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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4294 (5): 573
–
585
http://www.mapress.com/j/zt/
Article
573
https://doi.org/10.11646/zootaxa.4294.5.6
http://zoobank.org/urn:lsid:zoobank.org:pub:D2E6BE18-6CC3-43DD-B509-36AE0D78DE51
Neotype designation of Aphanius iconii, first reviser action to stabilise the usage
of A. fontinalis and A. meridionalis and comments on the family group names of
fishes placed in Cyprinodontidae (Teleostei: Cyprinodontiformes)
JÖRG FREYHOF
1
, MÜFİT ÖZULUĞ
2
& GÜLŞAH SAÇ
2
1
Leibniz-Institute of Freshwater Ecology and Inland Fisheries (IGB), Müggelseedamm 310, 12587 Berlin, Germany.
E-mail: j.freyhof@igb-berlin.de
2
Istanbul University, Science Faculty, Department of Biology, 34134 Vezneciler, İstanbul, Turkey.
E-mail: mozulu@istanbul.edu.tr, gulsahsac@gmail.com
Abstract
As First Revisers, we give priority to Aphanius fontinalis over A. altus and A. litoralis and to A. meridionalis over A. par-
vus. A neotype is designated from Lake Eğirdir for Aphanius iconii. Family groups in the deeply polyphyletic fish family
Cyprinodontidae are discussed. Aphaniidae, Cubanichthyidae and Orestiidae are proposed as distinct families, restricting
the family Cyprinodontidae to the New World genera Cualac, Cyprinodon, Floridichthys, Jordanella and Megupsilon.
Key words: Biodiversity, killifish, freshwater fish, taxonomy
Introduction
Akşiray (1948) published the description of one new species and 15 new subspecies of Aphanius from Turkey.
Villwock (1958, 1964) as well as Wildekamp (1993) and Wildekamp et al. (1999) did not accept any of these 16
taxa as valid. Wildekamp et al. (1999) placed A. cypris alexandri, A. cypris boulengeri, A. cypris orontis, A.
sophiae similis and A. sophiae mentoides in the synonymy of A. mento. They also synonymized A. burduricus
iconii, A. chantrei aksaranus, A. chantrei altus, A. chantrei flavianalis, A. chantrei fontinalis, A. chantrei litoralis,
A. chantrei maeandricus, A. chantrei meridionalis, A. chantrei obrukensis and A. chantrei parvus with A. anatoliae
and treated A. burduricus burduricus as a synonym of A. anatoliae sureyanus.
Hrbek et al. (2002) and Geiger et al. (2014) studied molecular characters in different Anatolian Aphanius
species and found A. anatoliae, as understood by Wildekamp et al. (1999), to be highly polyphyletic. Geiger et al.
(2014) therefore suggested treating A. iconii, A. fontinalis, A. maeandricus and A. meridionalis as valid species, a
view accepted by Pfleiderer et al. (2014).
The type localities of A. chantrei aksaranus, A. chantrei flavianalis and A. chantrei obrukensis are all situated
in the wider Lake Tuz basin and fishes from the respective type localities have been included by Hrbek et al. (2002)
and/or Geiger et al. (2014) in their molecular studies, demonstrating that these names are indeed likely to be
synonyms of A. anatoliae. Geiger et al. (2014) and Pfleiderer et al. (2014) identified fishes from Lake Işıklı basin
and springs in Dinar as A. maeandricus, those from Lake Eğirdir as A. iconii, those from springs in former Lake
Söğüt in the Taurus Mountains as A. meridionalis, and those from a spring near Lake Yarışlı near Burdur as A.
fontinalis. The situation in A. maeandricus needs no discussion as it is the only name available for this species. But
the situation is not as clear in the cases of A. iconii, A. meridionalis and A. fontinalis.
Akşiray (1948) described A. burduricus iconii based on a syntype series consisting of fishes from three
different origins: from Lake Eğirdir, from the area of Çumra and from Hotamış Lake. The last two locations are
situated in the Konya endorheic basin in Central Anatolia. All materials examined by Akşiray (1948) are lost.
Hrbek et al. (2002) found the fishes from the Lake Eğirdir basin to be clearly distinguished from other Anatolian
Aphanius but those from the Konya endorheic basin to be very similar to A. anatoliae.
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Akşiray (1948) described A. chantrei meridionalis from Lake Söğüt and A. chantrei parvus from the
geographically close Lake Gölhisar basin. Hrbek et al. (2002) and Geiger et al. (2014) studied A. meridionalis from
the Lake Söğüt basin and supported it status as a valid species but could not include specimens from Lake Gölhisar
basin as these seemed to be extirpated. The molecular study by Hrbek et al. (2002) revealed another clade
including fishes from the Lake Burdur and Lake Salda basins. Akşiray (1948) described A. chantrei fontinalis and
A. chantrei litoralis (Lake Burdur basin) and A. chantrei altus (Lake Salda basin) in the same publication.
To stabilise the names in Aphanius, we act as First Revisers to give priority of A. fontinalis over A. altus and A.
litoralis whenever the two latter names, either of them or both, are considered to be synonyms of A. fontinalis; and
of A. meridionalis over A. parvus whenever the two names are considered to be synonyms; and we designate a
neotype for A. iconii.
Working on Aphanius, we also reviewed the molecular studies on cyprinodontiform phylogeny and found
strong disagreements between published family assignment and phylogenies of killifish and livebearers. While
phylogenetic studies published in high-ranked journals do rarely commit to taxonomic conclusions, the revision of
the family group Cyprinodontidae is more than overdue. Here we propose a new family structure for the killifishes
previously placed in the family Cyprinodontidae.
Material and methods
After anaesthesia, fishes were fixed in 5% formaldehyde and stored in 70% ethanol or directly fixed in 99%
ethanol. Measurements were made with a dial caliper and recorded to 1 mm. All measurements were made point-
to-point, never by projections. Methods for counts and measurements follow Kottelat & Freyhof (2007). Standard
length (SL) is measured from the tip of the snout to the posterior extremity of the hypural complex. The length of
the caudal peduncle is measured from behind the base of the last anal-fin ray to the posterior extremity of the
hypural complex, at mid-height of the caudal-fin base. The last two branched rays articulating on a single
pterygiophore in the dorsal and anal fins are counted as "1½". Simple rays of dorsal and anal fins are not counted as
they are deeply embedded. The neotype is included in the calculation of means and standard deviations (SD) of
males.
Collection codes: FSJF, Fischsammlung J. Freyhof, Berlin; IUSHM, Istanbul University, Science Faculty,
Hydrobiology Museum, İstanbul; ZMH, Zoologisches Museum Hamburg, Hamburg. Authors of species and family
names are listed in Table 1.
First Reviser actions to stabilise the use of A. fontinalis and A. meridionalis
The First Reviser action is the principle that in cases of conflicts between simultaneously published names, the first
subsequent author can decide which has precedence (ICZN 1999: Article 24.2). Akşiray (1948) published the
description of one new species and 15 new subspecies of Aphanius from Turkey. Neither Villwock (1958, 1964),
Wildekamp (1993), Wildekamp et al. (1999) or any later author accepted any of these 16 taxa as valid. Recently,
Geiger et al. (2014) and Pfleiderer et al. (2014) accepted A. fontinalis and A. meridionalis as valid species but did
not discuss the other available names for these species published by Akşiray (1948).
Akşiray (1948) described in the same publication A. chantrei fontinalis from a spring near Lake Yarışlı, A.
chantrei altus from the spring Karapınar near Yeşilova, and A. chantrei litoralis from Lake Bahçeözü (Lake
Karataş). Hrbek et al. (2002) include in their molecular study fishes from the spring Karapınar near Yeşilova (A.
chantrei altus), Pınarbaşı near Kemer in Lake Karataş basin (A. chantrei litoralis), Lake Burdur (A. sureyanus) and
Düger (a spring in Lake Burdur basin). In the trees shown by Hrbek et al. (2002), all these populations form one
group of very closely related populations. Geiger et al. (2014) include also A. sureyanus and fish from a spring near
Lake Yarışlı (A. fontinalis, Fig. 1) in their study, both being also very closely related. We see no reason to reject the
results by Hrbek et al. (2002) and Geiger et al. (2014) and treat all these populations, except those from Lake
Burdur, as conspecific. All these fish are distinguished by having a wide black anal-fin margin in males (vs. narrow
in A. anatoliae, A. maeandricus, A. meridionalis, A. danfordii, A. marassantensis and A. villwocki) and the body
being completely covered by scales (vs. scales reduced in A. transgrediens, A. saldae, A. splendens and A.
sureyanus).
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As First Revisers, we give precedence of A. fontinalis over A. chantrei altus and A. chantrei litoralis. Aphanius
sureyanus is treated as distinct from A. fontinalis despite these two species sharing the molecular character states
examined so far (Hrbek et al. 2002, Geiger et al. 2014). Aphanius fontinalis is distinguished from A. sureyanus by
being completely covered by scales (vs. scales reduced in A. sureyanus). There is a large variability in scale
patterns within A. sureyanus (Villwock, 1963). It cannot therefore be fully excluded that A. fontinalis might turn
out to be a synonym of A. sureyanus; further studies are needed to clarify the situation.
Akşiray (1948) described in the same publication A. chantrei meridionalis from Lake Söğüt (Fig. 2) and A.
chantrei parvus from the Lake Gölhisar basin (Fig. 3). Until now, only A. meridionalis has been accepted as a valid
species, by Geiger et al. (2014). Aphanius parvus has not yet been studied for molecular characters as it has not
been recorded in the last 25 years (Wildekamp et al. 1999) or perhaps even since 1957. It cannot be fully excluded
that this population might be extirpated. We examined preserved material from Lake Gölhisar (Fig. 3) collected in
1957, including some captive-breed fishes from this stock, likely to have been preserved a few years later (ZMH
3502). As First Revisers, we give A. meridionalis precedence over A. parvus. These populations are considered
conspecific as the adult males of both have a completely black dorsal fin (vs. dorsal fin with a white or hyaline base
in A. anatoliae, A. fontinalis, A. saldae, A. splendens, A. sureyanus and A. transgrediens, dorsal fin with white or
hyaline blotches or bands above the dorsal-fin base in A. maeandricus, A. iconii, A. danfordii and A.
marassantensis) and a narrow black anal-fin margin (vs. wide margin or anal fin almost completely black in A.
fontinalis, A. saldae, A. splendens, A. sureyanus and A. transgrediens). Field work is encouraged around Lake
Gölhisar to clarify whether this population might still be present or has been extirpated. Aphanius meridionalis,
however, might be more widespread. While Hrbek et al. (2002) and Geiger et al. (2014) found it also in the Lake
Avlan basin, we identified as A. meridionalis a population (Fig. 4) from a small reservoir south of Yeşilova in the
Lake Salda basin.
TABLE 1. Family names and species mentioned in this study, and their authors
Aphaniidae Hoedeman, 1949
Cubanichthyidae Parenti, 1981
Cyprinodontidae Wagner, 1828
Orestiidae Bleeker, 1859
Valenciidae Parenti, 1981
Aphanius anatoliae (Leidenfrost, 1912)
Aphanius burduricus Akşiray, 1948
Aphanius burduricus iconii Akşiray, 1948
Aphanius chantrei aksaranus Akşiray, 1948
Aphanius chantrei altus Akşiray, 1948
Aphanius chantrei flavianalis Akşiray, 1948
Aphanius chantrei fontinalis Akşiray, 1948
Aphanius chantrei litoralis Akşiray, 1948
Aphanius chantrei maeandricus Akşiray, 1948
Aphanius chantrei meridionalis Akşiray, 1948
Aphanius chantrei obrukensis Akşiray, 1948
Aphanius chantrei parvus Akşiray, 1948
Aphanius cypris alexandri Akşiray, 1948
Aphanius cypris boulengeri Akşiray, 1948
Aphanius cypris orontis Akşiray, 1948
Aphanius danfordii (Boulenger, 1890)
Aphanius marassantensis Pfleiderer, Geiger & Herder, 2014
Aphanius mento (Heckel, 1843)
Aphanius transgrediens (Ermin, 1946)
Aphanius saldae (Akşiray, 1955)
Aphanius sophiae mentoides Akşiray, 1948
Aphanius sophiae similis Akşiray, 1948
Aphanius splendens (Kosswig & Sözer, 1945)
Aphanius sureyanus (Neu, 1937)
Aphanius villwocki Hrbek & Wildekamp, 2003
Chriopeoides pengelleyi, Fowler, 1939
Yssolebias martae (Steindachner, 1875)
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FIGURE 1. Aphanius fontinalis, FSJF 3456, male, 42 mm SL; Turkey: Yarışlı.
FIGURE 2. Aphanius meridionalis, FSJF 2460, female, 38 mm SL, male, 33 mm SL; Turkey: Söğüt.
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FIGURE 3. Aphanius meridionalis, ZMH 3502, from the top, 39 mm SL, 28 mm SL, 27 mm SL, 25 mm SL; Lake Gölhisar.
FIGURE 4. Aphanius meridionalis, FSJF 3669, male, 29 mm SL; Turkey: reservoir south of Yeşilova.
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Neotype designation for Aphanius iconii Akşiray, 1948
Akşiray (1948) figured nine syntypes of Aphanius iconii, of which five originate from Lake Eğirdir (Plate 3: 21–
25) and four from the surroundings of Konya (Plate 4: 43–46). Because Akşiray (1948) did not explicitly mention a
holotype in his original description of A. iconii, all nine specimens included by Akşiray (1948) become syntypes
(ICZN, 1999: Art. 72.2 and Art. 73.4). Data presented by Hrbek et al. (2002) already indicate that two species are
hidden under the name A. iconii. While the populations from the surroundings of Konya can be identified as A.
anatoliae, the populations from the Lake Eğirdir basin represent a distinct species. We could examine preserved
materials from both populations and also found specimens from a place between Konya and Çumra (ZMH 3495)
identical to A. anatoliae and distinct from those from Lake Eğirdir. As two species are involved under the name A.
iconii, a neotype designation is needed to secure the name A. iconii to one of the identified species. Following
ICZN (1999; Art. 74.1), we designate here as lectotype the male specimen shown by Akşiray (1948) on plate 3 as
number 23 (36 mm total length), which was collected in Lake Eğirdir. This male lectotype shows the diagnostic
wide, white band above a black base in the dorsal fin. As this lectotype is lost, we designate the specimen IUSHM
2017-1272 as neotype of A. iconii (Fig. 5).
ICZN (1999: Art. 75.3) prescribes seven qualifying conditions to designate a neotype. Art. 75.3.1. requires a
statement that the neotype is designated with the express purpose of clarifying the taxonomic status or the type
locality of a nominal taxon. As two species are involved in the syntype series, the neotype designation clarifies the
type locality and the taxonomic status of the species. Art. 75.3.2. requires a statement of the characters that the
author regards as differentiating the species from its congeners. The diagnosis given below satisfies this
requirement. Art. 75.3.3. requires data and description sufficient to ensure recognition of the specimen designated.
The diagnosis and description given below satisfies this requirement. Art. 75.3.4. requires that the author's reasons
for believing the name-bearing type specimen(s) to be lost or destroyed be stated, together with the steps taken to
trace it or them. Akşiray (1948) is not known to have preserved any material and no museum material has ever
been claimed to owe its provenance to Akşiray’s (1948) descriptions of Aphanius. We also searched in ZMH and
IUSHM again without success. Art. 75.3.5. requires evidence that the neotype is consistent with what is known of
the former name-bearing type from the original description. The most important diagnostic character of the species
(see below) is visible in the lectotype and in the neotype. Art. 75.3.6. requires that the neotype comes from a
location as close as practicable to the original type locality (of the lectotype), meaning from Lake Eğirdir, which it
does. Art. 75.3.7. requires a statement that the neotype is the property of a recognized scientific or educational
institution, cited by name, that maintains a research collection, with proper facilities for preserving name-bearing
types, and that makes them accessible for study. IUSHM fulfil this requirement.
The name Aphanius iconii is now definitively restricted to the species known from Lake Eğirdir and the use of
the name stabilised. A description of the neotype and some individuals collected with the neotype is given below.
Aphanius iconii Akşiray, 1948
(Figs. 5–7)
Neotype. IUSHM 2017-1272, male, 25 mm SL; Turkey: Isparta prov.: spring Karaot at shore of Lake Eğirdir,
about 4 km north of Yenice, 38°08.094'N 30°54.443'E.
Additional material. FSJF 2325, 16, 25–32 mm SL; IUSHM 2017-1273, 10, 23–28 mm SL; same data as
neotype.—FSJF 2476, 6, 28–39 mm SL; IUSHM 2017-1274, 13, 26–34 mm SL; Turkey: Isparta prov.: lower
stream Çayköy at Koysazı bridge, southeast of Eğirdir, 37°50.488N 30°53.493'E.—FSJF 2271, 16, 18–35 mm SL;
IUSHM 2017-1281, 7, 19–32 mm SL; Turkey: Konya prov.: spring Eflatun Pınarı at Sadıkhacı, 37°49.507'N
31°40.458'E.
Diagnosis. Aphanius iconii belongs to the A. anatoliae group in which the following species are recognised as
valid: A. anatoliae, A. fontinalis, A. maeandricus, A. meridionalis, A. danfordii, A. marassantensis, A.
transgrediens, A. saldae, A. splendens, A. sureyanus and A. villwocki. Adult male A. iconii are distinguished from
adult males of all these species by having a black dorsal-fin base followed by a wide, white or hyaline band at
about below the middle of the dorsal fin (vs. dorsal fin with a white or hyaline base or base and proximal part of
dorsal fin hyaline in A. anatoliae, A. fontinalis, A. saldae, A. splendens, A. sureyanus and A. transgrediens, dorsal
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fin with white or hyaline blotches shortly above the black dorsal-fin base in A. maeandricus, A. danfordii and A.
marassantensis). Adult male A. iconii are also distinguished from adult male A. fontinalis, A. saldae, A. splendens,
A. sureyanus and A. transgrediens by having a narrow black anal-fin margin (vs. margin wide or anal fin almost
completely black).
Description. See Figures 5–7 for general appearance and Table 2 for morphometric data of neotype and 10
additional individuals. Dorsal head profile concave. Dorsal profile convex from nape shortly behind head to end of
dorsal-fin base or to mid-height of back. Ventral profile gently convex or almost straight from breast to anal-fin
origin. Profile straight along caudal peduncle. Body compressed. Body deepest at about dorsal-fin origin. Greatest
body width at pectoral-fin base. Lower jaw sharply upturned, oriented nearly perpendicular to body axis. Caudal
peduncle compressed laterally, 1.6 times longer than deep in neotype, 1.4–1.7 in males 1.7–2.0 in females. No
axillary lobe at base of pelvic fin. Pectoral fin rounded, reaching 80–90% of distance between pectoral- and pelvic-
fin bases. Pelvic-fin origin four predorsal scales in front of vertical of dorsal-fin origin. Pelvic fin not reaching or
reaching to anus. One large scale between pelvic-fin bases. Anus situated slightly in front of anal-fin origin, tissue
around genital papillae swollen in nuptial female. Anal fin with convex posterior margin. Dorsal fin roundish in
females, reaching to vertical of middle of last anal-fin ray, elongated in males, reaching to of vertical of tip of anal
fin. Extremity of dorsal fin rounded in both sexes. Caudal fin rounded to truncate. Largest individual examined 39
mm SL.
TABLE 2. Morphometric data of Aphanius iconii (neotype IUSHM 2017-1272, IUSHM 2017-1273, n=5 males and 5
females). Means, range and SD of males include neotype.
Dorsal fin with 10½ (8), 11½ (4), branched rays. Anal fin with 9½ (9)–10½ (3) branched rays. Caudal fin with
7+7 (8), 7+6 (1), 7+8 (1) or 8+8 (2) branched rays. Pectoral fin with 13 (3), 14 (9) and pelvic fin with 6 rays. Scales
neotype other individuals
male males (5) females (5)
mean range SD mean range SD
Standard length (mm) 26 28–29 23–29
In percent of standard length
Head length 30.5 30.2 29.1–30.9 0.6 30.3 28.4–32.5 1.5
Body depth at anal-fin origin 26.4 25.5 23.6–26.4 1.0 21.6 20.8–23.0 1.0
Predorsal length 58.1 56.8 55.4–58.1 1.0 57.3 56.5–58.1 0.7
Preanal length 65.7 65.1 64.0–66.1 0.9 67.3 65.5–69.1 1.5
Prepelvic length 51.0 49.4 48.2–51.0 1.0 50.4 48.4–52.0 1.5
Distance between pectoral and pelvic-fin origins 20.4 19.5 17.9–20.4 1.0 19.9 18.9–20.8 0.9
Distance between pelvic and anal-fin origins 16.4 17.4 16.4–18.0 0.6 17.8 16.0–18.9 1.4
Depth of caudal peduncle 16.5 15.7 14.8–16.6 0.9 14.0 12.9–15.6 1.0
Length of caudal peduncle 25.9 24.6 23.2–26.1 1.2 25.4 23.2–26.8 1.4
Dorsal-fin base length 19.3 20.0 17.6–22.8 1.9 17.2 15.1–20.3 2.0
Anal-fin base length 12.7 13.5 12.1–15.0 1.0 10.9 10.2–11.3 0.4
Pectoral-fin length 18.0 17.4 16.4–18.5 0.9 17.4 15.5–19.9 1.7
Pelvic-fin length 14.1 14.4 14.0–15.4 0.5 13.5 13.0–14.0 0.4
In percent of head length
Head depth at eye 56 57.1 55–59 1.4 55.4 28–32 2.8
Snout length 25 26.3 24–29 1.8 25.5 21–23 1.5
Eye diameter 37 35.1 33–37 1.5 35.6 56–58 1.7
Postorbital distance 46 46.6 45–51 2.2 46.0 65–69 1.1
Maximum head width 59 60.6 58–64 2.4 58.1 48–52 2.0
Interorbital width 34 34.7 31–37 1.9 33.5 19–21 1.7
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large and cycloid. Trunk and head entirely scaled. One scale row on upper part of opercule. Flank with 27 (8), 28
(8), 29 (1) scales along lateral series. Two (15) or three (2) rows of small scales on anterior caudal-fin base. Lateral
line incomplete, with 4–13 pores, scales pored mostly behind vertical of pelvic-fin origin. Teeth tricuspid, median
cusp longer than laterals.
FIGURE 5. Aphanius iconii, IUSHM 2017-1272, 25 mm SL, neotype; Turkey: Karaot.
FIGURE 6. Aphanius iconii, FSJF 2325, female, 30 mm SL; Turkey: Karaot; FSJF 2476, male, 25 mm SL; Turkey: Çayköy.
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FIGURE 7. Aphanius iconii, FSJF 2271, 30 mm SL; Turkey: spring Eflatun Pınarı.
Coloration. See Figures 5–7 for general appearance. Males in life: Head and flank silvery or yellowish brown.
Cheek, breast and belly whitish or pale yellow. Many small black spots on head and flank. Flank with 7–11 wide,
brown or yellowish-brown bars confluent with brown back. Interspaces silvery, much narrower than bars. Scales on
back with silvery or yellowish margin. Top of head pale brown. Pectoral fin hyaline or yellow. Pelvic fin hyaline or
yellow with narrow black margin. Anal fin hyaline or yellow with narrow black margin, absent in some individuals
and in juveniles, and few black blotches on posterior part in most individuals. Dorsal fin black with a black base
and a wide, hyaline or white band below middle of fin. Dorsal fin grey distally in some individuals. Caudal fin
hyaline or yellow with 1–2 bold black bands. Yellow colour faded in preserved fish.
In life and in preserved females: Body pale grey. Ventral part of head and belly silvery. Head and flank with
numerous, small, dark-brown or black blotches, usually horizontally elongates, irregularly set and shaped. Paired
fins hyaline, unpaired fins with dark-brown or black blotches, irregularly set and shaped, at base.
Distribution. Aphanius iconii is widely distributed in Lake Eğirdir basin and is known also from Lake
Kovada, which is close to Lake Eğirdir.
Remarks. Hrbek et al. (2002) as well as Geiger et al. (2014) included an Aphanius population from Eflatun
Pınarı in Lake Beyşehir basin in their analyses. In the study by Hrbek et al. (2002), these fish cluster with A.
anatoliae. They form a group slightly differentiated but closely related to A. anatoliae in the tree shown by Geiger
et al. (2014). We identify the Aphanius population from Eflatun Pınarı as A. iconii as they show the diagnostic wide
hyaline band above a black base in the dorsal fin in males (Fig. 7). It is beyond the aims of this study to analyse this
population further to resolve the disagreement between the molecular placement of these fishes and their male
colour pattern. We speculate, that introgressive hybridisation of A. anatoliae into A. iconii might have had occurred
in the Lake Beyşehir basin. An in-depth study of the Aphanius of Lake Beyşehir basin might help to resolve this
question in the future.
Aphaniidae as a valid family name for Western Palaearctic killifishes
While working on the Aphanius species discussed above we questioned the retention of this group of fishes in the
otherwise American family Cyprinodontidae. Recent phylogenetic studies place Aphanius outside of
Cyprinodontidae and more close to Valenciidae and Poeciliidae (Pohl et al. 2015, Helmstetter et al. 2016, Reznick
et al. 2017). The killifishes of the Western Palaearctic are usually placed in two families, Valenciidae and
Cyprinodontidae (Myers 1928, Parenti 1981, Kottelat & Freyhof 2007, Van der Laan et al. 2014). While the family
Valenciidae is well accepted as a distinct family endemic to Europe, Parenti (1981) and Van der Laan et al. (2014)
included four quite distinct groups of fishes in the family Cyprionodontidae. These are five related genera (Cualac,
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Cyprinodon, Floridichthys, Jordanella, Megupsilon) from North, Central and northern South America, one genus
(Aphanius) from the Western Palaearctic and North to East Africa, one genus from the high Andes (Orestias), one
genus from the Caribbean and Colombia (Cubanichthys). Parenti (1981) opposed the allocation of Cubanichthys to
a separate subfamily (Cubanichthyinae) as distinct from the other genera (Cyprionodontinae), distinguished two
tribes in Cyprionodontinae (Cyprionodontini, Orestinii) and joined Aphanius and Orestias in the tribe Orestinii,
which is diagnosed by an "extremely robust lower jaw caused by a median extension of the lower dentary": Parenti
(1981:521).
Recently, Pohl et al. (2015), Helmstetter et al. (2016), and Reznick et al. (2017), published comprehensive,
multigene phylogenetic analyses of cyprinodontiform fishes, demonstrating that the family Cyprinodontidae, as
proposed by Parenti (1981), is clearly polyphyletic. As the family structure of cyprinodontiform fishes was not the
focus of these molecular studies, they did not draw any taxonomic conclusions. All three molecular studies found
Aphanius to be more closely related to Val e nc i a , the livebearers of the families Anablepidae and Poeciliidae, and
the African lampeyes (Aplocheilichthys and related genera), than to other genera in Cyprinodontidae. We thus see
no argument to retain Aphanius in Cyprinodontidae. The New World genera of the family Cyprinodontidae are
related to the killifish families Fundulidae, Profundulidae, and the livebearers of the Mexican family Goodeiidae.
Also, the genera Cubanichthys and Orestias are highly isolated and there is no indication that they belong to
Cyprinodontidae.
Pohl et al. (2015) and Helmstetter et al. (2016) recovered Aphanius as a sister group to Vale n c i a and suggested
that they might be included in the family Vale n c i i d ae . Parenti (1981:500) diagnosed Valenciidae by having a
unique, very long, attenuate dorsal process of the maxillary (vs. dorsal process rounded or absent in Aphanius and
other families discussed here). The divergence between Valencia and Aphanius is postulated to have occurred in the
Late Eocene (Helmstetter et al. 2016) and both are here accepted as separate families, Valenciidae and Aphaniidae.
Aphanius was previously placed in the tribe Orestinii (Parenti 1981) or treated as contained in a separate tribe
(Aphaniini) in the cyprinodontid subfamily Cyprinodontinae (Van der Laan et al. 2014). Moreover, we propose that
the two Neotropical killifish genera Cubanichthys and Orestias be allocated to their own families, Cubanichthyidae
and Orestiidae, respectively.
Cubanichthyidae, previously treated as a subfamily of Cyprinodontidae (Parenti 1981, Van der Laan et al.
2014), was well diagnosed by Parenti (1981). We follow Costa (2015) and Reis et al. (2003) in accepting
Yssolebias martae and Chriopeoides pengelleyi as species of Cubanichthys.
We cannot avoid to discussion of the stem of Orestias (see Van der Laan et al. 2014 for details). Maurice
Kottelat was consulted to help us in this case (M. Kottelat, pers. comm.). Gill (1896:224) used Orestiad-, Jordan &
Evermann (1896:631) used Oresti- and Myers (1931) used Orestiat- as a stem of Orestias. Orestias is a feminine
word in classical Greek (a mountain nymph). The genitive singular is orestiados and the stem is orestiad-. Thus, the
family name should be Orestiadidae, as already observed by Gill (1896: 224). But ICZN art 29.5 says "if a spelling
of a family-group name was not formed in accordance with article 29.3 but is in prevailing usage, that spelling is to
be maintained, whether or not it is the original spelling [...]". (prevailing usage: Oresti-idae; original spelling
Orestias-idae; according to 29.3: Orestiad-idae). It is far beyond the aim of this study to count how often different
stems have been used, but it is evident that all have been used several times. It seems that the stem Oresti- is in
prevailing usage, as used by Parenti (1981); many other widely accepted publications too, use this name (Reis et al.
2003, Nelson et al. 2016, just to name few). Therefore, we use Orestiidae as the family name of the fishes of the
genus Orestias.
Following the concept proposed here, all five family groups discussed are monophyletic. The extant genera
included and the diagnostic characters derived mostly from Parenti (1981) are given below.
Aphaniidae: Aphanius
Based on osteological data presented by Parenti (1981:521), Aphaniidae is distinguished from Orestiidae by
having a vomer (vs. absent), a cartilaginous mesethmoid (vs. ossified) and an ossified interhyal (not ossified). The
generic diagnosis is valid also as a family level diagnosis.
Cubanichthyidae: Cubanichthys
Parenti (1981:519) diagnosed the family Cubanichthyidae (as Cubanichthyinae) by its possession of several
rows of teeth in the upper and lower jaw, an enlarged supraoccipital crest, an elongate dorsal process of the
autopalatine, a supraorbital sensory-pore pattern consisting of a large third pore, and lacking an ossified ventral
limb of the posttemporal. The subfamily-level diagnosis is valid also as a family level diagnosis.
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Cyprinodontidae: Cualac, Cyprinodon, Floridichthys, Jordanella and Megupsilon
Parenti (1981:526) diagnosed the family Cyprinodontidae (as Cyprinidontini) by its having a derived form of
the attachment of the first vertebra to the skull and the pharyngobranchial teeth being organised in discrete rows.
The tribe-level diagnosis is valid as a family-level diagnosis.
Orestiidae: Orestias
Parenti (1981:526) diagnosed the genus Orestias by its lacking the vomer and the first postcleithrum. The
generic diagnosis is valid also as a family-level diagnosis.
Valenciidae: Valencia
Parenti (1981:500) diagnosed Valenciidae as a distinct family against all other groups in Cyprionodontiform
fishes by possession of a very long, attenuate dorsal process of the maxillary.
Comparative material
Aphanius anatoliae: FSJF 2483, 15, 19–36 mm SL; IUSHM 2017-1280, 48, 10–30 mm SL; Turkey: Aksaray
prov.: spring in Köşk Park within Sultanhanı, 38°14'29"N 33°32'45"E.—FSJF 2485, 38, 25–45 mm SL; IUSHM
2017-1282, 30, 21–40 mm SL; Turkey: Konya prov.: Meyil Lake , about 3 km southwest of Esentepe, 37°59'10"N
33°21'5"E.—FSJF 2527, 9, 21–30 mm SL; IUSHM 2017-1283, 25, 16–31 mm SL; Turkey: Konya prov.: stream
north of Sarıyayla, draining to former Lake Samsam, 39°7'8"N 32°45'33"E.—FSJF 2610, 4, 21–38 mm SL;
IUSHM 2017-1279, 21, 12–46 mm SL; Turkey: Konya prov.: stream at Gölyazı at road from Eskil to Cihanbeyli,
38°33'9"N 33°12'3"E.—ZMH 3489, 10, 27–37 mm SL; Turkey: Niğde prov.: Niğde.—ZMH 3495, 28, 28–39 mm
SL; Turkey: Konya prov.: stream 20 km north of Çumra.—ZMH 3490, 10, 34–45 mm SL, Turkey: Konya prov.:
Aksaray.
Aphanius danfordii: FSJF 2601, 16, 23–42 mm SL; IUSHM 2017-1276, 30, 16–45 mm SL; Turkey: Kayseri
prov.: Spring Soysallı west of Soysallı, west of Develi, 38°23'25"N 35°21'56"E.—ZMH 3473, 1, 29 mm SL; ZMH
3474, 1, 32 mm SL; ZMH 3475, 8, 19–27 mm SL; Turkey: Kayseri prov.: Çayırözü, about 15 km northwest of
Develi, 38°25’2’’N 35°17’22’’E.—ZMH 3476, 21, 19–36 mm SL; Turkey: Kayseri prov.: Soysallı, 38°20’28’’N
35°21’47’’E.—ZMH 3477, 33, 19–43 mm SL; ZMH 3478; 1, 40.1 mm SL; ZMH 3479, 1, 46 mm SL; Turkey:
Kayseri prov.: Develi, İlipınar.
Aphanius fontinalis: FSJF 3456, 20, 22–38 mm SL; Turkey: Burdur prov.: spring in Lake Yarışlı basin,
37°34’50’’N 29°56’50"E.
Aphanius maeandricus: FSJF 1876, 37, 16–40 mm SL; FSJF 3027, 9, 27–40 mm SL; IUSHM 2017-1278, 9,
28–41 mm SL; Turkey: Denizli prov.: Işıklı spring at Işıklı, 38°19'17"N 29°51'4"E.—FSJF 2470, 5, 26–38 mm SL;
IUSHM 2017-1277, 3, 25–27 mm SL; Turkey: Afyonkarahisar prov.: Spring Düden, 5 km east of Dinar, 38°3'7"N
30°10' 32"E.
Aphanius marassantensis: FSJF 3455, 29, 28–43 mm SL; Turkey: Ankara prov.: Hirfanlı Reservoir,
39°09'09''N 33°38'12''E.—ZMH 3480, 86, 19–55 mm SL; ZMH 3484, 1, 33 mm SL; ZMH 3485, 1, 40 mm SL;
Turkey: Kayseri prov.: Kayseri, Karpuzatan, 38°46'25"N 35°27'11"E.—ZMH 3481, 33, 18–28 mm SL; ZMH
26068, 18, 23–35 mm SL; Turkey: Kırşehir prov.: Kırşehir: springs in Kırşehir.—ZMH 3482, 20, 17–35 mm SL;
Turkey: Samsun prov.: Lake Balık about 12 km east of Bafra, 41°34'45"N 36°5'00"E.
Aphanius meridionalis: FSJF 2460, 7, 21–36 mm SL; IUSHM 2017-1275, 22, 20–39 mm SL; Turkey:
Antalya prov.: small field canal south of Kırkpınar, north of Kızılcadağ, 37°8'21"N 29°55'5"E.—FSJF 3669, 3, 27–
29 mm SL; Turkey: Turkey: Burdur prov.: reservoir south of Yeşilova, 37.490766 29.743290.—ZMH 3502, 17,
23–39 mm SL; Turkey: Lake Gölhisar.
Aphanius saldae: ZMH 3507, 40, 32–38 mm SL; Turkey: Burdur prov.: Lake Salda.
Aphanius splendens: ZMH 3505, lectotype, 31 mm SL; ZMH 3506, 1, 38 mm SL; Turkey: Isparta prov.: Lake
Gölcük.
Aphanius sureyanus: ZMH 7850, 40, 16–34 mm SL; Turkey: Burdur prov.: shore of Lake Burdur at road from
Burdur to Yeşilova.
Aphanius transgrediens: ZMH 3509, 52, 23–33 mm SL; Turkey: Afyonkarahisar prov.: spring system of
Acıgöl, 37°49'N 29°55'E.
Aphanius villwocki: FSJF 2626, 4, 31–55 mm SL; IUSHM 2017-1285, 6, 24-48 mm SL; Turkey:
FREYHOF ET AL.
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Afyonkarahisar prov.: spring about 11 km east of Emirdağ, 39°02'53''N 31°19'38''E.—FSJF 3091, 12, 20–35 mm
SL; Turkey: Konya prov.: Ilgın Lake, 38.32075N 31.89029E.
Acknowledgments
We are pleased to thank Rainer Sonnenberg (Plön) for his helpful comments on killifish families and Ralf Thiel
(ZMH) for giving JF access to his collection and for help in some collection details. Special thanks to Maurice
Kottelat (Delémont) for his help with the stem of Orestias. We thank the two reviewers for useful comments. They
took the burden to read the manuscript.
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