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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.782.26797" ID-GBIF-Dataset="c62d1cd5-20b2-45c9-abe3-8a8b98bb44fa" ID-PMC="PMC6160864" ID-Pensoft-Pub="1313-2970-782-95" ID-PubMed="30275721" ID-ZBK="2B276BC250E94244A32A524E2B266DB3" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-782-95" ModsDocOrigin="ZooKeys 782" ModsDocTitle="The genus Mercuria Boeters, 1971 in Morocco: first molecular phylogeny of the genus and description of two new species (Caenogastropoda, Truncatelloidea, Hydrobiidae)" checkinTime="1535418116552" checkinUser="pensoft" docAuthor="Boulaassafer, Khadija, Ghamizi, Mohamed &amp; Delicado, Diana" docDate="2018" docId="8356DA2F98A437585C733AEAB8679F3D" docLanguage="en" docName="ZooKeys 782: 95-128" docOrigin="ZooKeys 782" docSource="http://dx.doi.org/10.3897/zookeys.782.26797" docTitle="Mercuria tensiftensis Boulaassafer, Ghamizi &amp; Delicado, 2018, sp. n." docType="treatment" docUuid="42E0CFAF-0F49-48AB-BFC8-9323FEB6996D" docUuidSource="ZooBank" docVersion="5" lastPageNumber="109" masterDocId="E93FFFFB8C49FFD8E5404665FF8C7362" masterDocTitle="The genus Mercuria Boeters, 1971 in Morocco: first molecular phylogeny of the genus and description of two new species (Caenogastropoda, Truncatelloidea, Hydrobiidae)" masterLastPageNumber="128" masterPageNumber="95" pageNumber="105" updateTime="1668166182236" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>The genus Mercuria Boeters, 1971 in Morocco: first molecular phylogeny of the genus and description of two new species (Caenogastropoda, Truncatelloidea, Hydrobiidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Boulaassafer, Khadija</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ghamizi, Mohamed</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Delicado, Diana</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2018</mods:date>
<mods:detail type="volume">
<mods:number>782</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>95</mods:start>
<mods:end>128</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.782.26797</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.782.26797</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-782-95</mods:identifier>
<mods:identifier type="ZBK">2B276BC250E94244A32A524E2B266DB3</mods:identifier>
<mods:identifier type="ZooBank">2B276BC250E94244A32A524E2B266DB3</mods:identifier>
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<treatment ID-GBIF-Taxon="147440668" LSID="urn:lsid:zoobank.org:act:42E0CFAF-0F49-48AB-BFC8-9323FEB6996D" httpUri="http://treatment.plazi.org/id/8356DA2F98A437585C733AEAB8679F3D" lastPageId="14" lastPageNumber="109" pageId="10" pageNumber="105">
<subSubSection pageId="10" pageNumber="105" type="nomenclature">
<paragraph pageId="10" pageNumber="105">
<taxonomicName LSID="http://zoobank.org/42E0CFAF-0F49-48AB-BFC8-9323FEB6996D" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria tensiftensis" order="Littorinimorpha" pageId="10" pageNumber="105" phylum="Mollusca" rank="species" species="tensiftensis">Mercuria tensiftensis</taxonomicName>
<taxonomicNameLabel pageId="10" pageNumber="105">sp. n.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="105" type="type material">
<paragraph pageId="10" pageNumber="105">Type material.</paragraph>
<paragraph pageId="10" pageNumber="105">
Holotype, MNCN 15.05/200018H (ethanol 80%), a ditch in Sidi Bouzid, Chichaoua, Morocco,
<geoCoordinate direction="north" orientation="latitude" precision="1" value="31.493555">31°29.6133'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-8.785193">8°47.1116'W</geoCoordinate>
, 28/11/2015, K.B., M.G. Paratypes MNCN 15.05/200018P, UGSB 17910, and MHNM 18 ZTMH4 (from the same lot).
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="105" type="materials_examined">
<paragraph pageId="10" pageNumber="105">Other material.</paragraph>
<paragraph pageId="10" pageNumber="105">
MOROCCO. MHNM 18 ZTMH19, UGSB 17910, ditch in Sidi Bouzid, Chichaoua, 28/11/2015 (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="31.493555">31°29.6133'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-8.785193">8°47.1116'W</geoCoordinate>
); MHNM 18 ZTMH5, UGSB 17914, a pond near Lahjar Spring, Essaouira, 28/11/2015 (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="31.645971">31°38.7583'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-9.584971">9°35.0983'W</geoCoordinate>
); MHNM 18 ZTMH6, UGSB 17918, ditch in Haddada Bouzerktoun, Essaouira, 28/11/2015 (
<geoCoordinate direction="north" orientation="latitude" precision="9" value="31.6325">31°37.95'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-9.584971">9°35.0983'W</geoCoordinate>
); MHNM 18 ZTMH7, UGSB 19944, ditch in Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
, 34 km S of Marrakesh, 20/02/2017 (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="31.384861">31°23.0917'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-8.12255">8°7.353'W</geoCoordinate>
); MHNM 18 ZTMH8, UGSB 19945, a spring near Lalla Takerkoust dam, 34 km S of Marrakesh, 20/02/2017 (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="31.375818">31°22.5491'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-8.1273">8°7.638'W</geoCoordinate>
); MHNM 18 ZTMH9, UGSB 19946, Talkount, 80 km E of Marrakesh, 21/02/2017 (
<geoCoordinate direction="north" orientation="latitude" precision="1" value="31.676292">31°40.5775'N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-7.2671633">7°16.0298'W</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="11" lastPageNumber="106" pageId="10" pageNumber="105" type="diagnosis">
<paragraph pageId="10" pageNumber="105">Diagnosis.</paragraph>
<paragraph lastPageId="11" lastPageNumber="106" pageId="10" pageNumber="105">
Shell ovate-conic, whorls 4-5; periostracum whitish, exceptionally yellowish; body whorl large, convex, occupying approx. two-thirds of total shell length; umbilicus narrow, not covered by the inner lip; aperture ovate; protoconch microsculpture grooved; central radula tooth formula (5)4
<normalizedToken originalValue="C4(5)/1">-C-4(5)/1-</normalizedToken>
1; bursa copulatrix elongate, with a short duct; one seminal receptacle elongate, with a short duct; penis
<pageBreakToken pageId="11" pageNumber="106" start="start">gradually</pageBreakToken>
tapering; penial appendix dark pigmented, rectangular, shorter than penis, base narrow and black pigmented, medially positioned on inner edge of penis; nervous system elongated (mean RPG ratio = 0.64), slightly pigmented.
</paragraph>
</subSubSection>
<subSubSection lastPageId="14" lastPageNumber="109" pageId="11" pageNumber="106" type="description">
<paragraph pageId="11" pageNumber="106">Description.</paragraph>
<paragraph pageId="11" pageNumber="106">
Shell ovate-conic, whorls 4-5, height 3-5.1 mm (Figure 13
<normalizedToken originalValue="AG">A-G</normalizedToken>
; Suppl. material 1: Table 1). Periostracum whitish. Protoconch ca. 400
<normalizedToken originalValue="µm">µm</normalizedToken>
wide, whorls 1.5; nucleus ca. 125
<normalizedToken originalValue="µm">µm</normalizedToken>
wide (Figure 13H); protoconch microsculpture grooved (Figure 13I). Teleoconch whorls convex, with deep sutures. Body whorl large, occupying approx. two-thirds of total shell length. Aperture ovate, often attached to body whorl on the top; inner lip thicker than outer lip; peristome margin straight narrow (Figure 13D). Umbilicus narrow, not covered by the inner lip.
</paragraph>
<caption pageId="11" pageNumber="106">
<paragraph pageId="11" pageNumber="106">
Figure 13. Shells and opercula,
<taxonomicName lsidName="M. tensiftensis" pageId="11" pageNumber="106" rank="species" species="tensiftensis">M. tensiftensis</taxonomicName>
sp. n. A Holotype MNCN 15.05/200018H, B Shell, ditch in Sidi Bouzid C, D Shell, ditch in Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
E Shell, ditch in Talkount F Shell, a pond near Lahjar Spring G Shell, a spring near Lalla Takerkoust dam J, K Opercula (inner, outer sides), ditch in Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
H, I Protoconch and detailed microsculpture of protoconch, ditch in Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
.
</paragraph>
</caption>
<paragraph lastPageId="12" lastPageNumber="107" pageId="11" pageNumber="106">
Operculum as for genus, orange to brownish, about two whorls; muscle attachment area oval and located near the nucleus (Figure 13
<normalizedToken originalValue="JK">J-K</normalizedToken>
). Radula length intermediate, ca. 900
<normalizedToken originalValue="µm">µm</normalizedToken>
long (25% total shell length), approx. eight times longer than wide; approx. 60 rows of teeth (Figure 14A). Central tooth formula (5)4
<normalizedToken originalValue="C4(5)/1">-C-4(5)/1-</normalizedToken>
1; central cusp V-shaped (Figure 14B, D, E). Lateral tooth formula (4)3
<normalizedToken originalValue="C">-C-</normalizedToken>
3(4); central cusp
<pageBreakToken pageId="12" pageNumber="107" start="start">long</pageBreakToken>
, tongue-shaped (Figure 14
<normalizedToken originalValue="BC">B-C</normalizedToken>
). Inner marginal teeth bearing 13-16 cusps and outer marginal with 15-21 cusps (Figure 14D, F; Suppl. material 1: Table 2).
</paragraph>
<caption pageId="12" pageNumber="107">
<paragraph pageId="12" pageNumber="107">
Figure 14. Radulae,
<taxonomicName lsidName="M. tensiftensis" pageId="12" pageNumber="107" rank="species" species="tensiftensis">M. tensiftensis</taxonomicName>
sp. n. A, E a spring near Lalla Takerkoust dam B, C a pond near Lahjar Spring D, F ditch in Sidi Bouzid. A Radular ribbon B, D Rows of radular teeth C Lateral tooth and inner marginal teeth E Central tooth F Outer marginal teeth.
</paragraph>
</caption>
<paragraph lastPageId="14" lastPageNumber="109" pageId="12" pageNumber="107">
Animal darkly pigmented except for neck and tentacles (Figure 15G). Ctenidium well-developed, with 23-27 gill filaments, occupying nearly entire length of pallial cavity. Osphradium elongate, positioned opposite middle of ctenidium (Figure 15A). Stomach slightly longer than wide, with two chambers almost equal in size; style sac longer than wide, surrounded by unpigmented intestine (Figure 15B; Suppl. material 1: Table 3). Glandular oviduct approx. three times as long as wide. Albumen gland
<pageBreakToken pageId="13" pageNumber="108" start="start">longer</pageBreakToken>
than capsule gland (Figure 15
<normalizedToken originalValue="DF">D-F</normalizedToken>
). Bursa copulatrix elongate, two to three times longer than wide, with a duct shorter than bursal length. Renal oviduct unpigmented, coiled, making 2-3 loops. Seminal receptacle elongate, with a short duct, joining renal oviduct just above the insertion point with bursal duct (Figure 15
<normalizedToken originalValue="EF">E-F</normalizedToken>
; Suppl. material 1: Table 4). Prostate gland approx. two times longer than wide, bean-shaped; seminal duct entering the posterior region; pallial vas deferens emerging close to its anterior edge. Penis gradually tapering, attached to the area close to the right eye. Penis dark pigmented, tapering. Penial appendix dark pigmented, shorter than penis, base narrow, medially position on inner edge of penis. Terminal gland occupying the whole distal end of the appendix (Figure 15
<normalizedToken originalValue="GI">G-I</normalizedToken>
; Suppl. material 1: Table 5). Nervous
<pageBreakToken pageId="14" pageNumber="109" start="start">system</pageBreakToken>
with black granules, elongate (mean RPG ratio 0.64; Suppl. material 1: Table 6); cerebral ganglia approx. equal in size; ganglia darker than connectives and commissures (Figure 15C).
</paragraph>
<caption pageId="14" pageNumber="109">
<paragraph pageId="14" pageNumber="109">
Figure 15. Anatomical structures,
<taxonomicName lsidName="M. tensiftensis" pageId="14" pageNumber="109" rank="species" species="tensiftensis">M. tensiftensis</taxonomicName>
sp. n. A, D, E, G, H ditch in Sidi Bouzid B, F, I a pond near Lahjar Spring C a spring near Lalla Takerkoust dam. A Ctenidium B Stomach C Partial nervous system D Pallial oviduct E, F Bursa copulatrix and seminal receptacle G Head with penis H, I Prostate gland. RO renal oviduct SR seminal receptacle.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="14" pageNumber="109" type="etymology">
<paragraph pageId="14" pageNumber="109">Etymology.</paragraph>
<paragraph pageId="14" pageNumber="109">
The name
<taxonomicName genus="Truncatelloidea" lsidName="Truncatelloidea tensiftensis" pageId="14" pageNumber="109" rank="species" species="tensiftensis">tensiftensis</taxonomicName>
refers to the hydrological basin (Tensift) where this species was collected.
</paragraph>
</subSubSection>
<subSubSection pageId="14" pageNumber="109" type="distribution">
<paragraph pageId="14" pageNumber="109">Distribution.</paragraph>
<paragraph pageId="14" pageNumber="109">This species was found in ponds, springs, and ditches in proximal localities of the Tensift River basin in northwestern Morocco.</paragraph>
</subSubSection>
<subSubSection pageId="14" pageNumber="109" type="remarks">
<paragraph pageId="14" pageNumber="109">Remarks.</paragraph>
<paragraph pageId="14" pageNumber="109">
Shells of this species vary in size (2.4-5.1 mm shell height) and, accordingly, two morphotypes can be distinguished in all populations. One morphotype comprises small to medium-sized shells (2.4-4.0 mm shell height), with slightly shouldered spire whorls and a thick aperture. This morphotype is found in Lahjar, Talkount, and Lalla Takerkoust with an average shell length of 3.5 mm. The second larger group (4.0-5.1 mm shell height) comprises shells with five sloping spire whorls and a thin aperture. This morphotype is well represented in the populations from Sidi Bouzid Springs and Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
with an average shell length of 4.1 mm. Despite this morphological variation within the species, the estimated genetic distance was 0% for COI.
</paragraph>
<paragraph pageId="14" pageNumber="109">
Two morphotypes of male reproductive organs were also observed in dissected specimens. The most represented is that with a long penis, large appendix, and large prostate gland (localities of Lahjar near Essaouira, Sidi Bouzid, and Lalla Takerkoust dam). However, other dissected males showed a smaller retracted penis and a small degraded prostate gland (localities of Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
and Talkount). We observed that this second group of males contained parasites known to cause castration in host snails (
<bibRefCitation author="Lim, H-K" editor="Dawes, B" journalOrPublisher="Molecular Biology and Evolution" pageId="19" pageNumber="114" title="Academic Press" url="https://doi.org/10.1016/S0065-308X(08)60175-X" year="1972">Lim and Heyneman 1972</bibRefCitation>
,
<bibRefCitation author="Combes, C" journalOrPublisher="Institut Pasteur d'Algerie" pageId="17" pageNumber="112" pagination="153 - 193" title="Control of biomedically important molluscs. Archives de l'Institut Pasteur d'Algerie." volume="55" year="1986">Combes and Cheng 1986</bibRefCitation>
,
<bibRefCitation author="Mouahid, A" journalOrPublisher="Annals of Tropical Medicine &amp; Parasitology" pageId="19" pageNumber="114" pagination="341 - 348" title="Interference of Echinoparyphiumelegans with the host-parasite system Bulinustruncatus - Schistosomabovis in natural conditions." url="https://doi.org/10.1080/00034983.1990.11812478" volume="84" year="1990">Mouahid and Mone 1990</bibRefCitation>
,
<bibRefCitation author="Ashby, B" journalOrPublisher="Evolution" pageId="16" pageNumber="111" pagination="2234 - 2244" title="Parasitic castration promotes coevolutionary cycling but also imposes a cost on sex." url="https://doi.org/10.1111/evo.12425" volume="68" year="2014">Ashby and Gupta 2014</bibRefCitation>
) and propose this as the cause of such variation (see Figure 16). According to our observations in parasitized specimens of different populations, male and female genitalia seem more affected than other organs.
</paragraph>
<paragraph pageId="14" pageNumber="109">
<taxonomicName class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria tensiftensis" order="Littorinimorpha" pageId="14" pageNumber="109" phylum="Mollusca" rank="species" species="tensiftensis">Mercuria tensiftensis</taxonomicName>
sp. n. is characterized by its long shell (the longest shells among Moroccan
<taxonomicName class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria" order="Littorinimorpha" pageId="14" pageNumber="109" phylum="Mollusca" rank="genus">Mercuria</taxonomicName>
species) and its large and gradually tapering penis with a terminal gland occupying the entire distal end of the penial appendix. The new species differs from
<taxonomicName lsidName="M. midarensis" pageId="14" pageNumber="109" rank="species" species="midarensis">M. midarensis</taxonomicName>
sp. n. in its shorter penis (two times vs. three times longer than appendix in
<taxonomicName lsidName="M. tensiftensis" pageId="14" pageNumber="109" rank="species" species="tensiftensis">M. tensiftensis</taxonomicName>
sp. n. and
<taxonomicName lsidName="M. midarensis" pageId="14" pageNumber="109" rank="species" species="midarensis">M. midarensis</taxonomicName>
sp. n., respectively) (Suppl. material 1: Table 5), from
<taxonomicName lsidName="M. targouasensis" pageId="14" pageNumber="109" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
in its more elongate bursa copulatrix (Suppl. material 1: Table 4) and from
<taxonomicName lsidName="M. similis" pageId="14" pageNumber="109" rank="species" species="similis">M. similis</taxonomicName>
in its longer shell and its larger and longer penis. These morphological and anatomical differences were supported by molecular data. Hence, the mean genetic distance between
<taxonomicName lsidName="M. tensiftensis" pageId="14" pageNumber="109" rank="species" species="tensiftensis">M. tensiftensis</taxonomicName>
sp. n. and
<taxonomicName lsidName="M. similis" pageId="14" pageNumber="109" rank="species" species="similis">M. similis</taxonomicName>
was 6.4% and between the former and
<taxonomicName lsidName="M. targouasensis" pageId="14" pageNumber="109" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
and
<taxonomicName lsidName="M. midarensis" pageId="14" pageNumber="109" rank="species" species="midarensis">M. midarensis</taxonomicName>
sp. n. were 6.0% and 6.9%, respectively.
</paragraph>
</subSubSection>
<subSubSection pageId="14" pageNumber="109" type="ecology">
<paragraph pageId="14" pageNumber="109">Ecology.</paragraph>
<paragraph pageId="14" pageNumber="109">
<taxonomicName class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria tensiftensis" order="Littorinimorpha" pageId="14" pageNumber="109" phylum="Mollusca" rank="species" species="tensiftensis">Mercuria tensiftensis</taxonomicName>
sp. n. was found in ditches used for irrigation, springs, and ponds, attached to stones or dead branches in the water. Most of these localities, including the type one, are small water bodies under risk of desiccation or destruction. Co-occurring species were
<taxonomicName class="Gastropoda" family="Lymnaeidae" genus="Galba" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galba truncatula" pageId="14" pageNumber="109" phylum="Mollusca" rank="species" species="truncatula">Galba truncatula</taxonomicName>
,
<taxonomicName class="Gastropoda" family="Melanopsidae" genus="Melanopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Melanopsis praemorsa" order="Neogastropoda" pageId="14" pageNumber="109" phylum="Mollusca" rank="species" species="praemorsa">Melanopsis praemorsa</taxonomicName>
, and
<taxonomicName class="Gastropoda" family="Physidae" genus="Physella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Physella acuta" pageId="14" pageNumber="109" phylum="Mollusca" rank="species" species="acuta">Physella acuta</taxonomicName>
.
</paragraph>
<caption pageId="14" pageNumber="109">
<paragraph pageId="14" pageNumber="109">
Figure 16. Head and prostate glands of non-parasitized and parasitized males,
<taxonomicName lsidName="M. tensiftensis" pageId="14" pageNumber="109" rank="species" species="tensiftensis">M. tensiftensis</taxonomicName>
sp. n. A, B Non-parasitized male, a spring near Lalla Takerkoust dam C, D Non-parasitized male, ditch in Sidi Bouzid E, F Parasitized male, ditch in Sidi Bouzid
<normalizedToken originalValue="GJ">G-J</normalizedToken>
Parasitized males, ditch in Agadir
<normalizedToken originalValue="Ntachraft">N'tachraft</normalizedToken>
. Pr prostate gland.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>