the tentacles in the opening of the inhalant siphon, a condition observed in donacids (Narchi, 1972, 1978). In tellinids there was a reduction of tentacles to six lobes distributed around both siphonal apertures (Levinton, 1972). However, in P. persica, whereas the incurrent siphonal aperture is composed of six lobes, the excurrent siphonal aperture has six well-developed digitiform tentacles. This feature was also observed in Macoma biota (Piffer et al., 2011; per‑ sonal observation). The excurrent tentacles in M. biota is cylindrical and slightly thicker than P. persica. Resumo Este estudo apresenta uma anatomia detalhada de um raro telinídeo do Oeste Atlântico, Phyllodina persica, sob um cenário comparativo. Algumas características são compartilhadas com outros telinídeos como os hemipal‑ pos grandes em comparação com as brânquias; brânquias com demibrânquia exterior apresentando lamela única, ausente na região pericárdica; o estômago do tipo‑V as‑ sociado com saco do estilete unido ao intestino proximal e intestino distal apresentando um grupo dorsal e outro ventral de circunvoluções, separados por músculos trans‑ versais. O estômago apresenta uma tiflossole lateralmente alargada, embora rasa, sem margens pronunciadas. Esta característica não é encontrado em outras espécies de te‑ linídeos. Outra característica notável no estômago é que ambas aberturas dos cecos gástricos são maiores do que a abertura da bolsa gástrica esquerda, sendo tão larga quanto a abertura do saco do estilete. Além disso, há a formação de um interessante processo na charneira an‑ terior, além de um par de músculos protratores oblíquos, 126 Marques, R.C. & Simone, L.R.L.: Anatomy of Phyllodina persica localizados posteriormente ao músculo retrator anterior do pé, sendo um novo tipo de musculatura íntrinseca ob‑ servada em bivalves. Além da anatomia, este estudo apre‑ senta um registro mais austral para P. persica, expandin‑ do sua distribuição para a região sudeste do Brasil. Key-Words: Tellinidae; Anatomy; Morphology; Phyllodina; Western Atlantic. References Abbott, R.T. 1974. American Seashells. 2. ed. New York, Van Nostrand. 663 p. Ansell, A.D. 1981. Functional morphology and feeding of Donax serra Röding and Donax sordidus Hanley (Bivalvia: Donacidae). Journal of Molluscan Studies, 47(1):59‑72. Ansell, A.D.; Harvey, R. & Gunther, C.P. 1999. Recovery from siphon damage in Donax vittatus (da Costa) (Bivalvia: Donacidae). Journal of Molluscan Studies, 65(2):223‑232. Arruda, E.P. & Domaneschi, O. 2005. New species of Macoma (Bivalvia: Tellinoidea: Tellinidae) from southeastern Brazil, and with description of its gross anatomy. Zootaxa, 1012:13‑22. Barón, P.J. & Ciocco, N.F. 1997. Anatomía de la almeja Tellina petitiana d'Orbigny, 1846. I. Organización general, valvas, manto, sifones, aductores, pie y branquias (Bivalvia, Tellinidae). Revista de Biología Marina y Oceanografía, 32(2):95‑110. Barón, P.J. & Ciocco, N.F. 1998. Anatomía de la almeja Tellina petitiana d'Orbigny, 1846. III. Sistema nervioso y gónada (Bivalvia, Tellinidae). Revista de Biología Marina y Oceanografía, 33(1):139‑154. Beukema, J.J. & Meehan, B.W. 1985. Latitudinal variation in linear growth and other shell characteristics of Macoma balthica. Marine Biology, 90(1):27‑33. Bloomer, H.H. 1905. On the anatomy of certain species of Solenidae. Journal of Malacology, 12(1):78‑85. Boss, K.J. 1966. Evolutionary sequence in Phyllodina (Bivalvia: Tellinidae). Report of American Malacological Union, Pacific Division, 1:21‑23. Boss, K.J. 1969. The subfamily Tellininae in the Western Atlantic (Part I). Johnsonia, 4(45‑47):217‑364. Bright, D.A. & Ellis, D.V. 1989. Aspects of histology in Macoma carlottensis (Bivalvia: Tellinidae) and in situ histopathology related to mine-tailings discharge. Marine Biological Association of U.K, 69(2):447‑464. Caddy, J.F. 1969. Development of mantle organs, feeding, and locomotion in postlarval Macoma balthica (L.) (Lamellibranchiata). Canadian Journal Zoology, 47(4):609‑617. Cauquoin, M. 1967. Lamellibranchiate molluscs – Tellinidae, Scrobiculariidae and Donacidae. Annales de l’Institut Oceanographique, 45(2):227‑223. Ciocco, N.F. & Barón, P.J. 1998. Anatomía de la almeja Tellina petitiana d'Orbigny, 1846. II. Sistema digestivo, corazón, riñones, cavidad y glándulas pericárdicas (Bivalvia, Tellinidae). Revista de Biología Marina y Oceanografía, 33(1):73‑87. Dall, W.H. 1900. Synopsis of the family Tellinidae and of the North American species. Proceedings of the United States National Museum, 23(1210):285‑326. Dall, W.H. & Simpson, C.T. 1901. The Mollusca of Porto Rico. Bulletin of the United States Fish Commission, 20(1):351‑52. Díaz, J.M. & Puyana, M. 1994. Moluscos del Caribe Colombiano. Un catálogo Ilustrado. Bogotá, Colciencias, Fundación Natura and Invemar. 367 p. Domaneschi, O. 1995. A comparative study of the functional morphology of Semele purpurascens (Gmelin, 1791) and Semele proficua (Pulteney, 1799) (Bivalvia: Semelidae). Veliger, 38(4):323‑342. Franc, A. 1960. Classe de Bivalvia. In: Grassé, P.P. (ed.) Traite de zoologie; anatomie, systematique, biologie. Paris, Masson et Co. p. 1605‑1802. Gilbert, M.A. 1977. The behavior and functional morphology of deposit feeding in Macoma balthica (Linné, 1758) in New England. Journal of Molluscan Studies, 43(1):18‑27. Keen, A.M. 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. 2. ed. Stanford, Ed. Stanford. 1064 p. Kennedy, V.S.; Lutz, R.A. & Fuller, S.C. 1989. Larval and early postlarval development of Macoma mitchelli Dall (Bivalvia, Tellinidae). Veliger, 32(1):29‑38. Levinton, J.S. 1972. Stability and trophic structure in depositfeeding and suspension feeding communities. American Naturalist, 106(4):472‑486. McLean, R.A. 1951. The Pelecypoda or bivalve mollusks of Porto Rico and the Virgin Islands. Scientific Survey of Porto Rico and the Virgin Islands, 17(1):1‑183. Narchi, W. 1972. On the biology of Iphigenia brasiliensis Lamarck, 1818 (Bivalvia, Donacidae). Proccedings of the Malacological Society of London, 40(2):79‑91. Narchi, W. 1978. Functional anatomy of Donax hanleyanus Philippi 1874 (Donacidae-Bivalvia). Boletim de Zoologia da Universidade de São Paulo, 3(1):121‑142. Owen, G. 1953. The shell in the Lamellibranchia. Quarterly Journal of Microscopical Science, 94(1):57‑70. Passos, F.D. & Domaneschi, O. 2004. Biologia e anatomia funcional de Donax gemmula Morrison (Bivalvia, Donacidae) do litoral de Sao Paulo, Brasil. Revista Brasileira de Zoologia, 21(4):1017‑1032. Piffer, P.R.; Arruda, E.P. & Passos, F.D. 2011. The biology and functional morphology of Macoma biota (Bivalvia: Tellinidae: Macominae). Zoologia, 28(3):321‑333. Pohlo, R.H. 1972. Feeding and associated functional morphology of Sanguinolaria nuttallii (Bivalvia: Tellinacea). Veliger, 14(3):298‑301. Purchon, R.D. 1955. The functional morphology of the rock boring lamellibranch, Petricola pholadiformis (Lam.). Journal of Marine Biological Association, 34(2):257‑278. Purchon, R.D. 1963. A note on the biology of Egeria radiata Lam. (Bivalvia: Donacidae). Proceedings of the Malacological Society of London, 35(2):251‑271. Purchon, R.D. 1987. The stomach in the Bivalvia. Philosophical Transactions of the Royal Society of London, B 316:183‑276. Rios, E.C. 1975. Brazilian marine mollusks iconography. Rio Grande, Fundação Universidade do Rio Grande. 331 p. Rios, E.C. 1994. Seashells of Brazil. 2. ed. Rio Grande, Fundação Universidade do Rio Grande. 368 p. Simone, L.R.L. & Dougherty, J.R. 2004. Anatomy and systematics of northwestern Atlantic Donax (Bivalvia, Veneroidea, Donacidae). Malacologia, 46(2):459‑472. Simone, L.R.L. & Guimarães, C.H. 2008. Comparative anatomical study of two species of Semele from Thailand (Bivalvia: Tellinoidea). The Raffles Bulletin of Zoology, 18(Suppl.): 137‑149. Simone, L.R.L. & Wilkinson, S. 2008. Comparative morphological study of some Tellinidae from Thailand (Bivalvia: Tellinoidea) The Raffles Bulletin of Zoology, 18(Suppl.): 151‑190. Smith, L.S. & Davis, J.C. 1965. Haemodynamics in Tresus nuttallii and certain other bivalves. Journal of Experimental Biology, 43(1):171‑180. Tenório, D.O. 1984. O gênero Tellina Linnaeus, 1758 (Mollusca, Bivalvia) na plataforma continental brasileira. Papéis Avulsos de Zoologia, 53(9), 2013 Trabalhos Oceanográficos, Universidade Federal de Pernambuco, 8(1):7‑138. Thiriot-Quiévreux, C. & Wolowicz, M. 1996. Etude caryologique d'une néoplasie branchiale chez Macoma balthica (Mollusca, Bivalvia). Comptes rendus de l’Académie des sciences. Série 3, Sciences de la vie, 319(10):887‑89. Turgeon, D.D.; Lyons W.G.; Mikkelsen P.; Rosenberg G. & Moretzsohn, F. 2009. Bivalvia (Mollusca) of the Gulf of Mexico. In: Felder, D.L. and D.K. Camp (eds.), Gulf of 127 Mexico-Origins, Waters, and Biota. Biodiversity. Texas A&M Press, College Sattion. p. 711‑744. Warmke, G.L. & Abbott, R.T. 1961. New York, Caribean Seashells, Dover Publications. 348 p. Yonge, C.M. 1949. On the structure and adaptation of the Tellinacea, deposit-feeding Eulamellibranchia. Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences, 234(609):29‑76. Aceito em: 22/03/2013 Impresso em: 30/06/2013 EDITORIAL COMMITTEE Publisher: Museu de Zoologia da Universidade de São Paulo. Avenida Nazaré, 481, Ipiranga, CEP 04263‑000, São Paulo, SP, Brasil. Editor-in-Chief: Carlos José Einicker Lamas, Serviço de Invertebrados, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42.494, CEP 04218‑970, São Paulo, SP, Brasil. E‑mail: editormz@usp.br. Associate Editors: Mário César Cardoso de Pinna (Museu de Zoologia, Universidade de São Paulo, Brasil); Luís Fábio Silveira (Museu de Zoologia, Universidade de São Paulo, Brasil); Marcos Domingos Siqueira Tavares (Museu de Zoologia, Universidade de São Paulo, Brasil); Sérgio Antonio Vanin (Instituto de Biociências, Universidade de São Paulo, Brasil); Hussam El Dine Zaher (Museu de Zoologia, Universidade de São Paulo, Brasil). Editorial Board: Rüdiger Bieler (Field Museum of Natural History, U.S.A.); Walter Antonio Pereira Boeger (Universidade Federal do Paraná, Brasil); Carlos Roberto Ferreira Brandão (Universidade de São Paulo, Brasil); James M. Carpenter (American Museum of Natural History, U.S.A.); Ricardo Macedo Corrêa e Castro (Universidade de São Paulo, Brasil); Mario de Vivo (Universidade de São Paulo, Brasil); Marcos André Raposo Ferreira (Museu Nacional, Rio de Janeiro, Brasil); Darrel R. Frost (American Museum of Natural History, U.S.A.); William R. Heyer (National Museum of Natural History, U.S.A.); Ralph W. Holzenthal (University of Minnesota, U.S.A.); Adriano Brilhante Kury (Museu Nacional, Rio de Janeiro, Brasil); Gerardo Lamas (Museo de Historia Natural “Javier Prado”, Lima, Peru); John G. Maisey (American Museum of Natural History, U.S.A.); Naércio Aquino Menezes (Universidade de São Paulo, Brasil); Christian de Muizon (Muséum National d’Histoire Naturelle, Paris, France); Nelson Papavero (Universidade de São Paulo, Brasil); James L. Patton (University of California, Berkeley, U.S.A.); Richard O. Prum (University of Kansas, U.S.A.); Olivier Rieppel (Field Museum of Natural History, U.S.A.); Miguel Trefaut Urbano Rodrigues (Universidade de São Paulo, Brasil); Randall T. Schuh (American Museum of Natural History, U.S.A.); Ubirajara Ribeiro Martins de Souza (Universidade de São Paulo, Brasil); Richard P. Vari (National Museum of Natural History, U.S.A.). INSTRUCTIONS TO AUTHORS - (April 2007) General Information: Papéis Avulsos de Zoologia (PAZ) and Arquivos de Zoologia (AZ) cover primarily the fields of Zoology, publishing original contributions in systematics, paleontology, evolutionary biology, ontogeny, faunistic studies, and biogeography. Papéis Avulsos de Zoologia and Arquivos de Zoologia also encourage submission of theoretical and empirical studies that explore principles and methods of systematics. All contributions must follow the International Code of Zoological Nomenclature. Relevant specimens should be properly curated and deposited in a recognized public or private, non-profit institution. Tissue samples should be referred to their voucher specimens and all nucleotide sequence data (aligned as well as unaligned) should be submitted to GenBank (www.ncbi.nih.gov/ Genbank) or EMBL (www.ebi.ac.uk). Peer Review: All submissions to Papéis Avulsos de Zoologia and Arquivos de Zoologia are subject to review by at least two referees and the Editor-in-Chief. All authors will be notified of submission date. Authors may suggest potential reviewers. Communications regarding acceptance or rejection of manuscripts are made through electronic correspondence with the first or corresponding author only. Once a manuscript is accepted providing changes suggested by the referees, the author is requested to return a revised version incorporating those changes (or a detailed explanation of why reviewer’s suggestions were not followed) within fifteen days upon receiving the communication by the editor. Proofs: Page-proofs with the revised version will be sent to e‑mail the first or corresponding author. Page-proofs must be returned to the editor, preferentially within 48 hours. Failure to return the proof promptly may be interpreted as approval with no changes and/or may delay publication. Only necessary corrections in proof will be permitted. Once page proof is sent to the author, further alterations and/or significant additions of text are permitted only at the author’s expense or in the form of a brief appendix (note added in proof ). Submission of Manuscripts: Manuscripts should be sent to the SciELO Submission (http:// submission.scielo.br/index.php/paz/login), along with a submission letter explaining the importance and originality of the study. Address and e‑mail of the corresponding author must be always updated since it will be used to send the 50 reprints in titled by the authors. Figures, tables and graphics should not be inserted in the text. Figures and graphics should be sent in separate files with the following formats: “.JPG” and “.TIF” for figures, and “.XLS” and “.CDR” for graphics, with 300 DPI of minimum resolution. Tables should be placed at the end of the manuscript. Manuscripts are considered on the understanding that they have not been published or will not appear elsewhere in substantially the same or abbreviated form. The criteria for acceptance of articles are: quality and relevance of research, clarity of text, and compliance with the guidelines for manuscript preparation. Manuscripts should be written preferentially in English, but texts in Portuguese or Spanish will also be considered. Studies with a broad coverage are encouraged to be submitted in English. All manuscripts should include an abstract and key-words in English and a second abstract and keywords in Portuguese or Spanish. Authors are requested to pay attention to the instructions concerning the preparation of the manuscripts. Close adherence to the guidelines will expedite processing of the manuscript. Manuscript Form: Manuscripts should not exceed 150 pages of double-spaced, justified text, with size 12 and source Times New Roman (except for symbols). Page format should be A4 (21 by 29.7 cm), with 3 cm of margins. The pages of the manuscript should be numbered consecutively. The text should be arranged in the following order: Title Page, Abstracts with Key-Words, Body of Text, Literature Cited, Tables, Appendices, and Figure Captions. Each of these sections should begin on a new page. (1) Title Page: This should include the Title, Short Title, Author(s) Name(s) and Institutions. The title should be concise and, where appropriate, should include mention of families and/or higher taxa. Names of new taxa should not be included in titles. (2) Abstract: All papers should have an abstract in English and another in Portuguese or Spanish. The abstract is of great importance as it may be reproduced elsewhere. It should be in a form intelligible if published alone and should summarize the main facts, ideas, and conclusions of the article. Telegraphic abstracts are strongly discouraged. Include all new taxonomic names for referencing purposes. Abbreviations should be avoided. It should not include references. Abstracts and key-words should not exceed 350 and 5 words, respectively. (3) Body of Text: The main body of the text should include the following sections: Introduction, Material and Methods, Results, Discussion, Conclusion, Acknowledgments, and References at end. Primary headings in the text should be in capital letters, in bold and centered. Secondary headings should be in capital and lower case letters, in bold and centered. Tertiary headings should be in capital and lower case letters, in bold and indented at left. In all the cases the text should begin in the following line. (4) Literature Cited: Citations in the text should be given as: Silva (1998) or Silva (1998:14‑20) or Silva (1998: figs. 1, 2) or Silva (1998a, b) or Silva & Oliveira (1998) or (Silva, 1998) or (Rangel, 1890; Silva & Oliveira, 1998a, b; Adams, 2000) or (Silva, pers. com.) or (Silva et al., 1998), the latter when the paper has three or more authors. The reference need not be cited when authors and date are given only as authority for a taxonomic name. (5) References: The literature cited should be arranged strictly alphabetically and given in the following format: • Journal Article - Author(s). Year. Article title. Journal name, volume: initial page-final page. Names of journals must be spelled out in full. • Books - Author(s). Year. Book title. Publisher, Place. • Chapters of Books - Author(s). Year. Chapter title. In: Author(s) ou Editor(s), Book title. Publisher, Place, volume, initial page-final page. • Dissertations and Theses - Author(s). Year. Dissertation title. (Ph.D. Dissertation). University, Place. • Electronic Publications - Author(s). Year. Title. Available at: <electronic address>. Access in: date. Tables: All tables must be numbered in the same sequence in which they appear in text. Authors are encouraged to indicate where the tables should be placed in the text. They should be comprehensible without reference to the text. Tables should be formatted with vertical (portrait), not horizontal (landscape), rules. In the text, tables should be referred as Table 1, Tables 2 and 4, Tables 2‑6. Use “TABLE” in the table heading. Illustrations: Figures should be numbered consecutively, in the same sequence that they appear in the text. Each illustration of a composite figure should be identified by capital letters and referred in the text as: Fig. 1A, Fig. 1B, for example. When possible, letters should be placed in the left lower corner of each illustration of a composite figure. Hand-written lettering on illustrations is unacceptable. Figures should be mounted in order to minimize blank areas between each illustration. Black and white or color photographs should be digitized in high resolution (300 DPI at least). Use “Fig(s).” for referring to figures in the text, but “FIGURE(S)” in the figure captions and “fig(s).” when referring to figures in another paper. Responsability: Scientific content and opinions expressed in this publication are sole responsibility of the respective authors. Copyrights: The journals Papéis Avulsos de Zoologia and Arquivos de Zoologia are licensed under a Creative Commons Licence (http://creativecommons.org). For other details of manuscript preparation of format, consult the CBE Style Manual, available from the Council of Science Editors (www.councilscienceeditors.org/publications/style). Papéis Avulsos de Zoologia and Arquivos de Zoologia are publications of the Museu de Zoologia da Universidade de São Paulo (www.mz.usp.br). Always consult the Instructions to Authors printed in the last issue or in the electronic home pages: www.scielo.br/paz or www.mz.usp.br/publicacoes. View publication stats