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Arratia 1983 Trichomycteruslaucaensischungaraensis
Arratia 1983 Trichomycteruslaucaensischungaraensis
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Trichomycterus chungaraensis
n. sp. and Trichomycterus
laucaensis n. sp. (Pisces,
Siluriformes, Trichomycteridae)
from the high Andean range.
a
Gloria Arratia F.
a
Museum of National History, University of Kansas,
Lawrence, Kansas, 66045, U.S.A.
Published online: 24 Jun 2010.
To cite this article: Gloria Arratia F. (1983): Trichomycterus chungaraensis n. sp. and
Trichomycterus laucaensis n. sp. (Pisces, Siluriformes, Trichomycteridae) from the high
Andean range., Studies on Neotropical Fauna and Environment, 18:2, 65-87
This article may be used for research, teaching, and private study purposes.
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Studies on Neotropical Fauna and Environment 18 (1983), pp. 65-87.
ABSTRACT
Trichomycterus chungaraensis n. sp. and Trichomycterus laucaensis n. sp. are described. These
new species are from isolated environment 4,300 m above sea level in the high Andes of the north
of Chile. Some new characters of TrichomycterusrivulatusValenciennes from Titicaca Lake and
Lauca River are given.
TAXONOMY
ORDER : SILURIFORMES
FAMILY : TRICHOMYCTERIDAE
SUBFAMILY : PYGIDIINAE
GENUS : Trichomycterus Valenciennes, 1833
NEW TRICHOMYCTERIDAE FROM HIGH ANDES 67
Diagnosis
Fishes with deep and strong laterally compressed caudal penduncle. Short
dorsal fin; its origin behind the middle of total length. Anal fin placed partially
below dorsal fin or behind it. With or without pectoral filament. Small pelvic
fin. Anus placed near the base of pelvic fin, or between pelvic and anal fins.
Short supraorbital bone with variable shape. Orbitosphenoid short. More than
12 pairs of ribs. Caudal fin truncated, slightly rounded or rounded. More than
32 caudal fin rays; with 6 + 7 pincipal caudal rays; with two segmented and
nonbranched caudal fin rays in both lobes. Upper hypural 3,4 and 5 fused, or
only hypural 4 or 5 fused.
Elongate fishes of small size, maximum length not exceding 120 mm, depth of
caudal peduncle much less than the maximum depth of trunk. Head approxi-
mately triangular in shape. Maximum depth of body in the middle of predorsal
length. (Body relationships in Table 1). Nasal barbel reaches to the anterior
Table 1. Body relationships of Trichomycterus chungaraensis n. sp.
border or just behind the posterior border of the orbit; the maxillary barbels as
the longest reaches until the interopercular ; submaxillary barbel a little shorter
than maxillary barbel.
Incisiform shaped denticles in the external row on the interopercular. Pec-
toral, pelvic, dorsal and anal fins with rounded margin. Origin of pelvic fin in
the middle of total length. Origin of dorsal fin slightly in front of anal fin or
before. Caudal fin with posterior margin slightly rounded. Anus near the base
of pelvic fin. Great number of small translucent papillae all over the body,
including the base of fins. Preural 1 with large and strong neural apophysis (=
epural).
Interopercular armed with 34 to 60 denticles; opercular armed with 6 to 16
denticles. 6 to 8 branchiostegals. Pectoral fin with 8 or 9 rays. First pectoral ray
not included in a filament. Pelvic fin with 5 rays. Dorsal fin with 10 to 12 rays.
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Anal fin with 10-12 rays. Caudal fin with 38 to 46 rays. 38 to 42 vertebrae; 9 to
15 precaudal vertebrae, and 24 to 34 caudal vertebrae. 12 to 16 pairs of ribs.
The specific name is derived from Chungará Lake, Chile.
Type locality: Streams of Vertiente de Mal Paso, Chungará Lake, 4500 m above sea level, North
Chile, South America.
Description
Elongate fishes, with rounded margin of the fins and with narrow and laterally
compressed caudal peduncle (Fig. 2). Head flattened, approximately trian-
gular, maximum width in the opercular region. The body proportions vary in
lern
wide ranges (Table 1). The eyes are dorsal; they are black and rounded and
their size varies strongly (Table 1); proportionally, the eye size of adult
specimens is smaller than that of juveniles of small total length. The subter-
minal and moderately wide mouth has fleshy lips with numerous translucent
papillae which cover the oral surface, also. The barbels are of different length.
The nasal barbels are the shortest; they reach to the anterior border of the eye
NEW TRICHOMYCTERIDAE FROM HIGH ANDES 69
or just behind the posterior border of the eye. The maxillary barbels reach
generally the anterior margin of the interopercular; the submaxillary barbels
have a similar length or are shorter than the maxillary ones.
The branchiostegal membranes are joined to the isthmus; number of bran-
chiostegal rays is variable from 6-6, 7-7, 7-8 to 8-8. The most frequent values
are 7-7 and 8-8; in some specimens a small anterior branchiostegal ray ap-
pears.
The cranium shows the pattern described for Trichomycterus fin Arratia et
al., 1978), with a narrow and elongate supraorbital (Fig. 3A) displaced for-
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ward, near the articulation of the nasal-ethmoid lateral complex with the
frontal. The supraoccipital (Fig. 4A) does not form lateral crests for the
insertion of muscles, and the lateroposterior part of pterotic is comparatively
shorter than in T. laucaensis n. sp., and T. rivulatus.
The premaxillary and dentary bear 5 of 6 row of teeth. The teeth of the
external row are incisiform and the inner rows have conical teeth.
The opercular and interopercular (Fig. 5) bear denticles, there are 7 to 16 on
the opercular, and 34 to 60 on the interopercular. The denticles of the internal
row on the interopercular are larger than the others, and some of them are
incisiform, and other have a curious shape, as a bludgeon. The number of
denticles of the opercular and interopercular varies from one side to the other
in one individual and these denticles may be replaced during the life of the fish.
There are 38 to 42 vertebrae (average: 39.9) not counting the first one which
is fused to the cranium. There are 12 to 16 pairs of ribs, 15 is the most frequent
value, exceptionally, there are 11 pairs.
The origin of the dorsal fin is behind the middle of the total length. The
origin of the pelvic fin is in the middle of the total length in approximately 50%
70 GLORIA ARRATIA F.
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of specimens (Graph. 1), in some of them is just in front of the middle of the
total length, and in the rest it is behind the middle of the total length. The origin
of the anal fin is behind the origin of the dorsal fin.
—T.laucaantla
T. chungarían •!•
Fig. 5. Interopercular and preopercular (with the external row of denticles, only).
A: Interopercular of T. chungaraensis n. sp.;
B: Opercular of T. chungaraensis n. sp.;
C: Interopercular of T. laucaensis n. sp.;
D: Opercular of T. laucaensis n. sp.
The pectoral fin is rounded, and the leading ray is not prolonged in a
filament. There are 8 to 9 pectoral fin rays, ocassionally 10. The pelvic fin is
rounded and small and has 5 rays and a pelvic splint.
The dorsal fin has 10 to 12 rays of which the first 2 to 4 may be small and
simple; there are 7 to 10 soft dorsal rays; the most frequent number is 9. The
anal fin has 10 to 12 rays of which the first 2 to 4 may be simple; there are 7 soft
rays, ocassionally 8.
The caudal fin has a slightly rounded margin. The supporting skeleton of the
caudal rays involves 6 to 9 vertebrae. Hypural 1 and 2 are fused and incom-
pletely separated from the parhypural by a suture (Fig. 6A). Upper hypural 4
and 5 are coossified; hypural 3 is free. In 8% of the studied specimens the
upper hypural H3 is fused with H4 + H5, too. The uroneural is wide anteriorly.
The preural arch 1 is complete and has a long strong neural spine which
functions as a epural; this situation is present in 72.9% of the studied indivi-
duals. A large epural is present in 4.2% of the specimens. The epural is small
GLORIA ARRATIA F.
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Coloration: Variable; mainly depending upon the type and upon the change of
the substrate. Most of the fishes have a brown-yellow colour with many
brown and dark spots of different size on the head, the dorsal region, and on
the flanks; the fins are pale yellowish or pale brown with small dark spots.
Other specimens show the grey or pale brown skin on the head, the dorsal
region of the trunk, and on the flanks, with a dark line in the middle of the
flanks. The ventral part of the body is whitish or yellowish.
72 70 68
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extends behind the posterior border of the eye until the opercular; maxillary
barbel reaches until the posterior border of the interopercular. Submaxillary
barbel a little shorter than the maxillary one. Incisiform shaped denticles in the
external row on the interopercular.
Pectoral, dorsal, anal and caudal fin with posterior margin truncated. Origin
of the pelvic fin about in the middle of total length. Preural 1 with short neural
spine. Caudal skeleton without epural, generally. Anus near the base of pelvic
fin. Great number of small translucent papillae all over the body, including
base of the fins.
Interopercular armed with 45 to 55 denticles; opercular armed with 14 to
20 denticles. 7 or 8 branchiostegals. Pectoral fin with 8 or 9 rays; first pectoral
ray generally not prolonged in a filament. Pelvic fin with 5 rays. Dorsal fin with
11 to 15 rays. Anal fin with 9 to 11 rays. Caudal fin with 44 to 59 rays. 37 to 40
vertebrae; 12 to 15 precaudal vertebrae, and 22 to 27 caudal vertebrae. 13 to
17 pairs of ribs.
The specific name is derived from Lauca River, Chile.
Holotype: No. 160878 A, Ichthyological Collection. Faculty of Forestry Sciences. Universidad de
Chile.
Type locality: System of Lauca River, Parinacota, 4,390 m above sea level, Northern Chile, South
America.
Description
Elongate fishes with truncated margin of pectoral, dorsal, anal and caudal fins,
and with deep and laterally compressed caudal peduncle (Fig. 8). Head flat-
tened, approximately trapezoidal. The body proportions vary in wide ranges
(Table 2). The eyes lie dorsally.
The barbels show variations in length; the longest is the maxillary barbel
reaching the anterior margin of the interopercular or extending just to the
posterior margin of this bone.
NEW TRICHOMYCTERIDAE FROM HIGH ANDES 75
filament in 97,4% of the studied specimens. A small filament has been ob-
served in some of the largest specimens of both sexes. The pectoral fin shows 8
or 9 rays, ocassionally 10. The pelvic fin is similar to that of T. chungaraensis n.
sp.
The origin of pelvic, dorsal and anal fin is situated similar to that in T.
chungaris n. sp., with some variations (Table 1; Table 2; Graph. 1).
The dorsal fin has 11 to 15 rays of which the first 3 to 5 may be small and
simple. There are 8 to 10 soft dorsal fin rays; the most frequent number is 9.
The anal fin has 9 to 11 rays of which the first 2 to 5 may be simple. There are 6
to 9 soft anal fin rays; the most frequent number is 7.
The caudal fin is truncated. The supporting skeleton of the caudal rays
involves 8 to 11 vertebrae. The pattern of fusion of hypurals is similar to that
described above for T. chungaraensis n. sp. (Fig. 6B); contrary to T. chun-
garaensis n. sp. the hypural 4 + 5 is fused and the hypural 3 is free in 85 % of the
population; all three upper hypurals are free in 15% of the individuals as in
Nematogenys inermis (Fig. 11 A). The uroneural (Fig. 6B) is narrow and not
sharpened as in T. chungaraensis n. sp.
There is no epural in 80.7% of the specimens; the neural apophysis 1 is long
(= epural) in 19.3% of the specimens. The variations of the hypurapophyses
are similar to those described for T. chungaraensis n. sp.
There are 44 to 5 9 caudal fin rays (average : 50.1 ). The distribution of caudal
fin rays is similar to that of T. chungaraensis n. sp.
The anus is placed close to the base of the pelvic fins. The urogenital papilla
is conical in males and rounded in females; it is difficult to distinguish the sexes
in young specimens based on the shape of the papillae. The body, with
exeption of the fins, is covered with numerous translucent papillae slightly
rounded or conical. They seem to be more numerous on the head.
different sizes and shapes on the head, dorsal region, and on the flanks. The
fins are pale yellow or brown with some small dark spots. The ventral region of
the body is pale yellow or whitish.
DISCUSSION
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II FO/ILM)
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Graph. 2. Values of peduncle depth in per cent of standard length (PD/SL (%)) of the individuals
in chance samples of Trichomycterus chungaraensis and T. laucaensis.
— T. laucaansi»
-—-T. ehungsraanal«
HW/HL<«)
Graph. 3. Values of head width in per cent of head length (HW/HL (%)) of the individuals in
chance samples of Trichomycterus chungaraensis and T. laucaensis.
x: mean.
Table 3. Number of vertebrae, branchiostegals, fin rays and denticles in Trichomycterus chun-
garaensis n. sp., T. laucaensis n. sp., and T. Rivulatus.
Relationships Individuals less than 60 mm long. Individuals more than 80mm long.
Total length/Caudal fin depth 4.6 5.7 5,8 5.2 6.1 7.7 4.7 5.2 5.6 5.4 6.5 8.2
Head length/Eye diameter 7.0 9.1 11.9 5.7 8.7 11.4 7.0 11.0 18.0 7.5 10.1 13.0
Head length/Standard length (%) 19.0 20.3 22.0 18.0 20.1 22.0 16.0 18.0 20.0 16.0 20.5 23.0
Peduncle depth/Standard length (%) 9.0 10.1 12.0 10.0 11.6 13.0 9.0 10.3 11.0 11.0 14.6 16.5
Head depth/Head length (%) 48.0 56.6 75.0 43.0 50.3 61.0 48.0 52.9 64.0 50.0 56.1 64.0
mm. : minimum
aver. : average
max. : maximum
Table 5. Some body characters and relationships of Trichomycterus rivulatus from Valenciennes (1848), Eigenmann (1918), Tchemavin
(1944), and this paper. 3
o
Valenciennes Eigenmann Tchemavin this paper
S
Standard length/Head length 4.5-5.5 4.C1-6.2 4.5- 5.9 o
Head length/Standard length (%) _ 16 - 2 1 16.8- 21.0 SC
Peduncle depth/Standard length (%) _ 10 - 1 7 12.9- 18.1
Predorsal length/Standard length (%) _ _ 58 - 6 8 60.5- 64.7
Prepelvic length/Standard length (%) _ 53 - 6 1 53.0- 58.0
Preanal length/Standard length (%) _ _ 63 - 7 4 63.0- 69.0
Interorbital width/Head length (%) _ _ 25 - 4 5 25.1- 36.0
Head depth/Head length (%) 32 - 6 1 55.0- 65.6
Head width/Head length (%) - 73 - 1 0 5 87.5- 106.7
Perú and Western Bolivia from the region of Lake Junin in the north to Lake
Poopó in the south (roughly from 10° to 20°S). It is not known whether it is
found in mountains of Chile at the same latitude".
T. laucaensis n. sp. (Fig. 8) is externally more similar to T. rivulatus (Fig. 9)
than to T. chungaraensis n. sp. (Fig. 2); and it is possible that Mann has
considered them as T. rivulatus because of the similar external aspect. I have
studied specimens of T. rivulatus from Titicaca Lake and Lauca River to solve
the problem of similarity of the fishes of Lauca River with those of Titicaca
Basin.
Valenciennes (1848) used mainly the number of dorsal and anal fin rays (Table
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PH
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NS1
epural in 55% of the specimens, 32% of the specimens have a functional long
epural; 11% of the specimens have an oval small epural; and 2% of the
specimens have a long neural apophysis 1 (= epural).
The number of the caudal fin rays is the highest within Trichomycteridae,
with 62 to 68 caudal fin rays (average: 64,5). The distribution of the caudal fin
rays is similar to that described above for T. chungaraensis n. sp.
Conclusion
Trichomycterus rivulatus differs (Table 3, and compare Table 5 and 1) from T.
chungaraensis n. sp. mainly in the maximum total length, depth of the caudal
peduncle, depth of the head, shape of the caudal fin, number of vertebrae,
number of precaudal and caudal vertebrae, number of caudal fin rays, and
number of denticles on opercular and interopercular. The skeleton shows
some differences also, for example: shape of supraorbital, presence or absence
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B. Caudal skeleton.
The caudal skeleton presents important characters with good evidence to
separate different taxonomic categories (Arratia et al., 1978) within the Tri-
chomycteridae.
Hypural fusion.
Lunberg & Baskin ( 1969) and Arratia et al. ( 1978) have postulated that one of
the evolutive trends within Siluriformes is the fusion of hypurals, and the
fusion of the parhypural with hypural 1, too.
T. chungaraensis n. sp., T. laucaensis n. sp., and T. rivulatus have a par-
84 GLORIA ARRATIA F.
hypural sutured with the fused hypural 1 + 2 (Fig. 11B, C and D). This is an
intermediate situation between Nematogenys (and Diplomystes) (Fig. 11 A)
and T. areolatus (Fig. 1 IF) and chiltoni.
Most of the specimens of T. chungaraensis n. sp., T. laucaensis n. sp., and T.
rivulatus show an incomplete fusion between the upper hypurals; the hypural 3
is free and the hypurals 4 and 5 are fused. It is interesting to see that T.
laucaensis retains the pattern of Nematogenys in the three upper hypurals in
15% of the specimens (within the studied specimens, two are similar to that of
T. areolatus (Fig. 1 IF). T. chungaraensis n. sp. shows the complete fusion of all
three upper hypurals in 8% of the specimens. The populations of the Chungará
Basin and of the Lauca River show some changes in their osteological con-
figuration, it could be the case that these changes are caused by some genetic
modification of these isolated species.
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Epurai.
The presence of a functional epural, or of a small oval epural, or the absence of
the epural is different within Diplomystidae and Trichomycteridae. Di-
plomystes and Nematogenys show an epural (functional) in 100% of the
specimens but there are variations within different species in Trichomycterus,
Hatcheria and Bullockia (Fig. 12). The trend goes towards loss of the epural. A
comparison between paleontological evidences from pholidophorids, Jurassic
teleosteans and extant teleosteans demonstrates that the trend is to reduce the
number of épurais (Patterson, 1968 ; Patterson & Rosen, 1977). The loss of the
epural within Trichomycteridae can be considered then as a derived feature.
The presence of a long functional neural spine 1 in T. chungaraensis, and
some specimens of T. laucaensis has not been described before as a pattern of
any Siluriformes species. Lunberg & Baskin (1969) established for Vandellia
cirrhosa "no epural; but one with a full preural spine 1 (= epural), (in Table
1)". I have observed only 3 specimens with such feature in T. areolatus (of
more than 1,000 studied caudal skeletons); and only one in Bullockia mal-
donadoi. I have not seen a long neural spine 1 (= epural) in any specimen of T.
chiltoni, T. heterodontum, T. borellii, T. corduvence, T. mendozensis and
Trichomycterus sp. I consider as abnormality the presence of a long neural
spine 1 as epural in some specimens. I suppose that the functional long neural
spine 1 in most of specimens of T. chungaraensis n. sp. is the results of a genetic
change which has affected this isolated species in one moment during the time
since the species has been isolated in the high Andean.
C. Local species.
Tchernavin criticized Eigenmann strongly because of the use of political
bounderies, and of local criteria to separate species of South American fishes.
Even the first aspect is a mistake, the second aspect is a reality in South
America. That can be shown, for example, iarPercilia irwini Eigenmann, for
Trichomycterus chiltoni (Eigenmann), tor Bullockia maldonadoi (Eigenmann)
(region close to the coast between Concepción and Angol, Chile; in Arratia et
NEW TRICHOMYCTERIDAE FROM HIGH ANDES 85
100,
I: fe
so
: ns1
EU. ne
%
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100,
SO
%
10 tt
50
Fig. 12. Intraspecifíc and interspecific variation (in %) of the epural in some trichomycterids.
A: Nematogenys inermis; B: Trichomycterus rivulatus;
C: T. chungaraensis n. sp.; D: T. laucaensis n. sp.;
E: T. mendozensis; F: T. areolatus and Bullockia maldonadoi; G: T. corduvence and
Hatcheria macraei; H.T. chiltoni. fe: functional epural; ne: no epural; nsl: neural spine
as functional epural; oe: small oval epural.
86 GLORIA ARRATIA F.
al., 1978), and for Trichomycterus mendozensis Arratia et al. (small streams in
high mountains in Mendoza, Argentina). A very clear example of a local
species within South American fishes is Gynocharacinus bergi Steindachner
from Valcheta Stream, Province of Rio Negro, Argentina (Pozzi, 1936; Rin-
guelet et al., 1967). It is possible to find many similar examples of local species
within the South American fauna.
Ringuelet et al., (1967) recognized that the South American ichthyofauna
shows a strong endemism which makes this fauna completely different to that
of the Neartic and of the African fauna. Therefore, Thernavins critic of the
work of Eigenmann is not correct in this aspect.
Even in this paper two other new species with very restricted geographical
distribution are described.
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ACKNOWLEDGEMENTS
The present investigation is part of the Research Program MAB 6 UNESCO, No. 1105-77-01.
I am especially grateful to Prof. Dr. H.-P. Schultze for having read and criticized the entire
manuscript, and Dr. A. Veloso for the fruitful discussions, and for the antecedents of the
environment of Chungará Lake and Lauca River.
I am indebted very much to Drs. A. Veloso, J. Navarro, N. Diaz and M. Salaverry for collecting
fish specimens; and Mr. H. Torres and R. Palma from CONAF, for his technical assistance.
RESUMEN
Se describen dos nuevas especies de peces del Altiplano Chileno, Trichomycterus chungaraensis
n. sp. y Trichomycterus laucaensis n. sp. Ambas habitan aislados ambientes sobre los 4.300 m de
altura en la Cordillera de los Andes, en el norte de Chile. Se aportan nuevos antecedentes sobre
Trichomycterus rivulatus del Lago Titicaca y del Sistema hidrográfico del Río Lauca.
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