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New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 1
New Bythinella (Gastropoda, Bythinellidae)
species from western Turkey
Mehmet Zeki Yıldırım1, Ümit Kebapçı2, Seval Bahadır Koca3, Arzu Yüce4
1 Faculty of Education, Mehmet Akif Ersoy University, Burdur, Turkey 2 Faculty of Arts and Sciences, Mehmet
Akif Ersoy University, Burdur, Turkey 3 Faculty of Fisheries, Süleyman Demirel University, Eğirdir, Isparta,
Turkey 4 Kocaeli University, Hereke O.I. Uzunyol Vocational School, Kocaeli, Turkey
Corresponding author: Ümit Kebapçi (kebapci@mehmetakif.edu.tr)
Academic editor: R. Hershler|Received 8 July 2014|Accepted 30 December 2014|Published 4 February 2015
http://zoobank.org/791823C0-1143-48FF-B6B4-6DA47934A6F2
Citation: Yıldırım MZ, Kebapçı Ü, Bahadır Koca S, Yüce A (2015) New Bythinella (Gastropoda, Bythinellidae) species
from western Turkey. ZooKeys 481: 1–13. doi: 10.3897/zookeys.481.8225
Abstract
Bythinella anatolica sp. n., Bythinella istanbulensis sp. n., Bythinella magdalenae sp. n., and Bythinella
wilkei sp. n. from western Turkey are described herein. Illustrations of the shell and genitalia of the newly
described taxa, together with comparisons with previously known Bythinella taxa and a key to the species
from western Turkey, are also provided.
Keywords
Bythinella, new species, freshwater, springs, Turkey
Introduction
Bythinella Moquin-Tandon, 1856, the sole genus of the caenogastropod family
Bythinellidae (Szarowska 2006; Wilke et al. 2013), is composed of small sized (1–3
mm) species occurring almost exclusively in springs (rarely in upper courses of nutrient
poor montane streams or caves) having relatively cold waters below 10 °C (Boeters
1998). Although typically characterized by cylindrical (sometimes ovate-conic) shells
ZooKeys 481: 1–13 (2015)
doi: 10.3897/zookeys.481.8225
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Mehmet Zeki Yıldırım et al. / ZooKeys 481: 1–13 (2015)
2
with rounded apertures, congeners are dicult to discriminate owing to intraspecic
variation and the morphostatic mode of divergence observed in the genus (Falniowski
et al. 2012). e genitalia of these snails is characterized by a penial appendix with
a agellum in the male (Glöer and Pesic 2010), and a J-shaped cylindrical bursa
copulatrix in the female (Falniowski et al. 2009b).
Bythinella contains 132 species and subspecies (Yıldırım et al. 2006; Georgiev 2009;
Glöer and Georgiev 2009; Bank 2013; Georgiev and Glöer 2013; Glöer 2013; Odabaşı
and Georgiev 2014; Georgiev and Glöer 2014; Glöer and Pešić 2014) and is among
the most species-rich genera in the Truncatelloidea. e geographic range of the genus
extends from northern Africa and the Iberian Peninsula through central Europe to the
Balkan countries, Ukraine and Turkey (Kristensen 1985; Haase et al. 2007). Until re-
cently, only a few species had been recorded in the eastern half of this range. All 21 of the
species in Bulgaria and 10 of 12 species in Romania were described within the last decade
(Falniowski et al. 2009 a,b; Georgiev and Glöer 2013, 2014); although only three species
are known from continental Greece (Bank 2013), the actual number is estimated to be
10 based on molecular data (Falniowski and Szarowska 2011). Together with diculties
in conchological identication mentioned above, the small number of local researchers
and scant material from the area have contributed to the recognition of only a few species
of Bythinella in its eastern range until recently. e case in Turkey, forming the eastern
limit for the genus, does not dier much in this regard. e rst mention of the genus
from this country was by Schütt (1965), who reported Bythinella opaca (Frauenfeld,
1857) from Belgrad Forest in Prov. Istanbul. e assignment of the species from Istan-
bul to B. opaca (Frauenfeld, 1857) was later refuted by Schütt (1980) and Yıldırım et al.
(2006), however the taxonomic status of this snail remains uncertain. ree additional
species from the Asian part of Turkey have also been described (Kebapçı and Yıldırım
2010; Odabaşı and Georgiev 2014): Bythinella turca Radoman, 1976 from Cire spring
(Prov. Isparta), B. occasiuncula Boeters & Falkner, 2001 from Kırkoluk springs (Prov.
İzmir), and recently described B. kazdaghensis Odabaşı & Georgiev, 2014 from Ayazma
Stream (Prov. Çanakkale).
In the present study, four new Bythinella species are described from western Tur-
key based on eld surveys between 2006 and 2013. ese species are discriminated
based on details of shell morphometry, head and cephalic tentacle pigmentation, peni-
al appendix, tubular gland, female genitalia, central tooth of radula and operculum. A
comparison with other species and an identication key to other congeners described
from Turkey are also presented.
Material and methods
Specimens were collected by hand-netting and preserved in 75% ethanol. Dissections
and measurements of the genital organs and the shells were carried out using a Olym-
pus SZ12 stereo microscope; photographs were taken with a digital camera system.
New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 3
Morphological terminology largely follows Radoman (1973, 1976) and Hershler and
Ponder (1998). e scale bars in the gures are 1 mm.
Abbreviations: SW: shell width, SH: shell height, rs: reseptaculum seminis, ah:
aperture height, aw: aperture width, Coll. Yıldırım: Collection of M. Zeki Yıldırım in
Zoological Museum of the Mehmet Akif University (Burdur, Turkey).
Systematics
Genus Bythinella Moquin-Tandon, 1856
Type species. Bulimus viridis Poiret, 1801.
Bythinella anatolica Yıldırım, Kebapçı & Bahadır Koca, sp. n.
http://zoobank.org/F4B75E78-F447-40C7-BCF0-5B2FA8D00A57
Figs 1, 2, 6a
Holotype (Coll. Yıldırım): SH 2.78 mm, SW 1.67 mm, ah 1.22 mm, aw 1.11 mm.
Coll. Yıldırım; TURKEY, Manisa, Çırpıcıdede hill on Spil Mountain, N 38°44.66',
E 27°24.30', 17. 07. 2006. Leg. M. Z. Yıldırım. Paratypes: 18 ex. (5 dissected), same
data and locality as holotype in Coll. Yıldırım.
Type locality. A spring below Çırpıcıdede hill on Spil Mountain and its small
outow stream down the hill, Manisa.
Etymology. Named after Anatolia.
Description. Shell thin, usually cylindrical-ovoid (SW/SH 57.19%), appearing
blackish (owing to darkly pigmented animal), with slightly rough surface; apex blunt,
depressed on the left side; having 3 1/2–4 very tumid whorls (more convex the left side),
last whorl more inated, sutures deep. Aperture ovoid or pear shaped, height of the
aperture usually greater than penultimate whorl, umbilicus relatively broad and deep,
sometimes covered by the outer lip, palatal lip margin not reected, columellar and
parietal margins broadly reected.
Head black; tentacles unpigmented. Operculum oval and with distinct growth lines.
Penis unpigmented and variable in natural position among individuals (straight to
bent or folded), though not in shape. Tubular gland thickened, penial appendix usu-
ally very short (varying according to the shell size of the individuals). Bursa copulatrix
narrow and elongated, rs1 large and globular in shape. Central tooth of radula with 9
pointed cusps, 1 median and 4 each on sides; lateral margin without any cusps, but
undulated; basal lip roundish (Figures 1, 2).
Measurements (n=19): SH: 2.79 mm (min 2.37/max 3.07), SW: 1.65 mm (min
1.37/max 1.92), SH/SW: 1.7 (min 1.59/max 1. 91), SW/SH : 0.58 (min 0.52/max
0.62), ah/SH: 0.44 (min 0.41/max 0.46).
Mehmet Zeki Yıldırım et al. / ZooKeys 481: 1–13 (2015)
4
Figure 1. Shell and anatomy of a Bythinella species (B. anatolica sp. n.). Scale bars = 1 mm.
Dierential diagnosis: Identied by its larger shell (except B. wilkei sp. n.) and ear–
like aperture having reected margins, a character state not observed in other Turkish
species.
Habitat. Specimens were collected from under the stones in the spring outow
down the hill.
Remarks. e new species is distinguished from other Turkish species by its larger
shell dimensions and ear like shell aperture. Bythinella turca Radoman 1976 is distin-
guished from B. anatolica sp. n. by its blunter and shorter shell (SH 2,42 ± 0,13; max.
2.81 mm). SW/SH is close to B. istanbulensis sp. n., from which it can be distinguished
by its oval shell with convex whorls. Bythinella occasiuncula and B. magdalenae, the
geographically most proximate congeners, have smaller shell dimensions, while B. oc-
casiuncula can be identied by the shorter and blunter shell shape and B. magdalenae
sp. n. by smoother periphery and shallow sutures.
According to our current knowledge of Peri-Aegean Bythinella species, the area
of the Aegean coast of Turkey is a center of diversity for the genus. e Eastern
Aegean Islands Ikaria, Kos, Lesbos, and Chios are inhabited by B. kosensis (Schütt
1980; Bank 1988), while within the provinces of Aydın, İzmir, and Manisa on ad-
jacent Anatolian mainland there are three species (B. anatolica sp. n., B. magdalenae
sp. n. and B. occasiuncula) geographically isolated from the remainder of the species
recorded in Turkey (Figure 7).
New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 5
Bythinella istanbulensis Yıldırım, Kebapçı & Yüce, sp. n.
http://zoobank.org/8958B919-4F97-4EAA-A6F8-EB7059A8FEB7
Figs 3, 6b
Bythinella opaca, Schütt, H., 1965 Zur Systematik und Ökologie Türkischer Süsswas-
serprosobranchier. Zoologische Mededelingen, 41: 43–71. (misidentication).
Bythinellaopaca”, Schütt, H., 1980 Zur Kenntnis griechischer Hydrobiiden, Arch.
Moll.110 (4/6):115.
Bythinella sp. A, Yıldırım et al. 2006 Supplement to the Prosobranchia (Mollusca: Gas-
tropoda) Fauna of Fresh and Brackish Waters of Turkey, Tr. J. Zool. 30: 197–204.
Holotype (Coll. Yıldırım): Shell height: 2.74 mm, width 1.43 mm (Coll. M.Z.
Yıldırım); TURKEY, Istanbul, Bahçeköy, a small spring at the entrance of Bahçeköy
in Belgrad Forest, N41°11.09', E28°59.5', 23.02.2013, leg. A. Yüce. Paratypes: 27
ex., same data and locality as holotype; 5 ex. Zoologisches Museum Hamburg (ZMH
79661), 5 ex. Naturhistorisches Museum in Wien (NHMW 109174), rest in the Coll.
Yıldırım. Additional material: 5 ex. in the University of Giessen (Coll. Prof. Wilke),
rest in the Coll. Yıldırım (5 dissected).
Figure 2. Shell and anatomy of Bythinella anatolica sp. n.: a shell b, c, d male head and various positions
of penis e, f, g penes h female genitalia. Scale bars = 1 mm.
Mehmet Zeki Yıldırım et al. / ZooKeys 481: 1–13 (2015)
6
Type locality. A small spring at the entrance of Bahçeköy in Belgrad Forest,
Bahçeköy, Istanbul.
Etymology. Named after the type locality.
Description. Shell broad cylindrical oval, yellowish horn-colored, surface smooth
and glossy covered by a thick, solid and rough encrustation due to chemical conditions
of the spring water; apex blunt; 3½–4 convex whorls, rapidly and regularly increasing
to form a cylindrical shape, last whorl about 2/3 of the shell height, sutures relatively
deep; aperture pear-shaped, height nearly the same as or slightly less than the penul-
timate whorl, columellar margin with an distinct inner lip reaching basal and apical
corners of the aperture; operculum orange, umbilicus narrow and slit-shaped (Figs.
2a, 6a).
Tubular gland relatively short and thick. Pallial roof unpigmented, head having
little pigmentation (Fig. 2b), rs1 small and attached to oviduct (Fig. 2c), thus not easily
discernible.
Measurements (n=28): SH: 2.67 mm (min 2.18/max 2.87), SW: 1.66 mm (min
1.37/max 1.94), SH/SW: 1.61 (min 1.39/max 1.84), SW/SH: 0.62 (min 0.54/max
0.72), ah/SH: 0.41 (min 0.37/max 0.50).
Dierential diagnosis. is new species is distinguished from other Turkish Byth-
inella species by its nearly cylindrical shell with broad and blunt apex. Small and at-
tached rs1, unpigmented and light coloured pallial roof are the other key characters.
Habitat. Specimens were collected from the stones in a small spring and its pond.
Remarks. Based on shell characters only, Schütt (1965) misidentied Bythinella
specimens from Belgrad Forest as Bythinella opaca (Frauenfeld, 1857), a species origi-
nally described from Carniola (Slovenia) and Italy. Bythinella angelitae was introduced
as a replacement name for the preoccupied name Paludinella opaca Frauenfeld, 1857, a
Figure 3. Shell and anatomy of B. istanbulensis sp. n.: a shell b male head and penis in natural position
c penis d female genitalia. Scale bars = 1 mm.
New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 7
homonym of Paludinella opaca M. von Gallenstein, 1848, by Haase et al. (2007). e
two closely related species B. angelitae and B. opaca can be morphologically separated
only by radular dentition, and the geographic range of Bythinella angelitae is restricted
to the border areas of Austria and Slovenia (Haase et al. 2007). Although they have
been recorded in previous studies (Wagner 1941; Grossu 1956; Angelov 1960, 2000),
Bythinella austriaca (Frauenfeld, 1857) and Bythinella opaca (M. von Gallenstein,
1848) have not been conrmed from the eastern Balkan countries in recent studies
(Falniowski et al. 2009a, b). erefore, there is a large distribution gap between the
ranges of Central European taxa and that of B. istanbulensis sp. n.
Despite the conchological similarities, the new species is anatomically distinct
from the Central European species B. austriaca and B. opaca. While the lighter pallial
roof colouration is also observed in Bythinella opaca (Glöer & Pešic, 2006), B. austriaca
has a dark pallial roof colouration. e new species diers from both of these taxa
in having a very short and broad penial appendix (cf. long trumpet-shaped penial
appendix in these species).
Bythinella magdalenae Yıldırım, Kebapçı & Bahadır Koca, sp. n.
http://zoobank.org/669E7842-7985-473B-837B-C06917BF0962
Figs 4, 6c
Holotype (Coll. Yıldırım): SH 2,5 mm, SW 1,62 mm, ah 1,12 mm, aw 1,06 mm;
TURKEY, Aydın, İncirliova, Karagözler Village, Karapınar spring above the village,
37°57,796'N, 27°49,375'E, leg. S. Bahadır Koca. Paratypes: 17 ex in Coll. Yıldırım,
same data and locality as holotype; 18 ex in Coll. Yıldırım, TURKEY, Aydın, İncirliova,
Karagözler Village, Çaycuk spring above the village, 37°57,829'N, 27°49,230'E, leg.
S. Bahadır Koca.
Type locality. Karapınar Spring, Karagözler Village, İncirliova, Aydın.
Etymology. Named after the late Polish malacologist Magdalena Szarowska
(1952–2013), who contributed greatly especially to the knowledge of the Balkan
Truncatelloidea.
Description. Shell ovate-conical and with 3–3,5 whorls. Apex truncated. Last whorl
slowly increasing and broader than previous whorls, sutures not deep and periphery
nearly at. Aperture roundish oval. Umbilicus small, hollow shaped and covered by the
lip. Operculum nucleus along left margin, oval shaped and with distinct growth lines.
Head having little amount of pigmentation, rather concentrated on the tips of the
tentacles and around the mouth. Tentacles slightly longer than snout. Penis unpig-
mented, tapering towards the tip, tip not very pointed at its distal end but club-shaped.
Tubular gland externally visible, thin, elongated and convoluted. Bursa copulatrix nar-
row and elongated, rs1 globular in shape. Central tooth of radula with 9 pointed cusps,
1 median and 4 each on sides; lateral margin without any cusps, but undulated.
Measurements: Karapınar Spring (n=18): SH: 2.55 mm (min 2.19/max 2.91),
SW: 1.64 mm (min 1.37/max 1.78), SH/SW: 1.55 (min 1.42/max 1.72), SW/SH :
Mehmet Zeki Yıldırım et al. / ZooKeys 481: 1–13 (2015)
8
0.65 (min 0.58/max 0.70), ah/SH: 0.45 (min 0.44/max 0.46). Çaycuk Spring (n=18):
SH: 2.41 mm (min 2.06/max 2.78), SW: 1.59 mm (min 1.31/max 1.87), SH/SW:
1.52 (min 1.37/max 1.66), SW/SH : 0.66 (min 0.64/max 0.67), ah/SH: 0.46 (min
0.46/max 0.48).
Dierential diagnosis. Bythinella magdalenae sp. n. is distinguished from B. co-
sensis Schütt 1980 (from Kos island) by the smaller number of shell whorls and the
diering number of cusps on the central tooth of radula. It can be distinguished from
other Turkish congeners by the at periphery of the whorls and shallow sutures.
Habitat. Specimens were collected from two small, closely proximal seepage springs.
Remarks. Çaycuk Spring is very close to the type locality, thus indeed B. mag-
dalenae sp. n. can be considered a single spot endemic as in the case of other species
known from Turkey.
Bythinella wilkei Yıldırım, Kebapçı & Bahadır Koca, sp. n.
http://zoobank.org/3C7814DB-62E3-458B-9FB9-E4A026F339DD
Figs 5, 6d
Holotype (Coll. Yıldırım): SH 2.75 mm, SW 1.84 mm, ah 1.28 mm, aw 1.12
mm; TURKEY, Kocaeli, Maşukiye, spring along the road to Kartepe, 40°40.603'N,
30°08.605'E, leg. S. Bahadır Koca. Paratypes: 29 ex in Coll. Yıldırım, same data and
locality as holotype.
Type locality. Spring along the road to Kartepe, Maşukiye, Kocaeli.
Etymology. Named after the malacologist omas Wilke (Justus Liebig Univer-
sity, Germany).
Description. Shell oval conical, light brown, but appearing blackish due to en-
crustation; having 3–3.5 tumid whorls; last whorl slowly increasing and broader than
previous whorls, sutures deep. Apex blunt. Aperture roundish oval, last whorl strongly
descending towards aperture and aperture projected forward, lip in some individuals
Figure 4. Shell and anatomy of B. magdalenae sp. n.: a shell b male head and penis in natural position
c penis d female genitalia. Scale bars = 1 mm.
New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 9
having a small protuberance in upper palatal margin. Umbilicus small, hollow or com-
pletely covered by the lip. Operculum ovate, translucent and with distinct growth lines.
Head having little pigmentation, pigment rather concentrated on bases of the ten-
tacles and around mouth. Tentacles short and broad. Unpigmented penis tapering
towards the tip, tip not pointed at its distal end; tubular gland thick and with two
convolutions. Bursa copulatrix narrow and elongated, rs1 elongate. Central tooth of
radula with 9 pointed cusps, 1 median and 4 each on sides; lateral margins lacking
cusps, but undulated.
Measurements (n=22): SH: 2.80 mm (min 2.56/max 3.03), SW: 1.88 mm (min
1.56/max 2.19), SH/SW: 1.50 (min 1.31/max 1.68), SW/SH: 0.66 (min 0.60/max
0.76), ah/SH: 0.47 (min 0.41/max 0.51).
Dierential diagnosis. Excluding B. anatolica sp. n. described from Manisa Prov-
ince, Bythinella wilkei sp. n. has a larger shell than other Turkish congeners. e nearly
circular aperture, which is strongly descending and projected forward, is also charac-
teristic of the new species.
Figure 5. Shell and anatomy of Bythinella wilkei sp. n.: a shell b male head and penis in natural position
c penis d female genitalia. Scale bars = 1 mm.
Figure 6. Bythinella species from western Anatolia: a Bythinella anatolica sp. n. b B. istanbulensis sp. n.
c B. magdalenae sp. n. d B. wilkei sp. n.
Mehmet Zeki Yıldırım et al. / ZooKeys 481: 1–13 (2015)
10
Habitat. e specimens were collected from the outow of a small spring having
little vegetation and a gravel substrate.
Remarks. e species is one of the two northernmost distributed congeners in
Turkey. Unlike B. istanbulensis sp. n. the new species is found in higher altitudes.
Identication key to the Bythinella species from Turkey
1 Shell having 3.5–4 whorls ........................................................................... 2
Shell having 3–3.5 whorls ........................................................................... 4
2 Shell elongated oval, aperture margins reected ........... Bythinella anatolica
Figure 7. Map showing the locations of Bythinella species in Turkey and the Eastern Aegean Islands:
Bythinella anatolica sp. n. (black triangle), B. istanbulensis sp. n. (closed square), B. kazdaghensis (dia-
mond), B. kosensis (asterix), B. magdalenae sp. n. (closed circle), B. occasiuncula (open circle), B. turca
(open triangle), B. wilkei sp. n. (open square).
New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 11
Shell nearly cylindrical, aperture margins not reected ................................3
3 Pallial roof black, penis shorter than penial appendix .......... B. kazdaghensis
Pallial roof light coloured, penis longer than penial appendix .... B. istanbulensis
4 Tubular gland thin, long and convoluted (more than 2 loops) ....................5
Tubular gland thick, short and less convoluted ...............................B. wilkei
5 Shell oval, SH<2.3 mm ........................................................ B. occasiuncula
Shell ovate-conic and usually SH>2.3 mm .................................................. 6
6 Periphery almost straight, sutures shallow, tentacles longer than proboscis .....
............................................................................................... B. magdalenae
Periphery convex, sutures deep, tentacles shorter than proboscis ...... B. turca
Acknowledgements
We thank Eike Neubert (Naturhistorisches Museum der Burgergemeinde Bern) for revis-
ing of the rst version of the manuscript and MSc student Estée Bochud for photography.
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New Bythinella (Gastropoda, Bythinellidae) species from western Turkey 13
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... The genitalia are characterized by a penial appendix with a flagellum in the male (Glöer and Pesic, 2010). This genus shows wide distribution from Central Europe to East Europe, Ukraine and Turkey, and North Africa to the Iberian Peninsula (Yıldırım et al., 2015). In Turkey, eight species have been described to date: Bythinella opaca Gallenstein, 1848; B. turca Radoman, 1976;B. ...
... magdalenae Yıldırım, Kebapçı & Bahadır Koca 2015;andB. wilkei Yıldırım, Kebapçı &Bahadır Koca 2015 (Yıldırım et al., 2015). Among these species, the presence of B. opaca Gallenstein, 1848 was accepted by Schütt (1965), but he later refuted that acceptance (Schütt, 1980); thus, the taxonomic situation of this species remains uncertain. ...
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... Biyolojik izleme programlarında en çok kullanılan ve SÇD kapsamında bir kalite unsuru olan taban büyük omurgasızları su kütlesi hakkında kapsamlı bir değerlendirme imkânı sunmaktadır (Karr, 1999;Kenney, Sutton-Grier, Smith, ve Gresens, 2009;Kazancı vd., 2010). Taban omurgasızları içinde önemli bir yeri olan tatlısu molluskları hakkında ülkemizde yapılan ilk çalışmalar Roth (1839) ile başlamış, müteakiben yerli ve yabancı çok sayıda bilimsel çalışmayla (Schütt, 1964;Schütt ve Bilgin, 1970;Bilgin, 1980;Schütt ve Şeşen, 1993;Yıldırım, 1999;Ustaoğlu, Balık ve Özbek, 2001Öktener, 2004;Yıldırım vd., 2006;Kebapçı ve Yıldırım, 2010;Kılıçaslan ve Özbek, 2010;Koşal-Şahin, Bahadır-Koca ve Yıldırım, 2012;Odabaşı ve Georgiev, 2014;Yıldırım, Kebapçı, Koca ve Yüce, 2015;Koşal-Şahin ve Zeybek, 2016;Gürlek, 2017;Gürlek, Koşal-Şahin, Dökümcü ve Yıldırım, 2019;Gözler ve Baytaşoğlu, 2020;Tomilova vd., 2020;Odabaşı, Akay ve Koyuncuoğlu, 2020;Sands vd., 2020) hız kazanmıştır. ...
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We studied morphology (shell, penis and flagellum, female reproductive organs) and 395 partial sequences of mtDNA COI and 93 of ITS-1 in Bythinella from continental Greece. Molecular techniques inferred ten molecularly distinct species: two in the Peloponnese; one in the Parnassus and Attica; two sympatric/parapatric at Volos; one in the Lefkas; four in northern Greece. The differentiation was probably not older than 3.5 MYA, most species originated in the last 2 MYA (Pleistocene). Haplotype diversity, nucleotide diversity and mismatch distribution indicated common bottlenecks followed by fluctuations in population size. A nested-clade analysis indicated allopatric fragmentation with instances of long distance colonization, and restricted gene flow with isolation by distance. The decrease in Bythinella species richness from the north to the south was due to the geological history, colonization and recolonization, and short and long-distance dispersal, to survive in the unstable environment.
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Two new species of the genus Bythinella Moquin-Tandon, 1856, i.e. B. marici n. sp. from Bosnia and Hercegovina, and B. istoka n. sp. from Kosovo were described. The holotypes of both species as well as the penis morphology are depicted. In addition a list and distribution map of the Bythinella spp. occuring in the Western Balkan Peninsula is provided.
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A new species of the genus Bythinella Moquin-Tandon, 1856 (Gastropoda: Risooidea: Bythinellidae), from a spring in Strandzha Mt, SE Bulgaria was described as new for science. By this there are 22 Bythinella species known from Bulgaria.
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The genus Bythinella comprises many species throughout Europe, but species delimitation, traditionally based on shell morphology and genital anatomy, is often a matter of debate. Out of an ongoing large-scale project on the phylogeny of the genus, we analyzed the relationships of species occurring in the south Austrian province Carinthia and in neighboring Slovenia as a model for similar cases of systematic and taxonomic ambiguity. Our analyses based on sequence data of a fragment of COI comprising 638 bp, morphological and anatomical investigations confirmed the presence of three species, B. opaca (Gallenstein, 1848), B. robiciana (Clessin, 1890) and B. angelitae nom. nov. for B. opaca (Frauenfeld, 1857). The latter, while genetically distinct, is morphologically and anatomically cryptic in that it can only be distinguished from B. opaca by the denticulation of the radular marginal teeth. B. robiciana, on the other hand, is morphologically well defined, but genetically not separable from B. opaca, its stem species. Thus, taxonomy in Bythinella has to be based on the integration of morphology, anatomy and genetics. Our phylogenetic analyses suggest that B. opaca has colonized Carinthia, which has largely been covered by glaciers during the last ice age, along two routes, one from the south and a second one from the southeast.
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The genus Bythinella comprises many species throughout Europe, but species delimitation, traditionally based on shell morphology and genital anatomy, is often a matter of debate. Out of an ongoing large-scale project on the phylogeny of the genus, we analyzed the relationships of species occurring in the south Austrian province Carinthia and in neighboring Slovenia as a model for similar cases of systematic and taxonomic ambiguity. Our analyses based on sequence data of a fragment of COI comprising 638 bp, morphological and anatomical investigations confirmed the presence of three species, B. opaca (Gallenstein, 1848), B. robiciana (Clessin, 1890) and B. angelitae nom. nov. for B. opaca (Frauenfeld, 1857). The latter, while genetically distinct, is morphologically and anatomically cryptic in that it can only be distinguished from B. opaca by the denticulation of the radular marginal teeth. B. robiciana, on the other hand, is morphologically well defined, but genetically not separable from B. opaca, its stem species. Thus, taxonomy in Bythinella has to be based on the integration of morphology, anatomy and genetics. Our phylogenetic analyses suggest that B. opaca has colonized Carinthia, which has largely been covered by glaciers during the last ice age, along two routes, one from the south and a second one from the southeast.