Unraveling a new lineage of Hydrobiidae genera (Caenogastropoda: Truncatelloidea) from the Ponto-Caspian region Author Diana Delicado Author ladimir Pešić Author Peter Glöer text European Journal of Taxonomy 2016 208 1 29 journal article 10.5852/ejt.2016.208 2c5cb62e-9bc7-4fba-a4cb-e5a2fda8c719 2118-9773 269306 urn:lsid:zoobank.org:pub:F807AC79-2615-463B-8272-82523B46210A Intermaria zagrosensis ( Glöer & Pešić, 2009 ) comb. nov. Figs 2–3 , Tables 2–3 Pseudamnicola zagrosensis Glöer & Pešić, 2009 : 37 , pl. 6, figs 4–6. New diagnosis Shell ovate-conic, yellowish, with pyriform aperture; protoconch microsculpture wrinkled; central radular tooth formula (5)4–C–4(5)/2–2; pyriform bursa copulatrix; one seminal receptacle elongate with a short duct; penis gradually tapering, with a small distal lobe on inner edge, end tapered and grayish pigmented; nervous system elongated (mean RPG ratio = 0.60) and slightly black pigmented. Fig. 2. Shell, operculum and radula of Intermaria zagrosensis (Glöer & Pešić, 2009) gen. et comb. nov. from Sar Pol Kangarar stream, Kermanshah Province, Iran. A–B . Shell in front view. C . Shell in lateral view. D . Protoconch and microsculpture. E–F . Internal and external side of the operculum. G . Rows of teeth of the radula. H . Central teeth. I . Detail of outer marginal teeth. Fig. 3. Anatomy of Intermaria zagrosensis (Glöer & Pešić, 2009) gen. et comb. nov. from Pol Kangarar stream, Kermanshah Province, Iran. A . Ctenidium and osphradium. B . Partial nervous system. C . Female genitalia. D . Bursa copulatrix and seminal receptacle. E–F . Head of a male and penis. G . Prostate gland. Anatomical abbreviations given in the Material and methods section. Table 2. Shell measurements (in mm) of the species: 1 . Intermaria zagrosensis (Glöer & Pešić, 2009) . 2 . Intermaria kermanshahensis (Glöer & Pešić, 2009) . 3 . Persipyrgula saboori (Glöer & Pešić, 2009) . 4 . Shadinia terpoghassiani (Shadin, 1952) . ANOVA results are shown through effect size F, degrees of freedom (df), residuals (r) and the resulting significance (p–value: *** when 0; ** when 0.001; * when 0.01; NS = no significant).

1	2	3	4
Mean ± SD; CV (Max–Min) (n = 11)	Mean ± SD; CV (Max–Min) (n = 7)	Mean ± SD; CV (Max–Min) (n = 7)	Mean ± SD; CV (Max–Min) (n = 20)	ANOVA (F df,r)
SL	3.58 ± 0.40; 0.11 (4.50–2.98)	3.30 ± 0.36; 0.11 (4.00–2.80)	2.90 ± 0.67; 0.23 (4.40–2.48)	3.02 ± 0.25; 0.08 (3.61–2.60)	F3,40 = 6.1**
SW	2.28 ± 0.33; 0.15 (3.00–1.74)	2.00 ± 0.13; 0.07 (2.70–1.77)	1.84 ± 0.52; 0.28 (3.00–1.48)	1.68 ± 0.14; 0.09 (1.96–1.41)	F3,40 = 10.4***
SL/SW	1.58 ± 0.10; 0.07 (1.71–1.45)	1.58 ± 0.05; 0.03 (1.66–1.50)	1.59 ± 0.08; 0.05 (1.70–1.48)	1.80 ± 0.10; 0.06 (2.05–1.63)	F3,40 = 19.4***
AH	1.82 ± 0.22; 0.12 (2.34–1.58)	1.62 ± 0.13; 0.08 (1.71–1.37)	1.31 ± 0.15; 0.11 (1.50–1.09)	1.45 ± 0.11; 0.08 (1.68–1.25)	F3,37 = 18.3***
SL–LBW	0.83 ± 0.08; 0.10 (0.97–0.67)	0.75 ± 0.08; 0.10 (0.83–0.61)	0.66 ± 0.05; 0.07 (0.75–0.62)	0.84 ± 0.11; 0.14 (1.08–0.66)	F3,37 = 5.9**
WBW	1.98 ± 0.19; 0.09 (2.26–1.58)	1.92 ± 0.12; 0.06 (2.07–1.76)	1.51 ± 0.07; 0.04 (1.62–1.44)	1.58 ± 0.13; 0.08 (1.88–1.37)	F3,37 = 26.3***
AL	1.77 ± 0.16; 0.09 (2.11–1.53)	1.67 ± 0.11; 0.07 (1.79–1.54)	1.27 ± 0.12; 0.09 (1.39–1.06)	1.41 ± 0.12; 0.08 (1.70–1.24)	F3,37 = 28.2***
AW	1.36 ± 0.09; 0.07 (1.50–1.18)	1.23 ± 0.10; 0.08 (1.35–1.06)	0.97 ± 0.05; 0.07 (1.06–0.90)	1.03 ± 0.08; 0.08 (1.13–0.86)	F3,37 = 41.9***
WPW	1.20 ± 0.12; 0.10 (1.41–0.98)	1.17 ± 0.05; 0.04 (1.24–1.10)	0.96 ± 0.05; 0.05 (1.02–0.90)	1.03 ± 0.09; 0.08 (1.24–0.9)	F3,37 = 13.9***
WAW	0.13 ± 0.02; 0.18 (0.19–0.11)	0.13 ± 0.02; 0.19 (0.18–0.11)	0.10 ± 0.02; 0.20 (0.13–0.07)	0.13 ± 0.03; 0.24 (0.23–0.08)	F = 1.4NS 3,37
NSW	4.15 ± 0.21; 0.05 (4.50–4.00)	4.37 ± 0.14; 0.03 (4.50–4.25)	4.20 ± 0.25; 0.06 (4.50–4.00)	4.13 ± 0.17; 0.04 (4.50–4.00)	F = 2.6NS 3,37

Material examined Holotype IRAN: ZMH 51406: 4.5 mm height, 3.0 mm width. Paratypes IRAN: ZMH 51407 (5 ex.) and P. Glöeris (68 ex.) collection. Type locality IRAN : Sar Pol Kangavar stream, Kangavar city , Kermanshah Province , 34º30' N , 47º55' E , 27 Jun. 2005 . Table 3. Ctenidium, osphradium. Female and male genitalia and nervous system measurements (in mm) of the species: 1 . Intermaria zagrosensis (Glöer & Pešić, 2009) . 2 . Intermaria kermanshahensis (Glöer & Pešić, 2009). 3 . Persipyrgula saboori (Glöer & Pešić, 2009) . 4 . Shadinia terpoghassiani (Shadin, 1952) . For ANOVA results effect size F, degrees of freedom (df), residuals (r) and the resulting significance (p–value: *** when 0; ** when 0.001; * when 0.01; NS = not significant) are shown.

1	2	3	4
Mean ± SD; CV (Max–Min) 3 ♀ – 2 ♂	Mean ± SD; CV (Max–Min) 1 ♀ – 3 ♂	Mean ± SD; CV (Max–Min) 2 ♀ – 2 ♂	Mean ± SD; CV (Max–Min) 5 ♀ – 3 ♂	ANOVA (F df,r)
CL	1.43 ± 0.14; 0.09 (1.56–1.28)	0.93 ± 0.08; 0.08 (1.00–0.84)	0.79 ± 0.05; 0.07 (0.85–0.72)	1.12 ± 0.08; 0.08 (1.25–1.00)	F3,15 = 32.9***
Os L	0.41 ± 0.09; 0.23 (0.40–0.27)	0.32 ± 0.02; 0.05 (0.34–0.30)	0.24 ± 0.06; 0.24 (0.33–0.20)	0.32 ± 0.06; 0.17 (0.40–0.27)	F3,15 = 4.8*
Os W	0.20 ± 0.00; 0.00 (0.20–0.20)	0.10 ± 0.00; 0.00 (0.10–0.10)	0.12 ± 0.03; 0.26 (0.16–0.09)	0.13 ± 0.03; 0.21 (0.15–0.08)	F3,15 = 11.8***
PoL	2.29 ± 0.20; 0.09 (2.44–2.06)	1.71	1.20 ± 0.11; 0.09 (1.28–1.13)	1.66 ± 0.22; 0.13 (2.00–1.40)	F3,7 = 12.6**
PoW	0.71 ± 0.08; 0.12 (0.80–0.63)	0.5	0.35 ± 0.03; 0.10 (0.38–0.33)	0.53 ± 0.06; 0.11 (0.57–0.43)	F3,7 = 12.6**
AgL	1.11 ± 0.13; 0.09 (1.23–0.98)	0.98	0.60 ± 0.01; 0.01 (0.61–0.60)	0.75 ± 0.04; 0.06 (0.80–0.68)	F3,7 = 22.9***
CgL	0.13 ± 0.05; 0.05 (1.17–1.07)	0.69	0.60 ± 0.14; 0.23 (0.70–0.50)	0.93 ± 0.09; 0.10 (1.00–0.77)	F3,7 = 14.7**
BC L	0.71 ± 0.11; 0.16 (0.84–0.63)	0.4	0.38 ± 0.11; 0.27 (0.46–0.31)	0.35 ± 0.07; 0.21 (0.42–0.25)	F3,7 = 10.7**
BC W	0.39 ± 0.04; 0.10 (0.43–0.36)	0.25	0.21 ± 0.06; 0.27 (0.25–0.17)	0.21 ± 0.05; 0.23 (0.27–0.15)	F3,7 = 10.3**
dBC L	0.40 ± 0.05; 0.11 (0.43–0.35)	0.45	0.38 ± 0.03; 0.07 (0.40–0.36)	0.46 ± 0.04; 0.10 (0.50–0.40)	F = 2NS 3,7
SR I L	0.28 ± 0.03; 0.11 (0.31–0.25)	0.24	0.08 ± 0.01; 0.18 (0.09–0.07)	0.26 ± 0.10; 0.41 (0.42–0.15)	F = 2.8NS 3,7
SR II L	absent	absent	absent	0.16 ± 0.04; 0.26 (0.22–0.12)	–
P L	1.37 ± 0.11; 0.08 (1.45–1.30)	1.00 ± 0.00; 0.00 (1.00–1.00)	0.78 ± 0.12; 0.15 (0.87–0.70)	1.09 ± 0.23; 0.21 (1.27–0.83)	F3,6 = 5.5*
P W	0.30 ± 0.00; 0.00 (0.30–0.30)	0.30 ± 0.02; 0.07 (0.32–0.28)	0.17 ± 0.03; 0.20 (0.20–0.15)	0.42 ± 0.06; 0.13 (0.47–0.36)	F3,6 = 17.7**
P L/Head	1.18 ± 0.14; 0.11 (1.28–1.08)	0.90 ± 0.14; 0.16 (1.06–0.80)	0.92 ± 0.11; 0.12 (1.00–0.84)	0.98 ± 0.15; 0.15 (1.10–0.81)	F = 1.8NS 3,5
ProL	1.10	0.84 ± 0.11; 0.13 (0.94–0.72)	0.68 ± 0.01; 0.01 (0.69–0.68)	0.76 ± 0.03; 0.04 (0.78–0.72)	F3,5 = 7.7*
ProW	0.55	0.38 ± 0.04; 0.10 (0.42–0.35)	0.34 ± 0.01; 0.04 (0.35–0.33)	0.37 ± 0.03; 0.08 (0.40–0.34)	F3,5 = 10.8*
RPG	0.61 ± 0.03; 0.05 (0.64–0.59)	0.50 ± 0.07; 0.13 (0.55–0.41)	0.52 ± 0.02; 0.05 (0.55–0.49)	0.45 ± 0.03; 0.06 (0.46–0.42)	F3,10 = 11***

Other localities IRAN: Darband stream in Darband village (Azna to Dorud road, ca 16 km to Azna city), Lorestan Province, 33º25' N , 49º17' E , ca 1800 m a.s.l., 23 Jun. 2005 . Description Shell ovate-conic with 4–4.5 whorls, height 3–4.5 mm ( Fig. 2 A–C, Table 2 ); periostracum yellowish; protoconch approximately 400 µm wide with 1.3 whorls and nucleus around 125 µm long; protoconch microsculpture wrinkled ( Fig. 2 D); body whorl about ⅔ of total length; rest of whorls slightly convex with deep sutures; aperture complete, pyriform, often attached to body whorl on the top; thin inner peristome but thicker than outer lip; peristome margin slightly sinuate ( Fig. 2 C). Operculum with ca 2 whorls ( Fig. 2 E–F) and muscle attachment area oval and located near the nucleus. Radula intermediate length (25% total shell length) bearing about 45 rows of teeth; central tooth formula (5)4–C–4(5)/2–2 ( Fig. 2 G–H); lateral teeth formula 4–C–4; inner marginal teeth having 18–20 sharp cusps; outer marginal teeth having 15–19 sharp cusps ( Fig. 2 I). Pigmentation and anatomy Head dark brown pigmented from snout to neck; tentacles brown pigmented except on ocular lobes; snout as long as wide, with medial lobation. Ctenidium extended across most of pallial cavity with 18–21 gill filaments; osphradium two times longer than wide and opposite middle of ctenidium ( Fig. 3 A, Table 3 ). Nervous system with black pigmentation and elongate (mean RPG ratio 0.60, Table 3 ); cerebral ganglia equal in size ( Fig. 3 B). Female pallial oviduct with a capsule gland longer than albumen gland ( Fig. 3 C, Table 3 ); pyriform bursa copulatrix with a duct longer than its length, and lying against the middle section of the albumen gland; renal oviduct white from the insertion point of bursal duct to the seminal receptacle and hereafter black making one or two loops; one elongate seminal receptacle with short duct ( Fig. 3 D). Male genitalia with penis gradually tapering with a small distal lobe on inner edge; distal end tapered, and grayish pigmented on the distal section ( Fig. 3 E–F); prostate gland twice as long as wide ( Fig. 3 G, Table 3 ). Remarks This species anatomically resembles I. kermanshahensis , snails of I. zagrosensis , however, are larger and present more cusps in lateral and marginal radular teeth, nervous system with higher RPG ratio, and relatively larger bursa copulatrix and penis. Ecology and distribution Recorded in streams and springs from the Kermanshah and Lorestan provinces (Iran).