Paraphanius Esmaeili, Teimori, Zarei & Sayyadzadeh, 2020

Paraphanius Esmaeili, Teimori, Zarei & Sayyadzadeh, 2020

Fig. 11 View FIGURE 11

Type species. Lebias mento Heckel, 1843

Diagnosis. Paraphanius is distinguished from all other genera in the family Aphaniidae by the unique nuptial male colour pattern, comprising a dark grey, bluish or almost black background colour with irregular or regular iridescent bluish, white or silvery spots, often arranged in vertical rows or narrow bars (vs. background colour silvery with brown or black bars in Anatolichthys , Aphanius , Kosswigichthys and most Aphaniops species, males with silvery vermiculation or roundish to ovoid silvery spots or blotches on grey background in other Aphaniops species). Male Paraphanius are further distinguished by absence of bars in the caudal fin and instead possess very narrow, blue-white or silvery rows of spots or small blotches forming bands on a grey, black or blue background (vs. bars presence in caudal fin of Anatolichthys , Aphanius , and all Aphaniops except A. furcatus ). Female Paraphanius are distinguished from other Aphaniid genera by presence of numerous silvery spots or small blotches on the flank (vs. absence, usually with narrow bars or brown or black blotches or spots) and absence of a bar or diamond-shaped to roundish bold black blotch at the centre of the caudal-fin base (vs. presence in Anatolichthys , Aphaniops , and Aphanius ).

Paraphanius is further distinguished by presence of a dermal sheath at the anal-fin base in the nuptial female (vs. absence in Aphaniops ), absence of head canals (vs. presence in Aphaniops and Aphanius ), and absence of black dorsal- and anal-fin margins in the male (vs. presence in Anatolichthys and Kosswigichthys , presence of black dorsal-fin margin only in Aphanius ), possession of a single row of tricuspid teeth (vs. three rows of conical teeth in Kosswigichthys ), and the body being completely covered by scales (vs. naked in Kosswigichthys ).

Included species. Paraphanius alexandri , P. boulengeri , P. mento , P. mentoides , P. orontis , P. similis , P. striptus .

Distribution. Paraphanius species are widespread from Antalya to the Seyhan, Ceyhan and Orontes drainages on the Mediterranean coast of Turkey ( Kara et al. 2010, Erk’akan & Özdemir 2011, Wildekamp et al. 1999), and southwards to the northern Dead Sea basin in the Levant. Paraphanius is also widespread in the Tigris and Euphrates river drainages from northern Syria to the Iraqi Marshes ( Krupp 1985). A translocated population of P. mentoides inhabits the crater lake Nemrut in eastern Turkey, and some landlocked populations exist in central Turkey ( Geiger et al. 2014) ( Fig. 12 View FIGURE 12 ).

Remarks. The colour pattern of Paraphanius species is clearly distinct from that exhibited by other Aphaniids, which typically comprises a series of bars on a silvery background. This pattern could be interpreted as being a reverse (silvery bars on brown background in Paraphanius vs. brown bars on a silvery background) of the situation found in other aphaniid genera. These bars can be partly (in some Aphaniops and Apricaphanius species) or completely (in some Aphaniops spp. and Apricaphanius saourensis ) dissociated into spots or a vermiculate flank pattern. In juvenile individuals the flank is silvery with a brown mottled pattern in all Aphaniids analysed for this character ( A. anatoliae , A. irregularis , A. maeandricus , A. marassantensis , A. meridionalis , A. villwocki , A. fasciatus , A. dispar , A. kruppi , A. stoliczkanus , P. mento , P. mentoides , P. orontis , P. similis , P. striptus . T. apoda , A. baeticus , A. saourensis , E. sophiae , E. isfahanensis , E. persicus , and E. vladykovi ). As they grow, male Paraphanius develop indistinct dark-brown bars with pale brown interspaces, which are often only visible in live individuals with aggressive mood. The brown bars later become wider, often partly confluent to each other, and silvery spots appear in the interspaces, which often form narrow bars. Paraphanius striptus is a valid species, and the only member of the genus to maintain a distinct pattern of wide dark-bluish bars with narrow silvery interspaces throughout ontogeny. In other Paraphanius species, the development of silvery spots is not restricted to males and or interspaces between the flank bars, and some species, particularly P. alexandri and P. mentoides , develop minute silvery spots over the entire flank which entirely mask the barred pattern. The silvery spots may be organised in vertical rows or narrow bars in juvenile individuals, but these typically dissociate in adults except in P. mentoides , P. striptus , and some populations of P. mento .

This pattern of ontogenetic development clearly demonstrates that the flank pattern of silvery bars or spots on a dark background exhibited by Paraphanius species is not a reverse condition of that seen in other Aphaniids. In reality, the dark-grey, brown or black bars are so wide that the silvery interspaces appear to form bars, or they become confluent, with the silvery spots formed by small gaps in the dark pigmentation. Comparable colour patterns are exhibited by Anatolichthys irregularis ( Fig. 3 View FIGURE 3 ), in which the adult male appear black with a pattern of yellowish blotches ( Yoğurtçuoğlu & Freyhof 2018), and some male Apricaphanius which appear blackish with whitish spots.

The distribution map of Paraphanius species ( Fig. 11 View FIGURE 11 ) suggests that P. mento is geographically separated from P. striptus ( Fig. 12 View FIGURE 12 ). However, Goren (1974) suggested that both species occur in Israel, and in some cases occur in sympatry. This situation has not been revisited, because we did not possess sufficient material to confirm the occurrence of P. mento in the Jordan River drainage within the framework of this study.

Esmaeili et al. (2020) clearly based their diagnosis of Paraphanius on Parenti (1981), because they examined only two juvenile individuals (24, 27 mm SL) from Beirut ( Lebanon). Parenti (1981) examined eight individuals, but it remains unclear of which species because her specimens originated from the aquarium trade. In AMNH I-28610 there are three cleared and stained individuals, and we expect that the published osteological characters were based only on these specimens.

Parenti (1981) stated that P. mento possesses an upturned lower jaw (vs. not upturned in other Aphaniid species). However, there exists a wide variation in mouth morphology throughout the family, particularly among Paraphanius and Anatolichthys species, rendering this character unsuitable for diagnosis of Aphaniid genera.

Esmaeili et al. (2020) distinguished Paraphanius from Aphaniops by possession of 9–14 (vs. 8–9) dorsal-fin rays. However, Villwock et al. (1983) counted 8–12 dorsal-fin rays in Aphaniops , while Akşiray (1948) counted 7– 11 branched dorsal-fin rays in some Turkish Paraphanius species. The large overlap in dorsal-fin ray counts indicate that this character cannot be used to distinguish these genera. Parenti (1981) stated that P. mento possesses a unique pattern of neuromasts on the head, but we did not find it distinctive from other Aphaniids lacking head canals.

Parenti (1981) also stated: “In A. mento , as well as in Orestias , Kosswigichthys , and Anatolichthys , the interhyal is cartilaginous and the urohyal is embedded ….”. Esmaeili et al. (2020), who treated Kosswigichthys and Anatolichthys as synonyms of Aphanius , misinterpreted this information when they wrote: “Also, according to Parenti [1981], Aphanius mento possesses several diagnostic features not found in other members of the genus Aphanius , including a cartilaginous interhyal (ossified in other Aphanius species), an embedded urohyal (not embedded in other Aphanius ),…”. However, Parenti (1981) clearly confirmed that she examined Paraphanius mento , Kosswigichthys asquamatus , Anatolichthys transgrediens and A. splendens , and that all share these two osteological characters. Teimori et al. (2018b) examined the urohyal bone in several aphaniid species and found Paraphanius to be nested within Aphaniops in a cladogram, but mention that Paraphanius mento can be distinguished from the studied Aphaniops species by morphometry of the urohyal bone. More research is needed to verify if the urohyal bone in Paraphanius is a unique shape.

Esmaeili et al. (2020) additionally distinguished Paraphanius by certain otolith characters and a thin or less developed epural (vs. thick and well developed in Aphanius and Aphaniops ) but give no details, which species were examined and how variable this character is. Esmaeili et al. (2020) only examined two juvenile Paraphanius. We recommend that these characters should be re-examined using a larger series of material.

While we fully support the conclusion of Esmaeili et al. (2020) to treat Paraphanius as a distinct genus, the only definitive diagnostic characters confirmed during the present study were elements of the unique colour pattern.

Material examined.

Paraphanius alexandri : FSJF 2318 , 21, 22–33 mm SL; Turkey: Hatay prov.: stream Arsuz east of Arsuz , 36.3992 35.8860. GoogleMaps — FSJF 2605 , 12, 34–46 mm SL; Turkey: Kahramanmaraş prov.: spring Çöçelli north of Çöçelli , south of Kahramanmaraş, 37.2812 37.1248. GoogleMaps — FSJF 3464 , 13, 22–39 mm SL; Turkey: Kahramanmaraş prov.: Elbistan , 38.1831 37.2194. GoogleMaps

Paraphanius boulengeri : FSJF 2506 , 12, 24–37 mm SL; GoogleMaps FSJF 3737 , 3, 32–39 mm SL; Turkey: Adıyaman prov.: River connecting lakes Gölbaşı and Azaplı south of Gölbaşı, 37.7903 37.6263. GoogleMaps

Paraphanius mento : NMW 59832 , 5 35–40 mm SL; Iraq: Mossul . — FSJF 2650 , 57, 16–52 mm SL; Syria: spring of stream Barada north-west of Damascus, 33.6753 36.0555. GoogleMaps — FSJF 2689 , 17, 29–44 mm SL; Syria: spring Al Fawwar , 33.2332 35.9248. GoogleMaps — FSJF 2701 , 21, 27–46 mm SL; Syria: Nahr al Tammasiyyar near Magsoofa, 33.2935 35.9707. GoogleMaps — FSJF 2657 , 8, 22–32 mm SL; Syria: River Orontes at Shayzar, 35.2716 36.5628. GoogleMaps — FSJF 2659 , 2, 31–43 mm SL; Syria: spring south of Qala’ at al Jarras, 35.3303 36.3106. GoogleMaps — FSJF 2683 , 16, 21–36 mm SL; Syria: River Orontes at Mashr’a al Bouz , 35.9508 36.3958. GoogleMaps — FSJF 2691 , 14, 22–34 mm SL; Syria: Reservoir of Nahr al Hawaiz , 35.3419 36.0156. GoogleMaps — FSJF 2758 , 54, 16–40 mm SL; Syria: River Orontes at Al Ghassaniyya, 34.5938 36.5336. GoogleMaps — FSJF 4001 , 2, 31–32 mm SL; Iraq: Shatt al Arab at Basra, 30.5395 47.832. GoogleMaps

Paraphanius mentoides : FSJF 2270 , 39, 26–61 mm SL; Turkey: Antalya prov.: spring Kırkgöz , 37.1098 30.5805. GoogleMaps — FSJF 3108 , 7, 35–42 mm SL; Turkey: Antalya prov.: stream at Döşemealtı , 37.0239 30.5918. GoogleMaps — FSJF 3678 , 26, 28–66 mm SL; Turkey: Bitlis prov.: Nemrut Lake , 38.6444 42.2367. GoogleMaps

Paraphanius orontis : FSJF 2431 , 9, 18–29 mm SL; Turkey: Hatay prov.: River Orontes at Sinanlı, 36.0974 36.0785. GoogleMaps — FSJF 3490 , 40, 25–37 mm SL; Turkey: Antalya prov.: Titreyengöl , 36.7524 31.4516. GoogleMaps

Paraphanius similis : FSJF 3126 , 14, 29–37 mm SL; Turkey: Konya prov.: Shallow pool east of Ereğli , 37.5390 34.0865. GoogleMaps — FSJF 2434 , 9, 15–24 mm SL; Turkey: Adana prov.: Seyhan below water regulation doors at Yüreyir , south of Adana , 36.9754 35.3354. GoogleMaps — FSJF 4012 , 5, 26–30 mm SL; Turkey: Niğde prov.: spring in Zengen , 37.8299 34.2499. GoogleMaps

Paraphanius striptus : FSJF 2680 , 1, 22 mm SL; Syria: Canal draining from spring at Al Asha’ari , 32.7389 36.0092. GoogleMaps — FSJF 3495 , 62, 19–31 mm SL; Israel: Einot Timsach east of Ma’agan Micha’el, 32.5510 34.9239. GoogleMaps — FSJF 3529 , 42, 23–36 mm SL; Israel: Nachal HaKibbutzim , 32.5031 35.5247. GoogleMaps