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Verlag Dr. Friedrich Pfeil
ISSN 0936-9902
Ichthyological Exploration
of Freshwaters
Volume 22
Number 4
An international journal for field-orientated ichthyology
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Volume 22
•
Number 4
•
December 2011
pages 289-384, 48 figs., 11 tabs.
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345
Ichthyol. Explor. Freshwaters, Vol. 22, No. 4
Ichthyol. Explor. Freshwaters, Vol. 22, No. 4, pp. 345-353, 5 figs., 3 tabs., December 2011
© 2011 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
Orestias gloriae, a new species of cyprinodontid fish
from saltpan spring of the southern high Andes
(Teleostei: Cyprinodontidae)
Irma Vila*, Sergio Scott*, Marco A. Mendez*, Felipe Valenzuela**,
Patricia Iturra** and Elie Poulin*
,
***
Orestias gloriae, new species, is described from the isolated springs that drain into the Carcote saltpan in the
southern high Andes, in the Chilean Altiplano, at an altitude of 3706 m a. s. l. (21°16'58.6" S 68°19'28.4" W). This is
the only fish species found in these saltpan springs characterized by scarce vegetation, aquatic insects, crustaceans,
and mollusks. The new species is separated from other species of Orestias by several unique characters such as a
truncated cephalic lyre-like pattern of neuromasts represented only by the rostral series followed by few and
large neuromasts irregularly placed; discontinuous series of infraorbital and preopercle-mandibular neuromasts;
thick and irregularly-shaped ornamented cycloid scales covering the skull roof; scales covering the posterior part
of the skull roof ankylosed into a plate. Additional characters are the presence of a protractile mouth and a char-
acteristic karyotype. Although differences in sizes is a common sexually dimorphic feature found in Orestias,
differences in the dentition on the fifth ceratobranchial distinsguish males and females of O. gloriae.
* Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa,
Santiago, Chile. E-mail: limnolog@uchile.cl
** Programa de Genética Humana. ICBM, Facultad de Medicina, Universidad de Chile, Independencia 1027,
Santiago, Chile. E-mail: piturra@med.uchile.cl
*** Instituto de Ecología y Biodiversidad, Facultad de Ciencias, Universidad de Chile. Las Palmeras 3425. Ñuñoa.
Santiago, Chile. E-mail: epoulin@uchile.cl
Introduction
The southern high Andean systems of the Alti-
plano range from 17° to 22° S and from 3000 to
about 4500 m in altitude. This region is hypoth-
esized to represent the remnants of remarkable
Plio-Pleistocene lake and river systems, which,
with the subsequent lack of rainfall, have evolved
into ponds and saltpans due to evaporation
mainly as the result of high temperatures in the
region and additionally, by human water use.
Currently, the Chilean Altiplano is represented
by many important water bodies such as Chun-
gará Lake in the north and Ascotán Salar in the
south (about 600 km between both; Chong, 1988;
Keller & Soto, 1998; Risacher et al., 2003; Placzek
et al., 2006).
The endemic ichthyofauna that inhabits the
aquatic systems of the Altiplano belongs to two
genera: the killifish Orestias and the catfish Tricho-
mycterus. The genus Orestias is known from 45
species, 23 of which live in Lake Titicaca where
346
it has had an explosive species radiation (Lau-
zanne, 1981; Parenti, 1984a
-
b; Loubens, 1989). In
addition to the Bolivian and Peruvian species, the
Orestias of the southern Altiplano in Chile were
first described by Arratia (1982) and Parenti
(1984a) in a revision of the genus. Within Orestias,
Parenti (1984a) defined four species complexes:
cuvieri, mulleri, gilsoni and agassii. The last complex
is characterized by (1) ontogenetic changes in
color pattern; (2) squamation of head and median
dorsal ridge with enlarged and smooth or granu-
lated scales (these scales are larger in comparison
to lateral flank scales); (3) presence of a lateral
shield formed by large and smooth or granulated
lateral scales positioned between opercle and a
point at about the posterior extent of the pectoral
fin; and (4) caudal peduncle relatively deep in
adults, reaching over 22 % of standard length
(SL).
At present, the agassii-complex includes 26
species, six of which have been described from
the southern Altiplano (Arratia, 1982; Parenti,
1984a
-
b; Vila & Pinto, 1986; Vila, 2006). Lüssen
et al. (2003) divided the agassii-complex species
group into two clades: one that lives exclusively
in Titicaca Lake basin (17°-20° S) and a second
clade that lives in the southern Altiplano region
(21°-22° S). Among the six species found in the
southern Altiplano, O. cf. agassii is the most
broadly distributed within the genus.
Recent fieldwork in the southern Altiplano
has revealed the presence of Orestias in the Carcote
saltpan (Fig. 1), a region that has not been previ-
ously investigated. The Carcote saltpan is a
remnant of a Pleistocene lake (Ochsenius, 1974)
at an altitude of 3706 m a.s.l. (21°16'58.6" S 68°19'
28.4" W). Due to evaporation, this aquatic system
evolved into a saltpan, but keeping small ponds
fed by springs in which live small populations of
Orestias of an unnamed species. Here, we describe
this new species and provide information about
its habitat.
Material and methods
Specimens were captured with hand nets and a
number of them were kept alive to obtain samples
used in chromosome studies. Specimens were
euthanized using 100 mg · l
-1
tricaine methanosul-
fonate. Morphometric and meristic data for all
the specimens were taken following the standard
methodology outlined in previous descriptions
of Orestias (e. g., Lauzanne, 1981; Parenti, 1984a;
Vila, 2006). All measurements were taken on
straight-line distances recorded with a precision
caliper reading to the nearest 0.1 mm. Cleared-
and-stained specimens were prepared following
the protocol of Taylor & Van Dyke (1985). They
were used in osteological studies of jaws, bran-
chial skeleton, and vertebrae and fin-rays counts.
The branchial apparatus was dissected and the
gill rakers of the left gill arches I to IV were
counted in 11 specimens. The terminology of the
head neuromast lines follows Coombs et al. (1988)
and Arratia & Huaquín (1995), with the exceptions
noted below.
The specimens were sexed by direct examina-
tion of the gonads under a stereomicroscope. The
number and morphology of chromosomes were
studied in four females and three males following
Vila et al. (2010). Institutional abbreviations follow
Leviton et al. (1985) except for: LLFS-UCH,
Laboratory of Limnology, Faculty of Sciences,
Universidad de Chile, Santiago.
Vila et al.: Orestias gloriae
Fig. 1. Location of the Altiplano in South America. Left
image: Southern Altiplano systems with Carcote Salar
(21°16'58.6" S 68°19'28.4" W), showing no connections
with other systems.
ı ı ı ı
70°W 69°W 68°W 67°W
18°S
–
19°S
–
20°S
–
21°S
–
22°S
–
N
0 100 km
Peru
Bolivia
C h i l e
P
a
c
i
f
i
c
O
c
e
a
n
Umaqui
Parinacota
Piacota
Isluga
Huasco
Chuviri
Cotacotani
Lauca
Collacahua
Carcote
Chuviri
Cotacotani
Lauca
Collacahua
Carcote
Ascotán
Chungará
347
Ichthyol. Explor. Freshwaters, Vol. 22, No. 4
Orestias gloriae, new species
(Fig. 2)
Holotype. MNHNC 7424, female, 70.4 mm SL;
Chile: Loa Province: springs of Carcote saltpan,
21°16'58.6" S 68°19'28.4" W, altitude 3706 m a.s.l.;
I. Vila & S. Scott, April 2006.
Paratypes. MNHNC 7425, 1 male, 21.9 mm SL;
MNHNC 7426, 1 female, 59.0 mm SL; LLFS-UCH
2006, 10 females 38.2-60.1 mm SL; LLFS-UCH,
7 males, 20.3-38.2 mm SL; collected with holo-
type.
Diagnosis. A relatively slender-bodied species,
Orestias gloriae is clearly distinguished from all
other species of Orestias by the following unique
characters: (1) The lyre-shaped pattern of dorsal
neuromasts, which is characteristics of Orestias is
reduced to the rostral and anterior supraorbital
lines formed by small neuromasts followed cau-
dally by few and large isolated neuromasts, which
are irregularly placed. (2) The infraorbital line of
neuromasts is represented by a discontinuous
series of few small neuromasts. (3) The supraor-
bital line of neuromasts is not connected with
both the infraorbital and preopercle-mandibular
lines. (4) The skull roof is covered by thick, ir-
regularly-shaped cycloid scales some of which
bear marked ridges, whereas other lack ornamen-
tation. (5) The large, irregularly-shaped scales
covering the posterior part of the skull roof are
ankylosed to each other into a thin, rigid plate.
In addition, O. gloriae presents a protractile
upper jaw reaching one-third of the head length
when extended. This value is higher than those
found in other species of Orestias with protact-
ibility and inhabiting the southern Chilean Alti-
plano. The new species has 29-31 vertebrae, a
value that represents the lowest range among
species of Orestias.
Description. Morphometric data for the holo-
type, and paratypes are summarized in Table 1.
Size not reaching beyond 80 mm SL, with a head
moderately large, length 26-31 % SL. Orbits not
reaching dorsal profile of head. Gently convex
dorsal profile between head and dorsal fin.
Slightly convex ventral profile between origin of
anal and caudal fins. Lateral profile notably ex-
panded just posterior to pectoral fin, decreasing
slightly posteriorly up to anal-fin origin. Caudal
peduncle relatively short and slender (see Ta-
ble 1). 29-31 vertebrae.
Dorsal series of neuromasts represented by
rostral and anterior section of supraorbital lines
(Fig. 3). Rostral line formed by continuous series
of approximately 8 to 10 relatively small and
rounded neuromasts on each side. No posterior
section of lyre-like pattern of dorsal neuromasts.
Infraorbital and preopercle-mandibular lines
formed by few, small and irregularly placed
neuromasts. Both infraorbital and preopercle-
mandibular lines of neuromasts not connected to
supraorbital line. Pores of both supraorbital and
infraorbital sensory canals difficult to observed.
Upper jaw protractile. Premaxilla as well as
dentary bearing mainly conical teeth, slightly
recurved medially, and with conspicuous acrodin
cup. Some teeth with slightly blunt tips. Premax-
illa with three main rows of teeth, and dentary
with two.
Preopercle and opercle laterally prominent.
Fig. 2. Orestias gloriae; Chile: Loa Province: springs of
Carcote saltpan. a, MNHNC 7424, holotype, 70.4 mm
SL; b, MNHNC 7425, paratype, 59.0 mm SL.
Table 1. Morphometric characteristics of Orestias glo-
riae.
holotype paratypes
Standard length (mm) 70.4
21.9-59.0
In percent of standard length
Head length 30.6
26.5-26.9
Head depth 21.9
20.1-21.9
Predorsal length 59.5
56.1-59.6
Preanal length 62.5
58.4-63.0
Caudal-peduncle length 28.9
22.2-23.7
Caudal-peduncle depth 15.0
13.3-18.3
Body depth 23.3
22.0-23.4
Eye diameter 9.2
7.3-9.7
Interorbital distance 9.3
4.4-9.0
Mouth width 7.0
7.0-7.1
a
b
348
Preopercle bordered anteriorly by thick, large,
and slightly rounded or oval scales.
Branchial apparatus (Fig. 4) characterized by
marked asymmetry between left and right sides,
otherwise branchial bones morphologically sim-
ilar to other Orestias, with following exceptions.
Fifth ceratobranchials forming a large isosceles
triangle and unfused at midline. Females (Fig. 4a)
with dorsal surface of ceratobranchial 5 densely
covered with two kinds of teeth; most teeth long
and conic, ending in a small conic acrodin cup
usually darker than base of tooth; most posterior
series of teeth formed by large, molariform teeth,
each bearing a small conic acrodin tip, character-
ized by its dark color. In males fifth ceratobranchi-
als bearing scarce teeth, discontinuously distrib-
uted on oral surface of bone. Most teeth molari-
form type (Fig. 4b).
Simple, partially ossified gill rakers present
on both sides of each arch (Fig. 4b). Anterior and
posterior margins of ceratobranchial 1 bearing
gill rakers. Only anterior row of gill rakers present
on anterior margins of hypobranchial and cera-
tobranchials 2 and 3. Ceratobranchial 5 bearing
few gill rakers on anterior margin. Number of gill
rakers on gill arches 1 to 4 ranging from 104 to
125.
Pectoral fin rounded, with 16-19 rays, both
first and last rays shorter than middle ones and
uniformly pigmented. Dorsal fin rather small,
with 14 or 15 rays, first principal rays (terminol-
ogy after Arratia, 2008) longer than remaining
rays giving fin a pointed shape. Anal fin pointed,
with 13-16 rays, anterior principal rays longer
than following ones. Caudal fin truncate, with
23-27 total rays.
Squamation. Head and trunk covered with cy-
cloid scales. Skull roof covered with scales (Fig. 3)
of different shapes and sizes, showing strong
bilateral asymmetry. Most anterior scales being
the smallest, generally slightly round- or oval-
shaped and with a few marked concentric striae.
Most posterior scales thick, large, irregularly
shaped and ankylosed between them forming a
thin, rigid plate covering skull roof. One elongate,
large scale positioned above each orbital margin
of skull. Each scale with concentric striae at its
mid section. Nine to 12 large and slightly round-
ed scales arranged in three rows positioned below
orbit, and preceded by a small dorso-anterior
cycloid scale with marked and protruding cir-
culi. Nine to 12 large, thick and rounded scales
positioned below orbit, on the cheek region, a
Fig. 3. Orestias gloriae, LLFS-UCH 2006, female, 50.9 mm SL; head in dorsal view. Thick lines and arrows between
scales represent area with ankylosed scales. Abbreviations: a.no, anterior nostril; i.ne, isolated neuromasts;
p.no, posterior nostril; ro.l, rostral line of neuromasts; sc, scales; sob.l, supraorbital line of neuromasts.
Vila et al.: Orestias gloriae
5 mm
i.ne
a.no
ro.l
p.no
sob.l
sc
sc
349
Ichthyol. Explor. Freshwaters, Vol. 22, No. 4
row of 6-8 bordering the preopercle. Commonly,
6-8 scales found at anterior preopercular mar-
gin.
Juveniles as well as adults fully scaled and
covered with cycloid scales small and of uniform
size over entire body. Ventral body region naked.
Median dorsal ridge scales barely visible and few.
Median dorsal ridge scales slightly larger than
those of anterior section of flank.
Scales begining at posttemporal region ar-
ranged in 3-5 horizontal rows, increasing from 8
to 10 in middle flank area, and decreasing to 3 or
4 on caudal peduncle. Flank scales decreasing in
size from rostrad to caudad. Exposed fields of
lateral body scales with striae and sclerites. 31-37
scales forming lateral series of middle flank in
males and females (see Table 2).
Karyotypes. Male and females karyotypes with
diploid chromosome number equal 48, consisting
of one pair of metacentric (m), three pairs of
submetacentric (sm), 20 pairs of subtelocentric
(st), and 11 pairs of telocentric (t) (Fig. 5). Funda-
mental arm number (NF) equal 56 (Fig. 5; Table 3).
Pair number 3 heteromorphic in all individuals.
No morphologically differentiated sexual chro-
Fig. 4. Orestias gloriae, LLFS-UCH 2006; branchial arches in dorsal view. a, female, 50.9 mm SL; b, male, 38.2 mm
SL (posterior part of ceratobranchial 5 teeth). Abbreviations: bb1-3, basibranchials 1-3; bb4?, possible basibran-
chial 4; bh, basihyal; cb1, cb4, cb5, ceratobranchials 1, 4 and 5; hb1, hypobranchial 1.
hb1
bh bb1 cb1 bb2−3bb4? cb4 cb5
1 mm
Table 2. Meristic and karyotype characteristics of species of Orestias from the southern Altiplano. 1, present
results; 2, Vila, pers. obs.; 3, Arratia (1982); 4, Vila & Pinto (1986); 5, Parenti (1984a); 6, Vila (2006).
size range
mm SL
2n
(NF)
vertebrae dorsal-fin
rays
anal-fin
rays
pectoral
fin rays
caudal fin
rays
scales in
lateral series
O. gloriae
21.9-58.9
48 (56)
29-31 14-15 13-16 16-19 23-27 31-37
O. cf. agassii
2
36.7-53.7
48 (54)
31-35 11-16 11-16 14-20 22-26 30-37
O. parinacotensis
2, 3
29.1-56.4
48 (54)
31-32 13-16 14-16 16-20 19-25 30-37
O. laucaensis
2, 3
41.7-90.0
50 (54)
33-35 14-15 14-15 16-17 24-28 31-39
O. chungarensis
2, 4
39.5-82.1
55 (54)
31-32 11-14 11-14 16-18 24-27 33-35
O. ascotanensis
2, 5
58.6-75.0
48 (54)
31-32 12-15 12-14 17-19 21-25 29-34
O. piacotensis
2, 6
37.4-71.7
52 (54)
32-34 12-14 12-14 14-16 21-27 30-34
a
b
350
mosomes. Karyotype of O. gloriae differentiating
from other southern Orestias in chromosomal
formula and presence of four biarmed pairs of
chromosomes (for comparison see Vila et al., 2010:
table 4).
Sexual dimorphism. Females reaching greater
lengths than males, a plesiomorphic condition in
killifishes after Parenti (1984a). Longest examined
female 70.4 mm SL, male 38.2 mm SL.
In females, oral surface of ceratobranchial 5
covered almost completely by elongate conic
teeth, and by massive molariform teeth placed at
its posterior border (Fig. 4a); conical teeth scarce
but molariform teeth more abundant in males
(Fig. 4b). Two or three projections or spiculae
present at posterior margin of scales covering
mid-flank lateral series of males. Spiculae absent
in females.
Coloration in life. Males and females with same
color pattern throughout ontogeny, and not
sexually dichromatic. Sexual dichromatism ob-
served only at spawning periods when males
present a yellow coloration in anal fin.
Individuals with uniform dark gray color on
head and flank because of numerous malano-
phores covering exposed field of scales. Dark gray
color turning into pale yellow in ventral region
of head and trunk. Pectoral, dorsal and anal fins
uniformly pigmented due to small but abundant,
star-shaped melanophores.
Habitat and reproduction. Habitat made of small
ponds and creeks formed by springs of the Carcote
saltpan near to Calama city (Fig. 1) at 3706 m
altitude, and representing an endorrheic isolated
system in the southern Altiplano that includes a
few endemic forms (Arratia, 1982; Parenti, 1984a;
Costa, 1997, 1998, 2003; Vila, 2006; Vila et al., 2007).
Water of small ponds with high salinity due to
concentrations of sodium chloride (Marquez et
Fig. 5. Orestias gloriae. a, chromosome metaphase plate; b, karyotype. Scale bar 10 µm.
Table 3. Relative length (%), Centromeric Index ± S.D.
and chromosome type of karyotype of Orestias gloriae.
M, metacentric; Sm, submetacentric; St, subtelocentric;
T, telocentric.
pair relative
length
centromeric
index
chromosome
type
1 4.53 45.65 ± 2.92 M
2 4.81 28.79 ± 2.32 Sm
3 5.90 36.03 ± 5.83 Sm
4 2.59 32.88 ± 3.60 Sm
5 5.27 19.91 ± 4.52 St
6 4.95 18.42 ± 4.21 St
7 4.68 18.60 ± 3.80 St
8 4.51 19.53 ± 4.10 St
9 4.32 19.23 ± 3.36 St
10 4.14 19.45 ± 3.82 St
11 3.91 18.83 ± 3.37 St
12 3.60 19.18 ± 3.25 St
13 3.00 20.70 ± 2.84 St
14 5.32 0 T
15 4.80 0 T
16 4.42 0 T
17 4.25 0 T
18 4.06 0 T
19 3.94 0 T
20 3.76 0 T
21 3.61 0 T
22 3.47 0 T
23 3.30 0 T
24 2.86 0 T
ab
M-SM
ST-T
1234
5678
910111213
14 15 16 17 18
19 20 21 22 23 24
Vila et al.: Orestias gloriae
351
Ichthyol. Explor. Freshwaters, Vol. 22, No. 4
al., 2009). Fauna of Carcote saltpan represented
by small populations of insects (Elmidae, Chri-
ronomidae, and Hydroptilidae), crustaceans
(Boeckellidae, Chidoridae, and Hyalellidae), and
mollusk (Biomphalaria).
Shallow waters of Carcote saltpan springs
inhabited by O. gloriae where small populations
are distributed among macrophyte mats. Obser-
vations in situ along with analyses of stomach
contents showed larvae of aquatic insects, am-
phipods and mollusks inside stomachs of O. glo-
riae.
Eggs moderately large, 1.9-2.3 mm in diam-
eter and surrounded completely by filaments.
Fishes protecting themselves from bird predation
by fixing eggs to vegetation in a manner similar
to that of other species of Orestias (Vila & Pinto,
1986; Pinto & Vila, 1987; Martínez et al., 1999; Vila,
2006).
Etymology. The specific epithet gloriae honors
the research work of Gloria Arratia on Chilean
fishes, and especially those of the Andean re-
gion.
Discussion
As described by Parenti (1981; 1984a,b), the genus
Orestias underwent an explosive sympatric spe-
ciation in Lake Titicaca. Nevertheless, most spe-
cies described for the southernmost region of the
Altiplano in Chile indicate a high degree of en-
demism with unique species in each hydro-
graphic system suggesting allopatric speciation
(Fig. 1). This may be a result of the geographic
isolation caused by the separation of the south-
western region from the rest of South America
by the lifting of the Andes during the mid Oli-
gocene (Placzek et al., 2006; Moreno et al., 2009).
During recent years salt content and temperature
have been increasing, as is revealed by the number
and extension of the saltpans in northern Chile
(Chong, 1988). The historical formation of the
saltpans is accelerated by climatic changes and
water extraction, two factors that diminish fresh-
waters and cause isolation of small populations
of Orestias in the springs that feed the salars
(Chong, 1988; Aceituno, 1997; Risacher et al., 2003;
Morales et al., 2011) and indirectly increases the
salinity of ponds associated to salars.
Lüssen et al. (2003) performed a phylogeo-
graphic analysis using haplotypes of mitochon-
drial D-loop, and reported that the species of the
O. agassii complex comprises two groups, one
corresponding to Lake Titicaca endemics and a
second one made mainly of the species inhabiting
the southern Altiplano, O. tschudii (from Huata-
jata, Bolivia and Puno, Perú) and O. agassii (from
Lake Titicaca). Recently, Scott (2010), using three
mitochondrial markers (D-Loop, ND2 and Cytb)
and meristic characters, completed a phyloge-
netic study of the Chilean species of the agassii-
complex, including the individuals from Carcote
saltpan studied herein. In this analysis, Scott
proposed the monophyly of the species of the
Chilean Altiplano and found that individuals
from Carcote belong to a lineage independent
from the other species, suggesting that they cor-
respond to a new species. The present study
supports Scott’s hypothesis that populations liv-
ing in Carcote saltpan represent a new species of
Orestias. As shown here, this new species is sup-
ported by several morphological features and its
characteristic karyotype.
According to Parenti, 1984a, one of the defin-
ing characters of Orestias is its unique squamation
and head pore pattern. The so-called “pore” pat-
tern does not correspond to pores of the cephalic
sensory canals but to series of neuromasts that
are present on the skull roof, around the orbit,
and on the preopercular and mandibular regions.
The neuromasts on the roof of the skull are or-
ganised in a lyre pattern (Fig. 3; Parenti, 1984a).
The unique morphology of the incomplete lyre
pattern of supraorbital neuromasts, reduced to
only the anterior portion of the supraorbital line
and the rostral line (Fig. 3), is a unique feature of
O. gloriae. In addition, in all species of Orestias
there is a continuous series of neuromasts around
the orbit (infraorbital line) and another in the skin
covering the mandible and preopercle (preoper-
cle-mandibular line) (Parenti, 1984a). Orestias
gloriae is unique in having both the infraorbital
and preopercle-mandibular series formed by few
and irregularly placed neuromasts.
According to Parenti (1984a: 123-124), Orestias
has a unique squamation “and the dorsal surface
of the head is incompletely covered by irregu-
larly distributed thickened scales that have been
referred to as scutes in Orestias (Tchernavin, 1944a,
1946) although the term scutes is considered inap-
propriate here”. Orestias gloriae presents a special-
ized squamation on the surface of the head (Fig. 3)
that agrees with the general features mentioned
by Parenti (1984a), but differs from other species
352
of the genus in that the posterior-most scales
covering the dorsal surface of the head are unor-
namented and are ankylosed to each other form-
ing a rigid plate.
The degree of protractibility of the premax-
illa in O. gloriae is the largest among species with
protractile jaws among southern species such as
O. cf. agassii, O. ascotanensis, and O. laucaensis.
Other species of the southern Altiplano such as
O. chungarensis do not have protractile mouths.
Comparative material. Orestias cf. agassii: MNHNC
P7394-P7395, 20, 36.7-53.7 mm SL; Chile: Tamarugal
Province: Huasco salt flat, 20°05' S 68°15' W.
O. ascotanensis: MNHNC P7401-402, 20, 58.6-
75.0 mm SL; Chile: Loa Province, Ascotán salt flat,
21°29' S 68°19' W.
O. chungarensis: MNHNC P7399-4000, 20, 39.5-
82.1 mm SL; Chile: Parinacota Province: Chungará Lake,
18°15' S 69°07' W.
O. laucaensis: MNHNC P7393, 20, 41.7-90.0 mm SL;
Chile: Parinacota Province: Lauca river, 18°05' S 69°15' W.
O. parinacotensis: MNHNC P7396-P7397, 20, 29.1-
56.4 mm SL; Chile: Parinacota Province: Parinacota
wetland, 18°10' S 69°20' W.
O. piacotensis: MNHNC P7398, 20, 37.4-71.7 mm SL;
Chile: Parinacota Province: Piacota Lake, 18°11' S
69°15' W.
Acknowledgements
This research has been supported by Grant Fondecyt
1080390, 1110243 and Proyecto Domeyko Biodiversidad,
Iniciativa Tranversal 1, Universidad de Chile. Sergio
Scott’s Doctoral Fellowship was supported by MeceSup
UCO-214. We acknowledge the comments by David
Véliz on an early version of the manuscript and the
critical comments of anonymous reviewers. Collecting
permits were issued by the National Fishing Service
(Subsecretaria de Pesca, Chile)
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Vila et al.: Orestias gloriae
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INSTRUCTIONS TO CONTRIBUTORS
Ichthyological Exploration of Freshwaters
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Asai, Toshinobu, Hiroshi Senou and Kazumi Hosoya: Oryzias sakaizumii, a new ricefish
from northern Japan (Teleostei: Adrianichthyidae) .............................................................. 289
Freyhof, Jörg, Füsun Erk’akan, Cevher Özeren and Anabel Perdices: An overview of the
western Palaearctic loach genus Oxynoemacheilus (Teleostei: Nemacheilidae) ................. 301
Costa, Wilson J. E. M.: Simpsonichthys margaritatus, a new seasonal miniature killifish from
the upper Paraná River basin, central Brazilian Cerrado (Cyprinodontiformes: Rivul-
idae) .............................................................................................................................................. 313
Kottelat, Maurice and Tan Heok Hui: Identity of Rasbora beauforti, a cyprinid fish from
Borneo (Teleostei: Cyprinidae) ................................................................................................. 319
Britz, Ralf, Maurice Kottelat and Tan Heok Hui: Fangfangia spinocleithralis, a new genus and
species of miniature cyprinid fish from the peat swamp forests of Borneo (Teleostei:
Cyprinidae) .................................................................................................................................. 327
Bragança, Pedro H. N., Wilson J. E. M. Costa and Cecile S. Gama: Poecilia waiapi, a new
poeciliid from the Jari river drainage, northern Brazil (Cyprinodontiformes: Cyprin-
odontoidei) ................................................................................................................................... 337
Vila, Irma, Sergio Scott, Marco A. Mendez, Felipe Valenzuela, Patricia Iturra and Elie Pou-
lin: Orestias gloriae, a new species of cyprinodontid fish from saltpan spring of the
southern high Andes (Teleostei: Cyprinodontidae) .............................................................. 345
Kullander, Sven O. and Tyson R. Roberts: Out of Lake Tanganyika: endemic lake fishes
inhabit rapids of the Lukuga River .......................................................................................... 355
Marshall, Bruce Gavin, Bruce Rider Forsberg, Laura Lorraine Hess and Carlos Edwar de
Carvalho Freitas: Water temperature differences in interfluvial palm swamp habitats
of Paracheirodon axelrodi and P. simulans (Osteichthyes: Characidae) in the middle Rio
Negro, Brazil ................................................................................................................................ 377
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Volume 22
•
Number 4
•
December 2011
Cover Photograph
Oxynoemacheilus paucilepis (photograph by Jörg Freyhof)
Jörg Freyhof, Füsun Erk’akan, Cevher Özeren and Anabel Perdices
(this volume pp. 301-312)
