q 2003 by the American Society of Ichthyologists and Herpetologists
Copeia, 2003(3), pp. 583?587
New Species of Creagrutus (Teleostei: Characiformes: Characidae) from
the Rio Uaupe?s Basin, Brazil
RICHARD P. VARI AND FLA? VIO C. T. LIMA
A new species of Creagrutus is described from the western portions of the Rio
Uaupe?s basin near the Brazilian-Colombian border region. The species is most close-
ly related to, and most similar externally to, Creagrutus amoenus and Creagrutus kun-
turus, but differs from those species in details of pigmentation, meristics, and mor-
phometrics.
Uma nova espe?cie de Creagrutus e? descrita da porc?a?o ocidental da bacia do rio
Uaupe?s, pro?ximo a? fronteira entre o Brasil e a Colo?mbia. A espe?cie e? mais proxi-
mamente relacionada, e mais similar externamente, a? Creagrutus amoenus e Creagru-
tus kunturus, mas difere destas espe?cies em detalhes da pigmentac?a?o, dados mer??s-
ticos e morfome?tricos.
THE recognized diversity in the Neotropical?sh genus Creagrutus has been increased
dramatically in recent years (Harold and Vari,
1994; Vari and Harold, 2001, and references
therein) but with major lacunae in the known
distribution of the genus across many regions of
South America (Vari and Harold, 2001: ?g. 18).
In many instances, the distributional gaps are
most likely artifacts of the lack of ichthyological
collecting efforts in a particular region rather
than indicative of the actual absence of species
of Creagrutus in those areas. Collecting efforts
by the second author in the upper Rio Tiquie?,
an ichthyologically poorly sampled tributary of
the Rio Uaupe?s, in the western portions of the
Rio Negro basin along the Brazilian-Colombian
border, yielded the ?rst samples of Creagrutus
from that river basin. These represent a previ-
ously unknown species of Creagrutus that is de-
scribed herein.
MATERIALS AND METHODS
Counts and measurements were taken ac-
cording to Harold and Vari (1994) with verte-
bral counts taken from radiographs. Institution-
al abbreviations follow Leviton et al. (1985). In-
stitutional catalog numbers are followed by the
number of specimens in the lot, range in stan-
dard lengths, and locality information.
Creagrutus tuyuka n. sp.
Figure 1
Holotype.?MZUSP 65679, 1 (65.8 mm SL), Bra-
zil, Amazonas, Rio Tiquie?, comunidade de Ca-
choeira Comprida, beach below the cachoeira
(0815944.00N, 70801905.20W). F. C. T. Lima et al.,
22 October 2000.
Paratypes.?MZUSP 78519, 6 (43.3?68.3 mm SL;
1 specimen, 51.1 mm SL, cleared and counter-
stained); USNM 369731, 2 (46.9?56.4 mm SL);
collected with holotype. MZUSP 64689, 2 (46.9?
56.4 mm SL); INPA 18658, 2 (51.8?53.1) mm
SL; Brazil, Amazonas, Igarape? Umari, tributary
of rio Tiquie?, comunidade de Sa?o Pedro
(08159410N, 698579230W), F. C. T. Lima et al.,
25?27 October 2000.
Diagnosis.?Creagrutus tuyuka is most similar in
terms of dentition, head and body form, and
other features to Creagrutus amoenus and Crea-
grutus kunturus with which is shares two syna-
pomorphies (see discussion under Phylogenetic
relationships below). Creagrutus tuyuka differs
from C. amoenus and C. kunturus in its lack of
the series of dark midlateral spots on the body
that progressively coalesce ontogenetically to
sometimes form a midlateral stripe in adults of
the latter two species and in the distance from
the snout to the anal ?n origin (27.2?30.7 vs
33.8?38.8 and 30.8?34.0, respectively), distance
from the snout to the pelvic ?n insertion (22.2?
25.6 vs 29.0?35.6 and 27.9?31.6, respectively),
and the caudal peduncle depth (8.9?9.8 vs
12.2?13.8 and 11.1?12.1, respectively). Creagru-
tus tuyuka further differs from C. amoenus in the
number of vertebrae (39?41 with 39 in four
specimens and 40 or 41 in 10 specimens vs 35
to 39 with 36?38 in 34 specimens and 39 in only
one of 36 radiographed specimens, respective-
ly) and from C. kunturus in the number of me-
dian predorsal scales (9 or 10 vs 12, respective-
ly).
Description.?Morphometric and meristic data
for the holotype and paratypes presented in Ta-
ble 1. Head and anterior portion of body mod-
584 COPEIA, 2003, NO. 3
Fig. 1. Creagrutus tuyuka, holotype, MZUSP 65679, 65.8 mm SL; Brazil, Amazonas, Rio Tiquie?, comunidade
de Cachoeira Comprida, beach below cachoeira.
TABLE 1. MORPHOMETRIC AND MERISTIC CHARACTERS OF HOLOTYPE OF Creagrutus tuyuka, NEW SPECIES. (A)
Holotype of C. tuyuka, MZUSP 65679; and (B) 13 paratypes of C. tuyuka. Standard length is expressed in
millimeters; measurements 1?14 are percentages of standard length; 15?18 are percentages of head length.
(A) (B)
Morphometrics
Standard length 65.8 43.3?68.3
1. Snout to anal-?n origin
2. Snout to pelvic-?n insertion
3. Snout to pectoral-?n insertion
4. Snout to dorsal-?n origin
5. Dorsal-?n origin to hypural joint
6. Dorsal-?n origin to anal-?n origin
7. Dorsal-?n origin to pelvic-?n insertion
8. Dorsal-?n origin to pectoral-?n insertion
9. Caudal peduncle depth
61.2
47.9
24.2
45.7
57.9
30.1
25.5
30.5
9.7
59.5?63.3
45.5?49.0
22.8?25.2
44.6?47.2
55.6?58.9
27.2?30.7
22.2?25.6
29.5?31.1
8.9?9.8
10. Pectoral-?n length
11. Pelvic-?n length
12. Dorsal-?n length
13. Anal-?n length
14. Head length
15. Postorbital head length
16. Snout length
17. Bony orbital diameter
18. Interorbital width
16.3
13.8
19.0
15.5
25.4
40.7
26.9
34.1
28.1
16.9?18.8
12.4?13.7
17.4?20.6
14.4?16.8
24.1?26.4
34.8?42.4
23.9?29.8
33.5?38.9
25.2?29.4
Meristics
Lateral-line scales
Scale rows between dorsal-?n origin and lateral line
Scale rows between anal-?n origin and lateral line
Predorsal median scales
Branched dorsal-?n rays
Branched anal-?n rays
Branched pelvic-?n rays
Pectoral-?n rays
Vertebrae
39
4
3
10
8
12
7
13
39
39?41
4?5
3?4
9?10
8
11?12
7
12?14
39?41
erately robust; region of body posterior of ver-
tical through anal-?n origin more slender.
Greatest body depth situated slightly anterior of
vertical through dorsal-?n origin. Dorsal pro?le
of head distinctly convex from tip of snout to
vertical through anterior nares, slightly convex
or straight from that point to vertical through
middle of orbit, and straight from that point to
rear of supraoccipital spine. Interorbital region
somewhat convex overall transversely, albeit ?at-
585VARI AND LIMA?RIO UAUPE? S CREAGRUTUS
tened medially. Predorsal pro?le of body slight-
ly convex. Ventral pro?le of head in many in-
dividuals with variably obvious obtuse angle at
anteroventral corner of dentary, pro?le straight
to gently curved from that region to isthmus.
Ventral pro?le of body convex to anal-?n origin
and slightly concave from anal-?n insertion to
caudal ?n.
Head pointed in lateral view and distinctly
pointed in dorsal view. Upper jaw distinctly lon-
ger than, and overhanging, lower jaw. Anterior
portion of snout ?eshy, with scattered papillae.
Papillae more concentrated along ventral mar-
gin of overhanging upper lip and on ?eshy folds
and plicae that extend between outer and me-
dial premaxillary teeth. Lower lip ?eshy anteri-
orly with scattered papillae along dorsal margin;
papillae more concentrated proximate to den-
tary teeth.
Infraorbital series moderately developed.
Central portion of ventral margin of third infra-
orbital horizontally straight and in contact with
horizontal limb of preopercle for distance equal
to, or slightly greater than, one-half of horizon-
tal length of third infraorbital. Posterior mar-
gins of third through ?fth infraorbitals distinctly
separated from vertical limb of preopercle, ex-
tent of gap decreasing gradually dorsally.
Premaxillary dentition in three series: pri-
mary row slightly curved, typically with six teeth,
?ve teeth present on one premaxilla in one
specimen, without pronounced gap between
?rst and second tooth of series but with antero-
medial tooth separated from contralateral
equivalent by distinct gap ?lled by ?eshy folds;
triangular cluster of three larger teeth lying me-
dial to primary row; and single tooth of form
similar to that of primary series situated lateral
to fourth tooth of primary premaxillary row.
Maxilla with two, or more commonly three, tri-
cuspidate teeth. Dentary with ?ve tricuspidate
teeth. First and second dentary teeth robust and
distinctly larger than remaining teeth in series,
with second tooth about one-third longer than
?rst two teeth and twice as long as third tooth;
fourth and ?fth teeth distinctly smaller than an-
terior teeth and compressed.
Dorsal-?n rays ii,8. Dorsal-?n origin located
slightly anterior of vertical through pelvic-?n in-
sertion. Pro?le of distal margin of dorsal ?n
very slightly concave. Anal-?n rays iii,11?12. Pro-
?le of distal margin of anal ?n ranges from
straight to slightly convex. Hooks typically pres-
ent on anal-?n rays in mature males of many
species of Creagrutus not found in examined
specimens. Pectoral-?n rays i,11?13. Tip of pec-
toral ?n falls 2 or 3 scales short of vertical
through pelvic-?n insertion. Pelvic-?n rays i,6,i
or i,7. Tip of pelvic ?n extends posteriorly to,
or falls slightly short of, anal-?n origin. Gill-rak-
ers 6?718?9.
Coloration in alcohol.?Overall ground coloration
of relatively freshly collected specimens light
tan. Dorsal surface of head with dense ?eld of
small to midsize dark chromatophores giving re-
gion distinctly dusky appearance. Intensity of
dark pigmentation more noticeable in region
over brain, presumably as consequence of deep-
lying pigmentation overlying membranes of that
organ in most species of Creagrutus. Larger spec-
imens with dark pigmentation especially dense
in region of fontanels. Dense surface pigmen-
tation continues anteriorly over snout and up-
per lip, often with pigmentation particularly in-
tense over snout. Crescent-shaped patch of
denser pigmentation anterior to nares obvious
in some smaller specimens, particularly those
with light overall coloration; patch continuous
with other dark pigmentation anterior to orbit
in more darkly pigmented smaller individuals
and all larger specimens. Regions immediately
anterior and ventral to orbit with variably de-
veloped ?eld of dark pigmentation. Lower por-
tion of cheek lacks dark chromatophores in all
but largest individuals that have scattered dark
chromatophores in that region. Variably devel-
oped pattern of dark pigmentation posterior to
orbit; pigmentation ranges from diffuse, irreg-
ular pattern to somewhat more distinct but,
nonetheless, overall diffuse horizontal stripe.
Dark pigmentation on cheek, when present,
and on opercle extends further ventrally in larg-
er individuals.
Scales of dorsal portion of body with small
dark chromatophores concentrated along pos-
terior margin of scales to form overall reticulate
pattern. Pigmented region on each scale more
extensive in larger individuals. Darker primary
portion of humeral mark rotund, with variably
developed, less intensely pigmented areas that
extend to varying degrees dorsal and ventral of
main body of mark in some individuals. Pig-
mentation in humeral mark absent or less de-
veloped along irregular horizontal line that
overlies lateral-line canal. Midlateral stripe ex-
tends from slightly posterior of rear of humeral
mark to rear of caudal peduncle. Stripe more
intensely pigmented in larger individuals. More
darkly pigmented individuals with scattered
dark pigmentation in region above base of anal
?n and across ventral portion of caudal pedun-
cle.
Dorsal ?n with membranes and margins of
anterior ?n rays with ?elds of black chromato-
phores on distal two-thirds of ?n. Pigmentation
586 COPEIA, 2003, NO. 3
does not extend to distal portions on ?n mem-
branes in smaller individuals but reaches ?n
margin in larger specimens. Intensity of pig-
mentation in that region increases ontogeneti-
cally and supplemented in larger individuals by
distinct elongate patches of dark pigmentation
on ?n membranes proximate to anterior mar-
gin of two or three ?n rays posterior to primary
?eld of dark pigmentation. Basal portions of
anal-?n rays outlined by small dark chromato-
phores in small to midsized individuals; pigmen-
tation extends further distally in larger speci-
mens. Anal-?n membranes unpigmented in all
smaller specimens and in larger individuals with
light overall pigmentation but with patches of
dark pigmentation that form obscure patch of
dark pigmentation on central portion of mem-
branes between ?rst and third to ?rst to ?fth
branched anal-?n rays in darkly pigmented in-
dividuals. Caudal-?n rays and membranes with
scattered small dark chromatophores; pigmen-
tation particularly developed in larger individ-
uals. More intensely pigmented specimens with
diffuse dark patches of pigmentation on basal
portions of each caudal-?n lobe, at base of mid-
dle caudal-?n rays, and on distal portions of
each ?n lobe. Pelvic and pectoral ?ns nearly hy-
aline in smaller individuals, with scattered dark
pigmentation along ?n margins in larger spec-
imens.
Ecology.?At the type locality of C. tuyuka, the
Rio Tiquie? is a moderate-sized river, about 12?
14 m wide, with black water that is not, however,
as dark as that in most of the Rio Negro basin
or even further downstream in the Rio Tiquie?.
The type series was collected along the edge of
a large pool immediately below a small fall pre-
ceded upstream by a long series of rapids. Water
depth in the collection area was approximately
0.20?0.50 m with a bottom composed of sand
and ?ne gravel. Other paratypes were collected
at Igarape? Umari, a clear water stream approx-
imately 0.4?0.6 m deep with a sandy bottom.
Stomach contents of the cleared-and-counter-
stained paratype consisted exclusively of benthic
aquatic insect larvae, speci?cally several entire
individuals of the Baetidae (Ephemeroptera)
and a head of Hydropsychidae (Trichoptera).
Underwater observations of individuals of C. tu-
yuka at a small beach about 1 km upriver from
the type locality indicated that individuals swim
in small schools of four to six ?sh close to the
substrate. These observations, along with the
stomach contents of the single paratype, indi-
cate that C. tuyuka is a benthic, insectivorous
forager, a feeding habit similar to that of C. brits-
kii from the upper Rio Tocantins basin and an
unidenti?ed species of Creagrutus from the up-
per Rio Paraguay basin (FCTL pers. obs.). Oth-
er species of Creagrutus are also insectivorous
(Vari and Harold, 2001)
Distribution.?Creagrutus tuyuka is only known
from Rio Tiquie?, a tributary of the Rio Uaupe?s,
in the Rio Negro basin of northwestern Brazil.
Phylogenetic relationships.?Creagrutus tuyuka
shares the series of synapomorphies for mem-
bers of Creagrutus proposed by Vari and Harold
(2001:39; clade C?synapomorphies 16 to 29)
and the synapomorphies for Clade E within that
genus that includes the majority of species of
Creagrutus (Vari and Harold, 2001:40; clade E?
synapomorphies 35 and 36). Vari and Harold
(2001:41) advanced three synapomorphies for a
clade (Clade I) that consists of C. amoenus and
C. kunturus. Creagrutus tuyuka shares two of
these features (Synapomorphy 57?primordial
ligament distinctly rotund in cross-section and
extends ventrally nearly to distal tip of maxilla;
and Synapomorphy 58?scapula lacks narrow
ring-like process that forms anterior border of
scapular foramen). Creagrutus tuyuka lacks the
third synapomorphy for C. amoenus and C. kun-
turus, the presence of midlateral body pigmen-
tation in the form of a series of dark spots that
progressively coalesce ontogenetically and
sometimes form a midlateral stripe in larger in-
dividuals. The latter feature, that is unique to
those two species in Creagrutus, is, thus, herein
considered to be a synapomorphy for C. amoe-
nus and C. kunturus. Creagrutus tuyuka is, in
turn, hypothesized to be the sister species of C.
amoenus and C. kunturus based on the common
possession in the three species of the two syna-
pomorphies noted above (Synapomorphies 57
and 58 of Vari and Harold, 2001).
Etymology.?The species name, tuyuka, is in ref-
erence to the Tuyuka tribe, an indigenous
group of the Rio Tiquie? who have long carefully
managed the subsistence ?shery of that region
(Chernela, 1989) and were of great assistance
to the second author during the expedition that
yielded the specimens that served as the basis
for this study.
COMPARATIVE MATERIAL EXAMINED
Creagrutus amoenus: Ecuador, Pastaza, R??o Danta,
tributary to R??o Conambo. USNM 164070, 1
(specimen cleared and counterstained). Ecua-
dor, Pastaza, R??o Danta, tributary to R??o Conam-
bo (18459030S, 768479040W), USNM 311297, 2
(64.2?68.0); USNM 311339, 3 (1 specimen
587VARI AND LIMA?RIO UAUPE? S CREAGRUTUS
cleared and counterstained). Ecuador, Napo,
R??o Cowi (08089100S, 768159180W), USNM
340985, 4. Ecuador, Sucumbios, R??o San Miguel
basin, R??o La Bermeja, in front of Communidad
Shuor Chari (approximately 08109N, 768259W),
USNM 340986, 4.
Creagrutus kunturus: Peru, Amazonas, Provincia
Condorcanqui, Cordillera del Condor, upper
R??o Comainas, 20 m upriver of Puesto de Vigi-
lancia No. 22 (38569300S, 788249200W), USNM
335146, 1 (paratype of Creagrutus kunturus).
Peru, Amazonas, Provincia Condorcanqui, Cor-
dillera del Condor, R??o Comainas, 30 m down
river of Puesto de Vigilancia No. 22 (38569300S,
788249200W), USNM 335148, 2 (paratypes of
Creagrutus kunturus). Peru, Amazonas, Provincia
Condorcanqui, Cordillera del Condor, Quebra-
da no. 3, tributary of R??o Comainas, Puesto de
Vigilancia No. 22 (38569300S, 788249200W),
USNM 335147, 2 (paratypes of Creagrutus kun-
turus; 1 specimen cleared and counterstained).
ACKNOWLEDGMENTS
The specimens that served as the basis for this
paper were collected during an expedition of
the ``Peixes e pesca no alto Tiquie?'' project, a
collaborative study by MZUSP, the Instituto So-
cioambiental, Federac?a?o das Organizac?o?es In-
d??genas do Rio Negro (FOIRN), and the Asso-
ciac?a?o das Tribos Ind??genas do alto rio Tiquie?
(ATRIART). We thank A. Cabalzar, F. Cabalzar,
M. Lopes, C. A. Ricardo, and the staffs of
FOIRN and ATRIART for diverse assistance dur-
ing the expedition. Particular thanks are owed
to the Tuyuka people from Cachoeira Compri-
da and Sa?o Pedro for their generous hospitality
during our visit. Research associated with the
project was supported, in part, by funding from
the Neotropical Lowland Research Project of
the Smithsonian Institution to RPV and by a
grant from FAPESP (grant 99/02402-0) to
FCTL. Pita?goras C. Bispo (MZUSP) identi?ed
stomach contents of the paratype of Creagrutus
tuyuka. Figure 1 was prepared by T. B. Griswold.
This paper bene?tted from comments and sug-
gestions of A. M. Zanata and T. A. Munroe.
LITERATURE CITED
CHERNELA, J. 1989. Managing rivers of hunger: the
Tukano of Brazil. Adv. Econ. Bot. 7:238?248.
HAROLD, A. S., AND R. P. VARI. 1994. Systematics of the
trans-Andean species of Creagrutus (Ostariophysi:
Characiformes: Characidae). Smiths. Contr. Zool.
551:1?31.
LEVITON, A. E., R. H. GIBBS JR., E. HEAL, AND C. E.
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VARI, R. P., AND A. S. HAROLD. 2001. Phylogenetic
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(RPV) SECTION OF VERTEBRATE ZOOLOGY (FISH-
ES), NATIONAL MUSEUM OF NATURAL HISTORY,
SMITHSONIAN INSTITUTION, WASHINGTON, DC
20560-0159; AND (FCTL) MUSEU DE ZOOLOGIA
DA UNIVERSIDADE DE SA?O PAULO, CAIXA POST-
AL 42594, 94299-970, SA?O PAULO, BRAZIL. E-
mail: (RPV) vari.richard@nmnh.si.edu; and
(FCTL) fctlima@usp.br. Send reprint requests
to RPV. Submitted: 3 Oct. 2002. Accepted: 14
Feb. 2003. Section editor: D. G. Buth.