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Published by he Hunga ian Na u al His o y Museum
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Ac a Zoologica Academiae Scien ia um Hunga icae 69(3), pp. 303–312, 2023
DOI: 10.17109/AZH.69.3.303.2023
RADIX RUFESCENS (J. E. GRAY, 1822)
(GASTROPODA: LYMNAEIDAE), A NEW SPECIES
FOR OMAN AND ARABIAN PENINSULA
And zej Falniowski1, Aleksand a Jaszczyńska1,2 and Sebas ian Ho man3
1Depa men o Malacology, Ins i u e o Zoology and Biomedical Resea ch,
Jagiellonian Uni e si y, ul. G onos ajowa 9, 30–387 K aków, Poland
E-mail: [email p o ec ed]; h ps://o cid.o g/0000-0002-3899-6857
2 Depa men o In e eb a e E olu ion, Ins i u e o Zoology and Biomedical Resea ch,
Jagiellonian Uni e si y, G onos ajowa 9, 30-387 K aków, Poland
E-mail: [email p o ec ed]; h ps://o cid.o g/0000-0002-9395-9696
3Depa men o Compa a i e Ana omy, Ins i u e o Zoology and Biomedical Resea ch,
Jagiellonian Uni e si y, ul. G onos ajowa 9, 30–387 K aków, Poland;
E-mail: [email p o ec ed]; h ps://o cid.o g/0000-0001-6044-3055
The auna o eshwa e gas opods o he A abian Peninsula, a dese land, is nei he ich
no ex ensi ely s udied. Only i e species o he amily Lymnaeidae ha e been eco ded
om his subcon inen so a . In Feb ua y 2023, a ew specimens o Radix u escens (J. E. G ay,
1822) ha e been collec ed om he s eam a Wadi Tiwi, NE Oman. The shell, emale pa o
he ep oduc i e o gans, p os a e, pa aepu ium and phallo eca a e p esen ed. Cy och ome
oxidase subuni I (COI) sequences con i med he assignmen o hese specimens o R. u es-
cens, known om Nepal, Bangladesh, and Myanma , no om he A abian Peninsula. Ei he
R. u escens is he six h lymnaeid species inhabi ing A abia, o a leas some o he eco ds o
R. na alensis (K auss, 1848), whose shells a e ha dly disce nible om he ones o R. u escens,
should be in e p e ed as misiden i ied R. u escens. Thousands o yea s o he ade ela ions
be ween Oman and India sugges acciden al anspo a ion o his snail by ade essels.
Keywo ds: shell, ep oduc i e o gans, cy och ome oxidase subuni I (COI), m DNA, mo-
lecula ela ionships
INTRODUCTION
Lymnaeidae Ra inesque, 1815, a globally dis ibu ed amily o eshwa-
e he e ob anch snails, o mo e han cen u y ha e been widely known as
showing inc edible a iabili y o hei shells. This was he eason why he lis
o synonyms in his g oup is ex emely long. The classic esea ch o Bake
(1911) on he No h Ame ican Lymnaeidae, and Roszkowski (1914) on he
lymnaeids om he p o undal o Lac Léman in oduced he lymnaeid axon-
omy based on he ana omical cha ac e s. Such s udies we e summa ized and
comple ed by Hubendick (1951), and con inued (e.g., Falniowski 1980a, b, c,
1981). Howe e , many ambigui ies s ill emained in he lymnaeid axonomy,
mos o hem sol ed la e wi h applica ion o he molecula da a (e.g., Co -
eal e al. 2010, Vina ski e al. 2020, Sai o e al. 2021).
Ac a Zool. Acad. Sci. Hung. 69, 2023
304 FALNIOWSKI, A., JASZCZYŃSKA, A. AND HOFMAN, S.
The A abian Peninsula inhabi s a unique assemblage o animal species,
he esul o he geologic and clima ic his o y o his egion. I s auna may
o igina e om Eas e n A ica, sou h-cen al Asia, o sou h-wes e n Asia.
The A abian Peninsula, a dese land in gene al, inhabi s a mode a ely ich
eshwa e auna (Ga cia e al. 2015). F om among he Lymnaeidae, only
i e species ha e been eco ded so a : Galba unca ula (O. F. Mülle , 1774),
Galba obus a Vina ski, 2018, Radix na alensis (K auss, 1848), Radix au icula -
ia (Linnaeus, 1758), and S agnicola palus is (O. F. Mülle , 1774) (Smi h 1894,
B own & Gallaghe 1985, Neube 1998, Vina ski 2018). Lymnaea a abica E.
A. Smi h, 1894 om Saudi A abia, s udied by Bin Dajem (2012), was ecen ly
conside ed a synonym o Radix na alensis (K auss, 1848). In Feb ua y 2023, a
ew specimens o Radix Mon o , 1810 and Melanoides Oli ie , 1804 we e col-
lec ed in Wadi Tiwi in Oman.
MATERIAL AND METHODS
The snails we e collec ed 4 h o Feb ua y 2023 a he s eam, om he s ones
and among he algae, a he lowe pa o Wadi Tiwi canyon in NE Oman, 22°48’15”N,
59°14’45”E, 20 m a.s.l. (Fig. 1). Wadi Tiwi is si ua ed in Sha kiyah, Eas e n Oman Moun-
Fig. 1. Wadi Tiwi, s eam in i s lowe pa
Ac a Zool. Acad. Sci. Hung. 69, 2023
305
RADIX RUFESCENS (GASTROPODA: LYMNAEIDAE) NEW FOR OMAN AND ARABIA
ains, ising o 2300 m al i ude. Wadi (A abic) means d y i e alley. Wadi Tiwi is abou
eigh km long. I s loo has e oded down h ough Eocene and Palaeocene o he op o he
ophioli e which is composed o highly se pen inised man le sequence ha zbu gi e. The en-
i e c us al sequence o he ophioli e has been e oded o , p io o deposi ion o he o e ly-
ing Palaeocene limes ones (Sea le 2019).The lowe pa o he Wadi, whe e he snails ha e
been ound, has some small blue pools, lined wi h palms and eed-beds.
The snails we e collec ed by hand and wi h a small sie e and ixed in 80% analy ically
pu e e hanol, changed wice. The shells we e pho og aphed wi h Canon EOS 50D came a
wi h Canon Mac o Lens. The snails we e dissec ed unde a Nikon SMZ18 mic oscope wi h
a da k ield, he ana omical s uc u es pho og aphed wi h Canon EOS 50D came a unde
a Nikon SMZ18 mic oscope wi h a da k ield. Measu emen s o he p aepu ium and penis
shea h we e done on he pho og aph wi h cu ime e . DNA was ex ac ed om whole
specimens; issues we e hyd a ed in TE bu e (3 × 10 min); hen, o al genomic DNA was
ex ac ed wi h he SHERLOCK ex ac ion ki (A&A Bio echnology), and he inal p oduc
was dissol ed in 20 μl o is-EDTA (TE) bu e . The ex ac ed DNA was s o ed a −80 °C
a he Depa men o Malacology, Ins i u e o Zoology and Biomedical Resea ch, Jagiel-
lonian Uni e si y in K aków (Poland). Mi ochond ial cy och ome oxidase subuni I (COI)
was sequenced. De ails o PCR condi ions, p ime s used, and sequencing we e gi en by
Sza owska e al. (2016).
In he phylogene ic analysis, sequences om GenBank we e used (Fig. 4). Sequences
we e ini ially aligned in he MUSCLE (Edga 2004) P og am in MEGA 7 (Kuma e al. 2016)
and hen checked in Bioedi 7.1.3.0 (Hall 1999). Unco ec ed p-dis ances we e calcula ed
in MEGA 7. The es ima ion o he p opo ion o in a ian si es and he sa u a ion es
o en i e da a se s (Xia 2000, Xia e al. 2003) we e pe o med using DAMBE (Xia 2013).
The da a we e analysed using app oaches based on Bayesian in e ence (BI) and Maximum
Likelihood (ml). Fo RAxML analysis, he jModelTes 2 ia he CIPRES Science Ga eway
(Mille e al. 2010) was used o ind he bes - i ing model, he model HKY+G was used.
The ML analysis was conduc ed in RAxML-NG . 0.8.0 (Kozlo e al. 2019) ia web se -
ice a ailable a h ps:// axml-ng. i al-i .ch/, wi h 10 andom and 10 pa simony s a ing
ees. In he BI analysis, he K81+G model o nucleo ide subs i u ion was applied in ee
econs uc ion. Model was selec ed using M ModelTes 2.4 (Nylande 2004). The analyses
we e un using MRBAYES . 3.2.7a (Ronquis e al. 2012) wi h de aul s o mos p io s. Two
simul aneous analyses we e pe o med, each wi h 10,000,000 gene a ions, wi h one cold
chain and h ee hea ed chains, s a ing om andom ees and sampling he ees e e y
1,000 gene a ions. The i s 25% o he ees we e disca ded as bu n-in. The analyses we e
summa ized as a 50% majo i y- ule consensus ee. Con e gence was checked in TRACER
.1.7.1 (Rambau e al. 2018), in all cases E ec i e Sample Size exceeded 200. FigT ee . 1.4.4
(Rambau 2010) was used o isualize he ees.
RESULTS
The shell (Fig. 2) wi h acu e spi e, high as o his genus, he ape u e
b oad and a iable in ou line; he shell esembles he ones p esen ed o R.
u escens in Vina ski e al. (2020). The albuminoid gland (Fig. 3A) la and
a he b oad, he co pus py i o mis sho and bulky, bu sa copula ix small
and wi hou a disce nible duc , p os a e na ow, sligh ly b oadened in i s dis-
al pa , he as de e ens ou le close o he ma gin o he e minal pa o
Ac a Zool. Acad. Sci. Hung. 69, 2023
306 FALNIOWSKI, A., JASZCZYŃSKA, A. AND HOFMAN, S.
he p os a e. The p aepu ium (Fig. 3B) p oximally cylind ical, dis ally g adu-
ally na owing, wi h spo s o black pigmen , he penis shea h pigmen less,
wi h mode a ely b oadened p oximal end. The leng h p opo ion be ween
he p aepu ium and penis shea h was 1.25, compa ed wi h 1.09–1.27 eco ded
o . R. u escens by Vina ski e al. (2020).
The maximum likelihood ee o COI (Fig. 4) clus e s ou Omanian speci-
men (boo s ap suppo 94%, Bayesian p obabili y 0.99) wi hin R. u escens, all
he o he specimens o his species a e om Nepal, Bangladesh, and Myanma .
The p-dis ance wi hin his g oup was 0.017. The Omanian specimen di e ed
om he o he R. u escens by p-dis ance 0.007–0.036. The sis e species o R.
u escens is R. na alensis. p-dis ance be ween hese wo species equals 0.092.
Fig. 2. Shells and man le pigmen a ion o Radix u escens. Scale ba : 5 mm
Ac a Zool. Acad. Sci. Hung. 69, 2023
307
RADIX RUFESCENS (GASTROPODA: LYMNAEIDAE) NEW FOR OMAN AND ARABIA
DISCUSSION
Ou da a clea ly iden i ied ou Omanian specimens as belonging o Radix
u escens. Gene ic di e ences be ween his specimen and he o he sequences
R. u escens we e much smalle han be ween R. u escens and i s sis e spe-
cies, R. na alensis. Also leng h p opo ion be ween he p aepu ium and penis
shea h con i med his species assignmen . This is he i s eco d o Radix u e-
scens no only in Oman, bu in all A abian Peninsula. Only i e species o he
Lymnaeidae, wo o hem ep esen ing he genus Radix, ha e been eco ded
(Smi h 1894, B own & Gallaghe 1985, Neube 1998): R. na alensis, molecu-
Fig. 3. Rep oduc i e o gans o Radix u escens: A = en ally, B–C = p aepu ium and phal-
lo heca (ag – albuminoid gland, cp – co pus py i o mis, hg – hae maph odi e gland, ng
– nidamen al gland, ph – phallo heca (penis shea h), p – p os a e, p ae – p aepu ium, u
– u e us, – agina, d – as de e ens). Scale ba : 1 mm
Ac a Zool. Acad. Sci. Hung. 69, 2023
308 FALNIOWSKI, A., JASZCZYŃSKA, A. AND HOFMAN, S.
la ly iden i ied so a only in A ica (Sai o e al. 2021), and R. au icula ia, wi h
a wide Eu asian ange. Su p isingly, R. euph a ica (Mousson, 1874), inhabi -
ing I an sepa a ed om Oman only by ela i ely na ow O muz S aigh , has
Fig. 4. Maximum likelihood ee compu ed o COI; boo s ap suppo s/Bayesian p ob-
abili ies and GB numbe s a e gi en
Fig. 5. Hypo he ical way o in asion o R. u escens, a e Sai o e al. (2021), modi ied
Ac a Zool. Acad. Sci. Hung. 69, 2023
309
RADIX RUFESCENS (GASTROPODA: LYMNAEIDAE) NEW FOR OMAN AND ARABIA
no been eco ded om he A abian Peninsula. Thus R. u escens would be
he six h lymnaeid species in he A abia. Howe e , he shells o R. u escens
a e p ac ically indis inguishable om he ones o R. na alensis (e.g., Vina ski
e al. 2020). Thus, ei he R. u escens inhabi s Wadi Tiwi (and some o he s ill
uns udied locali ies), o simply his is he i s p ope ly iden i ied specimen
o his species, and he eco ds o R. na alensis (a leas some o hem) e e o
R. u escens. On he o he hand, he supposed occu ence o R. na alensis in
Oman may be he esul o mig a ion om he Ho n o A ica Ho spo and he
Eas e n A omon ane Ho spo (Mi e meie e al. 2004).
The e olu iona y his o y o Radix and main coloniza ion ou es a e well
known (Sai o e al. 2021). The i s di e si ica ion a ea was si ua ed in eas In-
dia, he u escens/na alensis g oup spli hen in o wo lines. The ances o o R.
na alensis p obably mo ed in o wes e n India, and la e o A ica, R. u escens e-
mained in Nepal and Bangladesh bo h NE o he Indian Subcon inen , as well as
in Myanma a Malay peninsula. How R. u escens colonized Wadi Tiwi emains
enigma ic. One o he excellen maps o Sai o e al. (2021) we ha e comple ed
wi h ou inding (Fig. 5). The known a ea o occu ence o R. u escens may be
much wide han he molecula ly iden i ied indi iduals con i m, hus he pop-
ula ion disco e ed by us may be a pa o he ange co e ing pe haps he en i e
Indian Peninsula, sou he n Pakis an and I an, and a la ge pa o he A abian
Peninsula. Ob iously, mig a ion om he wes e n coas o India would be mo e
p obable. Some olde mig a ions may ha e been possible a he ime when he
Pe sian Gul was educed o a se ies o eshwa e lakes (Lambeck 1996).
Howe e , small gene ic di e ences be ween R. u escens om Oman and
om Nepal, Bangladesh, and Myanma sugges much younge episodes o
mig a ion. A abian Peninsula, especially Oman, since abou 2,200 BC (Paine
2013) was in ade ela ions wi h India. A abian dhow sailed wi h monsoon
winds (Miles 1919, LeBa on Bowen 1949, Villie s 2018), con eying ood,
imbe , e c., hus he acciden al anspo a ion o eggs o adul specimens o
aqua ic snails was p obable. Especially he ba els and cans wi h d inkable
wa e o he c ew, insed be o e e illing in he s eam in he ou le o Wadi
Tiwi (si ua ed no a om Su , o cen u ies he main po a his egion) we e
sui able o such anspo a ion. The an h opogenic anspo a ion o snails is
ex emely common (P es on e al. 2022). Also anspo a ion by bi ds canno
be excluded (Cha alambidou & San ama ia 2002, Figue ola & G een 2002).
*
Acknowledgmen s – The s udy was suppo ed by he g an N18/DBS/000012 om
he Ins i u e o Zoology and Biomedical Resea ch o Jagiellonian Uni e si y. We a e g a e-
ul o D M. V. Vina ski o his aluable co ec ions and addi ions o he o me e sion o
ou manusc ip .
Ac a Zool. Acad. Sci. Hung. 69, 2023
310 FALNIOWSKI, A., JASZCZYŃSKA, A. AND HOFMAN, S.
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