Phoxinus chrysoprasius ( Pallas, 1814 )

Bogutskaya, Nina G., Diripasko, Oleg A. & Palandačić, Anja, 2023, Novel data support validity of Phoxinus chrysoprasius (Pallas, 1814) (Actinopterygii, Leuciscidae), European Journal of Taxonomy 861, pp. 1-20 : 10-15

Phoxinus chrysoprasius ( Pallas, 1814)

Figs 5–6

Cyprinus chrysoprasius Pallas, 1814: 318 (mountain rivulets of the Crimea Peninsula).

Cyprinus phoxinus – Gablitz 1785: 182 (Salhir with tributaries Kuchuk-Karasu and Biyuk-Karasu).

Cyprinus chrysoprasius – Rathke 1836: 345 (Crimea). — Nordmann 1840: 483 (Crimea).

Phoxinus laevis – Kessler 1859: 539 (River Salhir at Simferopol’); 1877: 257 (partim: Crimea). — Dybowski 1862: 105 (partim: Crimea).

Phoxinus phoxinus – Berg 1912: 246 (partim: Crimea).

Morphological diagnosis

Among the species and clades examined of Phoxinus in this study, in the geographically close areas or drainages ( P. colchicus , P. csikii and P. strandjae ), P. chrysoprasius is distinguished by having a less scaled breast and anterior belly (type 3–6, commonly 3 or 4, vs type 3 to 12, commonly 5 to 11), the longest lateral line, with 55–91, commonly 71–90, pored scales, averaging 75.6 (vs 12–80, commonly 31–70, averaging 44.7–61.8), and the longest first uninterrupted section of the lateral line, 11–85 pored scales, commonly 21–60, averaging 47.1 (vs 2–71, commonly 11–50, averaging 15.2–36.0) pored scales. Phoxinus chrysoprasius is closest to the geographically distant Upper Danubian P. marsilii which is also characterised by a relatively less scaled breast and anterior belly (type 3–7, commonly 3 or 4) and a long lateral line (45–90, commonly 71–80, pored scales), but is distinguishable from the latter by a smaller difference between numbers of abdominal and caudal vertebrae, 3–6, averaging 4.8 (vs 1–4, averaging 3.0), a deeper caudal peduncle: caudal-peduncle depth 2.2–2.9, averaging 2.4, × in its length (vs 2.8–3.5, averaging 3.2), and no breeding tubercles on the breast scales in males (vs present).

Additionally, P. chrysoprasius is distinguished from other species or clades of Phoxinus ( P. krkae Bogutskaya et al., 2019 , P. lumaireul (Schinz, 1840) , P. septimaniae Kottelat, 2007 ) examined earlier ( Bogutskaya et al. 2019) by a combination of characters, none of which is unique, as follows: a deep caudal peduncle, minimum depth of caudal peduncle 2.2−2.9 × in caudal peduncle length; upper lip projecting beyond lower lip; rostral fold absent; mouth clearly subterminal, tip of mouth at level markedly below lowest point of eye; scales relatively small, total number of scales in lateral series 83−98; lateral line long, with 55−91 pored scales; lateral-line interruptions occurring in few places, commonly only on posterior body and caudal peduncle; scales on belly not extending over middle of the distance between pelvic- and pectoral-fin origins; patches of breast scales disconnected (commonly widely disconnected); total vertebrae commonly 40−41 (most frequent vertebral formulae 22 + 18 and 23 + 18); 4−6, commonly 5, anal-fin proximal pterygiophores anterior of first haemal spine of first caudal vertebra.

Molecular diagnosis (CO1 barcoding)

The single CO1 haplotype of P. chrysoprasius is presented in Supp. file 2. It is 33 mutational steps distant from the genetically closest mitochondrial clade (Clade 8: unnamed Phoxinus from Lake Ohrid) and 55 mutational steps distant from the geographically closest mitochondrial clade (Clade 19: unnamed Phoxinus from River Kuban’).

Etymology

The etymology is not explained in the original description. The species epithet, the adjective “ chrysoprasius ”, apparently refers to a characteristic feature of the colouration being a combination of “ chrys ” (Greek for “gold or yellow”) and “ prase ” (from the Greek for “leek”, referring to some greenish colouring also present).

Type series

Specimens belonging to the type series of the species, Cyprinus chrysoprasius Pallas, 1814: 318 , cannot be located at present and were apparently not preserved by Pallas (at least we are unaware of any published indications of their existence). The original description ( Pallas 1814: 318) does not unambiguously distinguish the species.

Material examined

Neotype ( Fig. 5A ) UKRAINE • ♂, 67.1 mm SL; upper reaches of the river Kuchuk-Karasu at Krasna Sloboda village, a tributary of Biyuk-Karasu, a tributary of Salhir; 44°58ʹ26ʺ N, 34°44ʹ19" E; 16 Jun. 2003; collectors N. Bogutskaya, O. Diripasko, A. Naseka and J. Freyhof leg.; ZFMK 93640–59 .

Remarks

It was examined morphologically and genetically (CO1) and designated herein as neotype. The entire set of data of the neotype in provided in Supp. file 2 (rhodopsin and CO1 data), and in Table 1 and Supp. files 3–4 (counts, scalation and morphometrics). The neotype has a moderately long lateral line and scalation pattern on the breast and anterior belly of type 3 (scale patches widely disconnected); scales in total lateral series, 86; scales in total lateral line, 55; scales in first uninterrupted section of lateral line, 39; predorsal abdominal vertebrae, 15; total vertebrae, 40; abdominal vertebrae, 22; and anal-fin pterygiophores anterior of first haemal spine, 5.

Comments on neotypification

Since the 1920s, Crimean freshwater drainages have been altered considerably by constructions of diverse network of irrigation channels and reservoirs, including the Severo - Krymsky (North Crimean) irrigation canal, which connected Crimea with the Dnieper drainage. As a result, 19 non-native taxa appeared in the Crimean inland waters ( Karpova & Boltachev 2011; Karpova 2017). Therefore, we cannot exclude anthropogenic translocations or invasions, although not yet recorded, of non-native Phoxinus on the peninsula, as has occurred with some clades of Phoxinus in other regions of Europe (reviewed in Palandačić et al. 2020). Neotypification provides nomenclatural support for the name Phoxinus chrysoprasius to be irrefutably applied to the native common minnow of the Salhir River in the Crimea Peninsula in the modern concept based on molecular data.

Accordingly, a specimen, examined morphologically and genetically (CO1), is designated herein as neotype for P. chrysoprasius under the conditions stipulated in art. 75.3 of the International Code of Zoological Nomenclature ( ICZN 1999). This act satisfies the provisions of art. 75 of the Code by clarifying its taxonomic identity in the modern concept (art. 75.3.1); nominating the above-mentioned combination of its phenotypic characters and CO1 haplotype as diagnostic features (art. 75.3.2); providing data and description sufficient to ensure recognition of the designated specimen (art. 75.3.3); giving reasons and references for believing that original type material is lost (art. 75.3.4); selecting a neotype consistent with the original description of the species (art. 75.3.5); choosing a neotype from the originally cited type locality, the Crimea Peninsula (art. 75.3.6); and recording that the neotype is the property of a recognised scientific institution, the Zoological Research Museum Alexander Koenig in Bonn (art. 75.3.7).

Description

The general appearance of Crimean Phoxinus is shown in Figs 5–6 ; meristic characters, types of scalation pattern on the breast and anterior belly and relative measurements of 29 adult specimens of both sexes (SL 49.5–87.2 mm), including the male neotype of P. chrysoprasius (designated above), are given in Supp. files 3–4.

Longest examined specimen (female) 87.2 mm SL ( Fig. 5B ). Body moderately stout, with deep caudal peduncle (minimum depth of caudal peduncle 2.2−2.9 × caudal peduncle length), dorsal and ventral profile roughly symmetrical with dorsal profile sometimes slightly more convex, and small hump at nape present in some larger specimens. Snout markedly stout and rounded. Mouth subterminal, tip of mouth at level to slightly to markedly below lowest point of eye, mouth cleft short. Upper lip projecting beyond lower lip, rostral fold absent.

Dorsal fin with 3 unbranched and 7½ branched rays. Distal dorsal-fin margin slightly concave, dorsal-fin origin markedly behind pelvic-fin insertion. Anal fin with 3 unbranched and 7½ branched rays, distal anal-fin margin is about straight or slightly convex.

Lateral line well developed though incomplete and interrupted in, commonly, posterior section only. First uninterrupted section of lateral line variable (11–85 scales), interruptions occurring in a few to many places in posterior region of body and caudal peduncle, lateral line containing 55–91 scales in total, commonly reaching beyond anal base. Total lateral series (including scales at caudal-fin base) 83–98 scales.

Scales on belly occasionally extending forward to beyond middle of distance between pelvic-fin base and pectoral-fin origin, but usually less. Patches of breast scales widely or narrowly separated (types 3 and 4) in 19 out of 29 examined specimens; few scales in between patches (type 5) and patches connected by 1–3 scales (type 6) found in 4 and 6 specimens, respectively.

Total vertebrae 39–42, commonly 40–41; abdominal vertebrae 22−23; predorsal vertebrae 14–15 (16 in one specimen); most common vertebral formulae 22 + 18 and 23 + 18; anal-fin pterygiophores anterior of first haemal spine not numerous, 4−6.

COLOURATION. In formaldehyde-preserved specimens ( Fig. 5 ), basic colour pattern consisting of 10–16 blackish mid-lateral bars or blotches, sometimes fused, not extending below horizontal of lower half of eye, pale lower half of flank and belly, and dark marking on back; black spot at middle of caudal-fin base. Dark bars or blotches are not apparent when the fish are alive, when the overall colouration is golden bronze or brown with a wide mid-lateral stripe of varying golden intensity overlaid on blackish stripe ( Fig. 6 ). Golden colouration first recorded as early as the 18 th century by Gablitz (1785) who wrote about the “gold” stripe on the sides of Crimean minnow, while three decades later, Pallas (1814) named the species referring to its “golden” colour. No extensive areas of pink, orange or red observed in adults collected in June 2003, immediately after spawning period, apart from a slight pink tinge to the bases of the pelvic and anal fins. Also, iridescent blue patches observed in larger adult males (in June) on the snout, gill covers and behind the head.

SEXUAL DIMORPHISM. Not observed with regard to the breast scalation type or meristic characters, though statistically significant for 10 morphometric characters, and prominent for % SL of pectoral-fin length (greater in males, reaching or almost reaching pelvic-fin origin), pelvic-fin length (greater in males, reaching or almost reaching anal-fin origin), anal-fin depth (greater in males), maximum body depth (greater in females), caudal-peduncle width (greater in females), head width (greater in females), and the % cranium roof length of cranium width between margins of pterotics (greater in females). DFA analysis clearly differentiated males from females in morphospace (DFA statistics values: Wilks’ Lambda 0.00006, approx. F (40.18) = 59.386, p <0.0000, perfect discrimination). Pectoral-fin in males broad and markedly rounded ( Fig. 5A ) with thickened rays vs narrow pointed fin without thickened rays in females ( Fig. 5B ). Genital papilla in males well developed, located just behind the anus and often protruding beyond the latter vs absence of genital papilla or a poorly developed papilla located at some distance from the anus in females. Mature males bear breeding tuberculation on head (size of the tubercles varies significantly; we roughly grouped them as ‘large’, ‘medium’, ‘small’ and ‘very small’). Breeding tubercles arranged in a regular manner and in larger sized (> 55 mm SL) males ( Fig. 5A ). Paired rows (number of tubercles on both sides of the head): in front of the naris (1 or 2, medium to large), between nares (2, large), in front of and below the orbit (0–4, commonly 0 or 1; commonly small if present) and along dorsal rim of the orbit (3–6, medium to large). Unpaired rows: a group of tubercles in interorbital space between rows along the dorsal rim of the orbit (3–7, small to large) and at the nape (7–28, small to medium). Tubercles along the temporal sensory canal and just behind the head along the lateral line, with variable number (0–13, very small to medium if present); the most numerous tubercles in this area – 6 on the left and 13 on the right – were found in the largest male of 71 mm SL. Tubercles were apparent neither on pectoral fin nor on breast scales.

Distribution

Type locality The type locality from the original description ( Pallas 1814: 318) reads “In rivulis saxosis montosae orae Chersonesi taurica” (“at rocky banks of mountainous rivulets of the Tauric Chersonese [Crimea

Peninsula]”).

Note on type locality

The type locality of P. chrysoprasius is commonly misinterpreted because of incorrect translation of the original Latin text. For example, in Eschmeyer’s Catalog of Fishes, the type locality is given as “hill streams on Mount Chersones, Crimea ” ( Fricke et al. 2021) but there is no Mount Chersones in the Crimea. The word “chersonese” (ancient Greek “χερσόνησος”) translates as “a peninsula”, and “Chersonesi taurica” literally means “the peninsula of Tavria”. The toponym Tauric Chersonese, applied to the whole of the Crimean Peninsula, should not be confused with the name Chersonesos, an ancient Greek colony, which aptly describes the site (a peninsula) on which the colony was established.

Peter Simon Pallas, a native German, worked in Russia from 1767 on and performed extensive natural history studies in various regions of the Russian Empire. During the period 1793–1794, he conducted expeditionary research in its southern regions, as a result of which he published, in German, a twovolume book “ Bemerkungen auf einer Reise in die südlichen Statthalterschaften des Russischen Reichs in den Jahren 1793 and 1794 ” (Observations made during a trip to the southern governorships of the Russian state in 1793–1794) ( Pallas 1801). The second volume contains a description of the journey to the Crimea. Even from the first paragraphs of this description, it is clear that Pallas was well acquainted with the history of the peninsula, using the word “Crimea” (derived from the Tatar toponym Qirim or Qyrym), and at the same time gives its ancient name, the Tauric Chersonesus, for example, “Der erste Wohnplatz der eigentlichen Krym oder des Alters so genannten Taurischen Chersones, ist Perekop, oder Or-Kapi, wie es die Tataren nennen” (The first dwelling place of the actual Krym or of the age of the, socalled Taurian Chersonese, is Perekop, or Or-Kapi as the Tatars call it) ( Pallas 1801: 4). While staying in the vicinity of Sevastopol, Pallas paid attention to the description of the ancient town of Chersones, and it is clear that he distinguished between the name of the ancient Greek colony and the name of the peninsula.

Comments on distribution

The Crimean minnow was first reported by Gablitz (1785: 182) in the upper reaches of the Salhir River and its tributaries Kuchuk-Karasu and Biyuk-Karasu (= Malaya Karasevka and Bol’shaya Karasevka)

under the name Cyprinus phoxinus . Pallas visited Salhir during his trips and described its fishes ( Pallas 1801: 20) but later Pallas (1814) did not specify the names of the rivers where he observed the species described as C. chrysoprasius , perhaps because he took for granted the occurrence of the species in other rivers of Crimea with a similar biotope. Rathke (1836: 345) and Nordmann (1840: 483) also used the name Cyprinus chrysoprasius in their studies of the fauna of Crimea, and also provided no indication of the rivers where the species occurred, though Nordmann (1840: 483) mentioned that it was “abundant in all rivers of Crimea, especially in the Salhir”. In later publications on the freshwater fish fauna of the Russian Empire, researchers indicated the distribution of C. chrysoprasius Pallas, 1814 (synonymised with Phoxinus laevis Agassiz, 1832 or Phoxinus phoxinus (Linnaeus, 1758)) as ‘Crimea’: Kessler (1859: 539– 540) described specimens of the Salhir drainage in the Simferopol region; Dybowski (1862: 105) reported it “in the Crimean mountain rivers”; and Kessler (1877: 257) and Berg (1912: 260), as “occurring in the Crimea”. Tseeb (1929) was apparently the first to limit the range of ‘ Phoxinus phoxinus ’ on the Crimea Peninsula to the rivers of the Salhir drainage (Sea of Azov basin), emphasising that it did not occur in the rivers of the Crimea flowing to the Black Sea. Berg (1949: 590) noticed this significant clarification, and corrected the range of Crimean Phoxinus to “in the Crimea, but only in the Salhir drainage”.

The historical and present distribution of the common minnow in Crimea has been thoroughly revised and analysed in recent publications ( Miroshnichenko 2003; Karpova & Boltachev 2011, 2012; Karpova 2017) clearly indicating that it only occurs in the Salhir River drainage.

Figures

Fig. 5. A. Neotype of Phoxinus chrysoprasius (Pallas, 1814), ♂ (ZFMK 93640-59). B. ♀ (ZFMK 93640- 59), 87.2 mm SL, same locality and date as the neotype.

Fig. 6. Phoxinus chrysoprasius (Pallas, 1814), living specimen, spawning male, Angara River, Salhir drainage, 44°51ʹ12ʺ N, 34°18ʹ47ʺ E; 16 Jun. 2003.