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<mods:title id="DD35D10F1FB91A93B2CEC150974437E5">The genus Mercuria Boeters, 1971 in Morocco: first molecular phylogeny of the genus and description of two new species (Caenogastropoda, Truncatelloidea, Hydrobiidae)</mods:title>
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<mods:namePart id="1631AC98D1D6742473035347D6EBEA9C">Boulaassafer, Khadija</mods:namePart>
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<mods:namePart id="E1CA9FD407BE79722CD5E5B787A366D3">Ghamizi, Mohamed</mods:namePart>
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<mods:namePart id="41914D3662C1009677BCA67D7E4ABA2A">Delicado, Diana</mods:namePart>
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<treatment id="3800B2819E78453D93DAA5440B9AF549" ID-GBIF-Taxon="147440669" LSID="urn:lsid:plazi:treatment:3800B2819E78453D93DAA5440B9AF549" httpUri="http://treatment.plazi.org/id/3800B2819E78453D93DAA5440B9AF549" lastPageId="7" lastPageNumber="102" pageId="3" pageNumber="98">
<subSubSection id="B755847BA0545AD676E86AD95F00BBFF" pageId="3" pageNumber="98" type="nomenclature">
<paragraph id="05EF77B460D9DC9B1FCBEDB8958B74D1" pageId="3" pageNumber="98">
<taxonomicName id="51C50058982B0D710F7B011BE2F10BCF" ID-CoL="3ZTHS" authority="Gloeer, Boeters &amp; Walther, 2015" authorityName="Gloeer, Boeters &amp; Walther" authorityYear="2015" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria targouasensis" order="Littorinimorpha" pageId="3" pageNumber="98" phylum="Mollusca" rank="species" species="targouasensis">
Mercuria targouasensis
<normalizedToken id="31989B9A6A881C8820A68AB716171845" originalValue="Glöer">Gloeer</normalizedToken>
, Boeters &amp; Walther, 2015
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="E366346C853AE4F5A7C18C71E0C5F91B" pageId="3" pageNumber="98" type="reference_group">
<paragraph id="EFE4861CB8DDD9D0FD693192519DC3ED" pageId="3" pageNumber="98">
<taxonomicName id="FDFBD16310142DF5B8C88659A38F8B60" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria confusa" order="Littorinimorpha" pageId="3" pageNumber="98" phylum="Mollusca" rank="species" species="confusa">Mercuria confusa</taxonomicName>
Backhuys &amp; Boeters, 1974: 113
</paragraph>
</subSubSection>
<subSubSection id="AF1C9B6889784F9B4FBD498AB0A64DB5" pageId="3" pageNumber="98" type="materials_examined">
<paragraph id="20054C6B7629A717662E28AE5E8E2D7C" pageId="3" pageNumber="98">Material.</paragraph>
<paragraph id="A0DE2FD0B19197AAFA2D383F44F94E8C" pageId="3" pageNumber="98">
Examined material. MOROCCO. MHNM 18 ZTMH10, UGSB 17912, Oum Rbii Springs, N of Khenifera, 01/06/2015 (
<geoCoordinate id="23E40EB02700BF6712D02690970DA5A1" direction="north" orientation="latitude" precision="1" value="33.053432">33°3.2059'N</geoCoordinate>
,
<geoCoordinate id="3F824F13DDB7503F16A43191AC078406" direction="west" orientation="longitude" precision="1" value="-5.414662">5°24.8797'W</geoCoordinate>
); MHNM 18 ZTMH11, UGSB 17955, a small ditch in Mirleft, 02/02/2015 (
<geoCoordinate id="F071B47F6BCF12629581D3BCA0543E05" direction="north" orientation="latitude" precision="1" value="29.583612">29°35.0167'N</geoCoordinate>
,
<geoCoordinate id="352D938A19C9FB20BB9C0E2F69E9D84D" direction="west" orientation="longitude" precision="1" value="-10.03075">10°1.845'W</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection id="79FBE9C136A81F0281C6F4E10358B38F" pageId="3" pageNumber="98" type="revised diagnosis">
<paragraph id="C8368582ADE6D980F1F51E4420E6F1AB" pageId="3" pageNumber="98">Revised diagnosis.</paragraph>
<paragraph id="9F72A453275EB63D69CB754399FEF39E" pageId="3" pageNumber="98">
Shell ovate-conic, whorls 3-5; periostracum whitish; body whorl occupying more than three-quarters of total shell length; aperture ovate; umbilicus narrow, partially covered by the inner lip; operculum brownish to slightly orange; central radular tooth formula 3
<normalizedToken id="5903D55AC29C0A6E6D5D4F815911605F" originalValue="C3/1">-C-3/1-</normalizedToken>
1; bursa copulatrix elongate, with a short duct; one seminal receptacle pyriform, with a short duct; penis gradually tapering, grey; penial appendix shorter than penis, grey, base wide, medially positioned on inner edge of penis; nervous system extremely elongated (mean RPG ratio = 0.70), gently black pigmented.
</paragraph>
</subSubSection>
<subSubSection id="8F4A2F3EF9660AEA7A15FBFB6858636D" lastPageId="4" lastPageNumber="99" pageId="3" pageNumber="98" type="description">
<paragraph id="3DF00B26CEBD511924400D3B4CCD1DEC" pageId="3" pageNumber="98">Description.</paragraph>
<paragraph id="5F9CEBF40BDED4AC0A9FB1CEE8942379" pageId="3" pageNumber="98">
Shell ovate-conic, whorls 3-5, height 2.63-3.43 mm (Figure 7
<normalizedToken id="B3ABB3E09067F366D1257F905218E3C6" originalValue="AC">A-C</normalizedToken>
; Suppl. material 1: Table 1). Periostracum whitish. Protoconch with two whorls, diameter ca. 600
<normalizedToken id="92310A1D3CEB6F5CAAC111FFCC0C660B" originalValue="µm">µm</normalizedToken>
; nucleus ca. 140
<normalizedToken id="E7D5A11FF0DF3394B90FB0632C7B36AF" originalValue="µm">µm</normalizedToken>
wide (Figure 7H); protoconch microsculpture granulated (Figure 7I). Teleoconch whorls convex, separated by deep sutures. Body whorl large, occupying three-quarters of total shell length. Aperture ovate, complete, in contact with the body whorl; inner lip thicker than outer lip; peristome margin straight (Figure 7D, E). Umbilicus narrow, partially covered by the inner lip.
</paragraph>
<caption id="A7DC8E4668742D16730928D0EB2B67F5" pageId="3" pageNumber="98">
<paragraph id="F5D000A90DF0BA557CE76C08C5E1DB09" pageId="3" pageNumber="98">
Figure 7. Shells and opercula,
<taxonomicName id="3C44E550CC459EB72BA1B941960341A6" lsidName="M. targouasensis" pageId="3" pageNumber="98" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
. A, B, E Shell, Oum Rbii Springs C, D Shell, a small ditch in Mirleft F, G Opercula (inner, outer sides), Oum Rbii Springs H, I Protoconch and detailed microsculpture of protoconch, Oum Rbii Springs.
</paragraph>
</caption>
<paragraph id="80A1D43CE0F276CA8CB6980E539EDAD1" pageId="3" pageNumber="98">
Operculum as for genus, light orange to brown, whorls 2; muscle attachment area near nucleus (Figure 7F, G). Radula length intermediate, ca. 800
<normalizedToken id="CC7460F595850FE12C571D3DDABB2709" originalValue="µm">µm</normalizedToken>
long (20% total shell length), seven times longer than wide, with approx. 50 rows of teeth (Fig. 8A; Suppl. material 1: Table 2). Central tooth formula 3
<normalizedToken id="71FF7FC058A6B708C976B257E7378716" originalValue="C3/1">-C-3/1-</normalizedToken>
1; central cusp tapered, long. Lateral teeth formula (4)3
<normalizedToken id="B59514691A571980BD495B264E9F8003" originalValue="C">-C-</normalizedToken>
3(4); central cusp wide, V-shaped (Figure 8B,
<normalizedToken id="6D850AE501D035F965831D28A4B358F5" originalValue="DF">D-F</normalizedToken>
). Inner and outer marginal teeth having 11-15 and 14-18 cusps, respectively (Figure 8C, F).
</paragraph>
<caption id="D7DC2E28C60A6173777D9BF39A8822D1" pageId="3" pageNumber="98">
<paragraph id="2693EA8BB6FC029CB23FEFAF10707A24" pageId="3" pageNumber="98">
Figure 8. Radulae,
<taxonomicName id="77A89095F41C60462729348B22449237" lsidName="M. targouasensis" pageId="3" pageNumber="98" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
.
<normalizedToken id="BAFCA3AD70CDF03451EFF3FB5BF9B4A0" originalValue="AD">A-D</normalizedToken>
Oum Rbii Springs E, F A small ditch in Mirleft. A Radular ribbon B, E Central tooth C, D, F Inner marginal, outer marginal, and lateral teeth.
</paragraph>
</caption>
<paragraph id="F666099604010661B1E1A70775CEC3AE" lastPageId="4" lastPageNumber="99" pageId="3" pageNumber="98">
Animal black pigmented except for pale area surrounding eye lobes and neck (Figure 9G, H). Ctenidium well-developed, with 19-25 gill filaments, occupying almost entire length of the pallial cavity. Osphradium elongate, positioned approximate middle of ctenidium (Figure 9A). Stomach nearly as long as wide; style sac slightly shorter than stomach, surrounded by an unpigmented intestine (Figure 9B; Suppl. material 1: Table 3). Glandular oviduct three times longer than wide. Capsule gland longer and thicker than
<pageBreakToken id="AF7E7F3DCCB199463BECE3C3D0C6995B" pageId="4" pageNumber="99" start="start">albumen</pageBreakToken>
gland. Bursa copulatrix pyriform to elongate, with a short duct. Renal oviduct unpigmented, coiled, making three loops. Seminal receptacle pyriform, with a short duct, joining renal oviduct above the insertion point with bursal duct (Figure 9D, E; Suppl. material 1: Table 4). Prostate gland bean-shaped, ca. 2.5 times longer than wide (Figure 9F; Suppl. material 1: Table 5); 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. Penial appendix slightly pigmented, shorter than penis, base wide, middle positioned on inner edge of penis. Terminal gland large, occupying the whole distal end of the appendix (Figure 9
<normalizedToken id="2616E77B1B58D7745664B7BE546921C0" originalValue="GJ">G-J</normalizedToken>
; Suppl. material 1: Table 5). Nervous system gently pigmented, extremely elongated (mean RPG ratio = 0.70; Suppl. material 1: Table 6); cerebral ganglia equal in size and shape (Figure 9C).
</paragraph>
<caption id="1982CCA8F35E59DF1382FF524CD9DE10" pageId="4" pageNumber="99">
<paragraph id="C8AD3F10C79F33288C58604EE77EF64B" pageId="4" pageNumber="99">
Figure 9. Anatomical structures,
<taxonomicName id="3F4E83AC82E6E9FD0C38726331CB79BD" lsidName="M. targouasensis" pageId="4" pageNumber="99" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
.
<normalizedToken id="50704FBDA989BE4F09FADEEC74DBB255" originalValue="AF">A-F</normalizedToken>
, H A small ditch in Mirleft G,
<normalizedToken id="E4FBFAC95B6BC947977D1B3F1404B77A" originalValue="IJ">I-J</normalizedToken>
Oum Rbii Springs. A Ctenidium B Stomach C Partial nervous system D Pallial oviduct E Bursa copulatrix and seminal receptacle F Prostate gland G, H Head with penis I, J Head and penis drawings. RO renal oviduct SR seminal receptacle.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="D6A9993863EA32A4BA9DB9A5CB988802" pageId="4" pageNumber="99" type="distribution">
<paragraph id="40CB9B8BD374BB45178FBA42EAAE2DF1" pageId="4" pageNumber="99">Distribution.</paragraph>
<paragraph id="BC899A2327A79632E29D087474F39005" pageId="4" pageNumber="99">This species was found in coastal streams in southwestern Morocco and in a spring-fed habitat in the Middle Atlas.</paragraph>
</subSubSection>
<subSubSection id="6FF41D0D39CD9D4DEA7C5A1ADCD1065A" lastPageId="7" lastPageNumber="102" pageId="4" pageNumber="99" type="remarks">
<paragraph id="9BEFA2C1D5ECECBF306A6586E2D28A7B" pageId="4" pageNumber="99">Remarks.</paragraph>
<paragraph id="14FC43B4577607D665D5B32F72448A78" lastPageId="6" lastPageNumber="101" pageId="4" pageNumber="99">
The morphological and anatomical descriptions presented here are based on specimens collected at two sites: one in the Mirleft region, 70 km from the
<pageBreakToken id="CDEF94671220209854597A622A79F345" pageId="5" pageNumber="100" start="start">type</pageBreakToken>
locality, (i.e., ford Oued Assaka), and another, in a more remote place in the Middle Atlas Mountains. The population collected in the surroundings of Mirleft may correspond to the species
<taxonomicName id="B5836EFED7D5BB29912D51FB05B8CB57" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria" order="Littorinimorpha" pageId="5" pageNumber="100" phylum="Mollusca" rank="genus">Mercuria</taxonomicName>
<normalizedToken id="D030655C43D569A9317A1BC4C2B59E1D" originalValue="mirlheftensis">'mirlheftensis'</normalizedToken>
(nomen nudum) from the same area suggested by
<bibRefCitation id="74D81CAEFAF37F55D0D8E4005620FC51" pageId="5" pageNumber="100">
<normalizedToken id="0EC2C8934EF52429323E38A2F70C2722" originalValue="García">Garcia</normalizedToken>
et al. (2010)
</bibRefCitation>
. However, the name
<taxonomicName id="3C67B3B2D4FF42C3BA48EC9997C1E912" genus="M." lsidName="M." pageId="5" pageNumber="100" rank="genus">M.</taxonomicName>
<normalizedToken id="D383AC5C45A1BA15D19236A08AAC71D8" originalValue="mirlheftensis">'mirlheftensis'</normalizedToken>
is not valid. Specimens collected in the Mirleft area resemble specimens from the type locality of
<taxonomicName id="15D061629244C8425A71FE9C530B5189" lsidName="M. targouasensis" pageId="5" pageNumber="100" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
regarding the shape of the penis and prostate, and also of the female genitalia, especially bursa copulatrix shape. Based on the geographic proximity of these two localities and the similarity in shell and anatomical characters of their specimens, we assigned the population from Mirleft to
<taxonomicName id="777F9B0222FF08F570F38CA015696C7A" lsidName="M. targouasensis" pageId="5" pageNumber="100" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
. Specimens from Oum
<pageBreakToken id="270459C34A7926E789C44C44D6841DED" pageId="6" pageNumber="101" start="start">Rbii</pageBreakToken>
(Middle Atlas) were also tentatively assigned to this species based on shell and morphological similarities and a short genetic distance (1.3%) between this and the Mirleft population. However, this assignment needs confirmation in future systematic studies on
<taxonomicName id="5BD3534164B84661B9BF9D557C0205F0" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria" order="Littorinimorpha" pageId="6" pageNumber="101" phylum="Mollusca" rank="genus">Mercuria</taxonomicName>
, which should include these and other Mediterranean species.
</paragraph>
<paragraph id="7C241F3A6178119E77E8DFEF88FE7533" lastPageId="7" lastPageNumber="102" pageId="6" pageNumber="101">
<taxonomicName id="50AB5D32599D661C5CB0570CB88CDDB5" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria targouasensis" order="Littorinimorpha" pageId="6" pageNumber="101" phylum="Mollusca" rank="species" species="targouasensis">Mercuria targouasensis</taxonomicName>
and
<taxonomicName id="82245BC7E8C6E8B42264FD70C9F6FD4F" lsidName="M. midarensis" pageId="6" pageNumber="101" rank="species" species="midarensis">M. midarensis</taxonomicName>
sp. n. are sister species and differ molecularly by 2.1%-3.4% (mean sequence divergence 2.8%). The two species are close in shell dimensions but differ in other shell features such as the relative size of the body
<pageBreakToken id="595DF067E4E2DECB19014F32513C70EB" pageId="7" pageNumber="102" start="start">whorl</pageBreakToken>
(larger in
<taxonomicName id="542E60C4261706308DACA2416473ECAD" lsidName="M. targouasensis" pageId="7" pageNumber="102" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
) or the umbilicus (wider in
<taxonomicName id="BB5B51A301E96D091257A693FAC2A98D" lsidName="M. midarensis" pageId="7" pageNumber="102" rank="species" species="midarensis">M. midarensis</taxonomicName>
sp. n.). They also differ anatomically;
<taxonomicName id="1D40787F90A61D884D73DBA74735FF76" class="Gastropoda" family="Hydrobiidae" genus="Mercuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mercuria midarensis" order="Littorinimorpha" pageId="7" pageNumber="102" phylum="Mollusca" rank="species" species="midarensis">Mercuria midarensis</taxonomicName>
sp. n. has typically a strap-like penis, 2.5 times longer than head length, a small penial appendix with narrow insertion into the penis, and an elongate bursa copulatrix, whereas in
<taxonomicName id="8C48BBE7F7AFD45CC80D3140FA2BE4F6" lsidName="M. targouasensis" pageId="7" pageNumber="102" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
, the penis is more often gradually tapering, equal or 1.5 times longer than head length, the penial appendix is larger with a wider insertion, and the bursa copulatrix is pyriform to elongate. These two species also differ in the number of cusps on radular teeth (Suppl. material 1: Table 2).
</paragraph>
</subSubSection>
<subSubSection id="B721E822E1C16F16DC50E7D71FAB78A2" pageId="7" pageNumber="102" type="ecology">
<paragraph id="8A05A48A8688A6C456AA0F9C409173A8" pageId="7" pageNumber="102">Ecology.</paragraph>
<paragraph id="1548042F526F53FCBE81B1ABD426B87A" pageId="7" pageNumber="102">
In the new localities of
<taxonomicName id="51989EAB63EC7D66E4BF355D1A1895F7" lsidName="M. targouasensis" pageId="7" pageNumber="102" rank="species" species="targouasensis">M. targouasensis</taxonomicName>
, this species was found attached to stones in a saltwater spring in the Middle Atlas (ca. 1,200 m a.s.l. altitude, and 37.9 PSU, practical salinity unit) and in the sediment of a ditch in the region of Mirleft cohabiting with
<taxonomicName id="1ACA6E84900E2587ED36CB57F6DD7DEF" class="Gastropoda" family="Melanopsidae" genus="Melanopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Melanopsis praemorsa" order="Neogastropoda" pageId="7" pageNumber="102" phylum="Mollusca" rank="species" species="praemorsa">Melanopsis praemorsa</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
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