A new species of tooth-carp, Aphanius mesopotamicus, from Iran and Iraq (Actinopterygii, Cyprinodontidae) Author Coad, Brian Canadian Museum of Nature, Ottawa, Canada text ZooKeys 2009 2009-12-28 31 3 149 163 journal article 10.3897/zookeys.31.131 fe006ef9-ac7d-4440-860c-56c73690a5c9 1313–2970 576586 9DB3A8C3-4419-4C6F-90ED-957B296E5C95 Aphanius mesopotamicus Coad , sp. n. urn:lsid:zoobank.org:act: 8B6BFE52-8D1E-45FE-AF41-9BE20C38538D Type material. Holotype : female, 29.3 mm SL, Iran , Khuzestan , canal branch of Karkheh River , 31°40´N , 48°35´E , 27 January 1978 , B. W. Coad and S. Coad ( CM- NFI 1979-0360 A). Paratypes : 37 (34 used in analyses, smallest male, smallest female and one deformed female not used in analyses), 14 males 17.1–23.9 mm SL and 23 females 14.6-29.1 mm SL, same locality as above ( CMNFI 1979-0360B). 6(4 used in analyses, smallest of each sex not used in analyses), 3 males 17.1–19.9 mm SL and 3 females 15.1-20.5 mm SL, Iran , Khuzestan , Karkheh River branch at Abdolkhan, 31°52'30"N , 48°20'30"E , 27 January 1978 , B. W. Coad and S. Coad ( CMNFI 1979–0364). Other, non-type material: 4, 2 males 22.7–24.3 mm SL, 2 females 20.2–25.3 mm SL, Iraq , Qarmat`Ali, Basrah, 30°34'N , 47°46'E , L. A. J. Al-Hassan ( BM (NH) 1982.9.2 :326-328). Table |. Meristic characters ( n = 30 for sophiae males, 23 for sophiae females, 16 for mesopotamicus males, and 23 for mesopotamicus females). SD = standard deviation.

Character	Species	Sex	Range	Mean	SD
Dorsal fin rays	sophiae	♁	12–15	13.1	0.82
mesopotamicus	♁	12–13	12.5	0.52
sophiae	♀	11–15	13.0	0.82
mesopotamicus	♀	11–13	12.4	0.58
Anal fin rays	sophiae	♁	10–13	11.8	0.82
mesopotamicus	♁	10–12	11.1	0.62
sophiae	♀	11–13	11.7	0.54
mesopotamicus	♀	11–12	11.5	0.51
Pectoral fin rays	sophiae	♁	14–18	16.1	0.84
mesopotamicus	♁	13–15	13.9	0.57
sophiae	♀	15–19	16.3	0.93
mesopotamicus	♀	14–15	14.1	0.29
Pelvic fin rays	sophiae	♁	5–6	5.9	0.35
mesopotamicus	♁	5–6	5.7	0.48
sophiae	♀	5–6	5.8	0.42
mesopotamicus	♀	5–6	5.8	0.39
Lateral series scales	sophiae	♁	27–31	28.6	1.16
mesopotamicus	♁	26–28	27.6	0.62
sophiae	♀	25–31	28.8	1.23
mesopotamicus	♀	26–29	27.3	0.82
Gill rakers	sophiae	♁	10–13	11.4	0.67
mesopotamicus	♁	10–12	11.0	0.52
sophiae	♀	10–12	11.5	0.59
mesopotamicus	♀	11–14	11.4	0.71
Caudal peduncle scales	sophiae	♁	14–20	16.5	1.20
mesopotamicus	♁	12–16	14.6	0.96
sophiae	♀	15–19	16.6	1.16
mesopotamicus	♀	12–16	14.1	1.04
Scales between lateral	sophiae	♁	4–6	5.3	0.58
series and dorsal fin	mesopotamicus	♁	4–5	4.3	0.45
sophiae	♀	5–7	5.2	0.60
mesopotamicus	♀	4–5	4.2	0.39
Scales between lateral	sophiae	♁	5–8	6.4	0.63
series and anal fin	mesopotamicus	♁	5–7	5.4	0.63
sophiae	♀	5–7	6.2	0.60
mesopotamicus	♀	4–6	5.2	0.60
Scales between lateral	sophiae	♁	6–8	7.4	0.67
series and pelvic fin	mesopotamicus	♁	5–7	6.3	0.58
sophiae	♀	7–9	7.6	0.59
mesopotamicus	♀	5–8	6.2	0.65
Total scales along flank	sophiae	♁	28–33	30.3	1.37
mesopotamicus	♁	27–29	28.7	0.60
sophiae	♀	27–32	30.5	1.31
mesopotamicus	♀	27–30	28.3	0.81

Character	Species	Sex	Range	Mean	SD
Precaudal vertebrae	sophiae	♁	11–13	11.9	0.61
mesopotamicus	♁	11–12	11.6	0.50
sophiae	♀	11–13	11.9	0.46
mesopotamicus	♀	11–13	11.8	0.49
Caudal vertebrae	sophiae	♁	15–17	16.2	0.55
mesopotamicus	♁	15–17	16.0	0.63
sophiae	♀	15–17	16.4	0.57
mesopotamicus	♀	14–17	15.4	0.65
Flank bars (males)	sophiae	♁	10–21	14.3	2.63
mesopotamicus	♁	10–15	12.4	1.36

Comparative material. Aphanius sophiae , material listed in Coad (1996 , 1998). Diagnosis . The new species is defined by pigmentation, distribution, meristics, and in multivariate morphometric and meristic space. Males of the new species have clear margins to the unpaired fins, no bars on the caudal fin and have 10–15 clearly defined flank bars. Females bear irregular blotches or spots on the flank. Pigmentation in A. mento and A. dispar , the two other and well-known species in the southern mesopotamian basin of Iraq and Iran , is highly distinctive. A. mento adult males are a dark blue-black with iridescent blue-white to silvery spots. A. dispar males have a caudal fin with 2–3 dark and light blue alternating broad bars, the last bar being yellow. The new species has been confused with A. sophiae but this species is endemic to an endorheic basin of southern Iran . Females of A sophiae , however, have fine spotting on the flank. Twelve of 14 meristic characters are significantly different for males and 9 of 13 meristic characters for females, although ranges overlap in all cases (Table 5). Discriminant function analyses indicate that the variables with the best discriminating power are pectoral fin rays, scales to pelvic fin, postorbital length and caudal peduncle length in males and pectoral fin rays, scales to pelvic fin, scales to dorsal fin, total scales and predorsal length in females. The new species is also distinguished from related species in western and southern Iran . Males of A. isfahanensis , a species endemic to an endorheic basin in west-central Iran , have very dark dorsal and anal fin margins. Females of A. persicus , a species endemic to an endorheic basin of southern Iran have thin, distinctive flank bars. Two other species, A. ginaonis , a hot spring endemic of southern Iran , and A. vladykovi , found in the high Zagros Mountains of Iran , are distinguished by non-overlapping meristic characters, respectively higher lateral scale counts (36–47; Coad 1988 ) and lower dorsal fin ray counts (5–7; Coad 1980 ). Description . Meristic characters are summarised in Table 1 and morphometric characters in Table 2 in comparison with A. sophiae . Twelve of 14 meristic characters are significantly different (p<0.05) for males, although ranges overlap, the characters not significantly different being pelvic fin rays counts and precaudal vertebrae. Nine of 13 meristic characters are significantly different (p<0.05) for females, although ranges overlap, the characters not significantly different being anal and pelvic fin rays counts, gill rakers and precaudal vertebrae. Tests for normality and heteroscedasity show that 8 morphometric characters can be compared between species as ratios with t-tests in females but only one in males. Males are more similar morphometrically than females. The 8 significantly different (p <0.05) characters in females are predorsal length, head depth and prepelvic length, all in standard length, and interorbital width, postorbital length, mouth width and anal fin length, all in head length, and head depth in head length. Th e sole male character is head width in head length. Table 2. Morphometrics for Aphanius sophiae and A. mesopotamicus . SD = standard deviation.

Number	Range	Mean	SD
Standard length/ Head length	A. sophiae	♁ ♀	30 23	3.2–3.6 3.2–4.0	3.5 3.6	0.11 0.18
A. mesopotamicus	♁ ♀	16 23	3.2–3.6 3.3–3.7	3.4 3.5	0.12 0.09
Standard length/ Predorsal length	A. sophiae	♁ ♀	30 23	1.6–1.8 1.6–1.7	1.7 1.7	0.05 0.04
A. mesopotamicus	♁ ♀	16 23	1.6–1.7 1.5–1.7	1.7 1.6	0.03 0.04
Standard length/ Prepelvic length	A. sophiae	♁ ♀	30 23	3.2–3.6 3.2–4.0	3.5 3.6	0.11 0.18
	A. mesopotamicus	♁ ♀	16 23	3.2–3.6 3.3–3.7	3.4 3.5	0.12 0.09
Standard length/ Preanal length	A. sophiae	♁ ♀	30 23	1.5–1.6 1.4–1.5	1.5 1.5	0.03 0.04
A. mesopotamicus	♁ ♀	16 23	1.5–1.7 1.4–1.6	1.6 1.5	0.04 0.05
Standard length/ Body depth	A. sophiae	♁ ♀	30 23	2.7–3.9 3.0–3.9	3.2 3.4	0.24 0.23
A. mesopotamicus	♁ ♀	16 23	3.1–3.7 3.2–3.7	3.4 3.5	0.19 0.13
Standard length/ Head depth	A. sophiae	♁ ♀	30 23	3.6–4.4 3.9–4.7	4.1 4.3	0.19 0.23
A. mesopotamicus	♁ ♀	16 23	4.0–4.7 4.0–4.7	4.3 4.4	0.16 0.17
Head length/ Head width	A. sophiae	♁ ♀	23 30	1.3–1.7 1.4–1.7	1.5 1.5	0.09 0.07
	A. mesopotamicus	♁ ♀	16 23	1.5–1.7 1.3–1.6	1.6 1.5	0.06 0.07
Head length/ Head depth	A. sophiae	♁ ♀	30 23	1.1–1.2 1.1–1.3	1.2 1.2	0.05 0.06
A. mesopotamicus	♁ ♀	16 23	1.1–1.3 1.1–1.3	1.3 1.2	0.05 0.04
Head length/ Orbit diameter	A. sophiae	♁ ♀	30 23	2.9–3.6 2.0–2.4	3.2 2.1	0.18 0.12
A. mesopotamicus	♁ ♀	16 23	2.1–3.2 3.0–3.4	2.9 3.1	0.25 0.11
Head length/ Snout length	A. sophiae	♁ ♀	30 23	3.6–4.2 3.2–4.2	3.9 3.8	0.18 0.25
A. mesopotamicus	♁ ♀	16 23	3.2–4.1 3.3–4.5	3.7 3.8	0.24 0.28
Head length/ Interorbital width	A. sophiae	♁ ♀	30 23	2.2–2.7 2.3–2.7	2.5 2.5	0.13 0.12
	A. mesopotamicus	♁ ♀	16 23	2.2–2.6 2.3–2.5	2.4 2.4	0.12 0.08

Number	Range	Mean	SD
Head length/ Postorbital length	A. sophiae	♁ ♀	30 23	2.0–2.4 2.0–2.3	2.2 2.2	0.11 0.08
A. mesopotamicus	♁ ♀	16 23	2.2–2.5 2.1–2.4	2.3 2.2	0.10 0.06
Head length/ Mouth width	A. sophiae	♁ ♀	30 23	2.8–3.6 2.8–3.5	3.1 3.2	0.19 0.20
A. mesopotamicus	♁ ♀	16 23	2.7–3.5 2.6–3.3	3.0 3.0	0.20 0.18
Head length/ Dorsal fin length	A. sophiae	♁ ♀	30 23	0.9–1.2 1.0–1.4	1.0 1.2	0.09 0.11
A. mesopotamicus	♁ ♀	16 23	0.8–1.2 1.2–1.4	1.0 1.2	0.09 0.07
Head length/ Anal fin length	A. sophiae	♁ ♀	30 23	1.1–1.5 1.3–1.6	1.3 1.4	0.09 0.08
A. mesopotamicus	♁ ♀	16 23	1.1–1.3 1.3–1.5	1.2 1.4	0.09 0.06
Head length/ Pectoral fin length	A. sophiae	♁ ♀	30 23	1.4–1.8 1.4–2.0	1.6 1.7	0.09 0.15
A. mesopotamicus	♁ ♀	16 23	1.5–1.8 1.5–1.9	1.6 1.7	0.10 0.09
Head length/ Pelvic fin length	A. sophiae	♁ ♀	30 23	2.5–3.5 2.7–3.7	2.9 3.2	0.24 0.33
A. mesopotamicus	♁ ♀	16 23	2.9–4.3 2.8–3.8	3.4 3.4	0.33 0.25
Dorsal fin length/ Anal fin length	A. sophiae	♁ ♀	30 23	1.1–1.4 1.0–1.4	1.3 1.2	0.08 0.10
A. mesopotamicus	♁ ♀	16 23	1.1–1.4 1.0–1.2	1.2 1.1	0.09 0.05
Pectoral–pelvic fin distance/	A. sophiae	♁ ♀	30 23	1.1–1.5 1.1–2.0	1.3 1.5	0.09 0.22
Pectoral fin length	A. mesopotamicus	♁ ♀	16 23	1.0–1.3 1.2–1.7	1.1 1.4	0.07 0.12
Pelvic–anal fin distance/ Pelvic fin length	A. sophiae	♁ ♀	30 23	1.1–1.7 1.1–2.0	1.4 1.5	0.14 0.22
A. mesopotamicus	♁ ♀	16 23	1.2–2.0 1.3–2.2	1.5 1.7	0.19 0.20
Caudal peduncle length/ Caudal peduncle depth	A. sophiae	♁ ♀	30 23	1.2–1.6 1.4–1.8	1.4 1.5	0.10 0.09
A. mesopotamicus	♁ ♀	16 23	1.4–1.8 1.5–1.9	1.6 1.7	0.10 0.10

Males are more distinct on the PCA (Fig. 2), where meristic and morphometric values are combined, than females ( Fig. 3 ). The first 5 eigenvectors explain over 57% of total variance for males ( Table 3 ) and also for females ( Table 4 ). Discriminant function analyses for males show the variables with the best discriminating power are pectoral fin rays, caudal peduncle length, scales to pelvic fin and postorbital length and for females are pectoral fin rays, scales to dorsal fin, predorsal length and total scales (Table 5). Description of pigmentation is based on preserved fish only (Fig. 1). Male pigmentation is as follows. The dorsal surface of the head and the upper flank are more heavily pigmented with melanophores than more ventral areas. Th e belly and lower Figure |. Aphanius mesopotamicus , holotype , female, 29.3 mm SL (CMNFI 1979-0360A) above; paratype , male, 21.7 mm SL (CMNFI 1979–0360B, below). 6 4 2 ) 2 ( FACTOR FACTOR (ļ) Figure 2. Principal components analysis for males of A. mesopotamicus s and A. sophiae k . head are unpigmented. Th e chin and snout have dense melanophores and a rim of melanophores underscores the eye. The dorsal, anal and caudal fins in males have wide clear margins. Th is is also seen in the material from Basrah, Iraq (BM(NH) 1982.9.2 :326–328). The caudal fin in the type series of the new species is darker just proximal to the clear margin, lighter in mid-fin and dark again at the base. Th e dorsal fin has irregular pigmentation on the membranes and, to a lesser extent, on the rays. Th e pigmentation may involve an overall darker colour in contrast to the light margin or may have some pattern to it. The pattern is often elongate and short blotches with no regular arrangement and sometimes may appear as up to 5 wavy and oblique bands. Dark pigmentation is found just behind the first ray on the fin membrane. Th e anal fin is darkest just proximal to the clear margin. Up to the last 6 membranes of the anal fin are dark and this pigment may be broken up in as many as 4 elongate bars along each membrane. A similar pattern is found in some dorsal fins and the general effect on both fins is that the postero-dorsal (anal fin) and postero-ventral (dorsal fin) parts of these fins are the darkest. The dorsal, anal and caudal fins generally have more pigment on the membranes than the rays and in some this is quite distinctive, making the rays stand out. The pectoral and pelvic fins in males are generally clear or somewhat milky and opaque and lack melanophores. The distal parts of the membranes between the last 5 rays of the pectoral fin and the small membrane area of the pelvic fins can be pigmented. Table 3. Loadings of eigenvectors on thirty-three components produced by principal components analysis of 14 meristic counts and 22 standardized measurements for male/female A. mesopotamicus and A. sophiae .

Character/Component	1	2	3	4	5
Total scales	-0.790	-0.044	0.353	-0.019	0.020
Lateral series scales	-0.744	-0.202	0.309	0.142	0.001
Pectoral fin rays	-0.721	0.385	0.290	0.238	-0.084
Scales to dorsal fin	-0.687	0.308	0.342	0.121	-0.167
Dorsal fin height	0.678	0.047	0.493	-0.250	-0.092
Scales to anal fin	-0.669	0.032	0.242	-0.150	-0.017
Anal fin height	0.663	-0.075	0.227	-0.236	-0.181
Scales to pelvic fin	-0.600	0.290	0.439	0.008	0.261
Body depth	0.594	0.569	0.180	-0.196	0.063
Interorbital width	0.553	-0.053	0.350	-0.275	0.012
Snout length	0.533	0.127	0.308	0.430	0.010
Prepelvic length	-0.075	0.811	-0.296	-0.038	-0.033
Pectoral to pelvic fin distance	-0.158	0.674	-0.398	-0.252	0.080
Preanal length	-0.185	0.621	-0.263	0.019	0.312
Head depth	0.251	0.616	0.110	0.042	0.191
Postorbital length	0.285	0.614	0.031	0.127	-0.275
Pelvic fin length	0.182	0.535	0.129	0.283	0.280
Caudal peduncle depth	0.231	0.524	0.134	0.088	-0.298
Orbit diameter	0.384	-0.098	0.532	0.455	-0.138
Caudal vertebrae	-0.317	0.059	-0.337	0.576	0.012
Caudal peduncle length	0.140	-0.487	0.270	0.517	0.095
Pelvic to anal fin distance	0.054	-0.211	0.249	0.012	0.690
Anal fin rays	-0.437	0.276	-0.255	-0.059	-0.546
Pelvic fin rays	-0.288	0.048	0.411	-0.013	0.118
Predorsal length	0.291	0.295	0.041	-0.375	0.292
Head width	0.107	0.382	-0.109	0.289	0.213
Precaudal vertebrae	-0.235	-0.047	0.479	-0.499	-0.175
Dorsal fin rays	-0.489	0.245	0.089	-0.047	-0.363
Caudal peduncle scales	-0.488	0.257	0.499	0.021	0.029
Gill rakers	-0.283	0.228	0.418	-0.303	0.170
Head length	0.453	0.461	0.174	0.352	-0.060
Mouth width	0.477	-0.064	-0.304	0.121	-0.401
Pectoral fin length	0.461	0.303	0.312	0.016	-0.021
Percent variance explained	21.250	14.031	9.816	6.726	5.564

Males have flank bars circling the caudal peduncle and reaching the anal fin base but fading ventrally on the lower part of the anterior flank, not reaching the ventral margin of the belly and becoming progressively shorter and less distinct the more anterior they are. Bars are 2–5 times broader than the pale interspaces. 6 4 2 ) 2 ( FACTOR 0