Revision of Phoxinus in France with the description of two new species (Teleostei, Leuciscidae)
Author
Denys *, Gaël P. J.
Author
Dettai, Agnès
Author
Persat, Henri
Author
Daszkiewicz, Piotr
Author
Hautecoeur, Mélyne
Author
Keith, Philippe
text
Cybium
2020
2020-09-30
44
3
205
238
journal article
10.26028/cybium/2020-443-003
7806280
Phoxinus fayollarum
,
new species
(
Fig. 9
)
Holotype
MNHN 2019-0263
,
m
, 58.1 mm SL;
France
:
Boron Stream
,
Pionsat
,
46°06.4’N
2°41.1’E
;
522 m
.
Paratypes
MNHN 2014-2729
,
5
m
,
3
f
, 57.5-68.1 mm SL;
France
:
Boron Stream
,
Pionsat
46°06.4’N
2°41.1’E
; alt.
522 m
.
MNHN 2014-2732
,
2
m
,
2
f
, 50.2-58.9 mm SL;
France
:
stream Nièvre
,
Dompierre-sur-Nièvre
,
47°14.1’N
3°15.0’E
; alt.
231 m
.
Additional material
MNHN 2010-0992
,
2
m
,
1
f
, 50.2-56.1 mm SL;
France
:
Ance du Nord Stream
,
Sauvessanges
,
45°23.1’N
3°53.6’E
; alt.
800 m
.
MNHN 2010-1055
,
2
m
,
1
f
, 48.9-54.8 mm SL;
France
:
Aurence Stream
,
Aixe-sur-Vienne
,
45°47.9’N
1°09.1’E
; alt.
205 m
.
MNHN 2011-0819
,
2
m
,
4
f
, 41.0- 60.5 mm SL;
France
:
Allier Stream
,
Saint-Haon
,
44°50.4’N
3°44.2’E
; alt.
734 m
.
MNHN 2011-0910
,
5
f
, 41.0-47.0 mm SL;
France
:
Varenne Stream
,
Soucé
,
48°28.1’N
0°39.5’W
; alt.
114 m
.
MNHN 2014-2733
,
4
m
,
11
f
, 48.1-67.0 mm SL;
France
:
Esves Stream
,
Ciran
,
47°02.6’N
0°53.6’E
; alt.
88 m
.
MNHN 2014-2740
,
1
m
,
4
f
, 59.1-73.6 mm SL;
France
:
Oua- tier Stream
,
Rians
,
47°10.2’N
2°35.6’E
; alt.
158 m
.
MNHN 2014-2745
,
2
m
,
8
f
, 43.4-70.1 mm SL;
France
:
Tusson Stream
,
Vancé
,
47°49.4’N
0°38.6’E
; alt.
83 m
.
MNHN 2014- 2767
,
4
m
,
4
f
, 41.2-51.9 mm SL;
France
:
Céphons Stream
,
Moulins-sur-Céphons
,
47°02.2’N
1°33.1’E
; alt.
121 m
.
Figure 9.
–
Phoxinusfayollarum
Holotype
m
(
A
), 58.13 mm SL from the Boron Stream, Pionsat 46°06.4’N, 2°41.1’E, MNHN 2019-0263; paratypes
m
(
B
) and
f
(
C
), same location as holotype, MNHN 2014-2729; nuptial colouration pattern for male (
D
) and female (
E
).
Material used for molecular analysis
MNHN 2019-0263
,
holotype
, tag 12474;
France
:
Boron Stream
,
Pionsat
,
46°06.4’N
2°41.1’E
; alt.
522 m
. (COI: Gen- Bank accession number:
MT975733
, 12S: GenBank acces- sion number:
MT975362
).
MNHN 2014-2729
,
paratype
,
1
, tag 12475;
France
:
Boron Stream
,
Pionsat
,
46°06.4’N
2°41.1’E
; alt.
522 m
. (COI: GenBankaccessionnumber:
MT975734
, 12S: GenBankaccessionnumber:
MT975363
)
.
MNHN 2010-0992
,
1
, tag 4209;
France
:
Ance du Nord Stream
,
Sauvessanges
,
45°23.1’N
3°53.6’E
; alt.
800 m
. (COI: GenBankaccessionnumber:
MT975768
, 12S: Gen- Bank accession number:
MT975381
)
.
MNHN 2010-1055
,
1
, tag 4237;
France
:
Aurence Stream
,
Aixe-sur-Vienne
,
45°47.9’N
1°09.1’E
; alt.
205 m
. (COI: GenBank acces- sion number:
MT975769
, 12S: GenBank accession num- ber:
MT975382
)
.
MNHN 2011-0819
,
1
, tag 10741;
France
:
Allier Stream
,
Saint-Haon
,
44°50.4’N
3°44.2’E
; alt.
734 m
. (COI: GenBankaccessionnumber:
MT975702
, 12S: Gen- Bank accession number:
MT975347
)
.
MNHN 2011-0910
,
1
, tag 10752;
France
:
Varenne Stream
,
Soucé
,
48°28.1’N
0°39.5’W
; alt.
114 m
. (COI: GenBankaccessionnumber:
MT975703
, 12S: GenBankaccessionnumber:
MT975348
)
.
MNHN 2014-2740
,
2
, tag 12347 and 12348;
France
:
Oua- tier Stream
,
Rians
,
47°10.2’N
2°35.6’E
; alt.
158 m
. (COI: GenBank accession numbers:
MT975724
,
MT975725
, 12S: GenBankaccessionnumber:
MT975358
)
.
MNHN 2010-0474
,
1
, tag 4029;
France
:
Dejoune Stream
,
Felletin
,
45°50.5’N
2°12.1’E
; alt.
578 m
. (COI: GenBank acces- sion number:
MT975752
)
.
MNHN 2010-1008
,
1
, tag 4189;
France
:
Semme Stream
,
Droux
,
46°09.1’N
1°08.7’E
; alt.
198 m
. (COI: GenBankaccessionnumber:
MT975766
)
.
MNHN 2011-0920
,
1
, tag 10755;
France
:
Sarthe Stream
,
Moulin-le-Carbonel
,
48°23.1’N
0°02.1’W
; alt.
133 m
. (COI: GenBankaccessionnumber:
MT975704
, COI: GenBank accession number:
MT975349
)
.
MNHN 2014-0015
,
2
, tag 18978 and 18979;
France
:
Sèvre niortaise River
,
Sainte- Eanne
,
46°22.9’N
0°09.4’W
; alt.
64 m
. (COI: GenBank accession numbers:
MT975735
,
MT975736
, 12S: GenBank accession numbers:
MT975365
,
MT975366
)
.
MNHN 2011- 0937
,
1
, tag 10777;
France
: Boralde
de Flaujac Stream
,
Espalion
,
44°32’N
2°47.5’E
, alt.
386 m
. (COI: GenBank accession number:
MT975707
)
.
Diagnosis
Phoxinus fayollarum
is distinguished from the other French
Phoxinus
species
by its unique nuptial colour pattern. The male has black bars in Z1 to Z2 and green in Z3 to Z4 (vs. black in Z2 to Z3). Z4 is bright yellow (vs. greyish of green) and the belly is red (vs. white, pinkish to light orange or greyish to black) (
Fig. 9B, D
). The female has a green stripe along Z3 (vs. black if present), the Z4 is yellowpinkish (vs. greyish or green), the belly red (vs. white), and the female has red spots at the pectoral, pelvic and anal fins bases (vs. pinkish) (
Fig 9C, E
).
Table V.
–
Morphometric data of
Phoxinus fayollarum
(holotype: MNHN 2019-0263; paratypes: MNHN 2014-2729, MNHN 2014- 2732; and additional materials (MNHN 2010-0992, MNHN 2010- 1055, MNHN 2011-0819, MNHN 2011-0910, MNHN 2014-2733, MNHN 2014-02740, MNHN 2014-2745, MNHN 2014-2767; n = 69). The calculations include the holotype.
| Holotype |
Holotype & other individuals Mean Min Max SD |
| SL (mm) |
58.1 |
53.5 |
41.0 |
73.6 |
8.0 |
|
In percent of standard length
|
| Head length |
27.6 |
26.0 |
23.6 |
28.2 |
1.0 |
| Predorsal length |
51.9 |
55.5 |
51.9 |
59.1 |
1.6 |
| Post dorsal length |
36.2 |
35.9 |
28.0 |
40.0 |
2.0 |
| Prepectoral length |
24.6 |
26.0 |
22.8 |
30.3 |
1.3 |
| Prepelvic length |
45.1 |
48.3 |
42.7 |
53.4 |
1.9 |
| Preanal length |
63.5 |
65.3 |
61.2 |
69.5 |
1.9 |
| Body depth |
21.5 |
21.3 |
15.2 |
28.3 |
2.3 |
| Distance dorsal anal |
23.7 |
22.2 |
16.8 |
27.0 |
2.0 |
| Distance pectoral pelvic |
17.1 |
20.8 |
16.2 |
25.0 |
2.2 |
| Pectoral fin length |
18.5 |
18.1 |
14.2 |
21.4 |
1.6 |
| Pelvic fin length |
15.7 |
14.8 |
12.1 |
18.3 |
1.4 |
| Dorsal fin length |
21.8 |
20.3 |
16.4 |
22.9 |
1.3 |
| Anal fin length |
19.3 |
19.9 |
16.6 |
23.5 |
1.4 |
| Caudal peduncle length |
25.9 |
25.5 |
20.9 |
28.9 |
1.6 |
| Caudal peduncle depth |
9.4 |
9.1 |
6.9 |
11.2 |
0.8 |
|
In percent of head length
|
| Head depth |
50.0 |
52.0 |
45.6 |
57.7 |
2.9 |
| Eye diameter |
22.3 |
26.4 |
21.5 |
32.3 |
2.6 |
| Snout length |
30.9 |
29.0 |
18.8 |
36.8 |
2.9 |
| Inter-orbital width |
32.1 |
32.5 |
26.1 |
38.3 |
2.7 |
Phoxinus fayollarum
is further distinguished from
P. bigerri
and
P. csikii
by a straight to slightly concave anal fin margin (vs. straight to convex), from
P. phoxinus
and
P. septimaniae
by a stout and blunt snout (vs. slightly point- ed, and two scaled patches at the pectoral fin bases separat- ed by a nonscaled area (vs. scaled breath between the pectoral fin insertions in some populations of
P. septimaniae
), and from
P. dragarum
by an incomplete lateral line usually reaching to the caudal peduncle (vs. usually reaching to the anal fin base or just slightly behind the last anal fin ray), and 56-84 pored scales (vs. 50-74).
The nuptial male
P. fayollarum
is further distinguished by its clearly distinct green bars crossing Z3 and Z4 (vs. indistinct in
P. phoxinus
), the red belly not extending to the caudal peduncle (vs. extending in
P. csikii
), having red pectoral, pelvic and anal fins bases (
vs
. pinkish in
P. bigerri
and
P. phoxinus
), orange pectoral, pelvic and anal fins (vs. yel- low in
P. csikii
and
P. dragarum
), no red disc on the inner side of the pectoral fin (presence in
P. septimaniae
), and the absence of a yellow dot between the mouth and the eye (vs. the presence in
P. septimaniae
). The nuptial female is distinguished by a white Z5 (vs. black in
P. phoxinus
,
P. septimaniae
and
P. dragarum
).
Description
Morphometric data is shown in table V. Largest examined specimen 73.6 mm SL. Body moderately elongated, dorsal and ventral profile about symmetric or ventral profile slightly more convex. Dorsal head profile markedly more convex than ventral one. Caudal peduncle length 2.2-3.5 times in its depth. Snout stout and blunt. Mouth slightly subterminal, upper lip and snout projecting beyond tip of lower jaw. Tip of upper lip at level or above level of lowest point of eye. In lateral view, eye not or almost flushed with dorsal profile. No supraorbital ridge.
Dorsal fin with 3 simple and 6½-8½ branched rays. Dis- tal dorsal-fin margin markedly convex, second branched ray longest, dorsal-fin origin behind pelvicfin insertion. Pectoral fin with 1 simple and 11-16 branched rays (usually 14-15), reaching to pelvicfin origin in male, not reaching in female. Pelvicfin with 1 simple and 6-9 branched rays (mainly 7-8), reaching slightly beyond anal fin origin in male, not reaching in female. Anal fin with 3 simple and 6½-8½ branched rays. Anal fin origin slightly in front of vertical through dorsal-fin origin. Anal fin margin straight to slightly concave, second branched ray longest.
Lateral line incomplete reaching caudal peduncle. 72-91 lateral scales, including 56-84 pored scales. Two scaled patches at pectoral fin origin separated by an unscaled area. No scale between pectoral fin bases. Nuptial tubercles present in both sexes, smaller in female.
Nuptial colouration
Male
. – Snout and top of head dark brown. Ablack bar on operculum reaching from top of head to branchiostegal rays. Cheeks white. Lips dark pink or red. Large yellow spot on operculum almost covering whole opercular and a white spot above at upper corner. Z1 brown to black. Z2 pale brown. Flank bars crossing Z1 to Z4, flank bars black from Z1 to Z2 and green from Z3 to Z4. Z4 bright yellow. Ablack line along Z5 between pectoral fin base and caudal peduncle, from pelvic fin base to caudal peduncle in some individuals. Pectoral, pelvic and anal fins yellow-orange. Pectoral, pel- vic and anal fin bases red. (
Fig. 9B, D
). Female: Nuptial col- ouration pattern similar to male but paler with black bars in Z1 to Z2, a green stripe along Z3, Z4 yellow-pinkish, and Z5 white. Lips pinkish and belly red. Presence of “male” characters in the female: white and yellow spots on operculum and red spots at pectoral, pelvic and anal fin bases (
Fig 9C, E
).
Colouration in preservation
Dark green, brown and black bars and dots remain after preservation in ethanol. Green colouration turns to dark brown, whereas yellow and red colourations are faded (
Fig. 9A
).
Molecular characterisation
On the COI marker,
P. fayollarum
has six diagnostic sites including two synapomorphies (Tab. II). For the 12S rDNA marker, 6 diagnostic sites characterise this species: C (vs. A or G) in position 122, C (
vs
. T) in position 357, T (vs. C) in position 454, C (vs. T) in position 649, G (vs. A) in position 675 and A (vs. G) in position 874.
Distribution
This species is endemic to the
Loire
drainage as well as the Sèvre Niortaise catchment (
Fig. 1
). It has been introduced in Garonne and Rhône drainages adjacent to the
Loire
catchment, for example in the stream Boralde de Flaujac and in the Chassezac River, Grevieres (
Geiger
et al.
(2014)
. This lineage was named
Phoxinus sp.
16 by
Palandačić
et al.
(2017)
.
Ecology
This species is common in the whole
Loire
drainage, and is especially abundant in the greyling zone (
Lasne
et al.,
2007
). It inhabits mostly clear and shallow streams with a slow discharge and a sandy or gravely ground. The spawing period is from April to May (
Mauduyt, 1848
) and in June- July in higher altitudes (Denys, pers. obs., 2014).
Etymology
The species name
fayollarum
is the genitive of a latinized name “fayolles”, fairies from of the
Auvergne region
folklore, in the headwater of
Loire
drainage. Fayolles are fairies who create spring waters. And we owe to one of them, called “Grande Fayolle”, the thermal waters of Vichy (
Ely and Tsaag Valren, 2013
).
Vernacular name
Loire
minnow (English), Vairon ligérien (French).
Nomenclatural note
De la Pylaie (1835: 533)
described
Leuciscus obtusus
as a little leuciscid or small minnow without spots (“Able ou petit Verdon sans taches” in French) in the Vendée department (so possibly in the Loire drainage). This name is a
nomen nudum
and potentially synonym of
P. phoxinus
(
Kottelat, 1997
)
. However, a minnow is characterised by the pres- ence of spots on the snout, the flanks and the tail. According to the original description (the absence of spots), this taxon could correspond to another leuciscid species but not to a
Phoxinus
, even if the ancient French word “verdon” means minnow. Moreover, no
type
specimen is known. Thus, we consider this taxon as an
incertae sedis
in
Leuciscidae
.