401 lines
45 KiB
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401 lines
45 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.752.24615" ID-GBIF-Dataset="bdaa607e-905b-4112-a1f7-2c435c253074" ID-Pensoft-Pub="1313-2970-752-1" ID-ZBK="780F7546CD364BAEA0A4F9B97D019AB5" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-752-1" ModsDocOrigin="ZooKeys 752" ModsDocTitle="A new species of the land planarian Issoca sheds light on the polyphyletic status of the genus (Platyhelminthes, Tricladida, Geoplaninae)" checkinTime="1524551626357" checkinUser="pensoft" docAuthor="Araujo, Ana Paula Goulart & Carbayo, Fernando" docDate="2018" docId="B68C8E6EECD16AFE551CA13EA0E66459" docLanguage="en" docName="ZooKeys 752: 1-15" docOrigin="ZooKeys 752" docSource="http://dx.doi.org/10.3897/zookeys.752.24615" docTitle="Issoca assanga sp. n." docType="treatment" docVersion="4" lastPageNumber="10" masterDocId="FC1BFFCD854759658C15627D643A407F" masterDocTitle="A new species of the land planarian Issoca sheds light on the polyphyletic status of the genus (Platyhelminthes, Tricladida, Geoplaninae)" masterLastPageNumber="15" masterPageNumber="1" pageNumber="1" updateTime="1643466238084" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A new species of the land planarian Issoca sheds light on the polyphyletic status of the genus (Platyhelminthes, Tricladida, Geoplaninae)</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Araujo, Ana Paula Goulart</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Carbayo, Fernando</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2018</mods:date>
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<mods:detail type="volume">
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<mods:number>752</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>15</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.752.24615</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.752.24615</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-752-1</mods:identifier>
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<mods:identifier type="ZBK">780F7546CD364BAEA0A4F9B97D019AB5</mods:identifier>
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<mods:identifier type="ZooBank">780F7546CD364BAEA0A4F9B97D019AB5</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="143666400" LSID="urn:lsid:plazi:treatment:B68C8E6EECD16AFE551CA13EA0E66459" httpUri="http://treatment.plazi.org/id/B68C8E6EECD16AFE551CA13EA0E66459" lastPageId="10" lastPageNumber="10" pageId="0" pageNumber="1">
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<subSubSection pageId="0" pageNumber="1" type="nomenclature">
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<paragraph pageId="0" pageNumber="1">
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<taxonomicName LSID="http://zoobank.org/FD7F2CF1-B799-4215-B75C-80E16FA90B1D" authority="sp. n." family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca assanga" order="Tricladida" pageId="0" pageNumber="1" phylum="Platyhelminthes" rank="species" species="assanga">Issoca assanga sp. n.</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1" type="synonymy">
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<paragraph pageId="0" pageNumber="1">Synonymy.</paragraph>
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<paragraph pageId="0" pageNumber="1">
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<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca" order="Tricladida" pageId="0" pageNumber="1" phylum="Platyhelminthes" rank="genus">Issoca</taxonomicName>
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sp. 1;
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<bibRefCitation author="Carbayo, F" journalOrPublisher="Zoologica Scripta" pageId="10" pageNumber="11" pagination="508 - 528" title="Molecular phylogeny of Geoplaninae (Platyhelminthes) challenges current classification: proposal of taxonomic actions." url="10.1111/zsc.12019" volume="42" year="2013">Carbayo et al. (2013)</bibRefCitation>
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.
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</paragraph>
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<paragraph pageId="0" pageNumber="1">Type material. Parque Estadual do Desengano (-21.87; -41.91), Santa Maria Madalena, Rio de Janeiro State, Brazil: Holotype F4085 (MZUSP PL. 1085): J. Pedroni et al. col., 13 August 2009: sagittal sections of copulatory apparatus on 28 slides. Paratype F4057 (MZUSP PL. 1082): J. Pedroni et al. col., 12 August 2009: sagittal sections of copulatory apparatus on 29 slides.</paragraph>
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<paragraph lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1">
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Reserva
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<normalizedToken originalValue="Biológica">Biologica</normalizedToken>
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Augusto Ruschi (-19.88; -40.54), Santa Teresa,
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<normalizedToken originalValue="Espírito">Espirito</normalizedToken>
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Santo State, Brazil: Paratype F2158 (MZUSP PL 1020): F. Carbayo et al. col., 26 March 2008: fixed in 80% ethanol. Paratype F2250 (MZUSP PL. 1023): F. Carbayo et al. col., 24 May 2008: fixed in 80% ethanol. Paratype F2266 (MZUSP PL. 1024): F. Carbayo et al. col., 24 May 2008: fixed in 80% ethanol. Paratype F2274 (MZUSP PL. 1025): F. Carbayo et al. col., 24 May 2008: sagittal sections of copulatory apparatus on 33 slides; transverse sections of cephalic region on 7 slides; horizontal sections of portion containing ovaries on 22 slides; sagittal sections of pharynx region on 34 slides; transverse sections of pre-pharyngeal region on 8 slides. Paratype F2309 (MZUSP PL. 1032): F. Carbayo et al. col., 24 May 2008: fixed in 80% ethanol. Paratype F2394 (MZUSP PL. 1037): F. Carbayo et al. col., 27 May 2008: sagittal sections of copulatory apparatus on 18 slides; sagittal sections of pharynx region on 20 slides; transverse sections of cephalic region on 4 slides; horizontal sections of portion containing ovaries on 8 slides; and transverse sections of pre-pharyngeal region on 6 slides. Paratype F2470 (MZUSP PL. 1042): F. Carbayo et al. col., 29 May 2008: transverse sections of cephalic region on 14 slides; horizontal sections of portion containing ovaries on 48 slides; sagittal sections of a portion posterior to ovaries on 23 slides; sections immediately before pre-pharyngeal region on 33 slides; transverse sections of pre-pharyngeal region on 16 slides; sagittal sections of pharynx region on 41 slides; sagittal sections of
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<pageBreakToken pageId="1" pageNumber="2" start="start">copulatory</pageBreakToken>
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apparatus on 61 slides. Paratype F2507 (MZUSP PL. 1045): F. Carbayo et al. col., 29 May 2008: preserved in 80% ethanol.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="type locality">
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<paragraph pageId="1" pageNumber="2">Type locality.</paragraph>
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<paragraph pageId="1" pageNumber="2">Parque Estadual do Desengano, Santa Maria Madalena, Rio de Janeiro State, Brazil.</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="diagnosis">
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<paragraph pageId="1" pageNumber="2">Diagnosis.</paragraph>
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<paragraph pageId="1" pageNumber="2">
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Species of
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<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca" order="Tricladida" pageId="1" pageNumber="2" phylum="Platyhelminthes" rank="genus">Issoca</taxonomicName>
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up to 97 mm in length. Widest dorsal pigment bands with 33-40% of body width. A few cutaneous muscle bundles in-sunk in the pre-pharyngeal region. Cephalic retractor muscle and the sub-neural parenchymal muscle not intersecting with each other. Copulatory apparatus relatively long. Prostatic vesicle extrabulbar, proximally dilated, and non-anastomosed. Ejaculatory duct thin, with its opening at the tip of the penis papilla; this papilla is conical with dorsal insertion posterior to the ventral one. Female atrium spacious with some lateral folds. Common glandular duct almost as long as the female atrium. Common muscle coat envelops male and female atria.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="external morphology">
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<paragraph pageId="1" pageNumber="2">External morphology.</paragraph>
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<paragraph pageId="1" pageNumber="2">
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The body is elongated with nearly parallel margins (Fig. 2
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<normalizedToken originalValue="A–B">A-B</normalizedToken>
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). Its cephalic region (5% of the body length) is sometimes slightly laterally dilated before converging to the rounded anterior extremity of the body, which is slightly concave ventrally (Fig. 2C). The posterior extremity is pointed. The dorsum is slightly convex, the ventral side is flattened, and the body margins are rounded so that the section of the body is elliptic. Six mature, fixed specimens measured 48-97 mm in length. Fixed, paratype F2470 is 71 mm in length, 7 mm in width and 1.7 mm in height. The creeping sole is as wide as 74-84% of body width at the pre-pharyngeal region (paratypes F2470 and F2394, respectively). The mouth lies at a distance from the anterior extremity equal to 55% of body length; the gonopore is at 80% (paratype F2274).
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</paragraph>
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<paragraph pageId="1" pageNumber="2">
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The dorsal color consists of a cream-colored (specimens from Ruschi, Fig. 2
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<normalizedToken originalValue="A–B">A-B</normalizedToken>
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) or yellowish (specimens from Desengano, Fig. 2D) median stripe (6.7% of body width). This stripe is longitudinally divided by a very thin black median line (2.2%), which can be very tenuous and discontinuous (Fig. 2B). The.median stripe is bordered on either side by a wide black band (33-40%). This wide band is externally bordered by a whitish stripe (4-6.7%). The median and the lateral stripes gradually pass into orange of cephalic region (Fig. 2C). A marginal zone is either pigmented with a black stripe (5%, specimens from Ruschi, Fig. 2
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<normalizedToken originalValue="A–C">A-C</normalizedToken>
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) or mottled with black pigment spots (8.3%; Desengano, Fig. 2D). The ventral side is whitish, passing into orange of the cephalic surface. This surface (5% of body length) extends along the margins of the body progressively occupying a wider surface towards the anterior extremity until they fuse at 1 millimeter from the anterior extremity of the body (Fig. 3A).
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</paragraph>
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<paragraph pageId="1" pageNumber="2">
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The eyes are formed by one pigmented cup 50
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<normalizedToken originalValue="µm">µm</normalizedToken>
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in diameter. They contour the anterior extremity in a row of 2-3 eyes along the first 2 millimeters (Fig. 2C); going backwards, they spread progressively on each side of the dorsum in a band with 33% of the body width until the end of anterior half of the body. Posterior to this region they are scarcer and the band narrows until posterior extremity of the body.
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</paragraph>
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<caption pageId="1" pageNumber="2">
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<paragraph pageId="1" pageNumber="2">
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Figure 2.
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<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca assanga" order="Tricladida" pageId="1" pageNumber="2" phylum="Platyhelminthes" rank="species" species="assanga">Issoca assanga</taxonomicName>
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sp. n. Living specimens. Cephalic region is orangish. A paratype F2266 B paratype F2309 C Cephalic region of paratype F2250 D paratype F4085. Scale bars not available.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="8" lastPageNumber="9" pageId="1" pageNumber="2" type="internal morphology">
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<paragraph pageId="1" pageNumber="2">Internal morphology.</paragraph>
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<paragraph lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
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The sensory pits are 30
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<normalizedToken originalValue="µm">µm</normalizedToken>
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deep and are distributed in a simple ventro-lateral row, from the very anterior extremity up to at least 38% of body length. In the prepharyngeal region (Fig. 3B), rhabditogen cells and cell glands producing eryth
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<pageBreakToken pageId="2" pageNumber="3" start="start">rophil</pageBreakToken>
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granules open through the dorsal epithelium (Fig. 3B); necks of these glands are thick - 20
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<normalizedToken originalValue="µm">µm</normalizedToken>
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in width. Three additional gland types discharge their content through dorsal and marginal epidermis. They become progressively more abundant from the midbody towards the body margins. These gland cells are as follows: one type of cell with cell neck 20
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<normalizedToken originalValue="µm">µm</normalizedToken>
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in diameter produces xanthophilic granules; the second type produces xanthophilic granules, and its neck is 8-10
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<normalizedToken originalValue="µm">µm</normalizedToken>
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in diameter; the third type produces cyanophilic granules and its neck is 8-10
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<normalizedToken originalValue="µm">µm</normalizedToken>
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thick. The ventral epidermis is pierced by two types of scarce cell glands, each of them secreting either erythrophilic or xanthophilic granules.
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</paragraph>
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<paragraph pageId="2" pageNumber="3">
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The cutaneous musculature comprises the three typical layers of
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<taxonomicName lsidName="" pageId="2" pageNumber="3" rank="subfamily" subfamily="Geoplaninae">Geoplaninae</taxonomicName>
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, i.e., a subepithelial circular layer followed by a double diagonal layer with decussate fibers, and a strongly developed longitudinal layer, 60-125
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<normalizedToken originalValue="µm">µm</normalizedToken>
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thick. The fibers of the latter muscle layer are gathered into compact bundles (Fig. 3
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<normalizedToken originalValue="B–D">B-D</normalizedToken>
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). A few small muscle bundles of the cutaneous longitudinal muscle are sunken in the pre-pharyngeal region (Fig. 3
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<normalizedToken originalValue="C–D">C-D</normalizedToken>
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). The number of these sunken muscle fibers increases towards the cephalic region. The cutaneous musculature thickness relative to body height at the pre-pharyngeal region is 16-18%.
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</paragraph>
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<caption pageId="2" pageNumber="3">
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<paragraph pageId="2" pageNumber="3">
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Figure 3.
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<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca assanga" order="Tricladida" pageId="2" pageNumber="3" phylum="Platyhelminthes" rank="species" species="assanga">Issoca assanga</taxonomicName>
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sp. n. A Ventral view of cephalic region of fixed paratype F2274. (B-L): Photomicrograph of histological sections B transverse section of left side of pre-pharyngeal region of paratype F2394 C Horizontal section of paratype F2394 at the level of the ovaries D transverse section of prepharyngeal region of paratype F2394 E, F horizontal section of cephalic region of paratype F2394.
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</paragraph>
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</caption>
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<paragraph lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
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The three usual parenchymal muscle layers are present throughout the body: a dorsal layer of diagonal decussate fibers (50
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<normalizedToken originalValue="μm">μm</normalizedToken>
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thick, or 5% of the body height,
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<pageBreakToken pageId="3" pageNumber="4" start="start">paratype</pageBreakToken>
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F2470), a transverse supraintestinal layer (40
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<normalizedToken originalValue="μm">μm</normalizedToken>
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, or 4%), and a transverse subintestinal layer (50
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<normalizedToken originalValue="μm">μm</normalizedToken>
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, or 5%). Additionally, there is a subneural layer (40
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<normalizedToken originalValue="μm">μm</normalizedToken>
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, or 4%) of transverse muscles. Dorso-ventrally oriented parenchymal fibers are abundant in the pre-pharyngeal region.
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</paragraph>
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<paragraph lastPageId="5" lastPageNumber="6" pageId="3" pageNumber="4">
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The muscular organization changes (Fig. 4) in the anterior region of the body with respect to that of the pre-pharyngeal region. The number of sunken ventral longitudinal cutaneous fibers increases at 5 millimeters from the anterior extremity of the body (equal to 10% of body length in paratype F2394), and bundles of both normal and
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<pageBreakToken pageId="4" pageNumber="5" start="start">sunken</pageBreakToken>
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ventral cutaneous muscles concentrate medially to give rise to the cephalic retractor muscle. The retractor is lens-shaped in cross-section at 4-2.6 mm from anterior extremity (Fig. 4
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<normalizedToken originalValue="A–B">A-B</normalizedToken>
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). It becomes elliptic at 2.0 mm (Fig. 4C) and roughly quadrangular (Fig. 4D) at 1.6 mm from anterior extremity. Muscle fibers of the retractor muscle are gathered in few but thick bundles. From this region towards anterior extremity,
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<pageBreakToken pageId="5" pageNumber="6" start="start">fibers</pageBreakToken>
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of the retractor muscle progressively detach in bundles that run to the body sides (Fig. 4
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<normalizedToken originalValue="D–F">D-F</normalizedToken>
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). As they detach, the retractor muscle becomes less apparent until it disappears close to the anteriormost extremity of the body. It could not be determined whether other fibers from this muscle run dorsally. Before disappearing the retractor muscle is elliptic in cross section (Fig. 4
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<normalizedToken originalValue="E–F">E-F</normalizedToken>
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).
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</paragraph>
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<paragraph pageId="5" pageNumber="6">In the cephalic region, diagonal, supraintestinal, and subneural parenchymal muscle layers are apparent and placed in the same position relative to the cutaneous longitudinal muscles. Even the subneural muscle and its fibers continue running over the retractor muscle (Fig. 4). The subintestinal parenchymal muscle layer is less apparent, and this layer is the first to disappear as it approaches the anterior extremity of the body. All parenchymal muscles fade out at the anteriormost body portion.</paragraph>
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<paragraph pageId="5" pageNumber="6">
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Dorso-ventral parenchymal muscle fibers are more abundant in the cephalic region than on the rest of the body, and they are frequently gathered in bundles of 3-10 fibers each. These fibers run approximately dorso-ventrally, connecting dorsal epidermis with the ventral glandular epidermis. Medially, these fibers run obliquely from the dorsal epidermis of one side of the body to anchor to the ventral glandular epidermis of the other body side, thereby rimming the retractor muscle (Fig. 4
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<normalizedToken originalValue="C–E">C-E</normalizedToken>
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).
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</paragraph>
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<caption pageId="5" pageNumber="6">
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<paragraph pageId="5" pageNumber="6">
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Figure 4. Serial transverse sections of the cephalic region of paratype F2470 at 4.0, 2.6, 2.0, 1.6, 1.3, and 1.0 mm from anterior extremity, respectively. Encircled region in F is enlarged in the inset.
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<normalizedToken originalValue="A–E">A-E</normalizedToken>
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at the same magnification as F.
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</paragraph>
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</caption>
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<paragraph pageId="5" pageNumber="6">
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The glandular surface of the ventral epidermis, orangish in color in living animals, widens towards anterior extremity of the body as the muscle fibers of the retractor concentrate medially. This surface is incompletely bipartite (Fig. 3A) and is richly pierced by gland cells (with 10-12
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<normalizedToken originalValue="μm">μm</normalizedToken>
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thick necks) producing erythrophilic granules, and by scarce glands (with 12-18
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<normalizedToken originalValue="μm">μm</normalizedToken>
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thick necks) producing cyanophilic granules.
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</paragraph>
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<paragraph pageId="5" pageNumber="6">
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The mouth is situated at a distance from the root of the pharynx equivalent to 25% of pharyngeal pocket length (Fig. 5). An esophagus is present with 13-20% of pharynx length. The pharynx is cylindrical, with dorsal insertion posterior to the ventral and located at mouth level. The lining epithelium of the pharyngeal pouch is squamous, non-ciliated, surrounded by a simple layer of circular fibers, followed by a layer of diagonal fibers (10
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<normalizedToken originalValue="µm">µm</normalizedToken>
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thick). The outer and inner pharyngeal epithelia are flat and ciliated. The outer epithelium is underlain by a longitudinal muscle (2.5
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<normalizedToken originalValue="µm">µm</normalizedToken>
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thick) followed by a circular muscle (17-75
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<normalizedToken originalValue="µm">µm</normalizedToken>
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), the latter with some longitudinal fibers interspersed. The inner epithelium is surrounded by a circular muscle (45-90
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<normalizedToken originalValue="µm">µm</normalizedToken>
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), followed by a longitudinal muscle (25
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<normalizedToken originalValue="µm">µm</normalizedToken>
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).
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</paragraph>
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<caption pageId="5" pageNumber="6">
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<paragraph pageId="5" pageNumber="6">
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Figure 5.
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<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca assanga" order="Tricladida" pageId="5" pageNumber="6" phylum="Platyhelminthes" rank="species" species="assanga">Issoca assanga</taxonomicName>
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sp. n. Photomicrograph a sagittal section of the pharynx of paratype F2394.
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</paragraph>
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</caption>
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<paragraph pageId="5" pageNumber="6">The central nervous system mainly consists of a ventral nerve plate. Cerebral ganglia (Fig. 4B) extend along the body from 1 millimeter to 4 millimeters behind the anterior tip (2% and 8%, respectively, paratype F2394).</paragraph>
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<paragraph lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
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The testes are located under the supraintestinal transverse muscle layer, partially between the intestinal diverticula. They extend from the level of the ovaries to nearly the root of the pharynx. The sperm ducts run between the subintestinal parenchymal muscle layer, dorsally to the ovovitelline ducts. They open into the antero-lateral portion of the prostatic vesicle. This vesicle is extrabulbar and proximally dilated, curves dorsally and penetrates the dorso-anterior aspect of the penis bulb. This vesicle is lined with a ciliated, columnar epithelium, showing an irregular free surface in its anterior portion. The prostatic vesicle receives fine erythrophilic granular secretions derived from glands
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<pageBreakToken pageId="6" pageNumber="7" start="start">in</pageBreakToken>
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the parenchyma. These penetrate the ciliated, columnar epithelium lining of the vesicle to discharge into the lumen. The vesicle is surrounded by intermingled decussate, circular, and longitudinal fibers. The prostatic vesicle passes into the relatively thin ejaculatory duct that is lined by a ciliated cuboidal epithelium surrounded by circular muscle. The ejaculatory duct is proximally sinuous and distally straight through the mid penis bulb, terminating at the tip of the penis papilla. The ejaculatory duct is lined with a cuboidal, ciliated epithelium and is surrounded by a circular muscle. The protrusible penis papilla is conical, slightly inclined ventrally, and with its dorsal insertion posterior to the ventral insertion (Fig. 6
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<normalizedToken originalValue="A–D">A-D</normalizedToken>
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). The penis papilla is as long as the male atrium, and is lined with a cuboidal, non-ciliated epithelium, and is surrounded by a circular muscle followed by a longitudinal muscle; some fibers of both muscles are intermingled. Numerous secretory cells located in the adjacent parenchyma produce erythrophilic granules (Fig. 6B) that are discharged along the length of the papilla.
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</paragraph>
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<paragraph pageId="6" pageNumber="7">
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The male atrium is mostly occupied by the penis papilla. It is more spacious in its anterior portion than in the posterior, and shows folded walls. One of these folds is a transverse, annular-shaped fold located halfway of the atrial length. From the roof of the distal portion of the male atrium, a large fold projects laterally and continues along the female atrium. The male atrium is lined with a columnar, non-ciliated epithelium, and is pierced by gland cells producing erythrophilic granules. A circular muscularis (5
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<normalizedToken originalValue="µm">µm</normalizedToken>
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||
thick) encircles the male atrium.
|
||
</paragraph>
|
||
<caption pageId="6" pageNumber="7">
|
||
<paragraph pageId="6" pageNumber="7">
|
||
Figure 6.
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca assanga" order="Tricladida" pageId="6" pageNumber="7" phylum="Platyhelminthes" rank="species" species="assanga">Issoca assanga</taxonomicName>
|
||
sp. n. A photomicrograph of a sagittal section of the copulatory apparatus of paratype F2470 B photomicrograph of a sagittal section of the penis papilla of paratype F2394 C photomicrograph of a sagittal section of the copulatory apparatus of paratype F2394 D diagrammatic representation of the copulatory apparatus of paratype F2394 from sagittal sections.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
|
||
The ovaries are 500
|
||
<normalizedToken originalValue="µm">µm</normalizedToken>
|
||
long in the antero-posterior body axis and 200
|
||
<normalizedToken originalValue="µm">µm</normalizedToken>
|
||
wide. They are located immediately above the ventral nerve plate, at a distance from anterior tip equivalent to 26% of body length (Fig. 3D). The ovovitelline ducts arise from the dorso-external surface of the anterior portion of the ovaries, and run backwards above the ventral nerve plate. They ascend laterally to the gonopore region, and subsequently unite dorsally to form a common ovovitelline duct just dorsal to the anterior section of the female atrium (Fig. 6D). The distal third of the ascending portion of these paired ducts receives shell glands. The ovovitelline ducts unite to form the common glandular ovovitelline duct, which runs caudally and progressively curving to the ventral side to communicate with the female genital canal. This canal is a projection of the postero-dorsal portion of the female atrium that runs dorsally and slightly anteriorly. The female atrium is an irregular, spacious cavity. Its walls are partially projected into
|
||
<pageBreakToken pageId="7" pageNumber="8" start="start">its</pageBreakToken>
|
||
lumen. One of these folds is continuous with a fold coming from the male atrium (Fig. 6D).
|
||
</paragraph>
|
||
<paragraph lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
|
||
The female atrium is lined with a columnar epithelium, which is lacunar in aspect in some parts. In the anterior portion of the female atrium, the simple columnar epithelium is 25
|
||
<normalizedToken originalValue="μm">μm</normalizedToken>
|
||
high; whereas the posterior portion is lined by a pseudostratified columnar epithelium (Fig. 7). The muscularis of the female atrium consists of two muscle layers; a simple (2.5
|
||
<normalizedToken originalValue="µm">µm</normalizedToken>
|
||
thick) longitudinal muscle followed by a circular muscle
|
||
<pageBreakToken pageId="8" pageNumber="9" start="start">(</pageBreakToken>
|
||
10
|
||
<normalizedToken originalValue="µm">µm</normalizedToken>
|
||
), both partially intermingled. The female atrium receives gland cells producing erythrophilic granules. The male atrium is 1.2 times longer than the female. The common muscle coat is well-developed, and wraps the male and female atria.
|
||
</paragraph>
|
||
<caption pageId="8" pageNumber="9">
|
||
<paragraph pageId="8" pageNumber="9">
|
||
Figure 7.
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca assanga" order="Tricladida" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="assanga">Issoca assanga</taxonomicName>
|
||
sp. n. Photomicrograph of a sagittal section of the lining epithelium of the posterior section of the female atrium of the holotype.
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection pageId="8" pageNumber="9" type="etymology">
|
||
<paragraph pageId="8" pageNumber="9">Etymology.</paragraph>
|
||
<paragraph pageId="8" pageNumber="9">
|
||
The specific epithet refers to the Tupi (indigenous Brazilian tribe) name
|
||
<taxonomicName lsidName="assanga" pageId="8" pageNumber="9" rank="species" species="assanga">assanga</taxonomicName>
|
||
, meaning dense, thick (
|
||
<bibRefCitation author="Tibirica, LC" journalOrPublisher="Traco, Sao Paulo" pageId="11" pageNumber="12" title="Dicionario Tupi-Portugues." year="1984">
|
||
<normalizedToken originalValue="Tibiriçá">Tibirica</normalizedToken>
|
||
1984
|
||
</bibRefCitation>
|
||
). It refers to the apparent cephalic glandulo-muscular organ of the new species.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="8" pageNumber="9" type="distribution">
|
||
<paragraph pageId="8" pageNumber="9">Distribution.</paragraph>
|
||
<paragraph pageId="8" pageNumber="9">
|
||
Parque Estadual do Desengano (Municipality of Santa Maria Madalena, Rio de Janeiro State) and Reserva
|
||
<normalizedToken originalValue="Biológica">Biologica</normalizedToken>
|
||
Augusto Ruschi (Municipality of Santa Teresa,
|
||
<normalizedToken originalValue="Espírito">Espirito</normalizedToken>
|
||
Santo State), Brazil.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9" type="discussion">
|
||
<paragraph pageId="8" pageNumber="9">Discussion.</paragraph>
|
||
<paragraph pageId="8" pageNumber="9">
|
||
The external aspect of
|
||
<taxonomicName genus="I." lsidName="I. assanga" pageId="8" pageNumber="9" rank="species" species="assanga">I. assanga</taxonomicName>
|
||
allows one to readily distinguish it from the remaining species of the genus by being considerably larger (97 mm in length after fixation) than the largest previously described species, i.e.
|
||
<taxonomicName genus="I." lsidName="I. jandaia" pageId="8" pageNumber="9" rank="species" species="jandaia">I. jandaia</taxonomicName>
|
||
, which is 50 mm in length alive, and 32 mm after fixation. Although the chromatic pattern of all species of the genus is striped, the stripes are as wide as 33-40% of the body width only in
|
||
<taxonomicName genus="I." lsidName="I. assanga" pageId="8" pageNumber="9" rank="species" species="assanga">I. assanga</taxonomicName>
|
||
sp. n. The widest colored stripe in other species of the genus is found in
|
||
<taxonomicName genus="I." lsidName="I. potyra" pageId="8" pageNumber="9" rank="species" species="potyra">I. potyra</taxonomicName>
|
||
, with a pair or paramedian bands with 17% of the body width each (
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="195 - 279" title="Sobre morfologia e taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciencias e Letras." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1954.120092" volume="19" year="1955">Froehlich 1955</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="93 - 121" title="On a collection of Brazilian land planarians. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1957.120237" volume="21" year="1958">1958</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Graff, LV" journalOrPublisher="Wilhelm Engelmann, Leipzig" pageId="10" pageNumber="11" title="Monographie der Turbellarien. II. Tricladida Terricola (Landplanarien). I-XII. Atlas von Achtundfuenfzig Tafeln zur Monographie der Turbellarien II. Tricladida Terricola (Landplanarien). Pls I-LVIII." year="1899">Graff 1899</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Marcus, E" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="5 - 215" title="Turbellaria brasileiros (9). Boletim da Faculdade de Filosofia, Ciencias e Letras. Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-4877.bsffclzoologia.1951.125221" volume="6" year="1951">Marcus 1951</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
|
||
Regarding the digestive system anatomy, the new species can be distinguished from the other species because the mouth is situated at a distance from the root of the phar
|
||
<pageBreakToken pageId="9" pageNumber="10" start="start">ynx</pageBreakToken>
|
||
equivalent to 25% of pharyngeal pocket, whereas the mouth in the other species is at a distance equal to 33% (
|
||
<taxonomicName genus="I." lsidName="I. jandaia" pageId="9" pageNumber="10" rank="species" species="jandaia">I. jandaia</taxonomicName>
|
||
) or>30% (
|
||
<taxonomicName genus="I." lsidName="I. rezendei" pageId="9" pageNumber="10" rank="species" species="rezendei">I. rezendei</taxonomicName>
|
||
and
|
||
<taxonomicName genus="I." lsidName="I. piranga" pageId="9" pageNumber="10" rank="species" species="piranga">I. piranga</taxonomicName>
|
||
, 50%;
|
||
<taxonomicName genus="I." lsidName="I. potyra" pageId="9" pageNumber="10" rank="species" species="potyra">I. potyra</taxonomicName>
|
||
, 67%;
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="195 - 279" title="Sobre morfologia e taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciencias e Letras." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1954.120092" volume="19" year="1955">Froehlich 1955</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="93 - 121" title="On a collection of Brazilian land planarians. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1957.120237" volume="21" year="1958">1958</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Marcus, E" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="5 - 215" title="Turbellaria brasileiros (9). Boletim da Faculdade de Filosofia, Ciencias e Letras. Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-4877.bsffclzoologia.1951.125221" volume="6" year="1951">Marcus 1951</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="9" pageNumber="10">
|
||
The new species can also be readily distinguished from
|
||
<taxonomicName genus="I." lsidName="I. rezendei" pageId="9" pageNumber="10" rank="species" species="rezendei">I. rezendei</taxonomicName>
|
||
by the general shape of the copulatory apparatus. Unlike the new species (with a relatively long copulatory apparatus, an extrabulbar prostatic vesicle, a horizontal penis papilla and a conspicuous female atrium), the copulatory apparatus in
|
||
<taxonomicName genus="I." lsidName="I. rezendei" pageId="9" pageNumber="10" rank="species" species="rezendei">I. rezendei</taxonomicName>
|
||
is relatively compact, the prostatic vesicle is intrabulbar, the penis papilla is vertical, and the female atrium is absent (
|
||
<bibRefCitation author="Marcus, E" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="5 - 215" title="Turbellaria brasileiros (9). Boletim da Faculdade de Filosofia, Ciencias e Letras. Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-4877.bsffclzoologia.1951.125221" volume="6" year="1951">Marcus 1951</bibRefCitation>
|
||
). The outlines of the copulatory apparatuses of the remaining species of
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca" order="Tricladida" pageId="9" pageNumber="10" phylum="Platyhelminthes" rank="genus">Issoca</taxonomicName>
|
||
are comparable to that of the new species: those being a relatively long and extrabulbar prostatic vesicle; and a penis papilla, or a papilla-like fold horizontal or moderately inclined (
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="195 - 279" title="Sobre morfologia e taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciencias e Letras." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1954.120092" volume="19" year="1955">Froehlich 1955</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="93 - 121" title="On a collection of Brazilian land planarians. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1957.120237" volume="21" year="1958">1958</bibRefCitation>
|
||
). However, the three species
|
||
<taxonomicName genus="I." lsidName="I. jandaia" pageId="9" pageNumber="10" rank="species" species="jandaia">I. jandaia</taxonomicName>
|
||
,
|
||
<taxonomicName genus="I." lsidName="I. piranga" pageId="9" pageNumber="10" rank="species" species="piranga">I. piranga</taxonomicName>
|
||
and
|
||
<taxonomicName genus="I." lsidName="I. potyra" pageId="9" pageNumber="10" rank="species" species="potyra">I. potyra</taxonomicName>
|
||
(
|
||
<taxonomicName genus="I." lsidName="I. spatulata" pageId="9" pageNumber="10" rank="species" species="spatulata">I. spatulata</taxonomicName>
|
||
is not in this comparison because its copulatory apparatus is yet unknown), differ from the new species in the following details. In contrast to
|
||
<taxonomicName genus="I." lsidName="I. assanga" pageId="9" pageNumber="10" rank="species" species="assanga">I. assanga</taxonomicName>
|
||
sp. n., the ejaculatory duct of
|
||
<taxonomicName genus="I." lsidName="I. jandaia" pageId="9" pageNumber="10" rank="species" species="jandaia">I. jandaia</taxonomicName>
|
||
opens into a cavity inside a penis papilla-like fold of the male atrium, the common glandular duct is relatively long, and the female atrium is intensely folded. The penis papilla of
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca piranga" order="Tricladida" pageId="9" pageNumber="10" phylum="Platyhelminthes" rank="species" species="piranga">Issoca piranga</taxonomicName>
|
||
occupies only the anterior half of the male atrium, and the muscle coat enveloping the male atrium is separated from that wrapping the female one (
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="195 - 279" title="Sobre morfologia e taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciencias e Letras." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1954.120092" volume="19" year="1955">Froehlich 1955</bibRefCitation>
|
||
), whereas in the new species the penis papilla is as long as the male atrium, and there is a common muscle coat wrapping male and female atria. Finally,
|
||
<taxonomicName genus="I." lsidName="I. potyra" pageId="9" pageNumber="10" rank="species" species="potyra">I. potyra</taxonomicName>
|
||
differs from the new species in that the wall of the prostatic vesicle is anastomosed, the ejaculatory duct is wide and irregular and it opens into the ventral side of the penis papilla, the male atrium is separated from the female one by a fold which is richly pierced by cyanophilic glands, the common glandular oviduct is relatively short, and the muscle coat enveloping the male atrium is separated from that of the female one (
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="93 - 121" title="On a collection of Brazilian land planarians. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1957.120237" volume="21" year="1958">Froehlich 1958</bibRefCitation>
|
||
), whereas in
|
||
<taxonomicName genus="I." lsidName="I. assanga" pageId="9" pageNumber="10" rank="species" species="assanga">I. assanga</taxonomicName>
|
||
sp. n., there are no anastomoses in the prostatic vesicle, the ejaculatory duct is relatively thin, and terminates at the tip of the penis papilla, the male atrial fold is not pierced by cyanophilic glands, the common glandular oviduct is relatively long, and the muscle coat wraps both male and female atria.
|
||
</paragraph>
|
||
<paragraph lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
|
||
One diagnostic feature of the genus is that the cephalic retractor muscle and the subneural parenchymal muscle are intersected, a condition present in the type species of the genus
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca rezendei" order="Tricladida" pageId="9" pageNumber="10" phylum="Platyhelminthes" rank="species" species="rezendei">Issoca rezendei</taxonomicName>
|
||
(see diagnosis in
|
||
<bibRefCitation author="Carbayo, F" journalOrPublisher="Invertebrate Systematics" pageId="10" pageNumber="11" pagination="449 - 468" title="Two new genera of geoplaninid land planarians (Platyhelminthes: Tricladida: Terricola) of Brazil in the light of cephalic specialisations." url="https://doi.org/10.1071/IT01035" volume="17" year="2003">Carbayo and Leal-Zanchet 2003</bibRefCitation>
|
||
). This feature is the only mismatching diagnostic trait of
|
||
<taxonomicName genus="I." lsidName="I. assanga" pageId="9" pageNumber="10" rank="species" species="assanga">I. assanga</taxonomicName>
|
||
sp. n. This condition could be polymorphic within the genus. However, two aspects support a different interpretation: (a) the polyphyly of
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca" order="Tricladida" pageId="9" pageNumber="10" phylum="Platyhelminthes" rank="genus">Issoca</taxonomicName>
|
||
has been highlighted (
|
||
<bibRefCitation author="Carbayo, F" journalOrPublisher="Zoologica Scripta" pageId="10" pageNumber="11" pagination="508 - 528" title="Molecular phylogeny of Geoplaninae (Platyhelminthes) challenges current classification: proposal of taxonomic actions." url="10.1111/zsc.12019" volume="42" year="2013">Carbayo et al. 2013</bibRefCitation>
|
||
); and (b) the general anatomy of the copulatory apparatus of
|
||
<taxonomicName genus="I." lsidName="I. rezendei" pageId="9" pageNumber="10" rank="species" species="rezendei">I. rezendei</taxonomicName>
|
||
is very different from that of the remaining species of the genus. These points suggest that rather than an interspecifically polymorphic character, the muscle intersection might reflect the polyphyletic status of the genus (see Fig. 1). Description of the muscular cephalic retractor muscle in the other species of
|
||
<taxonomicName family="Geoplanidae" genus="Issoca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Issoca" order="Tricladida" pageId="9" pageNumber="10" phylum="Platyhelminthes" rank="genus">Issoca</taxonomicName>
|
||
is limited. The muscular organization of the glandulo-muscular organ in
|
||
<taxonomicName genus="I." lsidName="I. jandaia" pageId="9" pageNumber="10" rank="species" species="jandaia">I. jandaia</taxonomicName>
|
||
was reported by
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="195 - 279" title="Sobre morfologia e taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciencias e Letras." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1954.120092" volume="19" year="1955">Froehlich (1955)</bibRefCitation>
|
||
as following the same structure as
|
||
<taxonomicName genus="I." lsidName="I. rezendei" pageId="9" pageNumber="10" rank="species" species="rezendei">I. rezendei</taxonomicName>
|
||
. The glandulo-muscular organ in
|
||
<taxonomicName genus="I." lsidName="I. piranga" pageId="9" pageNumber="10" rank="species" species="piranga">I. piranga</taxonomicName>
|
||
was described as being very similar to that of
|
||
<taxonomicName genus="I." lsidName="I. jandaia" pageId="9" pageNumber="10" rank="species" species="jandaia">I. jandaia</taxonomicName>
|
||
(see
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="195 - 279" title="Sobre morfologia e taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciencias e Letras." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1954.120092" volume="19" year="1955">Froehlich 1955</bibRefCitation>
|
||
).
|
||
<bibRefCitation author="Froehlich, CG" journalOrPublisher="Serie Zoologia" pageId="11" pageNumber="12" pagination="93 - 121" title="On a collection of Brazilian land planarians. Boletim da Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo." url="http://dx.doi.org/10.11606/issn.2526-3382.bffclzoologia.1957.120237" volume="21" year="1958">Froehlich (1958)</bibRefCitation>
|
||
mentioned that the glan
|
||
<pageBreakToken pageId="10" pageNumber="11" start="start">dulo-muscular</pageBreakToken>
|
||
organ (i.e., the cephalic retractor muscle plus associated viscid glands) in
|
||
<taxonomicName genus="I." lsidName="I. potyra" pageId="10" pageNumber="11" rank="species" species="potyra">I. potyra</taxonomicName>
|
||
"is similar to that of the other species of the genus". However, his diagrammatic reconstruction of the organ shows the cephalic retractor muscle underneath the subneural muscle layer, not intersected. Moreover, the retractor in
|
||
<taxonomicName genus="I." lsidName="I. rezendei" pageId="10" pageNumber="11" rank="species" species="rezendei">I. rezendei</taxonomicName>
|
||
diminishes by means of separating its fibers towards the body margins and the back (
|
||
<bibRefCitation author="Carbayo, F" journalOrPublisher="Invertebrate Systematics" pageId="10" pageNumber="11" pagination="449 - 468" title="Two new genera of geoplaninid land planarians (Platyhelminthes: Tricladida: Terricola) of Brazil in the light of cephalic specialisations." url="https://doi.org/10.1071/IT01035" volume="17" year="2003">Carbayo and Leal-Zanchet 2003</bibRefCitation>
|
||
), whereas the fibers in
|
||
<taxonomicName genus="I." lsidName="I. assanga" pageId="10" pageNumber="11" rank="species" species="assanga">I. assanga</taxonomicName>
|
||
were observed only running to the body margins. This lack of morphological details reinforces the need of a taxonomic revision of the clade LIS, as already suggested by
|
||
<bibRefCitation author="Carbayo, F" journalOrPublisher="Zoologica Scripta" pageId="10" pageNumber="11" pagination="508 - 528" title="Molecular phylogeny of Geoplaninae (Platyhelminthes) challenges current classification: proposal of taxonomic actions." url="10.1111/zsc.12019" volume="42" year="2013">Carbayo et al. (2013)</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |