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Trichomycterus chungaraensis n. sp. and

Trichomycterus laucaensis n. sp. (Pisces, Siluriformes,
Trichomycteridae) from the High Andean Range.

Article  in  Studies on Neotropical Fauna and Environment · January 1983

DOI: 10.1080/01650528309360621

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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

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 Studies on Neotropical Fauna and Environment 18 (1983), pp. 65-87.

Trichomycterus chungaraensis n. sp. and Trichomycterus

laucaensis n. sp. (Pisces, Siluriformes, Trichomycteridae) from
the High Andean Range.
Gloria ARRATIA F.
(Lawrence, Kansas)
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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.

The Titicaca Province of the Biogeographic Andean Region includes geogra-

phically the Poopó Basin in Bolivia, the Titicaca Basin (in Bolivia and Perú),
the southern part of the Andean region of Perú, Lauca River (in Chile and
Bolivia), Chungará Lake and other small water bodies in the north of Chile
(Fig. 1). All these aquatic systems are 4,000 m above sea level and they are
isolated at least since the last upraising of the Andean range, so they form
interesting habitats for studies.
For this province, Ringuelet (1975) cited the presence of three fish genera:
Trichomycterus, Astroblepus and Orestias. Valenciennes (1846) described
Trichomycterus rivulatus from specimens of Guasacona (apparently basin of
Titicaca Lake). Eigenmann (1918) considered many Andean trichomycterids
as synonyms of this species. Tchernavin (1944) critically revised T. rivulatus,
and he included also Trichomycterus oroyae and Trichomycterus quechorum as
synonyms in T. rivulatus. Tchernavin recognized ten species as synonyms of T.
rivulatus. His revision was mainly based on body proportions.
In the present work, two new species of Trichomycterus are described. Some
characters of Trichomycterus rivulatus are given, and the importance of some
structures present in the Trichomycteridae is discussed. The generic diagnosis
of Trichomycterus is modified from that established by Arratia (in Arratia et
al., 1978) so that the characters of these new species are included.
 66 GLORIA ARRATIA F.
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Fig. 1. Limits of the Titicaca province in South America.

MATERIALS AND METHODS

198 specimens of Trichomycterus chungaraensis n. sp. were studied. These fishes were caught in
Vertiente de Mal Paso, Chungará Lake, Chile. 84 specimens of Trichomycterus laucaensis n. sp.
were studied. They were collected in the system of Lauca River, Chile. More than 2,500 specimens
of Trichomycterus rivulatus from Titicaca Lake and Lauca River, T. mendozensis, T. borelli, T.
corduvence, T. heterodontum, T. areolatus, T. chiltoni, Trichomycterus sp.. Nematogenys inermis,
and Diplomystes chilensis were used for comparative studies. All specific diagnoses and redes-
criptions of all Trichomycterus species were revised, especially all of those considered to be
synonyms with T. rivulatus.
Meristic and morphometric data were obtained. A high number of these fishes were claired and
stained with alizarin according to the methods described by Hollister (1934) and Taylor (1967).
Body measurements were taken according to Tchernavin (1944); the depth of the caudal fin was
taken at the maximum depth of the posterior margin of the fin, however.
The specimens of T. chungaraensis n. sp. and T. laucaensis n. sp. were collected by A. Veloso,
J. Navarro and M. Salaverry. All studied fishes are kept in the Ichthyological Collection of the
Laboratory of Biology, Faculty of Forestry Sciences, Universidad de Chile.

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.

TRICHOMYCTERUS CHUNGARAENSIS N. SP.

Diagnosis
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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.

Relationships Minimum Average Maximum

Total length/Standard length 1.11 1.14 1.40
Total length/Predorsal length 1.72 1.86 1.91
Total length/Prepelvic length 1.83 1.98 2.11
Total length/Preanal length 1.52 1.65 1.74
Total length/Head length 5.05 5.84 6.86
Total length/Maximum depth 5.48 7.00 9.02
Total length/Peduncle depth 7.73 10.29 12.18
Total length/Head depth 7.30 10.52 13.18
Total length/Pectoral fin length 7.30 8.30 9.23
Total length/Dorsal fin length 6.23 7.49 8.91
Total length/Caudal fin depth 4.41 5.14 6.00
Standard length/Head length 4.42 5.12 6.00
Peduncle depth/Head depth 0.63 1.01 1.26
Maximum depth/Peduncle depth 1.25 1.46 1.74
Caudal fin depth/Peduncle depth 1.65 ••-• 1.98 2.19
Caudal fin depth/Maximum depth 1.08 1.18 1.60
Head length/Interorbital width 2.43 3.15 4.20
Head length/Preorbital length 1.48 2.41 2.75
Mouth width/Interorbital width 0.90 1.14 1.37
Head length/Eye diameter 6.95 11.18 18.00
Predorsal length/Standard length 59.0 61.8 66.0
Prepelvic length/Standard length 53.1 57.1 63.2
Preanal length/Standard length 64.0 68.1 79.0
Head length/Standard length 16.0 19.1 22.0
Peduncle depth/Standard length 8.0 10.5 14.2
Head width/Head length 85.0 97.5 107.0
Head depth/Head length (%)
44.0 53.3 75.0
Interorbital width/Head length 23.0 31.4 41.0
 68 GLORIA ARRATIA F.

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.

Holotype: No. 290878 A, Ichthyological Collection. Faculty of Forestry Sciences, Universidad de

Chile.
Paratypes: No. 290878 B, No. 181278A, No. 181278 B, No. 200579 A, No. 200579 B. Ichthyo-
logical Collection, Faculty of Forestry Sciences. Universidad de 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

Fig. 2. Lateral view of Trichomycterus chungaraensis n. sp.

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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Fig. 3. Supraorbital bones.

AiTrichomycterus chungaraensis n. sp.
B : Trichomycterus laucaensis n. sp.
C: Trichomycterus rivulatus.

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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Fig. 4. Posterior part of the cranium: dorsal view.

A: Trichomycterus chungaraensis n. sp.;
B : Trichomycterus laucaensis n. sp.
lc: lateral crest of supraoccipital; pt: pterotic; soc: supraoccipital; sp: sphenotic-prootic
complex; swc: swim bladder capsule.

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 •!•

Graph. 1. Origin of pelvic fin in Trichomycterus chungaraensis and T. laucaensis.

PPL: prepelvic length; TL: total length
 NEW TRICHOMYCTERIDAE FROM HIGH ANDES 71
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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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Fig. 6. Caudal skeleton.

A: Trichomycterus chungaraensis n. sp.;
B: Trichomycterus laucaensis n. sp. CP1: preural centrum 1; h: secondary hypurapo-
physis; Hl-5: hypurals 1-5; NS1: neural spine 1; PH: parhypural; U: uroneural.
and oval in 12.7% of the specimens, while 10.2% show no epural (Fig. 11C
and 12C). The presence or absence of hypurapophyses is as described for the
family in Arratia (1976). The hypurapophysis may be absent, but if it is
present, it is very small. The secondary hypurapophysis is developed slightly.
There are 38 to 46 caudal fin rays (average: 42.2), 6 + 7 principal caudal
rays in both lobes and 2 non-branched and segmented rays in both lobes.
34.5% of the individuals have 3 non-branched and segmented rays in one or
both lobes.
The anus is placed near to the base of the pelvic fins. There is a small conical
urogenital papilla which sometimes cannot be determined in females. It is
difficult to observe the papilla in juveniles of small length, too. Microscopical
examinations show numerous small papillae on the skin of head and trunk.

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.

Geographical distribution and preliminary ecological data: Trichomycterys

chungaraensis n. sp.. is endemic in the freshwater of the Andean (Fig. 7) where
 NEW TRICHOMYCTERIDAE FROM HIGH ANDES 73

72 70 68
17
AricoV \
20

23
>
26

29
j-jj

32

J olsontiogo
35
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38
ft)

41

44

47 '
i\ <
50

53

56

90

\ £¡
PoloVsur

Fig. 7. Geographical distribution of Trichomycterus chungaraensis n. sp., and T. laucaensis n. sp.

A: T. chungaraensis; •: T. laucaensis.
 74 GLORIA ARRATIA F.

it occupies a restricted environment at about 4,500 m above sea level. The

fishes were collected in small streams with stony bottom. The streams are
narrow and shallow (10 to 30 cm). The aquatic vegetation is scarce. These
fishes have been collected with tadpoles and adults of Telmatobius marmora-
tus.

TRICHOMYCTERUS LAUCAENSIS N. SP.

Diagnosis
Elongate fishes of small size, maximum length not exceding 141 mm. Deep
caudal peduncle, strong laterally compressed. Maximum depth of the body at
the origin of dorsal fin, generally. (Body relationships in Table 2). Nasal barbel
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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.

Paratypes: No. 160878B, 280878A, 280878B, 190579A, 190579B, 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

Fig. 8. Lateral view of Trichomycterus laucaensis n. sp.

Table 2. Body relationships of Trichomycterus laucaensis n. sp.

Relationships Minimum Average Maximum

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Total length/Standard length 1.09 1.13 1.25

Total length/Predorsal length 1.69 1.89 1.97
Total length/Prepelvic length 1.77 1.96 2.70
Total length/Preanal length 1.53 1.65 1.93
Total length/Head length 4.97 5.63 6.99
Total length/Maximum depth 5.33 6.75 7.88
Total length/Peduncle depth 7.10 8.76 10.88
Total length/Head depth 8.07 10.74 12.80
Total length/Pectoral fin length 6.50 8.97 9.88
Total length/Dorsal fin length 6.50 7.25 8.34
Total length/Caudal fin depth 4.63 6.12 8.14
Standard length/Head length 4.33 4.96 5.95
Peduncle depth/Head depth 0.92 1.23 1.58
Maximum depth/Peduncle depth 1.00 1.32 1.63
Caudal fin depth/Peduncle depth 1.04 1.49 2.08
Caudal fin depth/Maximum depth 0.91 1.10 1.40
Head Iength/Interorbital width 2.29 3.25 4.28
Head length/Preorbital length 1.92 2.54 3.77
Mouth width/Interorbital width 0.91 1.09 1.40
Head length/Eye diameter 5.70 9.44 13.70
Predorsal length/Standard length 58.0 62.6 66.0
Prepelvic length/Standard length 53.0 57.6 64.0
Preanal length/Standard length 58.0 68.2 72.4
Head length/Standard length 16.0 19.2 23.0
Peduncle depth/Standard length 10.0 12.8 16.5
Head width/Head length 76.0 96.5 112.0
Head depth/Head length 43.0 52.4 64.0
Interorbital width/Head length 23.0 36.2
30.5

The branchiostegal membranes are joined to the istmus. The number of

branchiostegal rays varies between 7-7, 7-8 or 8-8; the most frequent number
is 7-7.
The cranium shows the pattern described for the genus Trichomycterus
(Arratia et al., 1978). The supraorbital has a characteristic form (Fig. 3B). The
supraoccipital has prominent lateral crest; especially strong crest may be
 76 GLORIA ARRATIA F.

present in the largest specimens. The lateroposterior corner of the pterotic

(Fig. 4B) is strongly sharpened, it projects far outwards.
The premaxillary and dentary bear 4 or 5 rows of long teeth. The teeth of the
external rowe are incisiform, the other teeth are small and delicate.
The opercular and the interopercular (Fig. 5) bear denticles, there are 14 to
20 denticles on the opercular and 45 to 55 denticles on the interopercular. The
number of denticles is as variable as described above for T. chungaraensis n. sp.
There are 37 to 40 vertebrae in addition to the first one which is fused to the
cranium (average: 38.8 vertebrae). 12 to 16 precaudal vertebrae; the most
frequent number is 16. There are 22 to 27 caudal vertebrae, the most frequent
number is 22 to 23. There are 13 to 17 pairs of ribs.
The pectoral fin presents a truncated posterior margin, the shape of the fin is
similar to that of Hatcheria; nevertheless the first ray is not prolonged in a
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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.

Coloration: Variable, depending upon the type of substrate mainly. Most of

the fishes have a brown ground colour with numerous dark spots of different
sizes and shapes, on the head, the dorsal region, and on the fanks. Other
specimens show a very pale yellow ground colour with many brown spots of
 NEW TRICHOMYCTERIDAE FROM HIGH ANDES 77

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.

Geographical distribution and preliminary ecological data: Trichomycterus

laucaensis n. sp. is endemic to the freshwaters of the Andean (Fig. 7) where it
lies in the restricted and isolated environment offered by the system of Lauca
River about 4.390 m above sea level. The fishes have been collected together
with Orestias sp. of small maximal total length, and with tadpoles and adults of
the frog, Telmatobius marmomtus.

DISCUSSION
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Trichomycterus chungaraensis n. sp. is externally similar to Trichomycterus

areolatus Valenciennes but differs mainly in osteological characters of the
cranial bones, and caudal skeleton and in the number of fin rays (Arratia &
Chang, 1975; Arratia et al., 1978), the geographic distribution is different,
also.
Trichomycterus areolatus has 5 to 7 branchiostegal rays; 35 to 42 vertebrae;
13 to 17 dorsal fin rays; 9 to 12 anal fin rays; and 32 to 41 caudal fin rays. T.
areolatus has no epural and the upper hypurals are fused (Fig. 1 IF) and the
lower hypural fan includes the parhypural.
Trichomycterus areolatus is living from Illapel River (31°35' S) in the north,
to the aquatic systems of the South of Chile (42°S, approximately).
The differences are evidences enough to separate Trichomycterus chun-
garaensis n. sp. from Trichomycterus areolatus.
Trichomycterus chungaraensis n. sp. and T. laucaensis n. sp. differ mainly in
the shape of fins (Fig. 2 and 8), depth of caudal peduncle (Graph. 2), depth of
caudal fin, preorbital length, interorbital width, head width (Graph. 3), size of
eye, constitution of caudal skeleton (Fig. 6), and in some cranial structures as:
shape of supraorbital bone (Fig. 3), shape of pterotic, presence or absence of
lateral crest on the supraoccipital (Fig. 4), and number of fin rays and denticles
on the opercular and interopercular (Table 3).
Comparison between the largest specimens of both species shows a more
clear definition of some body relationships (Table 4); for example: those
relationships as total length/caudal fin depth, head length/eye diameter and
peduncle depth/standard length (%), show stronger differences between
adults of T. chungaraensis and T. laucaensis. In addition, the head of the largest
specimens of T. chungaraensis is shorter than that of juveniles while there is
not a real difference in T. laucaensis.
Mann (1954) established the presence of T. rivulatus Valenciennes for the
north of Chile, and de Buen ( 1958) mentionned this species even he did not see
the specimens determined by Mann as T. rivulatus. Tchervanin (1944: 271)
wrote: "Trichomycterus rivulatus is widely distributed in high mountains of
 78 GLORIA ARRATIA F.

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.

Structures T. chungaraensis T. laucaensis T. rivulatus

Vertebrae 38 to 42 37 to 40 37 to 39
Precaudal vertebrae 9 to 15 12 to 16 10 to 11
Caudal vertebrae 24 to 34 22 to 27 26 to 28
Branchiostegal rays 6 to 8 7 or 8 7 or 8
Pectoral fin rays 8 or 9 8 or 9 8 or 9
Pelvic fin rays 5 5 5
Dorsal fin rays 10 to 12 11 to 15 11 to 13
Anal fin rays 10 to 12 9 to 11 9 to 11
Caudal fin rays • 38 to 46 44 to 59 62 to 68
Interopercular denticles 34 to 60 45 to 55 31 to 40
Opercular denticles 7 to 16 14 to 20 10 to 14
 Table 4. Body relationships of individuals of two groups of different total length (less than 60 mm, and over 80 mm) of Trichomycterys
chungaraensis n. sp. and T. laucaensis n. sp.

Relationships Individuals less than 60 mm long. Individuals more than 80mm long.

T. chungaraensis T. laucaensis T. chungaraensis T. laucaensis

min. aver. max. min. aver. max. min. aver. max. min. aver. max.
Total length/Head length 5.1 5.5 8.9 5.1 5.4 6.1 5.6 6.2 8.3 4.9 5.6 7.0
Total length/Head depth 7.3 10.3 11.0 9.3 10.9 12.2 9.9 11.3 13.4 8.1 10.3 12.7 m
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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

Number of dorsal fin rays 8 13 16 1 1 - -13

Number of anal fin rays 7 11 + 4-6 9 - -11
Posterior margin of caudal fin rounded rounded not rounded rounded
 80 GLORIA ARRATIA F.

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.

A. Trichomycterus rivulatus Valenciennes.

Valenciennes (1848) used mainly the number of dorsal and anal fin rays (Table
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5) to separate T. rivulatus as a new species. Later, Eigenmann (1918) and

Eigenmann & Allen (1942) gave more characters of this species (Table 5).
Tchernavin (1944) considered ten formely described species as names or
synonyms of T. rivulatus* and he writes that the "study of variations of this
species is obscured by the changes with growth, the rates of which are ap-
parently different in different localities and in different series of specimens"
(p. 269). Tchernavin established some body relationships of T. rivulatus stu-
ding more than 300 specimens from different localities (Table 5).
The examinations of T. rivulatus shows that the supraoccipital (Fig. 10) is
more similar to T. laucaensis (Fig. 4B), with a lateral crest for the insertion of
muscles; the lateral crest is more prominent in T. rivulatus. The posterior end
of the supraorbital is sharpened (Fig. 3C). The premaxillary and dentary have
more than 4 rows of conical teeth. The opercular is armed with 10 to 14 conical
denticles, and the interopercular bears 31 to 40 denticles. Most of denticles on
the opercular and interopercular are conical; some of them are incisiform in
some specimens. There are 7 or 8 branchiostegal rays. There are 37 to 39
vertebrae (Table 3). There are 13 to 18 pairs of ribs.
The origin of pelvic, dorsal and anal fins are similar to those of T. chun-
garaensis and T. laucaensis. All fins have a rounded posterior margin. Valen-
ciennes (1846) and Eigenmann (1918) considered a rounded caudal fin as
characteristic for T. rivulatus; Tchernavin (1944) described a not rounded
caudal fin for this species; my observations are in agreement with those of
Valenciennes and Eigenmann.
The first pectoral fin ray is not prolonged in a filament, generally, only some
specimens are exceptional, and this variation has been observed already by
Eigenmann (1918), Eigenmann & Allen (1942), and Tchernavin (1944). The
pectoral fin has 8 or 9 rays as in T. laucaensis and T. chungaraensis. The pelvic
* Species considered as synonyms of Trichomycterusrivulatus:Trichomycterus incae Valencien-
nes; Trichomycterus gracilis Valenciennes; Trichomycterus barbatulo Valenciennes; Tri-
chomycterus pictus Castelnau; Trichomycterus eigenmanni Boulenger; Trichomycterus poeya-
nus Cope; Pygidium oroyae Eigenmann and Eigenmann; Pygidium quechuorum Steindachner;
Pygidium tiraque Fowler; and Pygidium atochae Allen.
 NEW TRICHOMYCTERIDAE FROM HIGH ANDES 81

Fig. 9. Lateral view of Trichomycterus rivulatus.

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Fig. 10. Posterior part of the cranium of Trichomycterusrivulatus;dorsal view..

lc: lateral crest of supraoccipital; pt: pterotic; soc: supraoccipital; sp: sphenotic-prootic
complex; swc: swim bladder capsule.

fin has 5 rays and a small splint as is common in Trichomycteridae in general

(Arratia et al., 1978).
The dorsal fin has 11 to 13 rays of which the first 3 to 5 may be small and
simple, and 7 to 9 are soft dorsal fin rays. The total number of dorsal fin rays
(counted in claired specimens stained with alizarin) is different from that of
Valenciennes (1848), Eigenmann (1918), and Tchernavin (1944) (Table 5).
Tchernavin (1944) established 16 dorsal fin rays, exceptionally 17, as
diagnostic for T. rovulatus, a number higher as observed here. Tchernavin
separated the branched dorsal fin rays (5-7) from all others in his work as a
way to give additional characters for this species. I found the same number of
branched dorsal fin rays. The difference to Tchernavin lies within the number
of simple dorsal fin rays and simple-segmented dorsal fin rays.
The anal fin has 9 to 11 rays which the first 3 to 4 are simple, and 6 or 7 are
soft anal fin rays; this total number shows some differences from those of other
authors (Table 5).
The support of the caudal skeleton involves 10 to 12 vertebrae. The pattern
 82 GLORIA ARRATIA F.

of hypural fusion is similar to that of T. chungaraensis and T. laucaensis (Fig.

1 IB, C. D), with fusion of hypural 4 and 5, and the hypural 3 free. There is no

PH
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NS1

Fig. 11. Semidiagramatic caudal skeleton of some South American trichomycterids.

A: Nematogenys inermis; B: Trichomycterus rivulatus; C: T. chungaraensis n. sp.;
D: T. laucaensis n. sp.; E: T. mendozensis n. sp.; F: T. areolatus. EP: epural; h:
secondary hypurapophysis; Hl-5: hypurals 1-5; NS1: neural spine 1; PH; parhypural;
U: uroneural.
 NEW TRICHOMYCTERIDAE FROM HIGH ANDES 83

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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of lateral supraoccipital crests, shape of pterotic, and presence or absence of

epural.
These differences are evidence enough to separate Trichomycterus chun-
garaensis n. sp. from Trichomycterus rivulatus.
Trichomycterus rivulatus differs (Table 3, and compare Table 5 and 2) from
T. laucaensis n. sp. mainly in the shape of pectoral, dorsal, anal, and caudal fins,
number of precaudal and caudal vertebrae, number of caudal fin rays and
number of denticles on opercular and interopercular. There are some differ-
ences in some osteological features, too. For example: the shape of the supra-
orbital, the development of a lateral supraoccipictal crest, the presence or
absence of epural.
The maximum length of T. rivulatus is about 350 mm (Eigenmann &
Eigenmann, 1890: 331); the new species from the high mountains of northern
Chile are comparatively smaller, the total length seems to be connected with
the environment the fishes occupy. Tchernavin (1944) studied series of speci-
mens from different localities, and he concluded that the populations of T.
rivulatus show different maximum length if they are living in lakes or in
streams, (p. 269: "the specimens from lakes and rivers are larger than those
from small streams").
The comparison between T. rivulatus and T. laucaensis n. sp. shows that
there are anatomical evidences to consider T. laucaensis as a new species.

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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Dr. Gloria Arratia F. Museum of National History, University of Kansas

Lawrence, Kansas 66045, U.S.A.

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