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Three New Brazilian Species of the Land Planarian Choeradoplana (Platyhelminthes: Tricladida: Geoplaninae), and an Emendation Of

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Three New Brazilian Species of the Land Planarian Choeradoplana (Platyhelminthes: Tricladida: Geoplaninae), and an Emendation Of Journal of Natural History Vol. 46, Nos. 19–20, May 2012, 1153–1177 Three new Brazilian species of the land planarian Choeradoplana (Platyhelminthes: Tricladida: Geoplaninae), and an emendation of the genus Fernando Carbayoa* and Eudóxia Maria Froehlichb aFernando Carbayo: Laboratório de Ecologia e Evolução, Escola de Artes, Ciências e Humanidades, Universidade de São Paulo – USP, Av. Arlindo Bettio, 1000, CEP 03828000, São Paulo, SP, Brazil; bEudóxia Maria Froehlich: Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, 101, 05508090, São Paulo, Brazil (Received 21 December 2011; final version received 11 January 2012; printed 5 April 2012) Three new Brazilian species of the neotropical land planarian genus Choeradoplana (Platyhelminthes: Tricladida: Continenticola: Geoplaninae) are described, making a total of nine species within the genus. All the new species share unique derived characters typical of the genus. Two of the new species exhibit important features representing morphological variations that were previously unknown for the genus: the dorsal cutaneous longitudinal muscle layer, as well as the ventral one, par- tially sunken into the parenchyma in one species, and the common ovovitelline duct approaching the copulatory apparatus ventrally in the other. As a consequence of these morphological variations, an emendation of the genus is proposed. Keywords: Flatworms; Turbellaria; Continenticola; neotropical; taxonomy; Brazil Introduction The subfamily Geoplaninae is restricted to the Neotropical region, including, with only a few exceptions, all the native species of the region. Most of the species are described from Brazil, although we are still far from attaining a comprehensive estimate of Brazil’s terrestrial flatworm richness (Carbayo and Froehlich 2008). Choeradoplana Graff, 1896 is one of the 17 geoplaninid genera presently recog- nized. The genus was established by Graff (1896) and thereafter reviewed by Froehlich (1955), who restudied in detail Choeradoplana iheringi Graff, 1899 and emended some important points of Graff’s description, such as the true nature (i.e. cutaneous sunken fibres) of the supposedly parenchymal longitudinal muscle fibres. He indicated C. iheringi as the type species, and diagnosed the genus “as Geoplana, but with two glandular cushions at anterior end. Part of the ventral subepidermal longitudinal muscle bundles internal to submuscular nerve plexus. These bundles originate the retractor of the glandulo-muscular organ (glandular cushions)”. In their index, Ogren and Kawakatsu (1990) transcribed Froehlich’s redefiniton of the genus adding to it, as a note: “female canal enters genital antrum dorsally”. They also commented on the presence/absence of the penial papilla in the different species. *Corresponding author. Email: [email protected] ISSN 0022-2933 print/ISSN 1464-5262 online © 2012 Taylor & Francis http://dx.doi.org/10.1080/00222933.2012.657699 http://www.tandfonline.com 1154 F. Carbayo and E.M. Froehlich Later Carbayo and Leal-Zanchet (2003) added to Froehlich’s diagnosis some other important morphological aspects that Froehlich had noticed but did not include in the diagnosis. So, the present diagnosis is as follows: Geoplanidae of elongated body; cephalic region with a glandulomuscular organ composed of two cushions; eyes and sensory pits absent in the anterior apex; creeping sole broad, more than a third of the body width; strong cutaneous longitudinal musculature, ventrally with a portion internal to the subcutaneous nerve plexus all throughout the body and, in cephalic region, entirely sunk into the parenchyma constituting the glandulomuscular retrac tor organ; some dorsal longitudinal cutaneous muscle fibres oriented dorsoventrally in the cephalic region; true parenchymal longitudinal musculature, weak or absent; bodies of rhabditogen cells located between the retractor muscle and the ventral epidermis of the cephalic region; dorsal testes; sensory papillae absent; copulatory apparatus without adenodactyls. As a contribution to the knowledge of the turbellarian fauna inhabiting the Brazilian Atlantic Rainforest, three new species of the genus Choeradoplana are herein described. Materials and methods The specimens studied were collected manually, mainly during the night, in Atlantic Rainforest areas in the State of São Paulo, Brazil, during 20082009. The specimens were photographed, and then killed with boiling water. A small piece of tissue was immediately cut off, measured and preserved in absolute ethanol for DNA extraction; it is standard practice in our laboratory to measure unidentified specimens or those belonging to species that are poorly known. The worms were fixed with 10% forma lin and subsequently stored in 80% ethanol. The anterior region of the body (from extremity up to the prepharyngeal region) was cut into a number of pieces: cephalic region, anterior region 1, anterior region 2, etc., besides pieces with the pharynx and the copulatory apparatus. The pieces were embedded in Paraplast, sectioned at 7-µm thickness, and stained with Mallory/Cason trichrome stain (Romeis 1989). Drawings were prepared using a camera lucida. The following abbreviations are used throughout the text and figures: a, anterior end of the body; cm, common muscular coat; CMI, cutaneous musculature thick ness relative to body height at the prepharyngeal region; co, common glandular ovovitelline duct; cr, creeping sole; dd, dorsodiagonal parenchymal muscles; de, dor- sal epidermis; dv, dorsoventral muscles; e, eye; fa, female genital atrium; g, gonopore; i, intestine; ls, normal longitudinal cutaneous muscles; m, muscles; ma, male geni tal atrium; mo, mouth; MZUSP, Museu de Zoologia da Universidade de São Paulo; ov, ovovitelline duct; ph, pharyngeal pocket; pv, prostatic vesicle; sb, subintestinal transverse muscles; sc, subcutaneous nerve net; sd, sperm duct; sg, shell glands; sk, sunken longitudinal cutaneous muscles; sp, supraintestinal transverse muscles; t, testis; v, vagina; vi, vitellaria; vn, ventral nerve plate. Systematic account Order TRICLADIDA Lang, 1884 Suborder CONTINENTICOLA Carranza et al., 1998 Family GEOPLANIDAE Stimpson, 1857 Journal of Natural History 1155 Subfamily GEOPLANINAE Stimpson, 1857 Choeradoplana von Graff, 1896 Choeradoplana bocaina sp. nov. (Figures 1–8; Tables 1, 2) Type material Holotype. MZUSP PL 997: Parque Nacional da Serra da Bocaina, São José do Barreiro/SP, Brazil, c.22◦45′08′′ S, 44◦37′19′′ W, F. Carbayo et al. col., 8 September 2008. Cephalic region: sagittal sections on 10 slides; prepharyngeal region: transverse sections on three slides; pharynx: sagittal sections on 13 slides; copulatory apparatus: sagittal sections on six slides. Paratypes. MZUSP PL 998: ibid., 22◦45′08′′ S, 44◦37′19′′ W, F. Carbayo et al. col., 7 September 2008. Anterior region 2: horizontal sections on seven slides; anterior region 3: transverse sections on four slides; pharynx plus copulatory apparatus: sagittal sections on 14 slides; MZUSP PL 999: ibid., 22◦45′08′′ S, 44◦37’19′′ W, F. Carbayo et al., col., 10 February 2008. Cephalic region: transverse sections on five slides; pharynx and copulatory apparatus: sagittal sections on five slides. Type locality Parque Nacional da Serra da Bocaina, São José do Barreiro/SP, Brazil, covered with primary Atlantic Rainforest. Etymology The specific epithet refers to the TupiGuarani (Indigenous Brazilian tribe) name of the National Park where the type material was collected. Diagnosis Choeradoplana species with small dark pigmented spots dispersed on the dorsum; pharynx bellshaped; extrabulbar portion of prostatic vesicle proximally twoforked, dishshaped distally; posterior section of the female atrium narrow. Description The largest live specimen (paratype MZUSP PL 998) measures up to 48 mm in length and 4 mm in width (see Table 1 for size of fixed specimens). The body is slender (Figure 1). Dorsum highly convex in crosssection; body margins rounded; ventral side slightly convex. The cephalic region, which bears the glandular cushions typical of the genus, is kept rolled up in the living worm when creeping and resting. The posterior end is pointed. The creeping sole is as wide as 74% of body width at the prepharyngeal region. Mouth 19.5 mm from anterior end, gonopore at 23 mm (holotype). Ground colour of dorsum is yellowishbrown; ventral side pale brown in the cephalic region, cream in rest of the body. Dorsum of holotype and of paratype MZUSP PL 999 with dark brownish spots of different size, more densely concen trated paramedianly, so delineating an irregular longitudinal stripe on each side. In the 1156 F. Carbayo and E.M. Froehlich Table 1. Choeradoplana bocaina sp. nov. – body measurements of fixed specimens (in mm and % of body measurements). Specimen Holotype Paratype MZUSP PL 999 Paratype MZUSP PL 998 Length (mm) 36 24 32 Width (mm) 3 2 3 Mouth to anterior tip 19.5 14.5 18.0 Relative M : L 54.2% 60.4% 53.1% Gonopore to anterior 23 19 23 tip Relative G : L 63.9% 79.2% 72.0% Creeping sole: width 73.8% ? 73.6% G, gonopore; L, length; M, mouth; W, width. Figure 1. Choeradoplana bocaina sp. nov. Holotype creeping in dorsal and lateral (inset) view. About 36 mm in length. paratype MZUSP PL 998 the dorsal region is almost completely covered by a dense web of brown pigment. There is medianly a darker longitudinal stripe of brownish pigment. Eyes formed by one pigmented cup, 40–45 µm in diameter. No clear halos around them. Eyes absent in the first 3.3 mm of body (paratype MZUSP PL 999). Posteriorly
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