A peer-reviewed open-access journal

Zookeys 733: 65-97 (2018)

doi: 10.3897/zookeys.733.20159 #7Z,00Ke y S

http:/ / ZOO keys -pen soft. net Launched to accelerate biodiversity research

Fishes of the Cusiana River (Meta River basin,
Colombia), with an identification key to its species

Alexander Urbano-Bonilla', Gustavo A. Ballen'?, Guido A. Herrera-R'?,
Jhon Zamudio!*, Edgar E. Herrera-Collazos', Carlos DoNascimiento’,

Saul Prada-Pedreros', Javier A. Maldonado-Ocampo'!

| Laboratorio de Ictiologia, Unidad de Ecologia y Sistemdtica (UNESIS), Departamento de Biologia, Facultad
de Ciencias, Pontificia Universidad Javeriana, Carrera 7 N° 43-82, Bogota, D.C., Colombia 2. Museu de Zoo-
logia da Universidade de Séo Paulo, Caixa Postal 42494, 04218-970 Séio Paulo, SP Brazil 3 Université Paul
Sabatier, 118 Route de Narbonne, 31062, Toulouse, France 4 Grupo de Investigaciones territoriales para el uso
y conservacién de la Biodiversidad, Fundacién Reserva Natural La Palmita - Centro de Investigacién, Carrera
4 N® 58-59, Oficina 301, Bogotd, D.C., Colombia 5 Colecciones del Instituto de Investigacién de Recursos
Biolégicos Alexander von Humboldt, Claustro de San Agustin, Carrera 8 N° 15-08. Villa de Leyva, Colombia

Corresponding author: Alexander Urbano-Bonilla (bio.ictiologia@gmail.com),

Javier A. Maldonado-Ocampo (maldonadoj@javeriana.edu.co)

Academic editor: D. Bloom | Received 9 August 2017 | Accepted 2 January 2018 | Published 26 January 2018
http://zoobank. ore/6B807294-6 1 6E-437F-95B6-2F FDAB1546F7

Citation: Urbano-Bonilla A, Ballen GA, Herrera-R GA, Zamudio J, Herrera-Collazos EE, DoNascimiento C,
Prada-Pedreros S, Maldonado-Ocampo JA (2018) Fishes of the Cusiana River (Meta River basin, Colombia), with an
identification key to its species. ZooKeys 733: 65-97. https://doi.org/10.3897/zookeys.733.20159

Abstract

The Cusiana River sub-basin has been identified as a priority conservation area in the Orinoco region in
Colombia due to its high species diversity. This study presents an updated checklist and identification key
for fishes of the Cusiana River sub-basin. The checklist was assembled through direct examination of speci-
mens deposited in the main Colombian ichthyological collections. A total of 2020 lots from 167 different
localities from the Cusiana River sub-basin were examined and ranged from 153 to 2970 m in elevation.
The highest number of records were from the piedmont region (1091, 54.0 %), followed by the Llanos
(878, 43.5 %) and Andean (51, 2.5 %). 241 species distributed in 9 orders, 40 families, and 158 gen-
era were found. The fish species richness observed (241), represents 77.7 % of the 314 estimated species
(95 % C]=276.1—394.8). The use of databases to develop lists of fish species is not entirely reliable; therefore
taxonomic verification of specimens in collections is essential. ‘The results will facilitate comparisons with other

sub-basins of the Orinoquia, which are not categorized as areas of importance for conservation in Colombia.

Copyright Alexander Urbano-Bonilla et al. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

66 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65—97 (2018)

Resumen

La sub-cuenca del rio Cusiana ha sido designada como una de las areas prioritarias para la conservacién
en la regién del Orinoco en Colombia debido a su alta diversidad de especies. Este estudio presenta una
lista actualizada y una clave de identificacién para los peces del area. Para ello se revisaron los especimenes
depositados en las principales colecciones ictiolégicas colombianas. Se examinaron un total de 2020 lotes de
167 localidades diferentes de la sub-cuenca del rio Cusiana, que oscilaron entre 153 y 2970 m en altitud. El
mayor numero de registros corresponde a la regidn del piedemonte (1091, 54.0%), seguida por los Llanos
(878, 43.5%) y Andina (51, 2.5%). Encontramos 241 especies distribuidas en 9 drdenes, 40 familias y 158
géneros. La riqueza de especies de peces observada (241), representan el 77.7% de las 314 especies estimadas
(IC 95% = 276.1-394.8). El uso de bases de datos para elaborar listas de especies de peces no es del todo
fable, por lo tanto, es esencial la verificacién taxondémica de los especimenes en las colecciones. Estos resul-
tados permitiran evaluar otras sub-cuencas de la Orinoquia, las cuales no estan categorizadas como Areas de

importancia para la conservacién en Colombia.

Keywords

Andean Orinoquia, Casanare, conservation, diversity, Llanos, Piedmont, species richness

Palabras clave

Diversidad, Casanare, Conservacién, Llanos, Orinoquia Andina, Piedemonte, Riqueza de especies

Introduction

The Orinoco River, with an estimated richness of 1002 species of freshwater fishes, is
the second most diverse drainage in the Neotropical region (Reis et al. 2016). None-
theless, the basin has been exposed to increasing threats due to human activities that
place the enormous fish diversity at risk (Barletta et al. 2010, Rybicki and Hanski
2013, Lasso et al. 2016). The systems draining the Andean region (western tributar-
ies of the Orinoco) are considered the most threatened at basin scale (Rodriguez
et al. 2007, Machado-Allison et al. 2010, Lasso et al. 2016). The rivers originating
in the Andes are heavily exposed to threats like habitat fragmentation, contamina-
tion, deforestation, the introduction of non-native species and mining (Machado-
Allison et al. 2010, Anderson and Maldonado-Ocampo 2011, Lasso et al. 2016).
Additionally, large gaps regarding the basic knowledge of fish diversity of the Andean
sub-basins are persistent, especially in Colombia (Maldonado-Ocampo et al. 2008,
Machado-Allison et al. 2010, Lasso et al. 2016). Filling those gaps are essential to
guide adequate conservation efforts for the freshwater ecosystems and therefore face
the threats above mentioned.

The Meta River basin, with headwaters on the Eastern Cordillera of Colombia, is
one of the major tributaries of the Orinoco River (Usma-Oviedo et al. 2016). Studies
on its fish diversity (e.g., Urbano-Bonilla et al. 2009, 2014, Villa-Navarro et al. 2011,
Maldonado-Ocampo et al. 2013, Urbano-Bonilla and Maldonado-Ocampo 2013),
and recent efforts (Zamudio et al. 2008, Urbano-Bonilla et al. 2016) have advanced
our understanding of the ecology of some species. The Cusiana River sub-basin is one
of the best-known Andean tributaries of the Meta River basin; the first inventories of

Fishes of the Cusiana River, with an identification key to its species 67

its fish diversity dated back to the 90’s with the establishment of oil companies in the
area. The Cusiana River sub-basin has been considered as a conservation priority area
for the Orinoco biodiversity due to its high diversity in several groups (Lasso et al.
2010, Trujillo et al. 2011), including fishes (Villa-Navarro et al. 2011).

Here an updated checklist and an identification key are presented for the fishes of
the Cusiana River sub-basin. We hope our results may establish a guideline that can be
replicated in other basins of the Orinoco drainage.

Materials and methods

The Cusiana River sub-basin has an extension of 7324 km? and 271 km in length,
originating at 3800 m asl on the eastern slope of the Eastern Cordillera in the Que-
bradas La Iglesia, Melgarejo, and Las Cafas, Boyaca Department (5°35'N, 72°47'W),
and empties at 150 m asl in the Meta river, Casanare Department (4°31'N, 71°51'W)
(IGAC 1999) (Fig. 1). The Cusiana River sub-basin was divided by altitudinal limits
in three distinctive regions based on Abell et al. (2008): Llanos (139-300 m asl), Pied-
mont (300-1235 m asl) and High Andes (1235-3000 m asl).

The checklist was assembled by examining specimens deposited in Colombian ichthyo-
logical collections. Acronyms used in the text follow Sabaj-Pérez (2016) except uncatalogued
material housed at Fundacién Universitaria del Tropico Americano (UNITROPICO). The
taxonomic list follows the classification system proposed by Reis et al. (2003) with recent
modifications proposed by Oliveira et al. (2011) for characiform families, Betancur-R et al.
(2016) at high-level groups for osteichthyans in general, and Thomaz et al. (2015) for gen-
era of the Stevardiinae. Valid species names were confirmed through queries on the Catalog
of Fishes of the California Academy of Sciences (Eschmeyer et al. 2017). Species were cat-
egorized as endemic (DoNascimiento et al. 2017), threatened (Mojica et al. 2012), migra-
tory (Usma-Oviedo et al. 2013), and species subject of conservation (Gonzalez et al. 2015).

Species richness interpolation and extrapolation was calculated following Chao et
al. (2014) and using the package iNEXT 2.0.12 (Hsieh et al. 2016) for R v.3.4.0 (R
Core Development Team 2017). The number of localities were obtained per Orinoco
basin from the data set of the “Catalogue of the Freshwater Fishes of Colombia” (Do-
Nascimiento et al. 2017).

To construct the key (for order, families and species), original descriptions of spe-
cies, taxonomic revisions, and direct examination of specimens were used. Finally, in
order to share the information produced herein, the dataset was uploaded to SiB Co-
lombia’s data portal (GBIF Colombia Node) in accordance with their protocols for
species lists. For the latter, the complete dataset was structured and standardized to
comply with the international biodiversity standard: Darwin Core standard (Wiec-
zorek et al. 2012). After mounting the dataset on a Darwin Core spreadsheet template,
it was uploaded to SiB Colombia's Integrated Publishing Tool for international visuali-
zation in their data portal. A DOI was provided by SiB Colombia for the shared data-
set available at http://doi.org/10.15472/er3svl, all the results, discussion and quantities
herein cited follow the version 1.8 of the published dataset.

68 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

73°0'0"W 72°0'0"W

5°0'0"N
5°0'0"N

Cusiana river sub-basin

=

Sampling location
© Andean Orinoquia
@ Piedmont
© Llanos

73°0'0"W 72°0'0"W

Figure I. Collection localities in the Cusiana River sub-basin.

Results

In total, 2020 lots from 167 different localities from the Cusiana River sub-basin rang-
ing from 152 to 2970 m asl were examined. Most of the records were found in the
piedmont (1091, 54.0%), followed by the llanos (878, 43.5%) and Andean Orinoquia
(51, 2.5%), suggesting sampling bias in elevation for this drainage, being inversely-
proportional to elevation (Fig. 2). The number of localities in Cusiana River sub-basin
represents the quantile 0.83 among the tributaries of the Orinoco drainage in Colom-
bia (Table 1). 241 species were found distributed in nine orders, 40 families, and 158
genera. The order Characiformes showed the highest richness with 106 species, fol-
lowed by Siluriformes (89), Gymnotiformes (20), and Cichliformes (15). The remain-
ing orders were represented by one to three species. The most speciose families were the
Characidae (54), Loricariidae (30), Cichlidae (15), Heptapteridae (15), Pimelodidae
(11), and Curimatidae (9), while the 34 remaining families were represented by 1 - 8
species. Extrapolation suggests that the expected richness for the Cusiana River sub-
basin is roughly 314 species (95% CI = 276.1—394.8) (Fig. 3); with the observed rich-
ness corresponding to around 77.7% the expected richness.

Concerning threatened species, five are currently categorized as Vulnerable (Pota-
motrygon motoro, Brachyplatystoma vaillantii, Pseudoplatystoma metaense, P orinocoense,

Fishes of the Cusiana River, with an identification key to its species 69

Distribution of sampling records
along the elevational gradient

oS
o _ o
a os |
4 iO _|
> 2 N
o 8H oe
c ise] 5 fay
2 =]
a ey go
= a
Te) So
g S7 Bene
* o
ite)
o ae
is A) * * mt * *
[ T T
0 500 1000 1500 2000 2500 3000 0 500 1000 1500 2000 2500 3000
Elevation (m asl) Elevation (m asl)

Figure 2. Distribution of sampling records along the Elevational gradient in the Cusiana River sub-

basin. Asterisks indicate categories with the lowest sampling along the elevational gradient.

Table |. Number of sampling sites per basin.

Basin Number of sampling locations
Guamal-Humadea oS
Guacavia-Guatiquia-Humea 174
Cusiana 167
Ariari 74
Upia 61
Guayuriba 57
Cravo-Norte 56
Pauto 56
Cravo-Sur 53
Arauca 43
Guayabero ap)
Tua 29

and Zungaro zungaro), and two as Nearly ‘Threatened (Potamotrygon orbignyi and Soru-
bim lima). There are 34 species endemic to the Orinoco drainage, 20 are migratory, and
8 are subjects of conservation. A total of 50 species are new records for the Cusiana Riv-
er sub-basin, while Cetopsorhamdia shermani and Rhamdia muelleri are also new records
for Colombia. Some species from the genera Andinoacara, Astroblepus, Ceratobranchia,
Cetopsorhamdia, Chaetostoma, Characidium, Corydoras, Creagrutus, Curimatopsis, Hy-
postomus, Imparfinis, Microglanis, Parodon, Parotocinclus, Pimelodella, Poecilia, Spatu-
loricaria, and Trichomycterus, require further revision by specialists.

70

Alexander Urbano-Bonilla et al. / ZooKeys 733: 65—97 (2018)

Sampling in the Cusiana basin

S.obs = 241
S.est = 314.5

Richness

) 20000 40000
Number of individuals

Figure 3. Species accumulation curve in the Cusiana River sub-basin. Abbreviations: S.obs = observed richness,

S.est = estimated richness. Continuous line represents interpolation, and dashed line represents extrapolation.

Key to orders, families, and species of fishes of the Cusiana River sub-basin

1

NX

Five pairs-of gills-in. ventral POsitionss..2.cs02y) .bssncuestuvscarpedecetdvacaeoceteanneaiaseaes
Pris Bas A ey A tea MYLIOBATIFORMES: Potamotrygonidae (2 species)
Two lateral gill openings, or just one gill opening under head... 2
Fyes-located Gn the:same:sid€ ot bod yin pew chia caetepunseteni gestae neva eesges
SS GW ne eA, . ree PLEURONECTIFORMES: Achiridae (2 species)
yes located.on Op pOSite Sides OliOU Vis Mein. ee Rar acest neha reteneraitzte: 3
Wiese tnitt a S61 esate eco eg coe eaves peti kcterie dante alec edu creel ocean geneween 4.
Leni Sallelil tye se tite Sha, aan nN Rs tah. tas Rook haar eka Ne Sante ks NS anaes 5
Anal bsentcone yerttal Gill peri Gaia sasnctiouadencanmewacsmonaguahaees
is SYNBRANCHIFORMES: Synbranchidae: Synbranchus marmoratus
Anal fin present, always long, two lateral gill openings... eee eeeeseeeeeeee
ee nee ee eT dis eR GYMNOTIFORMES (20 species)
Body naked or covered with bony plates; barbels present near the mouth.....
pac eaae: Bin lc anand «ot Wine be st Wa be ee STILURIFORMES (90 species)
Body totally or partially covered with scales; barbels absent.............. eee 6

Fishes of the Cusiana River, with an identification key to its species 71

6 Dorsal and anal fins with two or more spines; scales ctenoid.......... cesses 7
— Dorsal and anal fins without spines; scales cycloid (ctenoid in some groups)... 8
7 Leavetal Wirte ta terra p ted se... cee creessecoseeveceeeees CICHLIFORMES (15 species)
- Meare Ce eOMLIATVOLIS 84 baste te Marks rhodes phased: chisel iulduerasuhinsseresetad poeta testins phagh
ainsi achat fastest toner sthtnaiee ACANTHURIFORMES: Sciaenidae (2 species)
8 Mouth superior and somewhat protractile; adipose fin absent........... eee
Wi ud aera eval et estilo entle CYPRINODONTIFORMES (3 species)
us Mouth usually in terminal position, never protractile; adipose fin usually pre-
SEDER cca ce snare th tstos oa wtedTethv onsle th usgavonnieadeeeniens CHARACIFORMES (106 species)
MYLIOBATIFORMES
Potamotrygonidae
1 Dorsum brown with yellow or orange regularly distributed ocelli, ocelli rarely
fused; teeth of both jaws in adult males with cusps on the central axis, teeth
inptemales faccened Sint all LOWS -.5e.03cssassiedenvewacavarsnces Potamotrygon motoro
- Dorsum light brown with black or dark brown spots forming reticulate hex-
agonal lattices, mainly on the interorbital region; teeth small with triangular
slightly monocuspid crowns in males and trapezoid tricuspids in females......
Ueto dea dichanulnbi id lech lamas bhatt uses Lit es duatirbanicaleit at Potamotrygon orbignyi
PLEURONECTIFORMES
Achiridae
1 Pectoral fins present; gill openings wide and extending ventrally on both sides
OPTICAL, eae syd beneed be oi sabalant iabeinds oh aes ec siarebamad este Hypoclinemus mentalis
- Pectoral fins absent; gill openings limited to a narrow slit and never con-
nected ventrally to both sides of head... eee eeeeseerees Apionichthys sauli
GYMNOTIFORMES
1 Caudal finand. dorsal hlaiménr preseity ccoscccecsreeoerancnerceest Apteronotidae
- Caudal fiaranidsdersal ilanient alseint steko. bloc sSehoan tue pean ee taetneetase viaee eels 2
2 Mouth in upper position, body cylindrical... eee eeeeeseeeeeeeee Gymnotidae
= Mouth terminal or subterminal, body compressed ..........::.secesseeseeeseeeteeeseees 3
3 TSetlMPreseries MALS SPELLS EAR ta. c. et valtrata nella san cde ences Sternopygidae
- tleethabsenr anterior tia reso bbl Ula Te iaf.e Benes doseh gales acannieedolenieantercsaete
+: mbsehice of tmetital ACcessorccleetO-Or Gate... swam trvvees ta veentTaseecrnsneencetvecnca ses

Be ERE eer ee eer Hypopomidae: Brachyhypopomus brevirostris

Presence of mental accessory electric organ ........e eee Rhamphichthyidae

72 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)
Apteronotidae
1 Lower jaw with a distinct V-shaped median groove accommodating the
pointed decurved upper jaw.......ceeseeeeeseee Adontosternarchus devenanzii
— Lower jaw without a V-shaped median groove ........sceseeseseeeseeseeseeeeseseeseeees 2
2 Snoutitutbular can. a eo hese. ne vaee wcetabancemente sn tatahsee eats scRonauten. wadhionwaoeteatetyianceeet 3
- Shout obtuse ofclongatebut not tu bilapii tection teiih dua rventep retin rence sues 5
3 Absence of teeth on upper jaw... Platyurosternarchus macrostoma
- Preseree-Gireethy OT tS Pet AW cokecctr sn tscse deseees avsdh Revouen mrevavha dutewnatcasecBaredrs 4
“ Total anal-fin rays 212-242 wee Sternarchorhynchus oxyrhynchus
- otalkanal-finwrays: TO S= 21 Ot As. tsies aesisnatsoiss ensers Sternarchorhynchus roseni
5 Mice dorsalypale set iG-A DSO we Aare beaeae eet wees eters Compsaraia compsa
— Miid-dorsal: pale-stripe presetit.55 eo. cte t pees oe tees eenegee tee tease teens ee 6
6 Presence of two clear bands surrounding caudal peduncle.... Apteronotus albifrons
— Presence of a single clear band surrounding base of caudal peduncle........... 7
P More than 10 scales above lateral line 0. eee Apteronotus galvisi
10 or fewer scales above lateral line... eee Apteronotus bonapartii
Gymnotidae
Without a particular color pattern on body; anal fin confluent with tail .......
ye SP wb ove AWE + 0 Se teene | ie net sa 2: Electrophorus electricus
— Body color pattern formed by dark oblique bands alternating with pale bands;
aLalinUOt CONMivent willy tail. eects. Mae Same. ewe eee, eee 2
2 Obliquely-oriented dark bands or band pairs with straight or high-contrast
margins, 23 dark bands (usually more than 30) .......... Gymnotus cataniapo
- Obliquely-oriented dark bands or band pairs with irregular and wavy margins
By sahs catatsatsaheassnennd user auaeeninastae eutaahsad ca! Gymnotus carapo septentrionalis
Sternopygidae
1 Orbital margin free; all anal-fin rays simple; background color variable from
uniformly gray to black; humeral black blotch present, rarely diffused; white
narrow band under the lateral line, from the midbody to end of anal fin ......
sy Yat ot let pera Ys yale DMS alc Wig one Wn gh ented hat Sternopygus macrurus
- Orbital margin continuous; background color variable from translucent to
whites humeral blotchzabseinte csc cck -ccccsenceoesccdeencties she sdvasayttersd doves snenstesteaee 2
2 Anal fin either completely black or with a black margin; no horizontal dark

stripes on body; scales above lateral line 15-18......... Eigenmannia limbata
Anal fin hyaline; 2 or 3 horizontal dark stripes on body; scales above lateral
TPAGSAEMA TBs esubleteeeiSeneetons hipaa Den ineeasatet inset aeetp taae Eigenmannia sp.

Fishes of the Cusiana River, with an identification key to its species ,

Rhamphichthyidae
1 AL ErLOP Tiates LTSIC GUD DEE MD ccctec crepe cdereeecesie or qoewaleveu vecuterndsreectrenededs 2
— PURbenl Or sta reson es Cl ewl(aprela |1e .-.Aaa-Sevlaeeds ohtcategtdeh eteaer teaser tans toate pice 3
2 Abséncé-of sixthintraorbital BONE sie... cccsisicesansnveeoee Hypopygus lepturus
- Presence of sixth infraorbital bone, as a narrow tube, positioned vertically,
patallel no posterior border Gi eye... au ti:ececerceesdsbenndetnese Hypopygus neblinae
3 Number-of pectoral-tin rays fewer thati IG... os... seshesagereen ovecteotensnantereeentert
{dicate Galen aah oe Ce act du tad <eeronelt Sutoulounttenstoe pote Gymnorhamphichthys hypostomus
— Wumber-of pectoral-fin-rays moré-thaty 1G... este. fe iadioes teas cases cere enieaess A,
4 Atal ‘in ysually.cleat-or Hyalitie s,s s...SssBessewotstes Rhamphichthys apurensis
- Anal fin usually dark with a terminal dark band....Rhbamphichthys rostratus
CYPRINODONTIFORMES
1 Pectoral fin with 1-2 unbranched rays... eee Poeciliidae: Poecilia sp.
- Rectoral tin.with all cays branched... sssascanaunssme atau Cynolebiidae
Cynolebiidae
Scales on the ventral surface of the head «0.0... eee eeeeeeeeeeeee Anablepsoides sp.
- Scales absent on the ventral surface of the head........ Rachovia maculipinnis
SILURIFORMES
1 Mental barbels absent; ventral mouth in the form of a sucking disk with wide
Voy e etl ot Bisa tte cet teens teach tuaheata treaties dash t Anite testes denenh tinea screed 2
- Mental ibarbels presente retrial aT OWEN cnesa.ns idle nepneraortbinlosntnelba gah sndatesbeiltedychal' 3
2 Body tiated cut fica Poe Beers Ue tng be etait Seater Astroblepidae
- Bedy-coverediwithr bottyypla tess iit tectetunsnvateressecatabitectout net Loricariidae
3 Body depressed; skin covered with tubercles and completely keratinized.......
Sabet ete ce ee a cae eae a ge ala acai ed tear ce Aspredinidae
= Body not depressed; skin without. tubercles: .......dtezerseregnteeeagesdeecereseelneteeedse 4
~ Body covered awith-bony plates... accep cnermiwach ware seeneretha spread veesban ceeueeenapsher 5
- Bedy not covered. witht bommys pl ates «2 a-ce5 pa aca nen seapec eh octe Binnie te eaten 6
5 Sides of body covered with two rows of bony plates.............. Callichthyidae
Sides of body with a mid-lateral row of dermal plates armed with a lateral
CIPO TI cater ana ts diet oatns on woettae'cualt tnsteu toune'suston patente Patldtts pane des taatsaie anes tet eats Doradidae
6 Opercle and interopercle with odontodes.............eseeeeee: Trichomycteridae

- Opercle and sitréropercle witheumodoncod es... 20.20. :icteecd saat ch siphian eerboneet z

74 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)
a Datetalsine-system braiiched ot diead ¢ sates insets nce. Pimelodidae
- LateralFline‘systen-simple; notibranched on fiead ... .c22.enssneessueesareneretzes 8
8 Stborbital sults preserite: ouicvecves cecnwenmccveerassuenees cteeviwnceueeys Auchenipteridae
- Su Oi petal “SUM Ct sea USE libe ue setae hi shave Seubdae’s edtcatoatdebtue enue SecazhtMertaoneuhateysgha) 9
2 PPCLTIS OSC ALITA SEIN ates cach attest ve each tate banat ania eee ateaaneaceee aes Cetopsidae
AIDOSe ting SFeEseiit ss 27. Lees. ceeeece ecu Ate ete, BN ene 10
10 Eyes set on anteriormost half of head, thick skin........... Pseudopimelodidae
- Eyes set on posteriormost half of head, thin skin... Heptapteridae
Astroblepidae
1 ACI POse TrMApreseinte«, c.1dukene tana cssouteste se: <susatedeneeaedsteeuheate ed Astroblepus sp.
- HA CLUS OSE HR AISE ITE fet cases cab Yooth, Meaatct Baath Moca te th TN Astroblepus mariae
Loricariidae
| Wail'strorigly depressed «ite. dwcivsinceigavstonns deceranttecesttonsudtetenecustte clin 2
- Matlovalsiictosssectom. pee ee cet tee terete tees 12
2 Dorsal-fin insertion in front of the anal-fin insertion .........eeeeeeeeeeeeeeeeee 3
~ Dorsal-fin insertion anterior to the anal-fin insertion..........eeeeeeeeeseeeeeeeee 5
3 Three rows of abdominal plates... cess eeeeeeeeeeee Farlowella mariaelenae
- ilwe-tows-O Rab de mrtittal to laces. tai. osazion dines oneattone sotabeert ieee ee iets A saeten aus 4
“ Breeding odontodes on preorbital ridge present... Farlowella acus
- Breeding odontodes on preorbital ridge absent ................ Farlowella vittata
5 PES GE Peg Val COT Ege dh assent teae ah oeaaRtean ste dh steal bata cetanet dtedile SMoaes ebvoneed ssemnasbedt aie 6
- Teeth spoon-shaped, elongate, comb-shaped or even absent but never vil-
WEG EI beens sattancetsadsas sada deandcae Mss stsadmereatidadhtentiavsthea debendsastlaea Lens oc yea lonts 7
6 Pectoral snraye: be Lainnss tanwatialenas ameattisheueteunatidens Lamontichthys llanero
- ReGtona Haat ay Sal (OR jntidencatsldtesmt vit sedainbroneaen eadocteeedt Sturisoma tenuirostre
7 Premaxillary teeth larger than dentary teeth.............. Loricaria cataphracta
— Premaxillary teeth equal to or smaller than dentary teeth ........ eee eects 8
8 Snout round in outline; upper lip with long filaments covering mouth open-
TN Srey antcceun dceceewersncgvory scnvovecunwctesieencuet Mecsesexecueten cach Dentectus barbarmatus
= Snout acute to ovoid in outline; filaments covering mouth opening absent.... 9
9 Dentary teeth larger than premaxillary teeth....... Rineloricaria eigenmanni
- Dentary teeth smaller than or equal to premaxillary teeth... eee 10
10 A pair of digitiform papillae on rictal region of mouth and an additional pair
OTTO MCP TOO Le A sacsscatt seis uskicstienan betaltsamnte res asters utah hou Spatuloricaria sp.

Digitiform papillae on rictal region of mouth but absent on its mouth roof..

1]

2A

Zed

23

Fishes of the Cusiana River, with an identification key to its species is)

Anterior margin of abdominal plate cover oval in outline; fringes on upper lip

Gately plemented: veh. les weet hahesadnucdbanenet Loricariichthys brunneus
Anterior margin of abdominal plate cover irregular in outline; fringes on up-
SB Pye ICT CMS AE ici hag nat top abdcies hhoaswunsbdeedsplaablopuases Limatulichthys griseus
SHOURMAKEd whot-covercd (Wikly PlATES. nce: peuase sea nsseatuck ane semrenakenecoenaacet 13
Sriduricoveted av ot hyp la tesa ce. See areas, cee a ec 7

Fleshy tentacles on snout present; three series of lateral plate series on caudal
FOLelC C1 a cl aprgi, Naeh Man rde ce eR ait eh A, mn. Ee RTT RE Aa Ee Ancistrus triradiatus
Fleshy tentacles on snout absent; five series of lateral plate series on caudal
CCU Clee Fats sees cate anette a ee iew td iden dda ates edenee dca Spends Saedea’ peateatseaa tases 14
Cheek odontodes straight; fleshy excrescence on parieto-supraoccipital ab-
SEM ee ahe etoe ae oa Metent erent armed heme reuseteelet Chaetostoma dorsale

No dark spots on any fin but dark stripes present along rays in all fins; en-
larged second unbranched ray in anal fin bearing two fleshy ridges in mature
TIGNES « Arssnat eahcee Mreuns teas Saracen Roden abittsenss Mahan emmesn ea henk Chaetostoma formosae
Dark spots on fins; anal fin never bearing paired fleshy ridges in mature

Wel Ss Se re Rh a oR al ctl cach late RR Ee, Sth a Sorcha 16
Ventral portion of body posterior to pelvic-fin insertion light, never covered with
abundant dark spots; pectoral-fin spine with dark spots............ Chaetostoma sp.
Body uniformly spotted; pectoral-fin spine without spots... Chaetostoma joropo
Posterior serrae on pectoral-fin spine present......... Hypoptopoma machadoi
Posterior stirac on pectoral-fn-spine absent: os.c928 208 eestor dG desdesies 18
Spootieshapedateeth... esis. ca. 5 eoee cere ee es See ee 19
Waillito try Gest le a secsts ches oo Pae elds antics oud ata Meee Abdus asda eat cedcoanme ete 21
(Cheeks OU OMtOC ESA DSEME xe oa utes Picea ee eared Hypostomus plecostomoides
Cheekodontodesipresent anid erectile: fence Ae eee ieee cares 20

Dark background with yellow or white vertical and irregular bands; small
AAUULE SIZE R sash. dene sbourenths tah ooe abate ons umbeneernemaabuhs ups cbettaelnars Panaqolus maccus
Color pattern consisting of horizontal stripes; large adult size... eee eee
Nee Nadas Sagas Neat oes ren bea tn ealden oe on etna cere at case Saad dae Panaque nigrolineatus
Eyes visible in ventral view; adipose fin absent ............::ssseeeceeseeeeeeeeeeeeeeeee 22
Eyes not visible in ventral view; adipose fin present...........escesseeeeeeeeeeeeeees 23
Twenty-six or more premaxillary teeth, 21 or more mandibular teeth; teeth
slender, tightly spaced along the jaw rami, spacing between teeth equal to or
greater than the tooth shaft width, tooth cusps small... Otocinclus huaorani
Fewer than 25 premaxillary teeth, 20 mandibular teeth; teeth robust, spacing
between teeth equal to or greater than tooth shaft width, tooth cusps robust......
Peanut saan ea nrennnane Thnaaseeusescccaes enee eee vesnstsdiee Otocinclus vittatus
Coracoid bearing odontodes in ventral surface...............04 Parotocinclus sp.
Coracoid covered with skin and not bearing odontodes ......... cee eeeeeeeeeee 24

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Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

Setiform odontodes on sides of head; three series of lateral plates on caudal
[Sor [U's Yell kamera en eee Ma eee A regard. Lasiancistrus tentaculatus
No setiform odontodes on sides of head; five series of lateral plates on caudal
VDler (ETE tel raely SOV A ewan mer 1 RSPR ER PHAROS Ate, MOR NT Sone oMtCeL A Ue 25
Hypertrophied cheek odontodes surpassing base of pectoral fin; pectoral-fin
spine reaching or surpassing tip of pelvic-fin leading ray... eee eeeeeees
Pret RAE stg atin Ath APR L HEN nas Mas resi nas Raey a Dolichancistrus fuesslii
No hypertrophied cheek odontodes; pectoral-fin spine not reaching tip of
DelviC-hir lea dintestaye St vcac dxounnsteeuces th suctesPecbees se venteyPceaet meu ter vateunatee Eade 26
Multiplebuecali papillae: code cecesbecasetate rates Aphanotorulus ammophilus
Sinelemmecialuptiacal papilla .t; Maatevucantss Mani tosectes ests ts tte tants oatio, WSs 27:
More than 11 branched dorsal-fin rays........ Pterygoplichthys multiradiatus
Kress than Tl" branched dorsalsfnt raysscc eect cece sec seperate ee eee 28
Lower caudal-fin lobe dark and upper lobe light, no spots on caudal-fin; cau-
dal peduncle Slon Gated dete ccs ducpewteeaxerescos eeeenten Aphanotorulus emarginatus
Spots on caudal fin, no distinct dark background on lower caudal-fin lobe;
caudal wedunclesnos clone te dss.ais cuss Salleh venenta cv can csanantas necaveaansenhsnatanea des 29
Dark blotches closely-set on dorsal fin wees Hypostomus niceforoi
Dark spots arranged in two longitudinal series on interradial membranes, no
blotches on dorsal fin.............. Hypostomus argus (please note that these dif-
ferences in coloration are preliminary observations based on limited samples
along with type specimens, further and more extensive sampling will be re-
quired in order to test such differences between Hypostomus argus and H. nice-
foroi in the Orinoco basin of Colombia; G.A. Ballen & S. Reinales, in prep.)

Aspredinidae

1

Dorsal, ventral and lateral series of bony plates present on body; mouth infe-
rior; hard pectoral spine, without serrae on the anterior margin...
Pry eevee Wrest eee eee ree Ue ee eS. Hoplomyzon sexpapilostoma
No obvious bony plates on body; mouth terminal; hard pectoral spine serrate
on anterior and posterior Margins ........ cesses eeeeeee Bunocephalus aloikae

Callichthyidae

1

Snout depressed; maxillary barbel long, usually extending beyond gill open-
[fa Ba ar Facr Ar ee AT CENTLY RNC as rea Le, Ane tt AR eee IR 2
Snout compressed; maxillary barbel short, usually not extending beyond eye... 3
Caudal fin forked, without conspicuous dark bands; dorsal fin spine about
half the length of the first branched rays; six branched anal-fin rays..............
Ls Whe shes Meet ets RM Reh aE NRetE nai Rs LATS Hoplosternum littorale

Fishes of the Cusiana River, with an identification key to its species Fi

Caudal fin truncated, with dark transverse band and dark distal margin;

dorsal-fin spine long (52-64% dorsal-fin base length); four or five branched

ANTE IPAS: ce vouduss evecasu overuse vetonn eibicuetenenyeteuecjetens Aiezwene vedas Megalechis picta
3 Dorsal wWwathivs: arb Only tay sdarwarstiedsekacoroabagtongnbessenaat Corydoras sp.
- Darsalyinswitheutrdar kup lo te hes a-.521 cvs oscaich esr canes ones aapese threes btagreabeoda sae 4
+: A dark oblique stripe on body runs across dorsolateral body scutes to caudal-
fin base. No vertical bars on caudal peduncle or on caudal fin... eee
Raed Netix cesar tect ceca aS Maton Mba uct: Corydoras axelrodi
- A large dark blotch on the sides extending from slightly in front of dorsal fin to
nearly adipose fin, tapering off posteriorly. A vertical bar in the caudal pedun-
cle. Five to seven vertical bars on the caudal fin...... Corydoras septentrionalis
Doradidae
1 Meascillary batbel: branched vc. ieco- veneered nccoseracescceencssees Leptodoras nelsoni
— Maxillary. barbel-stiple prret-tamni fred: 5...2te.1) seus caterenhane sp detepa seasons siete! 2
2 Anterior and posterior dorsal-fin serrae absent................ Amblydoras affinis
- Anterior, posterior, or both dorsal-fin serrae present........sceeeseseeeseeseereeeees 3
3 Procurrent caudal-fin rays modified in fulcra; dark background with clear
lonenturdinal stripes i le sige wicca ae btanbaynintunpecenticee Platydoras armatulus
— Procurrent caudal-fin rays not modified in fulcra; color pattern without a
Clear n Stu ClinVal SERUIOC. fi. lovers yluceliy phaneds ot wnla ohicsels ohdshtogt alltteyteb is phakdaatiog tut ghnt 4.
= Thick lips, premaxillary and dentary teeth absent............ Oxydoras sifontesi
-. Thin lips, premaxillary and dentary teeth present..........cseesesessseseeeeeeeeceeees 5
5 sitiall Body, caudal iran cate: .sicscasevsccbetopsonscnsa tenes Amblydoras bolivarensis
— Large body, caudal emarginate with two distinct lobes...... Pterodoras rivasi
Trichomycteridae
1 Nasal barbel present (associated with anterior nostril), mouth subterminal.... 2
- INasalibatbelabsenex momenta vemtiall teat 4 ssee.bs sen sserebaeeh Miser ce beats cutspyate te a
2: Sides of body with a dark band or row of spots from just above the gill-opening
to the base of the upper caudal-fin rays.............. Trichomycterus dorsostriatus
- Sidestor DOdyUMik@ LAAT Y PIG NASMLCE A nc, sh skaters SiacePyabade von danrsengede soe ehdernapbaueen ple 3
3 Eight branched pectoral-fin rays (rarely nine rays); a single median epiphyseal
pore (rarely two asymmetrical pores)........eeseeeeeeeeeeeeeees Trichomycterus sp.
- Seven branched pectoral-fin rays (rarely six rays); a pair of symmetrical epi-
PE SCAl HOLES apie Bt ousiadlie a sontabueedieaica ath Pesiniostnbe st coved Trichomycterus migrans
+: Lower jaw rami not articulated medially, only joined by soft tissue...............

sa ican ate Med el ar oale OE a dane tid oh Sire PAN eA Dito Os Set aD aout PGR Vandellia beccarii

78

Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

Caudal peduncle with numerous procurrent caudal-fin rays, embedded in
wide dorsal and ventral membranes continuous with the caudal fin, caudal
fi POURed: Ot thUNCAted sec cccnciveceusvecneeestoveves ont Ochmacanthus alternus
Caudal peduncle with few procurrent caudal-fin rays, restricted to the origin
of the caudal fin, and lacking wide dorsal and ventral membranes continuous
with the caudal fin, caudal fin emarginated...........0... Schultzichthys bondi

Cetopsidae

1

First ray of pectoral and dorsal fins spinous, lateral line extending to over the
posterior portion of the base of the anal fin but falling short of the caudal
peauaticle.. Sore erates wsomicr etter tavern eeeuiiee: Cetopsidium pemon
First ray of pectoral and dorsal fins not spinous, lateral line either shorter or
extending posteriorly at least onto the posterior portion of the caudal pedun-
TCLS sce ten haste ds aegis ee ccwee ot gene eae eee acseaen ees earned intue te? Cetopsis orinoco

Heptapteridae

Eye margin free, first dorsal and pectoral-fin rays with a pungent spine.......2
Eye margin not free, first dorsal and pectoral-fin rays soft and segmented (at
Mase IGG 1 EAU NAL) RBA Fc 8 oe Sea ac Ole heed aces oR seb Saaaetu ona ne seoetae greece oat: oft 6
Occipital process in contact with the nuchal plate, posterior cranial fontanelle
OPW ovine enecetoheo asd eclwc¥ she's uve tonne thee ev'eaetinticghiveslsn tees eae o te ogee ndeene'eestptaetnnetrkie pitt 3
Occipital process not reaching posteriorly the nuchal plate, posterior cranial
fopitaric WetcloseCNor me CUse Gana cence. sehen teeth cuss Ase castns ok eceeb rare ate 5
Adipose-fin equals to one third of SL or shorter.............. Pimelodella metae
Adipose-tin, longer than: otie thirdiof Se viiz..astsecciceseacsoscsesetsabishiecdanbandiaeis 4
Upper caudal-fin lobe narrower and shorter than the lower caudal-fin lobe...
Eeteeh titteech abled tir bid cone atanahy dsdnch tou eck teachaboee ah ina datnlea aes Pimelodella cristata
Upper caudal-fin lobe about same width than lower lobe and distinctively
WTI C Ed ascluacuia eictsaee Mahalttacsettaane bathe eae beac Suettenic toon te enta's Pimelodella gracilis
Upper caudal-fin lobe distinctively shorter than the upper lobe; if undam-
aged, maxillary barbel surpassing the caudal peduncle..... Rhamdia muelleri
Caudal-fin lobes subequal, maxillary barbel not surpassing the caudal pedun-
CE snasans esuctornmurs saa tenance ebtunaiery enna asics erased mote es Rhamdia quelen
Lower caudal-fin lobe longer than upper lobe .... Phenacorhamdia taphorni
Lower caudal-fin lobe equal or shorter than upper lobe... eee eeeeeteeeeeees ff
Lateral-line canal incomplete, extending only to dorsal-fin base............. eee
Hrecastestealenuten tus tees Hatten tuatea hh eualt Sa vous ettsustoveatendterPatdasts datectn ty Nemuroglanis mariai
Lateral-line canal complete, extending to caudal-fin base... eee eee 8

10

14

14

Fishes of the Cusiana River, with an identification key to its species 72

Lateral-line canal posteriorly interrupted as a series of short canal segments..
ett tae Ue dan attatl ys uaul Geant rnetertae hata ut et Manabe Imparfinis microps
WarétalMiiie 0 ntl UO US: cares Srrepeeee coene onsite ete den cece cengeneeieeeeneestaest 9
PNCIP Os CaM PLCCHATIOUIAL te bath rsshad tas chimes tdtede ch suid naoneins.otaiius camer decane 10
ERGO SE? ME SERL AMAIA» 3hs5 vonesbipGanlestigrb soa Son see saagten boororh-ne eoRITE aay: aaulbeTat ees 11
Anal-fin origin at vertical through adipose-fin origin, adipose-fin longer than
one fourth of SL, six pale bars across dorsum and sides of body.............:e10+
Aerts 8 eit SRE UO LER Pe a a ees, A Chasmocranus rosae
Anal-fin origin anterior to adipose-fin origin, adipose-fin shorter than one
fourth of SL, four pale bars across dorsum and sides of body ..........ceeeeeeeees
SLRS I TE, UBIO |.) RUE SORT Cetopsorhamdia orinoco
First dorsal and pectoral-fin ray extended as a long filament, maxillary barbel
surpassing anal-fin base... eeceseeseeeeeseeeees Imparfinis pseudonemacheir
First dorsal and pectoral-fin ray not extended as a long filament, maxillary
barBel ot reaching analAisOriGiy nine, eco vaa renee tr ases.cathoned ensncatenebe om svegtied 12
Blotched pattern of body pigmentation... eee eeeeceseeseeereeseeeeeseeaeeeees 13
UnitormedarkiparternyGh pIsrientaniOn, vost cossvs.s Sess coencatasecsrosecsennarebepeaeat 14
Pelvic-fin origin approximately at vertical through dorsal-fin origin, adipose-
fin base extending far posteriorly beyond distal end of anal-fin ....... ee
Ty Pavers Arca ean rh RRC Mile Ps Undescribed genus and species
Pelvic-fin origin slightly behind mid-length of dorsal-fin base, adipose-fin base
ending around same level of distal end of anal fin..... Cetopsorhamdia shermani
Snout conical and conspicuously projected beyond mouth opening, maxil-
lary barbel not surpassing distal end of pectoral fin, dorsal and ventral mar-
gins of end of caudal peduncle with a conspicuous white area, caudal fin clear
contrasting with dark caudal fin base... Cetopsorhamdia aff. picklet
Snout depressed, not projected conspicuously beyond mouth opening, maxil-
lary barbel surpassing distal end of pectoral fin, dorsal and ventral margins
of end of caudal peduncle dark, pigmented as remaining of body, caudal fin
dusky, not contrasting with caudal fin base wo... eee eee Imparfinis sp.

Pseudopimelodidae

1

Lateral line incomplete; premaxillary tooth patch without posterolateral pro-
JSC HOM SS INANS Fes. Getto ket te tan tan Maw tines Sha OBER etic oe Maltuepie eas ek 2
Lateral line complete; premaxillary tooth patch with posterolateral projec-
CLONES AL OS SIZE 2% edt woentgr Ase se Maulana ovine eetlanihgeiralir a tiaahg helene sali cele Landon ote Saontnee 3
Bar-shaped blotch on caudal-fin base; lateral line reaching a vertical through
ACU OSC PIT OTIC. i.2tuka éaseaetosiodecaica bobirestiia tue intaunicahietecad Microglanis theringi
Triangle-shaped blotch on caudal-fin base; lateral line reaching middle of
GGr sale tease .s sess csvset eae Lae bande tanhe kdadaeedseeheweus Microglanis poecilus

80

Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

Caudal fin lanceolate; pectoral-fin spine covered with thin skin; anterior nares
On theanareinvol hemor ain Slabs dereaed sans Cephalosilurus apurensis
Caudal fin forked; pectoral-fin spine covered with thick skin; anterior nares
behind the maxillar barbel insertion...............0... Pseudopimelodus bufonius

Pimelodidae

10

Upper jaw projected well beyond lower jaw with premaxillary tooth patch
EXP OSC ecks crates Seen act seeasas Seas Rees arate nee anaes Sorubim lima
Lower jaw just slightly or not projected at all, premaxillary tooth plate not
PECSN Ge) (oh el wales CLOLSoLe coors Reon IRN =A RORY. Ses Puen Tr, TER ENe SIE oan 2
Upper portion of caudal-fin base with well-defined dark blotch..........0...... 3
Caudal-fin base without well-defined dark blotch ....... cece eeeeeseeereeeeeeeee <
Leading pectoral-fin ray soft and not pungent or spinoUS.......... cece eeeeeees
i iracovsvcudecum Ma ccaseucenemuiveccestans coat chien cecatetce iad Megalonema platycephalum
Leading pectoral-fin ray strong, pungent, with anterior and posterior serrae .
ova a heater ivan api san ves saa AoaRe oer eokwn side nLmcnaae cae tape ena Platysilurus mucosus
Eye visible in ventral view, set below midline of head in lateral view; barbels
Hata WiGde aes. 2c eee et oem eae dete eee a See Hypophthalmus edentatus
Eye not visible in ventral view, set above midline in lateral view; barbels round
HEE CLOSS-SECLLONN, eines vererr Pa en'ett fe wn orev eu'eten sno ectev Pa estes th ectev'ds te Tgy ee egal Pace edlpentivestentew eaten’ 5
Body’color pattern consisting of dark vertical Dats.s.c..c0i0Reecchieediijereonseten oto: 6
Body color pattern consisting of dark spots or plain coloration, never with
VERTICAL DAS. A558 vais nla steerable ucte a atien ret ected abistenseel tothe toe saa anc gh te sesatesicBOe tla 7
Abundant (more than 50) spots on caudal-fin.... Pseudoplatystoma metaense
Few (less than 50) spots on caudal-fin.......... Pseudoplatystoma orinocoense
Body color pattern consisting of numerous spots or blotches, sometimes fuss-
ing so that a vermicular pattern appears; interorbital region flat... 8
Body color plain or striped, never with spots or blotches; interorbital region
CONVEX. ods atpeneas vedhraibaea venciscerneon stapes este benasiet peels thee eesey mec ehbaeesmmumessndetennnett 9
Blotches on body; round vomerine tooth patches separated medially as two
Ina ePeticleiie MINT SH Naa AY 1k LS Nei ee ah, Rist, Leiarius marmoratus
Spots on body; laterally-elongated vomerine tooth patches fused medially as
a single wide band, sometimes the medial contact between patches visible....
gb elias onus BE ashore a hos ta oi hleeak ut obs ye ehue se MMR e wits teats Zungaro zungaro
Each caudal-fin lobe with dark longitudinal dark stripe from base to tip of
caudal fin; distinct blotch on dorsal fin... eee eee eee Pimelodus ornatus
Nordariemiarke-ontcatdal sts. stuns Meretennater a sitensusotert ements rarer ational 10
Giant in adult size; eye small, its diameter much more than three times in
dorsal-fin base length; presence of a patch of teeth on roof of mouth............
s AURA Tae ue U End SS Rd SA a hs Se A SE Brachyplatystoma vaillantii
Small in adult size; eye large, its diameter about three times in dorsal-fin base
length, absence of a patch of teeth on roof of mouth ....... Pimelodus blochii

Fishes of the Cusiana River, with an identification key to its species 81

Auchenipteridae
1 Mental barbelsta bsen tyra erences anes onlun ho nuevanrnue menue aed nicneceeures 2
— Art eam ADAG Sn [Re SER Sere Aas ha ad eedees ob cate widgh tae anta Deer eth hen Pac shou bueedde 3
2 Waudaletinidecply dorked, |. 2.05.) ccnsmsukendiqancapbinalehsbagerd Ageneoisus ucayalensis
- GE ati all iit e NCA n8 anihetis wnentetencpste emo sets ictatauceee nnedtelee rcs Ageneiosus magoi
3 7 OPA a TAT AGS ota Rh oe cos dete rats reads caler te bc. Centromochlus romani
- Sl Oe AA LUN RAY G Aee rc Bekelen Metuce oe aac te Saat eeees ctSaawet utube Meenonet ade due ceeetetee te ote 4
4 Dorsal-fin spine shorter than pectoral-fin spine... Pseudepapterus hasemani
— Dorsal-fin spine longer than or equal to pectoral-fin spine... eee 5
5 Caudal fin-obliquely: truncated :s..0.:.0-.ndssssecseensesets. Trachelyopterus galeatus
_ BE Tira bles sh'g 10) a Care BAP ya Oem Un ERR Rar Ure rue Entomocorus gameroi
CHARACIFORMES
i Webi without-teethrss, 5.200. foseescwss, -kephate ania tees aananeh huang: Curimatidae
- Iiowieh wit i Reet lis 2.e usa e, toss setter a crwnsiehurueerneecieas teak tahest oes he 2
2 Wahine Teeth PreSei Gene ekieen ares inenduanceen ete Ronee ret needed 3
- GaiilcecctisADceEMgee ttt et em tet eet. eto: 6
ao) Adiposedin.absent; :catidal fOr ded vs scsatecesanietinnie elttes teeter cee Erythrinidae
- Adipese fingpresent; caudal Tino trou ded eee sar! Meena vente ecreontegaeePerecees &
- Dentary canine teeth hypertrophied; caudal fin truncated; rostrum shotrt.....
Lit Ae RN AR TR Ae oth A Wh PA AA AO Nes hy RL Ctenoluciidae
- Dentary without hypertrophied canine teeth; caudal fin bifurcated; rostrum
PAO ct sascdtlecrad raphe asl esaan Gases cSusaaateca al Teanabescash engt ndteda an'snede des da sedtts haven Peas ts 5
5 ISGP cial al 0 63 (a Rr ORD A a A XD er Ctenolucidae
- SASS CIC OTC ee. t see fet vhestisestces ie covasertemmn iby etrairteneeg rere seat Acestrorhynchidae
6 Upper jaw without teeth; upper lip with teeth ...... eee ee eeeeeeeeeseeeeeeeseeeneees 7
- Upper jaw with-teeths uppenilip without teeth ..5..etenentva-aeascucene tions seteees 8
7. Prédorsal process present; tiOurheventu ble. ct at ace dessethid dene ase eranesinmeeeresreseers
fA hcsaher dude inevntt Misoes tages Maden ester Prochilodontidae: Prochilodus mariae
- Predorsal process absent; mouth non-evertible........... ce Chilodontidae
8 Abdoniinal-serraredykeelpresen tli 8 7 sb.ainencseerteealeceebtepeads tee Serrasalmidae
- ADOT Ser lated Ke lA SCL te ax oreuav er ceeeessuu en sacey erence secueetenedeer ete eameeces 5
wy Branchiallmembranetused tosis, 22,0... th jen coadesstoe eegn bedeege= Anostomidae
- Branclrialumembratie Aree frond sthan sce ust veer seevote qecteveesie cept cuchs 10
10 Pectoral fins strongly developed and horizontally oriented «0.0... 11
- Pectoraltiiastyverticallysontcinted, cc. caterer cer cduneem st eecrucne eteeeer meee 12
11 Fewer than three unbranched rays in pectoral fin... eee Parodontidae
— Three unbranched tays ine pectoral itis. 2. ot. soto Peieeed vee cee Crenuchidae
12 Chest extremely compressed and expanded, forming a ventral keel........... 13
- @hesrnenexpanded te forma -velittal Keel ......2s..0: hss eserdcacbebrinntadeanteeeied’ 14

82

Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

Anal-fin origin anterior to dorsal-fin origin; ventral keel well developed........
Sree a tu eon cre cpt ote Wt Gasteropelecidae: Thoracocharax stellatus
Anal-fin origin posterior to dorsal-fin origin; ventral keel moderately devel-
OPCs oti these th ahaubee tuk fa baka Setu iy gharvees tai Ts pied slash topt alla sedate: Triportheidae
Upper lobe of caudal fin longer than lower lobe... Lebiasinidae
Bet lobes of-caudal. tin of equal size 8h ee eee Dh ee ns 15
Two kowsiotsteeth, iieth es dentary s,s .Jacereccteaiedrundecdue slerweiers. Bryconidae
Oe Fowrolstecth Tn-Centatiye-oroe, cased ee care reach eA seeccah tei al seceteie! 16
Anterior margin of maxilla greatly arched above and meeting premaxilla bor-
AGrat FeMt An eletss cca sutssia.ccsdeeis Iguanodectidae: Bryconops giacopinni
Anterior margin of maxilla not greatly arched above and not meeting premax-

HA BOTASR ALT OMAN ONS ipa waioadivneswntinhnnathwehtnienihh nem etam Characidae

Curimatidae

Lateral line incomplete, with only few pored scales............ Curimatopsis sp.
Lateral line complete with all or almost all lateral scales with pores............. 2
Caudal fin with scales reaching two thirds in length in both lobes ................
nae SRO Rr ha Scr cle ina Ey een arr ear Curimatella immaculata
Caudal fin with scales only on the base of caudal-fin rays.........scesseeseeeeeees 3
Lateral line with: 42 otsmore: Sales toon enters Bt eyed aetieee Mev ce pees eeeceet 4
Watetall litle withino Lakme Werrseales, 7085. 5032) b. ils titaeh bade tt acct tug Abbe ube OAs 8
Lateral-line'scales aiore than Oiiss...etatoescens Potamorhina altamazonica
Iesteralclinesseales: 4 2=6 05 Aes sete. ons. co eee ee ac oe ee 5
Mouth terminal; dorsal fin without dark spot .........ceeeeseeseeeseeeseeeeeeeeeeeeeees 6
Mouth inferior; dorsal fin with or without dark spot .........eeseseeecereeeeeeeeeee 7
Interorbital width 53-58% of HL; anal-fin rays iii,10-iii,12.; abdominal re-
gion, pectoral, pelvic and anal fins with red coloration in live specimens.......
Lertnaphfaecncel Waban Uae ssuat st oheepat sniat ens oaputtaatiy dies SouttagteeSutevateSaae Curimata cerasina
Interorbital width less than 52% of HL. Anal-fin rays iii,8-iii,9; abdominal
region, pectoral, pelvic and anal fins without red coloration in live speci-
BEVO TA Sap Sale sess cd farsa pe mee Naper ath ts welsh Ba tae Rr deat Psectrogaster ciliata
Dark spot on the dorsal fin; prepelvic region of body not flattened or only
obtusely flattened; 1 to 3 scales between posterior border of anus and anal-fin
COLIC UT g ep hake lcedoe held startet ein eSB tos ates bln Oe Steindachnerina pupula
No dark spot on the dorsal fin; prepelvic region of body distinctly flattened;
5 to 7 scales between posterior border of anus and anal-fin origin............0...
Ey ORR rer tr Dee rot echo UPN Se Oy. Steindachnerina hypostoma
Without a dark spot on the basal portion of the middle rays of the dorsal fin...
ssh tenance cantar tansy Su ea ea ag te otc Tau ech chaise toete gee eats Cyphocharax spilurus
With a dark spot on the basal portion of the middle rays of the dorsal fin.....
Ser Ts ERE een ADs Rie ens esbLER Cee Salts ENSROMR a ice IMME chit Steindachnerina argentea

Fishes of the Cusiana River, with an identification key to its species 83

Erythrinidae

1 Maxilla with 2—3 canine teeth; caudal fin usually spotted; dark lateral stripe
AD SEIT CMS E sos teh habe ae thas eae hsethaeeeatash rus Hoplias malabaricus

x Maxilla without canine teeth; caudal fin not spotted; dark lateral stripe very
WG AMAT KEG 25. se ene sot pawn entcaananceuattantire tasted A ehet Hoplerythrinus unitaeniatus

Cynodontidae

1 Dorsal-fin origin anterior to anal-fin origin... Hydrolycus armatus

- Dorsal-fin origin in line with anal-fin origin «0.0... Raphiodon vulpinus

Ctenoluciidae

1 In adults, all lateral-line scales perforated (82 or more); body without round
spots, except one at: the base.of caudal fin ..isscevseueas Boulengerella cuvieri

— In adults, only up to 25 lateral-line scales perforated; body with black spots.
ww tnk in ENS dro Arenson MEN ty he A bth te AP Boulengerella maculata

Acestrorhynchidae

1 Lateral line complete with 99-131 scales; dorsal-fin origin anterior to the
atral EA ZOP IGT tess te inets eaceigieotis ds anentcabnrnecauaeressr eines Acestrorhynchus microlepis

= Lateral line incomplete with 33-37 lateral-line scales; dorsal-fin origin poste-
rior to-theanal-iny ofl ettt soli sedsusroweethbevtesnest Gnathocharax steindachneri

Chilodontidae

1 10 or 11 branched anal-fin rays; anal-fin margin convex or straight; mouth

terminal or slightly superior, dorsal fin with a series of dark spots on posterior
rays; posterior margin of scales smooth ....... cece Chilodus punctatus
= 6 to 8 branched anal-fin rays; anal-fin margin somewhat concave in vast
majority of specimens; mouth subterminal; dorsal fin with dark pigmenta-
tion across distal portions of anterior rays but lacking dark spots on remain-
ing portions of fin; posterior margin of scales somewhat serrate..............044
catch bathed in ant athena lata mane MeaadMee aust Caenotropus labyrinthicus

84

Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

Serrasalmidae

Adipose-fin base long, its length longer than the distance between posterior
margin of dorsal-fin base to the adipose-fin origin ......... Metynnis argenteus
Adipose-fin base short, its length shorter than the distance between posterior
margin of dorsal-fin base to the adipose-fin Origin 1... eeeeeeseeseeeeeeeeeeeee 2
Two rows of teeth on the premaxilla 0. Mylossoma duriventre
One towroisteeth Giathepretrasll la . 25 shee uch eee A a cactsenn dace niet eta aee cate 3
Base of dentary teeth circular and separated from adjacent ones; lower jaw prom-
inent and projecting greatly forward from upper jaw.............. Catoprion mento
Base of dentary teeth flattened, in contact or very close to adjacent ones;
lower jaw-equal:or slightly antérior-to: Wp Per Jaw reivssievvede soteriinrsonsiebsicernbsin 4
29 or fewer anal-fin rays; snout, head, jaws, and body short and robust; dorsal
profile of head to posterior margin of eyes convex; prepelvic and abdominal
region red in life; black and conspicuous humeral blotch posterior to oper-
TSN sows ar cement enc can seve esc euh en aedan eu tes beg ce eatig caveats pivevubanes Pygocentrus cariba
30 or more anal-fin rays; snout, head, jaws, and body long and slender; dorsal
profile of head to posterior margin of eyes concave; prepelvic and abdominal
region ted only-durtng breeding SeasOt® «ec toctch ides Maes tal etree ees 5
Base of posterior premaxillary tooth (6th) approximately equal to adjacent
one (5th); ectopterygoid teeth unicuspid, generally 5 or fewer wo... cee eee
Pisosted tai ute gu et Reateatay on ous Pec'duct he voucguen deste qoutes Males toads Pristobrycon striolatus
Base of posterior premaxillary tooth (6th) wider than adjacent one (5"); ec-
topterygoid teeth bicuspid or tricuspid, generally 7 or more «0.0.0... esses 6
Posterior margin of caudal fin hyaline, with black vertical bar at base of cau-
Chal Fits Sie Se ice teary dock bet tase crete Me ae oo Serrasalmus irritans
Caudal fin completely black or with its posterior margin black... 7
Body width 1.7 or less in SL; lateral spots vertically elongated «0.0.0.0...
pista nucche tae outta Mes aacsauea cad Shsiesaheaaeih ded hon ha aU Meus ehS seoobas Serrasalmus altuvei

Anostomidae

Caudal fin with scales only on its base and without dark bars ............eeeee 2
Caudal fin with scales at least over two-thirds of both lobes. Caudal-fin with
ODitequercdatk Dans Fe. heMa 7 hho Ml elt hte hoee ae Leporellus vittatus
Dentary teeth tricuspid to pentacuspid; 7 branched anal-fin rays.............0.
A NOT fe nc RY PR en Sea ee Schizodon scotorhabdotus
Dentary teeth incisiform or bicuspid; 8 or more branched anal-fin rays...... 3
Body with seven dark vertical bars, the second with a “Y” shape............eeeee
seats sed 214s ca olaheg 2. sues tee JB te eta ed ic Rane gas A aad Leporinus y-ophorus

Fishes of the Cusiana River, with an identification key to its species 85

=: Body with 4—5 longitudinal stripes separated by white or yellow stripes........
seilbatns dees tanaladata saldaleandtetiat antral rhe eos eateaticen DARD oi guste uaredreinalaa eau Leporinus striatus

— Body with two rounded black spots at midline ......... Leporinus gt. friderici

Parodontidae

1 Dark vertical bars present; dark lateral stripe absent........ Parodon apolinari

- Dark vertical bars absent; dark lateral stripe present..... Parodon aff. suborbitalis

Crenuchidae
1 Chest naked without scales between pectoral fins... Characidium gr. boavistae
- Chest covered with scales between pectoral fins 2.0... eeeeeeeeseeeeeeeeeeeeeeseeeeee 2
2 Body pigmentation composed by irregular lines and dots... Characidium pteroides
- Body pigmentation composed by well-defined vertical bars 0... eee 3
3 Svordewer biahched Pectoral Lay 9? cevsm sic sieicevenerretha sermwety tesbs aneveneaeeahat 2
— LO%oraitére branched: pectoral -finttay 8 nubeciedc he earner enter Pees 5
4 All vertical bars always originating on dorsum; body depth 23% of HL or
VSB tian alata te anit at atlettiaiee a saltataea tot Seat etDeanicaseitiet Characidium gt. zebra
- Not all vertical bars originating on dorsum, thin vertical bars not reaching the
dorsum between vertical bars originating on dorsum; body depth 24% of HL
ST ESUNCG) ¢o tN AR RTT a Ann SR AeA End A, OO or oA NO Characidium sp.
5 Mid-lateral stripe diffuse or absent; vertical bars fragmented in rhomboidal
GAGE Saree Fes ret at gs eR Tad aac ate ie ds anne Characidium cf. steindachneri
- Mid-lateral stripe very well developed; vertical bars continuous and with dots
appearance at junction with mid-lateral line ................. Characidium chupa
Triportheidae
1 Pectoral fin not surpassing ventral-fin origin.......... Engraulisoma taeniatum
Pectoral fin surpassing yenttal fOr iti.. csstaeseeobtevetseeteeerseretshetrareteneeubes 2
By 24-27 branched anal-fin rays; 5—6 scale rows between lateral line and dorsal-
PVC TCU Bah toe hart ohat tat pasty de her Ya ods ete heey) Sbload te Triportheus venezuelensis
Ds 28-32 branched anal-fin rays; 7 scale rows between lateral line and dorsal-fin
OAH Aa ce Uoeianehleaaniercnnttntta col Mt oats aa sg antedy sensctgnishileaaSeste Triportheus orinocensis
Lebiasinidae
1 IACIPIOSE TEN TESCIME i510. 2. ieenadtos Bscpian neesitse Mesninateste et Lebiasina erythrinoides

= GNGUIS CG) SESIN UN AIS OT CLE: seals ooh Fe etna at Ae Seabees ache a tbat ara dratn staat Ae etait eet ae 2

86 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)
2 One row of teeth in the premaxilla; maxilla S curved; dark lateral stripe com-
plete and well-marked (from mouth to caudal fin)............... Copella arnoldi
= Two rows of teeth on the premaxilla; maxilla not S curved; dark lateral stripe in-
complete and thin (from mouth to few scales after head)......Pyrrhulina lugubris
Bryconidae
i (Fwo-rows-olaeethnintale-premtasdlla? .. j.iccd. totes ser Soepec eos aeteone eee Salminus sp.
Z Thréeirows:et réeth-tm thes prema sills... tos..scezecsucsaceas camel vastegeeteag-e<stetoseess 2
2 Dark lateral stripe present, from opercle to tip of middle caudal-fin rays; up-
pemcaridalhndlobe aan pigmented |i cesnsenscegh eden snponenapturtornnie Brycon whitei
— Dark lateral stripe absent; upper caudal-fin lobe pigmented.... Brycon falcatus
Characidae
1 Rareralitire tion plete sp. 2.8. hen sedate tue: Mooctalate werk srestes corm 2
— Latetall line comipletes.co.4 tice occ cceniene tae BR nweenominer eee eedra ae 16
2 Doarsaltinicwithtesblackiblotchiong emer cette eee A a eae 3
- Lorca Stat ew Otp LOL CHOSE oo hs stan ede ernaietstendtetontn cee pne tear eqteetanetabe geataee’ A
3 Body height 26.6 t0:29.3% of SL -.2.:..t2ccedeee: Hyphessobrycon dorsalis
- Body height4$'34.5- 10: 36-9 V0OP SL wissecnscssasgeres tas Hyphessobrycon sweglesi
4 Oretowror prcriamil lary Coot Rs 2 yo. 2s Search atesehones tei oueke tak tace ea deities Reb ea ale 5
- Jl we-rows-olprentaxdl latyvtee tin. Locececanataterd ee a. trac. eee os na ee ars 7
5 Dorsal-fin origin posterior to the anal-fin origin ...... Paragoniates alburnus
- Dorsal-fin origin anterior to the anal-fin Origin... cele eeeeseeseeeeeeeeeeeeee 6
6 A dark embedded crescent-shaped mark on base of each caudal lobe; no red
edloron- caudal fit scisecwadin ccctiaidsest teas Microschemobrycon casiquiare
= Without such dark pigmentation on the base of caudal fin; caudal fin red in
lhe ar ae Eh et Ree een Re RINE TLIC Aphyocharax alburnus
7 Caudal fin with black pigmentation in both lobes... eee eeeeeeeeeeeeeeees
ah dapat Nai edn stte at ead dere eh tte da atts ented Soe kai Hyphessobrycon otrynus
Caudal fin-withoutiblack: pigmentation.s....coseecesbeercetssaenvsbierersboceacestanatabes 8
8 Anal-fin base with a conspicuous dark band....... Hemigrammus barrigonae
~ Anal-fineBase with out-dark PlomentaOn.s.,isrionseviserieesscescoonsesspenreatonndenetetne 9
9 25-27 branched anal-fin rays; caudal peduncle red or rose in life... eee
i honcbphigedinwai beat phe sra'decieeaphsirapir singh one sazinenlint le odetigh'eenienisectdipe® Hemigrammus stictus
~ 14—22 branched anal-fin rays; no red or pink color on caudal peduncle....10
10 A small dark line of black chromatophores along the anal- and caudal-fin

BASES ei catet cas bar cteesatrs on nseatPaeteudlt inven desas'snstor eaten tes Pctan pusedectae Cyanogaster noctivaga
Caudal fin without dark coloration or with a well-developed spot at its base
Gio MCA Ale tA © DOG e.Fticee Alon lasctisegeennse Aasntenteult ietetp yinicalsd beer ideokdis. Mean 11

1]

Fishes of the Cusiana River, with an identification key to its species 87

IMithrat corispicueuUs, Calida spotters ne eiee seu occagtecd sane ara: 12
Without a caudal spot or with a lateral stripe continuing along middle cau-
dal-finsrays) but notforming acatdal Spot oc ccisicesceecivnccveseceswsncgeteseceescwess 14
Dark melanophores present on sides of body and between anal fin and lateral
WHER Pesctecskapiga ce nsn ne oR annua MA RaW ntact Mn etch eet Hyphessobrycon metae
No dark melanophores present on sides of body and between anal fin and
Vater Bte tas Shere tare ete, Net eet a Pa ie ae Neh A OR, Saree Rae 13

Caudal spot triangle-shaped covering the central portion of caudal-fin base,
continuing along middle caudal-fin rays and usually not in contact with the
lateralis REISS: 2s. sess ac. cessor seen, wa dsant ee hens ate he Hemigrammus micropterus
A large caudal spot covering entire caudal-fin base, continuing along middle
caudal-fin rays and in contact with lateral stripe.... Hemigrammus newboldi
Body depth 27% or less of SL; 14-16 branched anal-fin rays; maxillary teeth
absent; black blotch on upper caudal-fin lobe .. Hyphessobrycon diancistrus
Body depth 28% or more of SL; 18-20 branched anal-fin rays; maxillary

teeth present; without dark blotch on upper caudal-fin lobe uw... eee 15
12 dentary teeth; 6—7 perforated lateral-line scales. Without a dark blotch at
CAUCAN PEA UNC EAE ees: eed eure eR nu cvigeoatelyiagh cars Hyphessobrycon taguae
14 dentary teeth; 9-10 perforated lateral-line scales. With a dark blotch at
fer Tivabidlge yea iteYal law ayy onharion en ene Aree eee Hyphessobrycon acaciae
Dorsal-fin origin at or posterior to vertical through anal-fin origin ........... Ly,
Dorsal-fin origin anterior to the anal-fin Origin... eee eeeeseeeeeeeeeeees 22
ON Grand etnray Gear cs tuc tibiae ubhag bheunnpoliunls shad Buactton as Xenagoniates bondi
D5 Sein alciiivetalys tt. b ies oO Ate ele tush esol Re Ete 18

Length of maxilla equal or shorter than vertical diameter of eye; adult males
with hypertrophied caudal-fin squamation on lower caudal-fin lobe.............
Pellet alow eeten rs sktieal ute tse arsaluse en seeetcnilttn DS ox Gephyrocharax valencia
Length of maxilla longer than vertical diameter of eye; adult males without

hypertrophied: caudal-finrsquatnation..2:c6c2s tsseccottaces sees atl arecneene 19
Norexteriaaltmatimilife tir teeth. -s.ccect. sect meestaae ae tenors Se Charax sp.
External mammiliform teeth present in the maxilla and premaxilla........... 20
50=65, lateral-liné-scales: Ma.7-5.t. Soe Roeboides dientonito
Go-ormmere) lateral Mines cal es xaos es wlan nettle eae nsdaoe Meath needa paasiitasle 21
66-70 perforated lateral-line scales; 12-14 gill rakers on lower arm of the first
Gi AB CIN a cuvevun ctiah ici uaten uk Meek weeks eta evacesccb ice luden tig tecnae: Roeboides sp.
70-88 perforated lateral-line scales; 7—11 gill rakers on lower arm of the first
rc Lge tigcl seme Ene, Be ee REN, RTO RAPE Roeboides affinis
PO.ort less total atial finnetayes ence ess. concen ornpkere shee: = sets ov ahoes ats orsrnn de ash oteore 23
Worst tee Coca larvae tulsa Sree ts erer oc eaters eater once toketeee sera eee rece erneesore 27

Premaxillary teeth arranged in 2 rows; Anterior triad of larger teeth with
FOULED AaSe A DSC Msn aiee cue ta seee eae saree tec dee se Neces bene Ceratobranchia sp.
Premaxillary teeth arranged in 3 rows; Anterior triad of larger teeth with
iROTeE te held MO: Woe 8) ReiSLel fl tos RA BPSD r eRe a SN et Re 24

34

2)

36

Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

Dorsal-fin origin anterior to pelvic-fin insertion... Creagrutus sp.
Dorsal-fin origin posterior or aligned with pelvic-fin origin wee 25
Third infraorbital poorly developed, leaving a broad gap between its posterior
margin and the horizontal limb of preopercle ............... Creagrutus bolivari
Third infraorbital well-developed contacting or with a small gap from hori-
zontal limb ot, pred perc be. coae ...Rercccetaa oe nse a eae nn. hee cee eee 26
38—42 lateral-line scales; interorbital distance 34.0—37.9% of HL ................
FE ee eRe, Re nner Ae be SBR 7 MER Se aE Creagrutus atratus
36—38 lateral-line scales; interorbital distance 28.6—35.4% of HL .............08.
sc ARSC One EN Saeed snc su dbSnd att Sa UST SOAR Ss dy Creagrutus cf. taphorni
AQ-oF moreso talramaletiny rays 5.08 osges. Bere oseeteeSeeeeae rece sedans neeteol aateek 28
DeOLMeSs Mota Katia L-LMal Ay Srectinssdl va. snadernsoarstemadusnevactonatraeecteenkrataen: 33
Cyeloidtseales-oir body: sides? 22k. nvieuescrte<octece trend on aeesectie testes ee taewes 29
Ctenoid scales on body sides, or at least in preventral area... eters 30
Maxilla shorter than vertical diameter of eye; cleithrum without a notch on
posteroventral portion; body sides with dark sinuous lines..... Markiana geayi
Maxilla longer than vertical diameter of eye; cleithrum with notch on poster-
oventral portion, near base of posteriorly directed spiniform projection; body
sides-withidatk-sitauous Lites: 2. 5208 Jai See aod desea ust ade Charax metae
58 or fewer lateral-line scales; premaxillary and dentary teeth multicuspid;

ctenoid:scales:only on preventral area c.2.1.-n0:tsateesenecees Ctenobrycon spilurus
59 or more lateral-line scales; one or more conical or canine teeth; ctenoid
Sica SHO IMELL CAR OGL. ah. the posed cdetna tha, cus steals Shake platens ontgvenssites shouleg eae ue 31
POO= 1. WOdatetal-line:scales. 2. thet s.stelduatee hues Cynopotamus bipunctatus
70284 laters Plinerscales > tyres See eater cee we, Wipe cron ore NN om er 32
35-38 branched anal-fin rays; 73—76 lateral-line scales ..........ceeceesceesteeeneeeees
SANIT A ht TR eI ROIs re ON PRO ER Acestrocephalus sardina

39-45 branched anal-fin rays; 79-84 lateral-lines scales....Galeocharax gulo
A pair of cuspidate teeth on premaxilla pointing forward on labial sides of
upper jaw; two large black blotches on body sides, one anterior to dorsal-fin

origin and another on caudal peduncle 0.0... eee Exodon paradoxus
No cuspidate teeth on premaxilla; different color pattern than that described
2 YO opty. are lee come algae 7h Roe ee Re PR ence ne es Peer te i oP oC 34
One tow -onpremearal lakyaheetiis. 0.0, Aty ie On Mer SNe Be ere Dee teat 35
shworows.ol premaxt laty, teetlnt.ec.cre.skecsescqeescdeter si eesece ets goes eet sedenereetes: 38

Premaxillary teeth with five cusps, the outer cusps very small; dentary teeth
with five cusps, the three central cusps flat and approximately equal in size
and the.owtercusps very-sitiall) <r siew,nacespnndnemddentplaet Cheirodontops geayi
Premaxillary teeth with nine cusps; dentary teeth heptacuspid, with central
GUS PalON COP Ha Me OLNCL Or ca clacaunestatlacssa Rate duine savas subscsc sale Ota Meatetientslvalnes 36
A remarkable elongation of the second unbranched dorsal-fin ray in males;
maxilla somewhat triangular, short, with mid-length portion deeper and
gradually narrowing to the posterior tip ...... cee eeeeeeeeeeee Odontostilbe pao

2)

48

49

Fishes of the Cusiana River, with an identification key to its species 89

No elongation of the second unbranched dorsal-fin ray in males; maxilla
somewhat spatula-like shaped, short or elongate, with a deep mid-length re-
gion, narrowing abruptly at posterior tip ....... cee eeeeeeeeeseeseeeeeeseeeeeeeeneceees 37
Adipose-fin origin at vertical through second or third last anal-fin ray inser-
tion; mature males with hooks on first to sixth-seventh branched anal-fin
PAYS ee ste a ROS. Pe CR ca cee PL Odontostilbe splendida
Adipose-fin origin at vertical through last anal-fin ray insertion; mature males
with hooks on first to twenty second branched anal-fin rays «00... ceeeeeeeeeeeeee
Faceted te SDR edt teeetiactetatod epieaete Mca te Odontostilbe pulchra
Presence of a predorsal spine; anterior three anal-fin rays black...
LA We ed chs Mel ee ta ORs Sic ENGR Se Ms, su EN a Poptella compressa

No predorsal spine; three anterior anal-fin rays unpigmented ...............04 39
39=38 totaltatial fins tays ..ciac coset ee eesener nec oe Sern eee cant ated cowl daseret ants 40
DD Otylessstotal Aiialti li; FAVS a, uussereete dabuassurhesetonbenenmastast on bash ove sale dna pyiantionse.t 42

Body depth 28.0-37.0% of SL; maxilla with 20-30 conical teeth; 6 scale
rows between lateral line and dorsal-fin origin; one humeral spot..................
Lac ealaeaa dcuensnehtel mune sued st Seat chats eRonapiitsenag teseaies ia Phenacogaster maculoblonga
Body depth 38.0% or more of SL; maxilla with 1-3 multicuspidate teeth; 7-9

scale rows between lateral line and dorsal-fin origin; two humeral spots......... 41
Caudal peduncle with a large dark spot.............. Tetragonopterus argenteus
Caudal peduncle without spot ...... cs .eeseseeseeseerees Gymnocorymbus bondi
G29 Osteotanlbatiicll Bitte t Ayes aoee re oieyoses sae reverses eee day oo vestes ore sory eoeones re setponticssny Severe 43
2-0 POP Al ALIA EIDE AY She aero tdeleg Chacon decade Shaler Bedtteg oat eorsitee MACY eget uc ce Ae.
LPS arial BAAS 508,258 cates Ses gsnses tenses asueeret Knodus deuterodonoides
MGrarial=biny: PAv6 lace. cekees deen ptsasnwee user, Bryconamericus cf. cismontanus
Spine-like pelvic bones projecting anteriorly from pelvic-fin base, with distal
Pip ieeeel relma SCAT ECS ees Seeger ase ee ee soctnee Jupiaba polylepis
Nos pineshikeipelwicibonies, tet. t eee ot ee et eee ad hee eee eden. 45
4eeth on -the-lnmer. row. of premtaxillay cass. tess. csieseesceupe cesta ees aseee 46
D:teethiOn,the snticterowror preriayt WA os... scmeecn: where sasedetg eaeee hteetas shen tase 48
SON tos hes Peco uesals 60.01 |e geen cee pee ay A eet eee Knodus alpha
Moresthian ‘thee snéethvon iataxi la i ete ctices cairn deseo teanlon teas dea tdapinen Saveenete 47

5 or fewer maxillary teeth in adult individuals (>5 cm SL); second premaxil-
lary tooth of inner row pentacuspid; 12-13 predorsal scales; fins of live speci-
ANNENS FOACIGI Sy ots res vconsuevuceds vctenv user queucetgutvenesiceegeveenrdtts Hemibrycon loisae
6 or more maxillary teeth in adult individuals (>5 cm SL); second premaxil-
lary tooth of inner row heptacuspid; 14-16 predorsal scales; fins of live speci-

WIETIS. TOE PECISIN eh Slee dance hs acta es negencsetgh nes sntenishgee dis Hemibrycon metae
One or both caudal-fin lobes with dark coloration .............cccccccccceeeececeeeees 49
Both caudal-fin lobes without distinct pigmentation... eee eeeeeeeeeee 50

Only upper caudal-fin lobe with dark pigmentation... eee eeseereeeee
lied heees beset Aas wens vanes eaten ee Stas at da gata nese Anat Moenkhausia lepidura
Both caudal-fin lobes with black pigmentation........ Moenkhausia dichoura

90 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)
50 Middle caudal-fin rays without dark coloration.............. Moenkahusia copei
- Middle caudal-fin rays with dark coloration .........cceesseeseeseeeeeeeeeeeeseeeeaes 51
51 Predorsal-line naked yen. sccccseecceenncevsvcenermcnsruccasel Astyanax bimaculatus
— Predmrsaldline wiciesealest: che oi. sa nthe te of ge vad tyenanie tata t anh ut uae el 52
a2 Base of anal fin with an oblique dark stripe extending across caudal peduncle
and onto middle and upper caudal-fin rays... eee Astyanax metae
- Base-of analthn-without-obliqnedatk Stripe. gu csde daze: eccetapdenctieeccest ot 53
53 #S-oramorerlatertal cline SCales v.00 siadsncevdeneetinsdooa rans soncainnues Astyanax integer
- Fewer than 45 lateral-line-scales........ ee eeeeseeseeseeeeeees Astyanax venezuelae
Cichlidae
1 African lips (posterior portion of the lower lip not covering part of the up-
per lip); three roundish ocellar blotches in adults (large and oval blotches in
UOT CS wets caceoe cor ecessease ceossnunmestectevecceip ez aeesicest: Cichla orinocensis
American lips (posterior portion of the lower lip covering part of the upper
lip) diferent color parte Lan ADOVE a twaced ve vsccuwoneitaad cai tenesitns ctathost tee Z
2 Bone expansion in the upper region of the first gill arch forming a well-
develojpedttleshyaGberrer aot e eae teeta et eee eee 3
-- Rirstgill arch. -withoutsstieh- al OW es.2.1..-spbtncesobanueraata tes tna renter. tects Setanta dee 6
3 Lower pharyngeal bone without teeth along its margin... eee eeeeeees
VEEL SRM TIWs BUENA Airtine rs Ae Moet ee Ory Sena AN An Wy Dy Mikrogeophagus ramirezi
- Lower pharyngeal bone with teeth along its margin... eile ceeeeseeeeeeee 4
4 Lateral spots on body present; two pectoral spots... Apistogramma macmasteri
= No lateral spots on body; with or without a single pectoral spot .............0... 5
Dy Pectoral fin base with spot; with distinct abdominal stripes... eee
Lidia geisoatce ara G Postel enutu sateen i cet Miaka ge ebeet Peat Apistogramma hongsloi
= Without spot on pectoral fin base; without abdominal stripes ..........e eee
Ppahshegh soebeautnn BA cts hesah. woken beseaekedeneaisanennas aohewersenante Apistogramma hoignei
6 Irrecularpredorsal. seal Apart cased erennonste eds sees crabs amass upp dndivamen teenie Th
Wniserialorrisetial predorsal'seale patter £2504. eo Se 11
7 Six or seven anal-fin spines; 24-30 scales in upper lateral line series; vertical
bars 6 and 7 parallel; body deep (50.7-55.6%of SL)..... Mesonauta egregius
= Three anal-fin spines; 40 or more scales in upper lateral line series; elongate
and somewhat cylindrical body (less than 50.0% of SL) .....eeseeseeeseeeseeeeeees 8
8 Rlumeraltb lapel presGnt (otic cs scesser ac mach avis penerotase mez meee 9
- Eitrmera lt bleteh absent 2ais5.5.20cat.20 aase. eset eee sk OE Bn BE ee iI
a Lateral line crossing middle portion of humeral blotch....... Crenicichla alta
— Lateral line crossing upper portion of humeral blotch... eee 10
10 Chain of blotches along lateral lines; length of posterior dorsal-fin spine

9.6—10.6% of SL; caudal peduncle length at ventral part 10.0—11.5% of SL;
length of ventral fin 19.4—20.4% of SLi eeeeeeeeeeeee Crenicichla sveni

Vt

14

15

Fishes of the Cusiana River, with an identification key to its species 91

No chain of blotches along lateral lines; length of posterior dorsal-fin spine
10.8—11.2% of SL; caudal peduncle length at ventral part 11.8—12.2% of SL;
length of ventral fin 17.3-18.2% of SLi. eeeeseeseeseeeeeeees Crenicichla sp.
Maxilla extending posterior to the anterior margin of eye; caudal-fin length
(from caudal-fin base to tips of middle caudal-fin rays) 16.2-24.6% of SL;
head depth at orbit level 16.7—21.6% SL; caudal peduncle depth where least
IR Eee BO! 0) a] I Decora a I A re hare SI Crenicichla geayi
Maxilla only reaching anterior margin of eye; caudal-fin length (from caudal-
fin base to tips of middle caudal-fin rays) 22.7—24.7% of SL; head depth at
orbit level 15.1-15.6% SL; caudal peduncle depth where least 9.34—9.83%
2) hoy DERG SOOT A, API OL AOE OETA RON I Crenicichla gr. wallacti
Winisetiallredoisal Seale (Atte nir sj neuter snedsbas meterteaerememepest Leese meeae 13
‘Trisetialipredorsaljscale pattetta s425 022. ott a eeea. eden acres ee ee ee ce 14
8 branched anal-fin rays; caudal fin rounded; less than 8 scales in the lower
hater lWliries sa eG Red athe ee Sere eee oe eee oe, ad Andinoacara sp.
7 branched anal-fin rays; caudal fin subtruncate-truncate; more than 8 scales
dir the- lower Matera Wie. is Sexi sil aipeteheas odd dosanasuSeaveaes Bujurquina mariae
Dark stripe extending from posterodorsal margin of eye to lower angle of
PROGISCRE Le ot oct crest a cd eekinen Bs ea teediien oor lau Ae Sy os Aequidens metae
Dark stripe restricted to a suborbital blotch only «0... eee eeeeeeeeeeeneeeeees 14
Head sides with bluish or greenish iridescent stripes (in live specimens)........
becsuet tat acter gua enc Paataal guatess eat iualt on tisuett siren ten gyoutestets estes se testa Pewee Aequidens diadema
Head sides without bluish or greenish iridescent stripes (in live specimens) ..
PME sch Seca Mee teh wsuleOR cutee eats NoRNE asks WSL ONE genset Aequidens tetramerus

Sciaenidae

Lateral-line scales not covered by smaller scales; body depth 3.8-4.3% of SL;
predorsal distance 3.0—-3.1% of SL; anal-fin base 5.9-9.3% of SL; caudal
peduncle length 3.4—3.7% of SL; postorbital length 2.0—2.4% of SL; length
of second anal-fin spine 1.8—2.3 % of HL... Pachyurus gabrielensis
Lateral-line scales covered by smaller scales; body depth 3.0—3.8% of SL;
predorsal distance 2.7—3.0% of SL; anal-fin base 11.2-17.1% of SL; caudal
peduncle length 3.8-4.7% of SL; postorbital length 1.6—2.0% of SL; length

of second anal-fin spine 2.6-6.0% of HL......... Plagioscion squamosissimus

Discussion

Regional checklists of freshwaters fishes become dynamic over time as studies in fresh-

water fish taxonomy and distribution advance for the Neotropical region (Reis et al.
2016). The most recent checklist for the Meta River basin reported 258 species for

92 Alexander Urbano-Bonilla et al. / ZooKeys 733: 65-97 (2018)

the Cusiana River sub-basin (Usma-Oviedo et al. 2016); it is a higher number of spe-
cies than reported herein because re-identification and taxonomic updating processes
of specimens excluded 59 of the 258 nominal species reported by Usma-Oviedo et al.
(2016). For example, most of the undetermined (e.g., Azcistrus sp., Aphyocharax sp,
Microglanis sp., and Odontostilbe sp.) and erroneous records (e.g., Hemibrycon cristiani,
Pyrrhulina brevis, Schultzichthys gracilis, and Steindachnerina guentheri) originally count-
ed as independent species were merged with other recorded species after our verification
of the data. This is not surprising, taxonomy proceeds at a faster pace than the institu-
tional ability to maintain updated records. For this reason, there is a need to account for
validation of species identification when regional checklists are assembled from multiple
secondary sources in order to avoid errors due to outdated or unverified data.

Extrapolation suggests that the drainage could have a richness of roughly 314 spe-
cies, indicating that the number of species found in the present study represents 77.7%
of the expected richness in the area (Table 1). However, this estimate represents a rough
estimate because sampling efforts have not been uniform across the drainage. A his-
torical sampling-specific bias is not expected in the Cusiana River sub-basin, with the
possible exception of an elevational bias (as is true for the whole Orinoco drainage in
Colombia). Given the non-uniform nature of sampling in comparable river systems,
we suggest that our extrapolations of species richness may be useful for comparisons
among drainages using collection records as the input for rough estimation. This is par-
ticularly important since most checklist studies compare observed and not estimated
richness, with the latter a more appropriate measure because it incorporates (and even
overestimates) uncertainty from the samples into the estimation, and it also serves ac-
counts for sampling effort among drainages.

Among similar Orinoco Andean tributaries, the Cusiana is one of the best-sampled
sub-basins, exceeding in species richness other recently well-sampled sub-basins such
as Orotoy (113 spp.; Ramirez-Gil et al. 2011) and Pauto (182 spp.; Maldonado-Oc-
ampo et al. 2013). In fact, the Cusiana River sub-basin represents the quantile 0.83
among Orinocoan drainages, indicating that 83 % of the other drainages had fewer
than 74 localities represented in the collections. The importance of the Cusiana River
sub-basin is not only determined by its fish richness, but also because of its diverse and
extensive aquatic ecosystem richness (rivers, streams, lagoons, estuaries, palm swamps,
riparian forests, and flooded savannas) that provide important areas for fish repro-
duction, shelter and food. Because aquatic ecosystems have dynamic ecological and
environmental processes (Teresa et al. 2015, Ribeiro et al. 2016, Toussaint et al. 2016),
management and conservation projects of sub-basins should be addressed at regional
(sub-basin) scales.

The documentation of the ichthyofauna in cis-Andean Colombian sub-basins has
been increasing during the last decade, but new records and species can likely still be
found in areas previously thought to be well-sampled (e.g., Ballen 2011, Vanegas-Rios
et al. 2015, Ballen et al. 2016a, 2016b, Burns et al. 2017, Garcia-Alzate et al. 2017).
Most of the sampling effort has been carried out in the piedmont and lowland areas

Fishes of the Cusiana River, with an identification key to its species 93

in the Cusiana as well as in other sub-basins, and exploration of High Andean areas
could lead to the discovery of local endemic species at the basin scale that usually are
underestimated (Carvajal-Quintero et al. 2015).

Sub-basins adjacent to the Cusiana draining along the eastern slope of the Eastern
Cordillera in the Orinoco region of Colombia (e.g., Guachiria, Casanare, Upia, Tua,
and Cravo Sur) have not been well sampled and their richness is surely underestimated
(Urbano-Bonilla et al. 2014). Continuous efforts are still to be carried out in order to
document the fish fauna present along this region; this information is crucial to better
understand how different anthropogenic activities (mining, oil extraction, agricultural,
and livestock practices) are affecting the environmental conditions of these areas and
as a consequence, the fish that live therein. Combination of this kind of information
and further environmental data is a necessary step in order to generate freshwater con-
servation strategies using different approaches and therefore go further toward effective
protection initiatives for species subject of conservation in the region.

Acknowledgments

Funding to conduct field work along the Orinoco piedmont were provided by the
Pontificia Universidad Javeriana (Project N° 5211411), Instituto de Investigacion de
Recursos Biolégicos Alexander von Humboldt, and Oleoducto Bicentenario (Project
N° 4400000166); cooperation agreement between Fundacién Reserva Natural La Pal-
mita, Centro de Investigacié6n and [AvH (N° 15-14-172-010CE), and Gobernacién
de Casanare (Consulting contract N° 1819-2014). GAB was funded through a doc-
toral scholarship and a BEPE internship by FAPESP (processes 2014/11558-5 and
2016/02253-1). To Dr. Francisco Villa-Navarro for providing fish database of the Zoo-
logical Collection of the University of Tolima (CZUT-IC). The authors thank Carlos A.
Garcia-Alzate, Donald Taphorn, Flavio Lima, Henry Agudelo, Jorge E. Garcfa-Melo,
Paulo Andreas Buckup, and Vitor Abrahao for their valuable time and comments on
taxonomic verification of species or keys, and Maria Paula Arango and Natalia Borray
for their valuable support in some phases of the laboratory analysis. We acknowledge
Donald Taphorn and Devin Bloom for checking the English of the manuscript.

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