A systematic revision of Tatia (Siluriformes: Auchenipteridae: Centromochlinae)

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Neo opical Ich hyology, 6(3):495-542, 2008
Copy igh © 2008 Sociedade B asilei a de Ic iologia
A sys ema ic e ision o Ta ia
(Silu i o mes: Auchenip e idae: Cen omochlinae)
Luisa Ma ia Sa men o-Soa es1,2 and Ronaldo Fe nando Ma ins-Pinhei o2
The auchenip e id ca ish genus Ta ia is e ised. Twel e species a e ecognized including h ee desc ibed as new. Ta ia is
diagnosed by he hyomandibula elonga ed an e odo sally, he anal- in base o adul males educed in leng h, and he caudal
peduncle la e ally comp essed and deep wi h a middo sal keel. Ta ia aulopygia occu s in he Madei a i e d ainage and is
dis inguished by he educed c anial on anel in adul s and male modi ied anal in wi h middle ays educed in leng h. Ta ia
boemia, known om he uppe U uguay i e d ainage, is dis inguished by i s unique colo pa e n wi h da k ch oma opho es
on he sides o body. Ta ia b unnea om i e basins in Su iname and F ench Guiana and he Neg o i e d ainage, Amazon
basin, is ecognized by i s wide head and mou h and by he male modi ied anal in wi h sha ply poin ed ip. Ta ia dunni, om
he uppe Amazon basin, is ecognized by i s na ow head, long pos clei h al p ocess in some specimens, and body colo a ion
wi h i egula blo ches o s ipes. Ta ia galaxias, endemic o he O inoco i e basin, is dis inguished by i s la ge eye and sho
snou . Ta ia gy ina, dis ibu ed in he uppe and cen al Amazon basin and in no he n Su iname, has a uniquely educed
mese hmoid, sligh ly p o uding lowe jaw, second nuchal pla e wi h sligh ly conca e la e al bo de s, hi d nuchal pla e
educed, small p e ome , low numbe o ibs, low numbe o e eb ae and sexual dimo phism ega ding in umescen male
geni al papilla. Ta ia in e media, eco ded om cen al and lowe Amazon basin, Tocan ins i e , and coas al d ainages in
Guyana, Su iname, F ench Guiana, and eas e n Pa á S a e, B azil, is dis inguished by he sho pos clei h al p ocess, small eye
and long snou . Ta ia nei ai, om he uppe Pa aná i e , Pa aguay i e and uppe Pa aíba do Sul i e basin, is dis inguished
by i s unique e eb al coun and caudal- in colo a ion consis ing o ans e se da k ba s. Ta ia s iga a, om cen al Amazon
basin and Neg o i e , is dis inguished by i s ho izon ally s iped colo pa e n and he modi ied male anal in wi h middle ays
educed in leng h. Ta ia caxiuanensis, a new species desc ibed om he Cu uá i e , lowe Amazon basin, is ecognized by i s
wide c anial on anel and dis inc i e anal in in ma u e males. Ta ia meesi, a new species desc ibed om he Essequibo i e
basin, Guyana, is dis inguished om congene s by he c anial on anel wi h wo sepa a e openings and hin nasal bone. Ta ia
nig a, a new species desc ibed om he cen al Amazon basin, is dis inguished by i s sho pos clei h al p ocess, low numbe
o e eb ae, and da k colo pa e n. All wel e species o Ta ia a e desc ibed o edesc ibed and a key o species is p o ided.
O gêne o Ta ia de auquenip e ídeos é e isado. Doze espécies são econhecidas incluindo ês desc i as como no as. Ta ia é
econhecido pelo hiomandibula o emen e endido an e odo salmen e, base da nadadei a anal de machos madu os eduzida
em amanho, e pelo pedúnculo caudal la e almen e comp imido e al o com uma quilha médio-do sal. Ta ia aulopygia oco e na
d enagem do io Madei a e é dis inguida pela on anela c aniana eduzida em adul os e pela nadadei a anal em machos madu os
endida, pela edução em amanho dos aios medianos. Ta ia boemia, conhecida da d enagem do al o io U uguai, é econhecida
po seu pad ão de colo ido único com c oma ó o os escu os pelas la e ais do co po. Ta ia b unnea, de bacias hid og á icas no
Su iname e Guiana F ancesa e ainda da d enagem do io Neg o na Amazônia, é econhecida pela ampla la gu a da cabeça e boca
e pela nadadei a anal modi icada em machos com ex emidade pon iaguda. Ta ia dunni, do al o Amazonas, é econhecida pela
cabeça es ei a, pelo p ocesso pós-clei al longo em alguns espécimens, e pela colo ação do co po com manchas i egula es ou
aixas cla as. Ta ia galaxias, endêmica da bacia do io O inoco, é dis inguida pelos g andes olhos e ocinho cu o. Ta ia
gy ina, com oco ência pelo al o e médio Amazonas, e pelos ios do no e do Su iname, possui mese móide eduzido, mandíbula
le emen e p ogna a, segunda placa nucal com bo do la e al es ei o, e cei a placa nucal eduzida, p é- ome pequeno, eduzido
núme o de cos elas, pequeno núme o de é eb as e dimo ismo sexual onde a papila geni al masculina é en umescida. Ta ia
1Labo a ó io de Ecologia de Peixes, sala 525, Pa ilhão Ha oldo Lisboa, Depa amen o de Ecologia, Ins i u o de Biologia Robe o Alcân a a
Gomes, Uni e sidade Es adual do Rio de Janei o. A . São F ancisco Xa ie , 524; Ma acanã, 20550-013 Rio de Janei o, RJ, B azil.
[email p o ec ed]
2Museu de Biologia P o . Mello Lei ão, Labo a ó io de Zoologia. A . José Ruschi, 4, Cen o, 29650-000 San a Te esa, ES, B azil.
[email p o ec ed]
A sys ema ic e ision o Ta ia
496
in e media é egis ada pa a o médio e baixo io Amazonas, io Tocan ins e ainda pa a d enagens cos ei as na Guiana,
Su iname, Guiana F ancesa e les e do Pa á no B asil. É dis inguida pelo p ocesso do pós-clei o cu o, olhos pequenos e
ocinho longo. Ta ia nei ai, do al o io Pa aná, io Pa aguai e al o io Pa aíba do Sul, é dis inguida pela con agem e eb al
exclusi a e pela colo ação da nadadei a caudal com ba as ans e sais escu as. Ta ia s iga a, dis ibuída pelo médio Amazonas
e io Neg o é econhecida pelo pad ão de colo ido com lis as ho izon ais i egula es e nadadei a anal modi icada endida em
machos, com aios medianos eduzidos em amanho. Ta ia caxiuanensis, no a espécie, desc i a pa a o io Cu uá no baixo
Amazonas, é econhecida pela on anela c aniana ampla e pela nadadei a anal de machos madu os dis in a. Ta ia meesi, no a
espécie, desc i a pa a o io Essequibo, Guiana, é di e enciada de seus congêne es pela on anela c aniana com duas abe u as
sepa adas e pelo osso nasal a ilado. Ta ia nig a, no a espécie, desc i a pa a o médio Amazonas, é dis inguida pelo p ocesso
pós-clei al cu o, pelo núme o eduzido de é eb as e pelo pad ão de colo ido escu ecido. Todas as doze espécies de Ta ia
são edesc i as ou desc i as e uma cha e de iden i icação é o necida.
Key wo ds: Sou h Ame ica, F eshwa e , Ca ish, Taxonomy, Cen omochlus.
In oduc ion
The sub amily Cen omochlinae comp ises small o me-
dium size auchenip e id ca ishes ha sha e de i ed anal- in
mo phology in males (Fe a is, 1988; Soa es-Po o, 1998). Fou
gene a, Cen omochlus,Ta ia,Glanidium, and Gelanoglanis,
comp ising 31 species, a e p esen ly ecognized as alid in
he sub amily (Fe a is, 2007).
Ta ia comp ises a g oup o ela i ely small (20-150 mm
s anda d leng h), noc u nal auchenip e ids [o
cen omochlins] ha eed on small ui s and in e eb a es
(H. A. B i ski, M. Goulding, pe s. comm.). They a e endemic
o Sou h Ame ica eas o he Andes and a e p esen in mos
o majo d ainages, including he O inoco, Amazon, Pa aná-
Pa aguay, and U uguay, as well as no he n coas al i e s
om he Essequibo o Amapá S a e and Ma ajó Island, B azil.
Ta ia is absen in he São F ancisco i e basin and small
i e ine basins along he B azilian eas coas . Ta ia is he
mos speciose genus in he sub amily, wi h 12 species
ecognized he e (Table 1). Ta ia species occu in len ic
sec ions o iga apés, i e s, and lakes, whe e hey emain
hidden in subme ged unks o ocky c e ices du ing he day
and eme ge only a nigh o o age (Lowe-McConnell, 1964,
1975, 1987; Soa es-Po o, 1995). A nigh , indi iduals o Ta ia
usually emain close o he su ace nea i e banks o in he
middle o he i e , swimming in a e y peculia way ( as and
e a ic), and cap u ing allen insec s (F.C.T. Lima, pe s. comm.).
Some species occu in black wa e i e s, bu none a e
es ic ed o hese en i onmen s. Many membe s o he genus
a e beau i ully colo ed and, consequen ly, e y a ac i e o
he o namen al ish ade. Ta ia species, howe e , a e no
well sui ed o cap i i y because o hei appa en in ole ance
o low oxygen condi ions. Mos do no su i e anspo o
Eu ope and No h Ame ica (Sands, 1984).
The gene ic name Ta ia was p oposed by Mi anda Ribei o
(1911: 360) o Cha les Ta e Regan o he B i ish Museum o
Na u al His o y, London, in hono o his many con ibu ions
o he knowledge o he Sou h Ame ican eshwa e ishes.
The g oup was e ec ed o include wo species p e iously
asc ibed o Cen omochlus:Ta ia in e media (S eindachne )
and T. aulopygia (Kne ), based on he p esence o a geni al
papilla o e he an e io anal- in ays (Mi anda Ribei o,
1911:353). A e Mi anda Ribei o’s (1911) desc ip ion he name
Ta ia was men ioned only in ca alogues (e.g. Gosline, 1945)
and p ac ically o go en. Subsequen au ho s such as Eigen-
mann & Allen (1942) and Fowle (1945a) desc ibed new spe-
cies in he genus Cen omochlus, wi hou men ioning Ta ia.
Boeseman (1953) e e ed o Ta ia, bu desc ibed a new spe-
cies as Cen omochlus c eu zbe gi. B i ski (1972) conside ed
Ta ia o be a junio synonym o Glanidium Lü ken due o
simila i ies o sexually dimo phic ea u es. In his s udy o
auchenip e ids p ima ily om Su iname Mees (1974) e-
desc ibed Ta ia and expanded he genus o 14 species, six o
which he desc ibed as new: T. b unnea, T. concolo , T.
galaxias, T. punc a a, T. e icula a and T.simplex. Fou addi-
ional species o Ta ia ha e been desc ibed subsequen ly
(Mees, 1988; Roye o, 1992; Soa es-Po o, 1995; Koch & Reis,
1996).
Rega ding in e ela ionships among auchenip e id gen-
e a, Fe a is (1988) and Cu an (1989) p esen ed independen
phylogene ic hypo heses based on cladis ic analysis. Fe a is
(1988) ecognized a monophyle ic basal clade wi hin
auchenip e ids “Cen omochlidae” and emo ed some spe-
cies p e iously assigned o Ta ia o wo undesc ibed gene a,
es ic ing he nomina e genus o: T. aulopygia, T. in e media,
T. dunni, T. galaxias, and one undesc ibed species. Cu an
(1989) ques ioned he monophyly o he speciose genus Ta ia.
The au ho conside ed Ta ia and Cen omochlus as only dis-
an ly ela ed; assigning Ta ia as close o Auchenip e ich hys.
Soa es-Po o (1998) s a ed Ta ia sensu s ic o as a mono-
phyle ic uni es ic ed o eigh species: T. aulopygia, T.
boemia, T. b unnea, T. c eu zbe gi, T. gy ina, T. in e media,
T. nei ai, and T. s iga a. O he species p e iously assigned
o Ta ia we e ans e ed o Cen omochlus, on he basis o
de i ed cha ac e s (Soa es-Po o, 1996, 1998). Mos ecen ly
Fe a is (2007) lis ed wel e alid species o Ta ia:T.
aulopygia, T. boemia, T. b unnea, T. c eu zbe gi, T. dunni, T.
galaxias, T. gy ina, T. in e media, T. musaica, T. nei ai, T.
simplex, and T. s iga a.
The p esen pape gi es a eapp aisal o Ta ia based on
examina ion o nume ous specimens in addi ion o hose a ail-
able o p e ious au ho s. Pa e ns o a ia ion wi hin and
be ween species a e conside ed h oughou hei dis ibu-
ions.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 497
Ma e ial and Me hods
Os eological ea u es we e examined in clea ed and s ained
(CS) specimens p epa ed acco ding o he p ocedu es o Tay-
lo & an Dyke (1985). P io o clea ing and s aining, speci-
mens we e dissec ed when possible o de e mine gu con-
en s, sexual ma u i y o gonads, and eco d myological in-
o ma ion. Os eological da a om some ypes o specimens
poo ly ep esen ed in ich hyological collec ions we e ob-
ained om adiog aphs (no ed as “R” in he Ma e ial Exam-
ined sec ion). Nomencla u e o os eological elemen s ollows
The Zeb a ish In o ma ion Ne wo k (ZFIN). Muscle names
ollow Sa men o-Soa es & Po o (2006). D awings we e en-
de ed om digi al pho og aphs p e e eably o clea ed and
s ained specimens. S aigh -line measu emen s we e made wi h
a digi al calipe , and eco ded in en hs o a millime e .
Measu emen s included: s anda d leng h (SL, om snou
ip o caudal- in base); body dep h (on nuchal shield, om
o igin o do sal- in spine o he belly); body wid h (wides
dis ance be ween la e al su aces o clei h a, aken be ween
an e io -mos ma gin o clei h al bone benea h pec o al- in
o igin); caudal-peduncle dep h (leas dis ance be ween do -
sal and en al su aces o caudal peduncle); caudal-peduncle
leng h ( om base o pos e io -mos anal- in ay o poin coin-
ciding wi h o igin o lowe unb anched caudal- in ay);
p edo sal leng h ( om snou ip o o igin o do sal in); p ea-
nal leng h ( om snou ip o anal- in o igin); p epel ic leng h
( om snou ip o pel ic- in o igin); do sal- in o igin o pec-
o al- in o igin; do sal- in o igin o pel ic- in o igin; pec o al-
in o igin o pel ic- in o igin; p epec o al leng h ( om snou
ip o pec o al- in o igin); do sal- in base ( om o igin o do -
sal- in spine o pos e io -mos base o do sal- in inse ion);
adipose- in base ( om o igin o adipose in old o i s pos e-
io -mos base); anal- in base (dis ance be ween geni al open-
ing and pos e io -mos base o anal- in inse ion, measu ed
unde he same c i e ia o bo h males and emales); do sal-
in spine leng h ( om base o spine o i s dis al ip); pec o al-
in spine leng h ( om o igin o spine o i s dis al ip, aken
wi h he spine e ec ed); pos clei h al (hume al) p ocess leng h
( om an e odo sal ma gin o exposed p ocess o pos e io -
mos ip o p ocess); i s b anched pel ic- in ay ( om base
o ip o i s b anched pel ic- in ay); longes anal- in ay
( om base o ip o i s b anched anal- in ay); head leng h
(HL, om snou ip o bony end o ope cle); head wid h (be-
ween do salmos ex en s o ope cula openings); snou dep h
(dis ance in median sagi al plane be ween pos e io nos il
and mid en al con ou o body); in e o bi al dis ance (leas
dis ance be ween do salmos ma gins o bony o bi ); le
in e na ial wid h (be ween le an e io and le pos e io nos-
ils); an e io in e na ial dis ance ( ans e se dis ance be-
ween an e io nos ils); pos e io in e na ial dis ance ( ans-
e se dis ance be ween pos e io nos ils); snou leng h ( om
snou ip o an e io mos ma gin o eye); maxilla y-ba bel
leng h ( om base o ip, wi h ba bel e ac ed); ou e men al-
ba bel leng h ( om base o ip, wi h ba bel e ac ed); inne
men al-ba bel leng h ( om base o ip, wi h ba bel e ac ed);
o bi al diame e (g ea es ho izon al dimension o eyeball);
mou h wid h (be ween co ne s o closed mou h).
Coun s o in ays and bony elemen s we e ob ained om
alcohol-p ese ed and clea ed and s ained specimens. Ve e-
b al coun s included all ib-bea ing cen a bu did no include
any o he an e io , complex cen um elemen s wi hou ibs,
ollowing Fe a is & Fe nandez (1987), and included he com-
pound caudal cen um (PU1+U1) as he las elemen . Hemal
spine coun s accompany he numbe s o co esponden e -
eb a, as shown in Fig. 1. Coun o b anchios egal ays we e
done only in clea ed and s ained specimens. The di ec ion o
do sal and pec o al- in spine se a ions a e e e ed o as
an o se o hose poin ing away om base o spine and e -
o se o hose ben owa ds base o spine. Male modi ied
Nominal species Mees (1974) Fe a is (2007) Assignmen he ein Taxonomic s a us
Cen omochlus aulopygius Kne , 1857 Ta ia aulopygia Ta ia aulopygia Ta ia aulopygia Valid
Ta ia boemia Koch & Reis, 1996 - Ta ia boemia Ta ia boemia Valid
Ta ia b unnea Mees, 1974 Ta ia b unnea Ta ia b unnea Ta ia b unnea Valid
Ta ia caxiuanensis, new species - - Ta ia caxiuanensis n.sp. Valid
Cen omochlus dunni Fowle , 1945 Ta ia in e media Ta ia dunni Ta ia dunni Valid
Ta ia galaxias Mees, 1974 Ta ia galaxias Ta ia galaxias Ta ia galaxias Valid
Cen omochlus gy inus Eigenmann & Allen, 1942 Ta ia gy ina Ta ia gy ina Ta ia gy ina Valid
Cen omochlus in e medius S eindachne , 1877 Ta ia in e media Ta ia in e media Ta ia in e media Valid
Ta ia meesi, new species - -Ta ia meesi n.sp. Valid
Glanidium nei ai Ihe ing, 1930 Ta ia nei ai Ta ia nei ai Ta ia nei ai Valid
Ta ia nig a, new species - -Ta ia nig a n.sp. Valid
Ta ia s iga a Soa es-Po o, 1995 - Ta ia s iga a Ta ia s iga a Valid
Cen omochlus al ae Fowle , 1945 Ta ia al ae Cen omochlus al ae Cen omochlus al ae Valid
Ta ia concolo Mees, 1974 Ta ia concolo Cen omochlus concolo Cen omochlus concolo Valid
Cen omochlus c eu zbe gi Boeseman,1953 Ta ia c eu zbe gi Ta ia c eu zbe gi Junio synonym o Ta ia gy ina Junio synonim
Ta ia musaica Roye o, 1992 - Ta ia musaica “Cen omochlus” musaicus [ince ae
sedis in Cen omochlinae]
Valid, ince ae sedis
in Cen omochlinae
Cen omochlus pe ugiae S eindachne , 1882 Ta ia pe ugiae Cen omochlus pe ugiae Cen omochlus pe ugiae Valid
Ta ia punc a a Mees, 1974 Ta ia punc a a Cen omochlus punc a us Cen omochlus punc a us Valid
Ta ia e icula e Mees, 1974 Ta ia e icula a Cen omochlus e icula us Cen omochlus e icula us Valid
Cen omochlus schul zi Rössel, 1962 Ta ia schul zi Cen omochlus schul zi Cen omochlus schul zi Valid
Ta ia simplex Mees, 1974 Ta ia simplex Ta ia simplex “Cen omochlus” simplex
[ince ae sedis in Cen omochlinae]
Valid, ince ae sedis
in Cen omochlinae
Table 1. Taxonomic lis o nominal species assigned o Ta ia by p e ious au ho s and he s a us o hose species as ecog-
nized in he p esen s udy.
A sys ema ic e ision o Ta ia
498
anal- in segmen ed ays (lepido ichia) may ha e adjacen
segmen s wi h cu ed bo de , o ming den icula ions he ein
e e ed o as an o se o e o se.
Ins i u ional abb e ia ions ollow Reis e al. (2003), wi h
he excep ion o MBML o Museu de Biologia P o esso
Mello Lei ão, San a Te esa, Espí i o San o, B azil and
NUPELIA o Núcleo de Pesquisas em Limnologia, Ic iologia
e Aquicul u a/ Fundação Uni e sidade Es adual de Ma ingá,
Ma ingá, Pa aná, B azil.
Resul s
Ta ia Mi anda Ribei o
Ta ia Mi anda Ribei o, 1911: 360 [ ype species: Cen omochlus
in e medius S eindachne , 1877, by subsequen designa-
ion by Jo dan, 1920: 545. Gende : eminine].
Diagnosis. Ta ia is dis inguished among he Cen omochlinae
by h ee uniquely de i ed ea u es. The hyomandibula is elon-
ga e an e odo sally, no con ac ing he na ow me ap e ygoid
and, ins ead, connec ed only o he apezoidal quad a e (Fig.
2); anal- in base o adul males is educed (anal- in base leng h
3.3-8.0% SL); and caudal peduncle is comp essed and deep
(caudal-peduncle dep h 10.1-18.6% SL), wi h a middo sal keel
pos e io o adipose in.
A unique combina ion o es ic ed cha ac e s aids in dis-
inguishing Ta ia: i s (an e io mos ) nuchal pla e p esen
(Fig. 3, n1); in ao bi al 1 bone sho , limi ed o an e io co ne
o o bi ; eye mode a ely la ge, be ween 17.4-42.9% HL; max-
illa sho e han o same leng h as au opala ine; e ac o
en aculi muscle absen , unc ionally subs i u ed by maxillo-
mandibula ligamen (Sa men o-Soa es & Po o, 2006); adul
males wi h modi ied anal in (all species), and i s unb anched
anal- in ay non-segmen ed o wi h segmen s used (excep
in T. b unnea).
Gene ic desc ip ion. Ta ia comp ises species wi h somewha
obus head; caudal peduncle much comp essed la e ally and
deep; well-de eloped adipose eye lid; eye mode a ely la ge,
and la e o-do sally loca ed. Following cha ac e s used o dis-
inguish species o Ta ia (illus a ions based on T. gy ina):
Head: c anial on anel always p esen (Fig. 3, o); p emaxilla
ans e sely elonga ed, mee ing i s coun e pa medially;
au opala ine ubula , o ien ed obliquely o longi udinal axis
o body; maxilla e y small, sho e han o same leng h as
au opala ine; h ee nuchal pla es p esen (Fig. 3, n1, n2, n3).
Epioccipi al p ocess small, connec ed o hi d nuchal pla e
by ligamen s (Fig. 3, pe). Mulle ian amus wi h dis al ip shaped
as sligh ly cu ed disc, i s conca e ace supe icially adjoined
o an e io wall o gasbladde . P e ome usually wi h well
de eloped a ow-shaped la e al p ocesses o wi h sho la -
e al p ocess. P e ome ian ee h p esen in T. galaxias and T.
in e media.
Suspenso ium (Fig. 2): Suspenso ium bones in Ta ia simi-
la among all species and dis inc i e among Cen omochlinae.
Hyomandibula na ow, elonga e, p ojec ed an e io ly as sho
memb anous lamina (Fig. 2, hy). An e io lamina p ojec ion
o hyomandibula su u ed only o quad a e, h ough ca ilage
en ally and deeply den a e su u e do sally. Me ap e ygoid
conical, do sally sha p, wi h li le lamina ex ension (Fig. 2,
m ), joined o quad a e ia ca ilage block only (T. gy ina, T.
caxiuanensis, T. meesi, T. s iga a) o ca ilage plus den a e
su u e (T. aulopygia, T. boemia, T. b unnea, T. dunni, T.
galaxias, T. in e media, T. nei ai, T. nig a). Suspenso ium
deeply no ched an e odo sally, hus me ap e ygoid no con-
ac ing hyomandibula. Quad a e apezoidal, wi h b oad base
(Fig. 2, qu), connec ed o p eope cle, hyomandibula and
me ap e ygoid; long p eope cle (Fig. 2, po) en al ma gins
su u ed o bo h quad a e and hyomandibula; sup ap eope cle
p esen as sho o obus canal bone (Fig. 2, sp); ope cle
lamina e and b oadly sub iangula (Fig. 2, op).
Hyoid a ch (Fig. 4): U ohyal na ow (Fig. 4, uh); sho
do sal hypohyal (Fig. 4, dh) associa ed wi h compa a i ely
la ge en al hypohyal (Fig. 4, h); an e io ce a ohyal well
de eloped (Fig. 4, ac), pos e io ce a ohyal smalle (Fig. 4,
pc); b anchios egal ay a icula ed o hyoid a ch;
b anchios egal ays 5-7; 3-5 on an e io ce a ohyal and wo
on pos e io ce a ohyal; pos e io mos usually la ges and
la ened. No b anchios egal ay associa ed wi h
in e ce a ohyal ca ilage be ween bones.
Pha yngeal (gill) a ches (Fig. 5): Basib anchial 1, in con-
ac wi h u ohyal (Fig. 5, ab), an e io o basib anchial 2;
basib anchial 2 o ming osseous od, b oades an e io ly (Fig.
5, bb2) and usually sepa a ed by gap om basib anchial 3;
basib anchial 3 sho e (Fig. 5, bb3), also b oades an e io ly;
basib anchial 4 la ge, la ened and comple ely ca ilaginous
(Fig. 5, bb4); basib anchial 2 bo de ed la e ally by ca ilagi-
nous head o hypob anchial 1; basib anchial 3 be ween ca i-
laginous head o hypob anchial 2 and ca ilaginous hypo-
b anchial 3; basib anchial 4 bo de ed la e ally by ca ilagi-
nous head o ce a ob anchial 4 and caudally by ca ilaginous
head o ce a ob anchial 5. Hypob anchial 1 mos ly osseous,
elonga e and expanded la e ally, sub iangula , wi h ca ilagi-
nous ends; hypob anchial 2 mos ly ca ilaginous,
sub iangula , wi h medial osseous pa ; hypob anchial 3 com-
Fig. 1. Schema ic ep esen a ion o pa ial axial skele on o Ta ia
boemia MCP 12949, 59.9 mm SL. Le side la e al iew. Abb e-
ia ions:a , anal in; d , dis al adials; hs15- hs18, hemal spines
15 and 18; p , pel ic in; p , p oximal adials; 8- 11, ibs 8 o
11; 8- 18, e eb al cen a 8 o 18. Scale ba = 1.0 mm.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 499
ple ely ca ilaginous, mo e apezoidal (Fig. 5, hb 1-3); hypo-
b anchial 4 absen . Fi e ce a ob anchials p esen , mos ly
ossi ied, wi h ca ilage on bo h ends (Fig. 5, cb 1-5). Fi s
ce a ob anchial suppo ing single ow o ake s (no illus-
a ed); i h ce a ob anchial expanded pos e o-medially o
suppo lowe pha yngeal oo hpla e wi h sho conical ee h.
Fi e epib anchials, all bu i h la gely ossi ied excep o ca -
ilaginous ends; epib anchials 1 and 2 od-like, wi hou ak-
e s; epib anchial 3 wi h pos e io uncina e p ocess;
epib anchial 4 b oad, wi h lamina ex ension; epib anchial 5
ca ilaginous, educed, loca ed in axil be ween ca ilaginous
ends o epib anchial 4 and ce a ob anchial 4. Pha yngob an-
chial 1 absen ; pha yngob anchial 2 sho , ca ilaginous, some-
wha ounded, placed be ween an e omedial ca ilaginous ips
o epib anchials 1 and 2; pha yngob anchial 3 elonga e, ossi-
ied, wi h expanded pos e io bo de ; pha yngob anchial 4
ossi ied, suppo ing uppe pha yngeal oo h pla e wi h sho
conical ee h.
La e al line sys em: In ao bi al 1 sho , wi h en o-la -
e al p ocess ei he clea ly es ic ed o an e io bo de o eye
o almos eaching en al bo de o eye; in ao bi als hin
and canalicula , o ming incomple e o comple e in ao bi al
se ies. La e al line on body s aigh , inconspicuous, wi h os-
si ied canal bones jus an e io ly.
Fins: Do sal in wi h majo spine p eceded by small do sal
locking spine and ollowed by 3-5 b anched ays; pec o al in
wi h one spine plus 3-5 b anched ays; pel ic in wi h one
unb anched plus 5 b anched ays, ma gin ounded; adipose
in abo e anal- in base; anal in wi h 3 unb anched plus 6-8
b anched ays; caudal in wi h 8+9 p incipal ays, o ked, lobes
wi h ounded ips; do sal caudal- in lobe sligh ly elonga e in
males o some species.
Ribs and e eb ae: Pleu al ibs 7-11; i s ib hicke han
o he s; ibs becoming p og essi ely smalle pos e io ly. Pos -
Webe ian e eb ae 30-39.
Sexual dimo phism. In ca ishes o sub amily
Cen omochlinae he anal in o sexually ma u e males is modi-
ied in o a spe m conduc o wi h he anal- in ays and p oxi-
mal adials di ec ed pos e io ly, aligned nea ly pa allel o e -
eb al axis (Fe a is, 1988; Soa es-Po o, 1998). In ma u e males
he anal in is s ongly modi ied such ha i is suppo ed by
enla ged and joined p oximal anal- in adials (Fe a is, 1988;
Soa es-Po o, 1998) and by pa ially ossi ied dis al adials.
Anal- in p oximal adials a e associa ed wi h he hickened
and leng hened hemal spines o adjacen e eb ae. Modi-
ied male anal in has h ee de eloped unb anched ays. The
i s unb anched ay is sho es , abou one-qua e leng h o
second unb anched ay, and wi h segmen s used (excep in
T. b unnea). Some species ha e he i s unb anched anal- in
ay p eceded immedia ely by a egumen a y keel. The second
and hi d unb anched ays a e hick. In he Cen omochlinae,
Ta ia is he only genus wi h species ha bea segmen s ha
cu e ou wa ds, o ming an o se den icula ions along he
an e io ma gin o he hi d unb anched anal in ay in adul
males.
Rema ks. Two o he nominal species assigned o Ta ia by
Fe a is (2007) a e no he ein ecognized as membe s o his
genus. Bo h Ta ia musaica Roye o and Ta ia simplex Mees
lack he de i ed cha ac e s lis ed abo e o Ta ia: hin,
an e odo sally elonga e hyomandibula ( s. b oad in bo h C.
musaicus and C. simplex); caudal peduncle la e ally com-
p essed ( s. ellipsoid); p opo ionally deepe caudal peduncle,
dep h 10.1-18.6% SL ( s. 7.9-9.6% SL in C. musaicus and 9.2-
10.0% SL in C. simplex); i s nuchal pla e p esen ( s. ab-
Fig. 2. Righ suspenso ium o Ta ia gy ina, INPA 20971, 28.0
mm SL. La e al iew. Abb e ia ions: aa, angulo-a icula ; dn,
den a y; hy, hyomandibula; io, in e ope cle; m ,
me ap e ygoid; op, ope cle; po, p eope cle; qu, quad a e; sb,
subp eope cle; sp, sup ap eope cle. Scale ba = 1.0 mm.
Fig. 3. Neu oc anium o Ta ia gy ina, INPA 20971, 28.0 mm
SL. Do sal iew. Abb e ia ions: ep, epioccipi al; o, single c a-
nial on anel, , on al; le, la e al e hmoid; me, mese hmoid;
na, nasal; n1, i s nuchal pla e, n2; second nuchal pla e; n3,
hi d nuchal pla e; pe, pos e io epioccipi al p ocess; ps,
pos empo al-sup aclei um; p , p e o ic; so, sup aoccipi al;
sp, spheno ic. Scale ba = 1.0 mm.

A sys ema ic e ision o Ta ia
500
sen ); maxilla sho e han o same leng h as au opala ine ( s.
longe han au opala ine). These wo species a e ince ae
sedis in Cen omochlinae (see Table 1), based in pa on in e -
ela ionships analysis by Soa es-Po o (1996), and a e he ein
p o isionally alloca ed unde Cen omochlus.
The monophyly and in e ela ionships among species o
Ta ia we e hypo hesized by Soa es-Po o (1998), who con-
side ed he clade T. gy ina plus T. c eu zbe gi as sis e o all
o he Ta ia species.
Insemina ing ep oduc i e mode (sensu Bu ns e al., 1997)
has been documen ed wi hin he Auchenip e idae ( on Ihe ing,
1937; Loi e al., 1989; Meisne e al., 2000). In he cen omochlin
ca ishes, insemina ing is supposed o ake place (Loi e al.,
1989; F anke, 1990), bu a ep oduc i e beha io is no ye
epo ed. Du ing ep oduc i e phase, he anal- in ays change
hei posi ion p obably channeling o discha ge o seminal
duc s, ac ing as a spe m conduc o du ing ep oduc ion. Such
a mo phological change was obse ed in some males o
Cen omochlus omani and also in T. nig a. The spe m p ob-
ably uns along he spoon shaped anal- in ays, and he ish
may p ess he in agains he belly o he emale, o insemina -
ing. Mo phological e idences sugges ing insemina ing ep o-
duc i e mode in Ta ia a e he in umescen emale geni al pa-
pilla (as in T. gy ina and T. nig a) and he spoon-shaped la e -
ally cu ed male anal in (obse ed in T. nig a). A de ailed analy-
sis o mo phological changes o he ep oduc i e ac needs
o be accomplished by his ological p epa a ions, and is ou o
he scope o p esen con ibu ion.
Dis ibu ion. Species o Ta ia a e b oadly dis ibu ed in he
egions eas o he Andean Co dille as (Fig. 6). Ta ia anges
om he Gua apiche i e , O inoco basin, in Venezuela, o he
Pelo as i e , U uguay d ainage, in B azil. The g oup is ound
in majo con inen al i e basins, such as he Amazon, O inoco
and Pa aná-Pa aguay; as well as in he Essequibo and small
coas al d ainages in F ench Guiana, Su iname and Amapá and
Ma ajó island in no he n B azil. Al hough well dis ibu ed in
cis-Andean Sou h Ame ica, he g oup is no known om ce -
ain d ainage basins, including he São F ancisco and small
coas al i e d ainages o no heas e n and a eas e n B azil.
Ta ia is ound, howe e , in headwa e s o Pa aíba do Sul ba-
sin, São Paulo S a e. The absence o Ta ia om ce ain a eas
may be a unc ion o inadequa e collec ions om hose i e
basins o may e lec ac ual disjunc dis ibu ions o he g oup
as whole.
Species Accoun
Ta ia aulopygia (Kne , 1857)
Figs. 7-9
Cen omochlus aulopygius Kne , 1857: 432, Pl. 8 ( ig. 26) [ ype
locali y: Guapo é i e ]. -S eindachne , 1876: 664-665 [no
locali y]. Eigenmann & Eigenmann, 1890: 270 [ci a ion].
Pea son, 1937: 110 [Mamo é i e d ainage].
Ta ia aulopygia. Mi anda Ribei o, 1911: 361 [gene ic desig-
na ion]. Gosline, 1945: 10 [lis ing]. Mees, 1974: 59-63 [no es
and dis ibu ion, in pa , Maciel, Guapo é i e ]. Sands,
1984: 38-39 [lis ing]. Lauzanne & Loubens, 1985: 112
[Mamo é i e d ainage]. Bu gess, 1989:242 [lis ing].
Soa es-Po o, 1995:204 [ci a ion]. Soa es-Po o, 1998: 331-
350 [ci a ion]. Lasso e al., 2001:97 [ci a ion]. Fe a is,
2003:476 [checklis ]. Fe a is, 2007: 77 [checklis ].
Cen omochlus in e medius. Fishe , 1917: 422 [in pa , Maciel,
Guapo é i e ]. Pea son, 1937: 110 [Mamo é i e d ainage].
Diagnosis. Ta ia aulopygia is dis inguished om all o he
species o Ta ia by ha ing a sho c anial on anel, wi h he
opening es ic ed o he on als (Fig. 8); a geni al papilla
Fig. 4. Righ hyoid a ch o Ta ia gy ina, INPA 20971, 28.0 mm
SL. Medial iew. Abb e ia ions: ac, an e io ce a ohyal; b ,
b anchios egal ays; dh, do sal hypohyal; ic, in e ce a ohyal
ca ilage; ih, in e hyal; pc, pos e io ce a ohyal; h, en al
hypohyal; uh, u ohyal. Scale ba = 1.0 mm.
Fig. 5. Pha yngeal a ches o Ta ia gy ina, INPA 20971, 28.0
mm SL. Do sal iew. Abb e ia ions: ab, an e io ( i s )
basib anchial; bb2, basib anchial 2; bb3, basib anchial 3; bb4,
basib anchial 4; cb1-5, ce a ob anchials 1 o 5 (an e io o
pos e io ); eb1-4, epib anchials 1 o 4 (an e io o pos e io );
hb1-3, hypob anchials 1 o 3; pb3, pha ingob anchial 3; pb4,
pha ingob anchial 4. Scale ba = 1.0 mm..
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 501
wi h hick lap o skin a ound he de e en duc in males (Fig.
9); and a no ched anal in in adul males (Fig. 9). The species
also is dis inguished om congene s by a combina ion o
cha ac e is ics: nasal bone pa ially su u ed o la e al ma gin
o mese hmoid; anal in wi h 7-8 b anched ays; 10-11 ibs;
38-39 pos -Webe ian e eb ae. Addi ional ea u es use ul o
dis inguishing his species include: hi d nuchal pla e well
p ojec ed la e ally wi h cu ed ip; and caudal- in lobes o
ma u e emales simila in leng h, bu ma u e males wi h sligh ly
elonga ed uppe lobe.
Desc ip ion. Measu ed adul specimens 75.2-159.0 mm SL;
mo phome ic da a p esen ed in Table 2. Body deep, head
sligh ly dep essed do so en ally. Head la ge, obus , ou -
line o head in do sal iew somewha ellip ical, b oade han
long. Do sal ou line o unk om do sal- in base o caudal
peduncle inc easingly comp essed pos e io ly. La e al p o-
ile o head om snou ip o abo e ope cula ma gin sligh ly
con ex o pec o al- in inse ion. Ven al p o ile o head and
abdomen la . Ven al p o ile o body gen ly cu ed, conca e
behind anal- in o igin. Head in egumen hin, c anial oo is-
ible; adipose eye lid well-de eloped; eye do sola e ally lo-
ca ed in an e io po ion o head; mou h e minal, uppe lip
ex ended pos e ola e ally as well-de eloped leshy ic al old;
an e io nos il ubula , loca ed on an e io bo de o snou ,
abo e lip; pos e io nos il la ge, ounded, limi ed by small
skin lap; ans e se dis ance be ween an e io nos ils sligh ly
sho e han dis ance be ween pos e io ones. Maxilla y ba -
bel sho , ex ending sligh ly beyond ip o pos clei h al p o-
cess, some imes sho e ; men al ba bels sho , ips no each-
ing pec o al- in base, a anged in a c along en al su ace o
jaw; inne men al ba bel abou 65.0-75.0% leng h o ou e
men al. Pos clei h al p ocess almos eaching e ical h ough
o igin o do sal in. Caudal peduncle deep, dep h abou 14.0-
15.8% SL.
Ros al bo de o c anium b oad wi h la ge mese hmoid;
Fig. 6. D ainage map o no he n Sou h Ame ica illus a ing he geog aphic dis ibu ion o he species o Ta ia. Symbols may
ep esen mo e han one locali y. Base map by Ma ilyn Wei zman.
A sys ema ic e ision o Ta ia
502
p emaxilla unde nea h wi h synchond al a icula ion; c anial
on anel sho , o oid, bounded by on al (Fig. 8); nasal ossi-
ied wi h wide medial langes pa ially su u ed o la e al ma -
gin o mese hmoid; au opala ine ubula , o ien ed obliquely
o longi udinal axis o body; maxilla e y small, sho e han
au opala ine; p e ome expanded an e io ly wi h well de el-
oped a ow-shaped la e al p ocesses; jaws o equal size; p e-
maxilla and den a y wi h ou o i e ows o conical ee h.
Fi s nuchal pla e somewha pen agonal; second nuchal pla e
deeply conca e along la e al ma gin; hi d nuchal pla e
cu ed, p ojec ed la e ally. Epioccipi al p ocess small.
Suspenso ium, hyoid a ch and ope cula bones as in ge-
ne ic desc ip ion. Sup ap eope cle p esen as sho obus
canal bone. Six b anchios egal ays a icula ed wi h hyoid
a ch: ou wi h an e io ce a ohyal and wo wi h pos e io
ce a ohyal.
Basib anchials 2 and 3 used oge he o ming osseous
od wi h b oad ca ilaginous an e io ip; basib anchial 4 la ge
la ened and comple ely ca ilaginous; used basib anchial 2
plus 3 bo de ed la e ally by ca ilaginous head o hypob an-
chial 1, ca ilaginous head o hypob anchial 2 and ca ilagi-
nous hypob anchial 3. Basib anchial 4 bo de ed la e ally by
ca ilaginous head o ce a ob anchial 4 and pos e io ly by
ca ilaginous head o ce a ob anchial 5. Hypob anchials,
ce a ob anchials, epib anchials and pha yngob anchials as
desc ibed in gene ic desc ip ion.
Fou in ao bi al bones in incomple e se ies. In ao bi al 1
b oad wi h sho en o-la e al p ocess on an e io bo de o
eye; emaining in ao bi als hin, educed o canalicula po -
ions. In ao bi al 2 smalles , close o in ao bi al 1, ollowed
by non-ossi ied po ion o canal below eye and wo pos e io
canal bones, one long and one sho , o ming pos e io o -
bi al im. La e al line on body wi h ossi ied canal bones pos-
e io ly o e ical h ough pel ic- in o igin.
Do sal in I,5 (n=7); do sal- in spine wi h 14-16 an o se
se a ions along en i e an e io ma gin; pos e io ma gin
smoo h. Pec o al in I,5 (n=7); pec o al- in spine wi h 21-24
an o se se a ions along an e io ma gin; small se a ions
close o spine base; 14-16 e o se se a ions along pos e io
ma gin; se a ions along bo h ma gins p og essi ely la ge
owa d spine ip. Pel ic in i,5 (n=7); ma gin ounded. Adi-
pose in small, o igin on e ical h ough end o anal- in base.
Anal in iii,7-8 (n=7); anal- in p e ygiopho es in eigh od-like
p oximal adials and se en ca ilaginous dis al adials. Cau-
dal in o ked, lobes wi h ounded ips, 8+9 p incipal ays, 18-
20 uppe p ocu en , 17-20 lowe p ocu en ays (n=7). Pleu-
al ibs 10-11 a ached o consecu i e e eb ae. Pos -
Webe ian e eb ae 38-39 (n=4).
Fig. 7. Ta ia aulopygia,(a) INPA 11080, male, 104.6 mm SL, Guapo é i e , Amazonas S a e, B azil. (b) ZMA 114.280, young
emale, 46.3 mm SL, Humai á, Amazonas S a e, B azil.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 503
Colo in alcohol. Head and body mo led wi h da k and ligh
b own pa ches o pigmen a ion, some imes o ming ain
blo ches i egula ly dis ibu ed o e sides o body (Fig. 7).
Lips and chin da k b own. Do sal in da k b own, pec o al in
usually pale b own o no pigmen ed. Adipose in pale b own,
caudal in usually wi h ans e se ba s bu young specimens
wi h whi ish spo s. Pel ic ins and belly whi ish.
Colo a ia ion. Body colo a ion in T. aulopygia is some-
wha a iable, usually wi h la ge bands o blo ches i egu-
la ly dis ibu ed o e sides o body. In some specimens col-
o a ion is uni o mly da k o pale b own, wi h caudal in mo led.
Some young specimens ha e small spo s o e en ola e al
pa s o body and a ba ed caudal in.
Mees (1974) no ed he p esence o longi udinal pale s eaks
on he body and obse ed i egula pa ches o pigmen a ion
on specimens om he Guapo é i e , bu all specimens a ail-
able o him a e now e y aded and unsui able o accu a e
desc ip ion o colo a ion. Recen expedi ions o he Guapo é
i e p o ided mo e adul specimens o T. aulopygia, on which
he abo e desc ip ion is based.
Sexual dimo phism. Based on examina ion o gonads, T.
aulopygia a ains sexual ma u i y abo e 80 mm SL. In ma u e
emales a geni al papilla is no e iden . The geni al papilla o
ma u e males is isible, wi h a hick skin lap a ound he de -
e en duc . The anal in o ma u e males (Fig. 9) is s ongly
modi ied, wi h he h ee unb anched and i s wo b anched
ays enla ged and hickened. The i s unb anched anal- in
ay is immedia ely p eceded by a egumen a y keel. The sec-
ond unb anched ay is in e media e in size be ween he neigh-
bo ing i s and hi d ays. The hi d unb anched ay is he
longes , o ming a minu e poin ed in ip (Fig 9, uiii); dis al
segmen s a e smalle , and an o sely cu ed (Fig. 9, ac). The
i s b anched ay is sligh ly cu ed owa ds he in ip, bea -
ing e o sely cu ed dis al segmen s (Fig. 9, c). The ou h
and i h b anched ays a e sho e , o ming a cen al no ch in
dis al ma gin o he anal in. The pos e io b anched ays a e
no mally de eloped and no educed.
The hemal spines in e digi a ing wi h anal- in
p e ygiopho es a e hick in males; bu hose hemal spines a e
undi e en ia ed in emales. Caudal- in lobes a e o compa-
able leng h in ma u e emales, whe eas uppe lobe is mo e
elonga ed in ma u e males.
Specimens o T. aulopygia his o ically housed in collec-
ions a e mos ly ju eniles, wi h only one syn ype bea ing a
modi ied anal in. In o ma ion on male anal- in mo phology
was imp o ed by ecen cap u es o adul specimens in he
Amazon basin.
Table 2. Mo phome ic da a o Ta ia aulopygia. SD = s an-
da d de ia ion.
Range Mean SD N
S anda d leng h 75.2-159.0 92.4 7
Pe cen s o s anda d leng h
Body dep h 15.2-20.3 17.8 1.69 7
Body wid h 18.7-23.5 20.4 1.96 7
Caudal peduncle dep h 14.0-15.8 14.6 0.82 7
Caudal peduncle leng h 23.4-28.4 25.9 1.98 7
P edo sal leng h 29.3-33.3 31.7 1.37 7
P eanal leng h 64.4-70.6 67.5 1.99 7
P epel ic leng h 46.7-51.8 49.4 1.69 7
Do sal o igin o pec o al o igin 19.1-24.2 21.8 1.89 7
Do sal o igin o pel ic o igin 26.0-29.5 27.2 1.25 7
Pec o al o igin o pel ic o igin 30.5-35.5 32.3 1.85 7
P epec o al leng h 17.8-22.5 20.0 1.60 7
Do sal- in base leng h 6.8-11.8 9.5 1.91 7
Adipose- in base leng h 7.9-12.7 10.1 1.92 7
Anal- in base leng h 3.3-10.8 7.4 3.39 7
Do sal- in spine leng h 16.0-21.2 18.6 1.92 7
Pec o al- in spine leng h 19.5-24.0 22.1 1.91 7
Pos clei h al p ocess lengh 15.8-19.4 16.5 1.86 7
Fi s b anched pel ic- in ay 10.7-15.3 13.2 1.67 7
Longes anal in ay 5.2-12.5 8.0 2.58 7
Maxilla y ba bel leng h 22.0-27.3 24.7 1.90 7
Ou e men al ba bel leng h 7.3-13.3 9.0 1.88 7
Men al ba bel leng h 4.1-10.0 5.6 1.92 7
Head leng h 21.8-26.6 23.4 1.80 7
Pe cen s o head leng h
Head wid h 75.2-80.2 78.1 1.84 7
Snou dep h 41.5-45.9 44.0 1.79 7
In e o bi al dis ance 53.2-58.9 57.7 1.94 7
Le in e na ial wid h 21.1-25.0 22.9 1.36 7
An e io in e na ial dis ance 38.4-43.8 40.8 1.81 7
Pos e io in e na ial dis ance 34.5-39.5 37.8 1.69 7
Snou leng h 36.6-41.2 38.5 1.89 7
O bi al diame e 23.1-26.5 24.6 1.51 7
Mou h wid h 50.4-55.5 53.1 1.94 7
Fig. 8. Neu oc anium o Ta ia aulopygia, INPA 11080, 80.0
mm SL. Do sal iew. Abb e ia ions: ep, epioccipi al; o, single
c anial on anel, , on al; le, la e al e hmoid; me, mese hmoid;
na, nasal; n1, i s nuchal pla e, n2; second nuchal pla e; n3,
hi d nuchal pla e; pe, pos e io epioccipi al p ocess; ps,
pos empo al-sup aclei h um; p , p e o ic; so, sup aoccipi al;
sp, spheno ic. Scale ba = 1.0 mm.
A sys ema ic e ision o Ta ia
510
usually wi h 3-5 sepa a ed de mal segmen s (lepido ichia). A
segmen ed i s unb anched ay was obse ed in mos
Su iname specimens, bu no in all, and may be associa ed o
egional di e en ia ion. The i s unb anched anal- in ay is
immedia ely p eceded by a egumen a y keel (Fig. 15, k). The
second unb anched ay has an in e media e size be ween he
neighbo ing i s and hi d ays. Thi d unb anched is he long-
es ay o ming a long sha p poin ed in ip oge he wi h he
i s b anched (Fig. 15, uiii). Thi d unb anched dis al seg-
men s a e an o sely cu ed (Fig. 15, ac). Fi s b anched ay
bea ing e o sely cu ed dis al segmen s (Fig. 15, c). Pos e-
io b anched ays a e no mally de eloped and p og essi ely
sho e ; wi h las ay educed (Fig. 15, b7).
Hemal spines 16-19 in e digi a e wi h he anal- in
p e ygiopho es; hemal spines 15-17 o 16-18 a e hickened in
ma u e males, bu undi e en ia ed in emales. The caudal- in
lobes ha e he same leng h in ma u e emales, whe eas uppe
lobe is elonga ed in ma u e males.
Dis ibu ion. Ta ia b unnea was desc ibed om he Su iname
and Ma owijne-Ma oni i e basins in Su iname. I was e-
co ded in F ench Guiana om he Ma oni and Sinnama y i e
basins. In B azil i occu s in he Neg o i e d ainage and a a
single locali y in Cen al Amazon basin, in he T ombe as
i e d ainage (Fig. 6).
Rema ks. Un il ecen ly, T. b unnea was hough o be e-
s ic ed o Su iname (Fe a is, 2003), al hough i was p e i-
ously eco ded in he Neg o i e as well (Wallace, 2002). The
o e all colo a ion o T. b unnea essembles ha o T. dunni,
om uppe Amazon. De ails in colo a ion o caudal in helps
o dis inguish be ween hese wo species, as in T. b unnea
he caudal in is whi ish wi h da k spo s o bands ( s. da ke
wi h whi ish blo ches in T. dunni). Addi ional dis inc ions
include a wide head in T. b unnea, 86.6-93.4% HL ( s. na ow
inT. dunni, 76.0-80.1% HL); a wide mou h, 54.0-59.7% HL ( s.
na ow, 48.1-52.3% HL); and a male modi ied anal in wi h
sha ply poin ed dis al ip ( s. sho ounded dis al ip). Ta ia
b unnea occu s in sympa y wi h T. in e media in some i -
e s in Su iname and F ench Guiana and also in he T ombe as
i e .
Ma e ial examined. 128 specimens (16.2-97.4 mm SL). Holo-
ype. Su iname: RMNH 26196, 56.5 mm SL, Compagnie s eam
(holo ype o Ta ia b unnea). Pa a ypes. Su iname: AMNH 58390,
1 (54.6 mm SL) (R), Kamaloe s eam, igh ma gin o Ma owijne
i e . RMNH 26197, 3 (47.6-60.0 mm SL), Compagnie s eam;
RMNH 26198, 3, 1 CS (35.1-40.3 mm SL), Kwambaolo s eam,
nea dam; ZMA 105.526, 4 (26.5-36.6 mm SL), G an i e , 63 Km
sou h o A obakka; ZMA 105.860, 1 (41.5 mm SL) (R), Sa a s eam,
abou 27 Km sou h o dam; ZMA 105.849, 7 (27.0-52.2 mm SL)
(R), Maka s eam, ibu a y o Lawa i e , Ma owijne dis ic
(pa a ypes o Ta ia b unnea). Non- ype specimens: B azil:
Amazonas: ANSP 165747, 2, iga apé Cas anho, Neg o i e ; CAS
76790, 2 (70.3-80.2 mm SL), Cuie as i e ; INPA 14228, 2 (96.9-
97.4 mm SL), U ubu i e , iga apé o Ga ião, Fa m Es eio, Neg o
i e basin; INPA 15989, 1 (66.6 mm SL), P esiden e Figuei edo,
U ubu i e , Neg o i e basin; INPA 16577, 1 (89.0 mm SL),
Jauape i i e , iga apé Cambina, Neg o i e basin; MCZ 52670, 2
(28.2-37.0 mm SL), Cuie as i e in isola ed pool; MZUSP 9352, 1
(18.6 mm SL), Cen al lake, le ma gin o Neg o i e be ween
Camanaú and Apeú i e s; MZUSP 31075, 1 (34.0 mm SL), Neg o
i e , Ba celos, island lake; MZUSP 44126, 1 (38.3 mm SL);
MZUSP 44258, 2 (23.4-36.7 mm SL), Neg o i e , Ana ilhanas
a chipaelago; ZMA 119.949, 1 (58.0 mm SL), Neg o i e and
ibu a ies; MZUSP 81139, 1 (78.0 mm SL), Tiquié i e , be ween
communi ies o Ca u u and Boca de Sal, Neg o i e d ainage; MZUSP
81177, 2 (36.9-59.4 mm SL), Tiquié i e , mou h o iga apé Açaí,
nea São Ped o communi y, Neg o i e d ainage; MZUSP 81250,
10, 1 CS (44.6-81.7 mm SL), Tiquié i e , be ween communi ies o
São Ped o and Ca u u ups eam om wa e alls, Neg o i e
d ainage; ZMA 119.949, 1 (58.0 mm SL), Neg o i e and ibu a ies.
Pa á: MNRJ 15332, 1 (32.7 mm SL), MNRJ 15333, 1 (44.4 mm
SL) and MNRJ 15334, 3, 1 CS (31.1-43.4 mm SL), iga apé
Sa acazinho, ibu a y o Ba a a lake, Po o T ombe as. F ench
Guiana: RMNH 28570, 3 (29.6-50.0 mm SL), Awahakiki i e ;
RMNH 28569, 2 (34.3-41.4 mm SL) and RMNH 28571, 4, 1 CS
(25-37 mm SL), Bala é s eam; RMNH 30494, 1 (70.0 mm SL),
Pe i -Sau , Sinnama y. Su iname: AMNH 58391, 2 (65.3-75.4 mm
SL), Su iname i e nea Bo opasi; RMNH 27530, 3 (26.7-28.0
mm SL), uppe Loë i e , ibu a y o Li ani i e ; RMNH 28568,
2 (42.5-45.0 mm SL), s eam below Aca ouany; RMNH 28654, 1
(51.5 mm SL), s eam below Bi ouac downs eam om Lombok
wa e alls; RMNH 28655, 1 (54.0 mm SL), ibu a y a igh ma gin
o Nicke ie i e below Blanche Ma ie Falls; RMNH 28656, 2 (50.0-
81.0 mm SL), ibu a y a igh ma gin o Kaboe i s eam, Co an jn
i e basin; RMNH 28658, 2 (45.1-74.0 mm SL), ibu a y a igh
ma gin o Kabalebo i e , abou 8 Km below A ana e o wa e alls;
USNM 226124, 1, s eam sou h o Ma api, Nicke ie dis ic ; USNM
226125, 5 (16.2-23.3 mm SL), ibu a y o Co an ijn i e , no h o
Tige Falls, Nicke ie dis ic ; ZMA 105.831, 3 (34.4-60.2 mm SL),
ibu a y o Nicke ie i e sou h o S ondansie Vallen.
Ta ia caxiuanensis, new species
Figs. 16-18
Holo ype.MPEG 9859, male (32.6 mm SL), B azil, Pa á: município
de Melgaço, Es ação Cien í ica Fe ei a Pena, Cu uá i e , Caxiuanã,
01o44’53”S 51o27’13”W, 1 No 1999, R. Ba hem and eam.
Fig. 15. Male modi ied anal in o Ta ia b unnea, RMNH
26196, holo ype, 56.5 mm SL. Le side la e al iew. Abb e ia-
ions: ac, an o sely cu ed den icula ion; b1, b anched i s
ay; b7, b anched se en h ay; dd, de e en duc ; c, e o sely
cu ed den icula ion; k, egumen a y keel; ui, unb anched
i s ay; uii, unb anched second ay; uiii, unb anched hi d
ay. Scale ba =1.0 mm.

L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 511
Pa a ypes. B azil, Pa á: MPEG 6201 (7, 1 CS, 29.3-40.7 mm SL);
MNRJ 28821 (2, 31.3-35.3 mm SL), all collec ed wi h he holo-
ype.
Diagnosis. Ta ia caxiuanensis is dis inguished om i s con-
gene s by a dis inc i e male modi ied anal in, wi h elonga e
ays, wi h no den icula ions on ay segmen s. The new spe-
cies is also dis inguished by he ollowing combina ion o
cha ac e s, as c anial on anel wide, wi h a single opening;
in ao bi al 1 wi h p onounced en al p ocess, almos each-
ing he an e o- en al bo de o eye im; nasal ossi ied as
ubes, wi h no medial langes o bone and no su u ed o
mese hmoid; i e b anched pec o al- in ays; and snou deep,
dep h mo e han 46.7% HL. Addi ional ea u es use ul o
dis inguishing T. caxiuanensis include caudal in lobes wi h
abou he same leng h in bo h adul emales and males; pos -
Webe ian e eb ae 32; and a smalle adul size, less han 50.0
mm SL.
Desc ip ion. Measu ed specimens 29.3-40.7 mm SL; mo pho-
me ic da a p esen ed in Table 5. Body slim, head sligh ly
dep essed do so- en ally. Head obus , ou line o head in
do sal iew almos ec angula , b oade han long. Do sal
ou line o unk om do sal- in base o caudal peduncle g adu-
ally comp essed la e ally. La e al p o ile o head om snou
ip o abo e ope cula ma gin sligh ly con ex un il pec o al-
in inse ion. Ven al p o ile o head and abdomen la . Ven al
p o ile o body gen ly cu ed behind anal in.
Head in egumen hick, c anial oo di icul o isualize;
well-de eloped adipose eye lid; eye la e o-do sally loca ed
in an e io po ion o head; mou h e minal, uppe lip ex ended
pos e o-la e ally as well-de eloped leshy ic al old; snou
ma gin ounded; an e io nos il ubula , loca ed on an e io
bo de o snou , abo e lip; pos e io nos il hin, ounded,
limi ed by small skin lap; ans e se dis ance be ween an e-
io nos ils p opo ionally he same as dis ance be ween
pos e io ones. Maxilla y ba bel o mode a e size, ex ending
beyond pos e io ip o pos clei h al p ocess, eaching e i-
cal h ough o igin o middle o do sal in; ou men al ba bels,
ips no eaching pec o al- in base, a anged in a c along en-
al su ace o jaw; inne men al ba bel abou 70.0-78.0% leng h
o ou e men als. Pos clei h al p ocess well de eloped, al-
mos eaching e ical h ough o igin o do sal in. Caudal
peduncle mode a ely deep, i s dep h abou 11.0-13.0% SL.
Ros al bo de o c anium b oad wi h mese hmoid b oade
han longe , p emaxilla unde nea h wi h synchond al a icu-
la ion; c anial on anel ellip ical, wi h single la ge opening,
bounded by mese hmoid and on al; nasal ossi ied, ubula ,
wi h no medial langes (Fig. 17). Au opala ine ubula , o i-
en ed obliquely o longi udinal axis o body; maxilla abou
same size o au opala ine; p e ome expanded wi h a well
de eloped a ow-shaped la e al p ocesses; jaws o equal size;
p emaxilla and den a y wi h h ee ows o conical ee h. Fi s
nuchal pla e sho , pen agonal; second nuchal pla e la e ally
conca e; hi d nuchal pla e p ojec ed la e ally, wi h p o-
nounced na ow ip. Epioccipi al p ocess small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion.Sup ap eope cle p esen
as long canal bone. Six b anchios egal ays a icula ed wi h
hyoid a ch: ou wi h an e io ce a ohyal and wo wi h pos e-
io ce a ohyal. Basib anchial 2 o ming osseous od wi h
b oad ca ilaginous an e io ip, sepa a ed om sho e
basib anchial 3.
Fou in ao bi al bones in incomple e se ies. In ao bi al 1
hin, wi h sho en o-la e al p ocess a ound an e io bo de
o eye; emaining in ao bi als hin, educed o canalicula
po ions. In ao bi al 2 smalles , close o in ao bi al 1, ol-
lowed by non-ossi ied po ion o canal below eye and by wo
pos e io canal bones much close o spheno ic, o ming ea
o o bi . La e al line on body wi h ossi ied canal bones only
nea head.
Do sal in I,5; do sal- in spine wi h 15-17 an o se se a-
ions along en i e an e io ma gin, pos e io ma gin smoo h.
Pec o al in I,5; pec o al- in spine wi h 17-20 an o se se a-
ions along an e io ma gin; 14-15 e o se se a ions along
pos e io ma gin; se a ions along bo h ma gins p og essi ely
la ge owa ds spine ip. Pel ic- in i,5, ma gin ounded. Adi-
pose in la ge, o igin on e ical h ough middle anal- in base.
Holo ype Range Mean SD N
MPEG 9859 (n=10)
S anda d leng h 32.6 29.3-40.7 32.8 10
Pe cen s o s anda d leng h
Body dep h 17.8 14.7-19.9 17.1 1.46 10
Body wid h 21.5 19.8-22.2 20.9 0.68 10
Caudal peduncle dep h 13.0 11.0-13.0 11.9 0.71 10
Caudal peduncle leng h 25.5 25.5-30.0 27.7 1.97 10
P edo sal leng h 30.7 28.5-33.8 31.0 1.52 10
P eanal leng h 72.4 62.1-72.4 65.9 3.68 10
P epel ic leng h 46.9 46.4-51.1 48.3 1.36 10
Do sal o igin o pec o al o igin 23.0 19.0-23.4 21.5 1.42 10
Do sal o igin o pel ic o igin 29.1 25.2-29.9 28.0 1.57 10
Pec o al o igin o pel ic o igin 30.4 30.0-35.6 33.0 1.92 10
P epec o al leng h 19.3 17.2-22.7 19.9 1.54 10
Do sal- in base leng h 10.1 8.6-11.4 10.1 0.89 10
Adipose- in base leng h 11.7 9.3-14.0 12.2 1.54 10
Anal- in base leng h 4.3 4.2-11.9 8.2 2.79 10
Do sal- in spine leng h 19.9 14.5-19.9 18.0 1.65 10
Pec o al- in spine leng h 20.2 16.9-22.9 21.1 1.69 10
Pos clei h al p ocess leng h 17.2 16.0-22.4 18.0 1.83 10
Fi s b anched pel ic- in ay 13.2 12.0-15.9 13.4 1.15 10
Longes anal in ay 10.7 7.4-12.3 10.5 1.38 10
Maxilla y ba bel leng h 34.4 30.0-35.5 33.4 1.97 10
Ou e men al ba bel leng h 8.9 7.3-10.6 8.9 0.86 10
Men al ba bel leng h 6.4 4.7-7.6 6.3 0.78 10
Head leng h 24.2 19.9-27.0 23.7 1.98 10
Pe cen s o head leng h
Head wid h 77.2 74.0-79.2 76.8 1.89 10
Snou dep h 48.1 46.8-50.7 48.7 1.75 10
In e o bi al dis ance 57.0 54.4-59.2 56.8 1.89 10
Le in e na ial wid h 24.1 21.5-27.6 23.4 1.74 10
An e io in e na ial dis ance 40.5 38.9-44.1 41.5 1.51 10
Pos e io in e na ial dis ance 41.8 38.3-43.4 40.4 1.52 10
Snou leng h 39.2 34.7-39.5 37.2 1.95 10
O bi al diame e 29.1 26.5-31.6 28.5 1.74 10
Mou h wid h 50.6 46.5-52.0 49.3 1.91 10
Table 5. Mo phome ic da a o Ta ia caxiuanensis. SD = s an-
da d de ia ion.
A sys ema ic e ision o Ta ia
512
Anal in iii, 7; anal- in p e ygiopho es in 8 od-like p oximal
adials and se en ca ilaginous dis al adials. Caudal in o ked,
lobes wi h ounded ips, 8+9 p incipal ays, 5-8 uppe p ocu en ,
5-6 lowe p ocu en ays. Caudal in lobes abou same leng h
in bo h adul emales and males. Nine pleu al ibs a ached o
consecu i e e eb ae. Pos -Webe ian e eb ae 32 (n=1).
Colo in alcohol. Do sal mid-line and la e o-do sal pa s o
body da k b own, wi h many small da k ch oma opho es, less
concen a ed o e lips and chin. Head and nuchal shield da k.
Ven ola e al pa s o body pale, belly whi ish. Do sal- in spine
da k pigmen ed, subsequen ays g ayish wi h da k base.
Pec o al, pel ic, anal and adipose ins hyaline. Caudal in
da k b own, wi h da k ch oma opho es on in ays.
Sexual dimo phism. Based on obse a ion o gonads, all ex-
amined T. caxiuanensis a e adul s. In emales a geni al papilla
is no e iden . The geni al papilla o ma u e males is isible,
wi h a skin ube a ound de e en duc . The anal in o ma u e
male is s ongly modi ied wi h he second and hi d un-
b anched plus i s b anched ays elonga ed and hickened,
ending in a ounded cu ed ip (Fig. 18, uiii, b1). No egumen-
a y keel p ecedes he i s unb anched anal- in ay. The sec-
ond unb anched ay is elonga ed, wi h an in e media e size
be ween he neighbo ing i s and hi d ays. The hi d un-
b anched and i s b anched ays a e he longes (Fig. 18, uiii,
b1). The dis al mos segmen s o he hi d unb anched ay a e
an o sely cu ed (Fig. 18, ac). The second b anched ay has
a na ow ip. Pos e io b anched ays a e p og essi ely sho e
and no educed.
Dis ibu ion. Known only om he ype locali y in Cu uá
i e , lowe Amazon basin, abou 330 km sou hwes om he
ci y o Belém, Pa á, B azil (Fig. 6).
E ymology. The speci ic name, caxiuanensis, is in e e ence
o he Flo es a Nacional de Caxiuanã, a p ese a ion a ea whe e
he ca ishes we e caugh .
Rema ks. Ta ia caxiuanensis is one o he smalles Ta ia,
eaching ma u i y a 29.3 mm SL. Only T. gy ina eaches ma-
u i y a a smalle size (28 mm SL).
Fig. 16. Ta ia caxiuanensis.(a) holo ype, MPEG 9859, male, 32.6 mm SL. (b) pa a ype, MPEG 6201, emale, 40.7 mm SL,
Caxiuanã, Pa á S a e, B azil.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 513
Ta ia dunni (Fowle , 1945)
Figs. 19-21
Cen omochlus in e medius. S eindachne , 1882: 4 [Ju aí and
Ja ua ana]. Eigenmann & Eigenmann, 1888: 156 [Tajapu u,
Te é, Ja ua ana, Icá, Ju aí, Lago Aleixo]. Eigenmann & Eigen-
mann, 1890: 269 [Amazonas, Solimões and ibu a ies].
Cen omochlus aulopygius. Eigenmann & Eigenmann, 1891:
34 [in pa , Amazonas, Solimões and ibu a ies]. Mi anda
Ribei o, 1968:10, ig. IX [Amazonas].
Ta ia in e media. Mi anda Ribei o, 1911: 360 [in pa , Ju aí,
Ju ua ana, Tajapu u, Te é, Içá, Lago Aleixo]. Gosline, 1945:
10 [Amazonas, Solimões and T ibu a ies]. Sands, 1984: 37
[ e e ence]. O ega & Va i, 1986:14 [ e e ence]. Bu gess, 1989:
242, pl. 113 [Amazon]. Soa es-Po o, 1998: 333 [ci a ion].
Cen omochlus dunni Fowle , 1945b: 111, igs. 11-13 [ ype
locali y: Colombia, Mo elia, Caque á i e d ainage].
Fowle , 1951: 462 [uppe Amazon, Colombia]. Rössel, 1962:
20 [no locali y]. Lüling, 1963: 50, ig. 15 [Quis o Cocha,
Iqui os]. Böhlke, 1984:24 [ e e ence].
Ta ia aulopygia. Mi anda Ribei o, 1962: 10 [Amazonas].
Ta ia dunni. Fe a is, 2003:476 [checklis ]. Fe a is, 2007: 77
[checklis ].
Diagnosis. Ta ia dunni is dis inguished by a unique combi-
na ion o cha ac e is ics: nasal ossi ied wi h na ow medial
langes pa ially su u ed o la e al ma gin o mese hmoid; anal-
in ays iii, 6-7; modi ied anal in in ma u e males wi h sho
ip; i s unb anched anal- in ay wi h segmen s used; las
b anched ay no mally de eloped, no educed (Fig. 21); ibs
10; and e eb ae 35. Addi ional ea u es use ul o dis in-
guishing T. dunni include de ails in colo a ion, as: con ou o
mou h da k, same colo as head; pos e io bo de o nuchal
shield usually da k; pec o al- in spine usually da k b own;
body colo a ion wi h i egula blo ches o s ipes, la ge speci-
mens wi h ain s ipes o blo ches; and caudal in da k wi h
whi ish blo ches.
Desc ip ion. Measu ed adul specimens 66.1-114.5 mm SL;
mo phome ic da a p esen ed in Table 6. Body deep, head
dep essed do so- en ally. Head obus , ou line o head in
do sal iew somewha ellip ic, sligh ly b oade han long.
T unk om do sal- in base o caudal peduncle becoming
g adually comp essed la e ally. Ou line o head in do sal iew
om snou ip o ope cula ma gin sligh ly con ex un il pec-
o al- in inse ion. Ven al p o ile o head and abdomen sligh ly
con ex. Ven al p o ile o body comp essed behind anal- in
o igin. Head in egumen hin, c anial oo isible; well-de el-
oped adipose eye lid; eye la e o-do sally loca ed in an e io
po ion o head; mou h e minal, uppe lip ex ended pos e o-
la e ally as well-de eloped leshy ic al old; an e io nos il
ubula , loca ed on an e io bo de o snou ; pos e io nos il
la ge, ounded, limi ed by small skin lap; ans e se dis ance
be ween an e io nos ils p opo ionally he same as dis ance
be ween pos e io ones in HL. Maxilla y ba bel sho , ex end-
ing close o pos e io ma gin o pos clei h al p ocess; men al
ba bel sho , ips no eaching pec o al- in base; inne men al
ba bel abou 60.0-90.0% leng h o ou e men als. Pos clei h al
p ocess almos eaching e ical h ough middle o end o
do sal in. Caudal peduncle deep, dep h abou 14.1-18.6% SL.
Ros al bo de o c anium b oad wi h mese hmoid b oade
han long; p emaxilla unde nea h wi h synchond al a icula-
ion; c anial on anel ellip ical, bounded by mese hmoid and
on al (Fig. 20); nasal ossi ied wi h na ow medial langes
pa ially su u ed o la e al ma gin o mese hmoid; au opala ine
ubula , o ien ed obliquely o longi udinal axis o body; max-
Fig. 17. Neu oc anium o Ta ia caxiuanensis, new species,
MPEG 6201, 29.3 mm SL. Do sal iew. Abb e ia ions: ep,
epioccipi al; o, single c anial on anel, , on al; le, la e al
e hmoid; me, mese hmoid; na, nasal; n1, i s nuchal pla e,
n2; second nuchal pla e; n3, hi d nuchal pla e; pe, pos e io
epioccipi al p ocess; ps, pos empo al-sup aclei h um; p ,
p e o ic; so, sup aoccipi al; sp, spheno ic. Scale ba = 1 mm.
Fig. 18. Male modi ied anal in o Ta ia caxiuanensis, new
species, MPEG 6201, 29.3 mm SL. Le side la e al iew. Ab-
b e ia ions: as, an o sely cu ed segmen ; b1, b anched i s
ay; b7, b anched se en h ay; dd, de e en duc ; ui, un-
b anched i s ay; uii, unb anched second ay; uiii, un-
b anched hi d ay. Scale ba =1.0 mm.
A sys ema ic e ision o Ta ia
514
illa abou same size o au opala ine; p e ome expanded an-
e io ly wi h well de eloped a ow-shaped la e al p ocesses;
jaws o equal size; p emaxilla and den a y wi h h ee o ou
ows o conical ee h. Fi s nuchal pla e apezoid; second
nuchal pla e sligh ly conca e along la e al ma gins; hi d
nuchal pla e cu ed, p ojec ed la e ally, wi h b oad ip.
Epioccipi al p ocess e y small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion. Sup ap eope cle p esen
as sho canal bone. Six b anchios egal ays a icula ed wi h
hyoid a ch: ou wi h an e io ce a ohyal and wo wi h pos e-
io ce a ohyal; las wo la ened and expanded.
Fi e in ao bi al bones in incomple e se ies. In ao bi al 1
la ened wi h sho en o-la e al p ocess; emaining
in ao bi als hin, educed o canalicula po ions. In ao bi als
2 and 3 close o in ao bi al 1, ollowed by non-ossi ied po ion
o canal below eye and by wo pos e io sho canal bones,
o ming pos e io o bi al im. La e al line on body wi h ossi ied
canal bones pos e io ly o e ical h ough pel ic in o igin.
Do sal in I,4-5, a ely I,4 (n=13); do sal- in spine wi h 13-
15 an o se se a ions along en i e an e io ma gin; pos e io
ma gin smoo h. Pec o al in I,5 (n=13); pec o al- in spine wi h
17-19 an o se se a ions along an e io ma gin; 11-13 e o se
se a ions along pos e io ma gin; se a ions along bo h ma -
gins p og essi ely la ge owa ds spine ip. Pel ic- in i,5
(n=13), ma gin ounded. Adipose in small, o igin on e ical
h ough end o anal- in base. Anal in iii, 6-7 (n=13); anal- in
p e ygiopho es in 8 od-like p oximal adials and se en ca i-
laginous dis al adials. Caudal in o ked, lobes wi h ounded
ips, 8+9 p incipal ays, 11-18 uppe p ocu en , 11-20 lowe
p ocu en ays (n=13). Pleu al ibs 10, a ached o consecu-
i e e eb ae. Pos -Webe ian e eb ae 35 (n=5).
Colo in alcohol. Colo a ion in p ese ed specimens umbe ,
wi h a he long longi udinal pale g ayish o whi ish i egula
blo ches; da k ma kings o m e icula ions; unde su ace o
head and abdomen whi ish; ins wi h da ke ma kings, mo e
con as ed on caudal (Fowle , 1945b: 111). Do sal su ace o
head almos da k b own; pos e io pa o nuchal shield ain ,
usually ligh b own. Con ou o lips and an e io nos ils usu-
ally da k. Ven al pa o head da k b own a ound chin ba -
bels. Caudal in da k wi h whi ish spo s o e lobes.
Colo a ia ion. Ta ia dunni usually has ain i egula bands
Fig. 19. Ta ia dunni.(a) ANSP 71705, holo ype, male, 83.3 mm SL, Caque á i e d ainage, Colombia. (b) FMNH 88187, emale,
77.5 mm SL, Conambo i e , Ecuado .
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 515
and spo s o e body. Some T. dunni, om he Pe u ian Ama-
zon, ha e a da k b own body and mo led anal in (INHS
43890). La ge specimens om he Pu us i e , in cen al
Amazonia, usually ha e a comple ely da k body (INPA 17182)
o a ligh ly spo ed colo a ion (INPA 17184).
Sexual dimo phism. Based on examina ion o gonads, T. dunni
a ains sexual ma u i y abo e 70 mm SL. In ma u e emales a
geni al papilla is no e iden . The geni al papilla o ma u e male
is isible, wi h an eme gen de e en duc . The anal in o ma-
u e male (Fig. 21) is s ongly modi ied, wi h h ee hickened
unb anched ays. The i s unb anched anal- in ay has seg-
men s used and is immedia ely p eceded by a sho egumen-
a y keel (Fig. 21, k). The second unb anched ay has an in e -
media e size be ween he neighbo ing i s and hi d ays. Thi d
unb anched is he longes ay o ming a sho ounded ip (Fig.
21, uiii); dis al segmen s smalle , an o sely cu ed (Fig. 21, ac).
Fi s b anched ay bea ing e o sely cu ed dis al segmen s
(Fig. 21, c). Pos e io b anched ays a e p og essi ely sho e ,
wi h he las b anched one somewha educed (Fig. 21, b6).
Hemal spines 15-18 in e digi a e wi h anal- in
p e ygiopho es; wi h hemal spines 15-17 hickened in ma u e
male, bu a e undi e en ia ed in emales. The caudal- in lobes
ha e he same leng h in ma u e emale, whe eas uppe lobe is
sligh ly elonga ed in ma u e male.
Dis ibu ion. Ta ia dunni has a wide dis ibu ional ange,
occu ing in he uppe Amazon, Ecuado and Colombia and
he cen al Amazon, B azil, in Solimões and Pu us i e s. I is
he species o Ta ia ha occu s a he highes ele a ions as i
has been eco ded a 1950 m abs, in Caque á i e d ainage,
Cauca moun ains Mo elia, Colombia (Fig. 6).
Rema ks. Ta ia dunni was placed in he synonymy T.
in e media by Mees (1974). The sha ed p esence o whi e
ma kings on da k body in bo h species was conside ed wi hin
he limi s o in aspeci ic a ia ion (Mees, 1974: 68). In spi e
o simila i ies in colo a ion, T. dunni is dis inguished by i s
long p eanal leng h, 70.0-75.0% SL ( s. 65.0-69.4% SL in T.
in e media) and sho in e o bi al dis ance, 54.1-59.4% HL
( s. 60.1-63.6% HL in T. in e media). The male modi ied anal
in di e s in bo h species. In T. dunni he las b anched ay is
ma kedly educed ( s. only sligh ly sho e han penul ima e
ay in T. in e media).
Ma e ial examined. 48 specimens (28.4-114.9 mm SL). Holo-
ype. Colombia: ANSP 71705, 1 (83.3 mm SL), uppe Amazonas,
Mo elia, Caque á i e d ainage (holo ype o Cen omochlus dunni).
Pa a ype. Colombia: ANSP 71706, 1 (50.1 mm SL), Al o Amazonas,
Fig. 20. Neu oc anium o Ta ia dunni, ANSP 71706, pa a ype,
50.1 mm SL. Do sal iew. Abb e ia ions: ep, epioccipi al; o,
single c anial on anel, , on al; le, la e al e hmoid; me,
mese hmoid; na, nasal; n1, i s nuchal pla e, n2; second
nuchal pla e; n3, hi d nuchal pla e; pe, pos e io epioccipi al
p ocess; ps, pos empo al-sup aclei h um; p , p e o ic; so,
sup aoccipi al; sp, spheno ic. Scale ba = 1 mm.
Range Mean SD N
S anda d leng h 66.1-114.5 82.5 15
Pe cen s o s anda d leng h
Body dep h 15.7-26.3 19.4 2.49 15
Body wid h 17.8-24.7 21.5 1.57 15
Caudal peduncle dep h 14.1-18.6 15.2 1.49 15
Caudal peduncle leng h 23.0-28.2 24.7 1.87 15
P edo sal leng h 29.2-36.7 31.9 1.79 15
P eanal leng h 70.0-75.0 72.0 1.98 15
P epel ic leng h 49.2-56.1 51.7 1.96 15
Do sal o igin o pec o al o igin 20.2-28.2 23.6 1.81 15
Do sal o igin o pel ic o igin 27.0-33.3 29.9 2.38 15
Pec o al o igin o pel ic o igin 30.7-38.9 34.7 2.02 15
P epec o al leng h 17.5-22.9 20.5 1.56 15
Do sal- in base leng h 7.8-12.4 10.0 1.41 15
Adipose- in base leng h 6.1-12.6 9.8 1.83 15
Anal- in base leng h 5.4-9.6 7.7 1.99 15
Do sal- in spine leng h 13.8-19.7 17.0 2.25 13
Pec o al- in spine leng h 19.7-25.3 22.3 1.60 13
Pos clei h al p ocess lengh 15.6-21.6 16.9 2.27 15
Fi s b anched pel ic- in ay 11.7-16.2 13.6 1.37 15
Longes anal in ay 5.2-14.1 8.9 2.15 15
Maxilla y ba bel leng h 27.3-32.9 30.1 2.17 15
Ou e men al ba bel leng h 6.6-14.2 9.5 1.93 15
Men al ba bel leng h 3.1-8.0 5.6 1.07 15
Head leng h 21.1-29.5 24.1 2.28 15
Pe cen s o head leng h
Head wid h 76.0-80.1 78.5 1.82 15
Snou dep h 47.0-51.7 48.7 1.81 15
In e o bi al dis ance 54.1-59.4 58.2 1.83 15
Le in e na ial wid h 18.3-25.5 22.1 2.10 15
An e io in e na ial dis ance 38.3-44.8 41.7 2.16 15
Pos e io in e na ial dis ance 36.1-42.0 38.6 1.75 15
Snou leng h 36.1-42.2 39.0 2.20 15
O bi al diame e 20.0-26.4 26.4 2.60 15
Mou h wid h 48.1-52.3 51.0 1.75 15
Table 6. Mo phome ic da a o Ta ia dunni. SD = s anda d
de ia ion.

A sys ema ic e ision o Ta ia
516
Mo elia, Caque á i e d ainage (pa a ype o Cen omochlus dunni).
Non- ype specimens. B azil: Amazonas: CAS 52124, 3 (28.4-40.8
mm SL), Cas elo B anco, lago G ande, Amazonas i e ; CAS 60681,
3 (81.0-97.2 mm SL), Amazon basin; INPA 3017, 1 (69.0 mm SL),
Ca alão, Solimões i e , iga apé Madalena; INPA 18477, 3 (72.0-
87.8 mm SL), Tupinamba anas Island; INPA 17182, 1 (91.7 mm
SL), Pu us i e ; INPA 17184, 2 (98.8-103.0 mm SL), Ja i Lake,
Pu us i e ; MZUSP 44127, 1 (93.3 mm SL), Pauini. Ecuado :
AMNH 58405, 3 (50.3-94.7 mm SL). AMNH 58406, 2 (66.7-70.8
mm SL), wes e n Ecuado ; FMNH 88187, 3 (66.1-84.2 mm SL),
Conambo i e , mou h o Shiona i e , eas e n Ecuado ; FMNH
92007, 1 plus 1CS, (96.9 mm SL), Conambo i e , mou h o Shiona
i e , eas e n Ecuado ; FMNH 92008, 1 (114.9 mm SL), Co ien es
i e , Napo, eas e n Ecuado ; FMNH 103964, 5, Zancudococha
lake, Agua ico i e d ainage; FMNH 103965, 2, Blanco i e close
o Tipu ini i e ups eam om b idge; FMNH 103966, 1, laguna
G ande de Cuyabeno and Queb ada de Ho migas, Agua ico i e
d ainage; FMNH 103967, 1, Cuyabeno i e , abou 1-3 km ups eam
o mou h o Agua ico i e ; FMNH 103968, 2, ibu a y o lowe
Cuyabeno i e ; FMNH 103969, 2, lake communica ing wi h
Laga ococha i e , abou 25 Km ups eam om mou h o Agua ico
i e ; FMNH 103970, 1, Aguas Neg as i e , abou 1-2 Km ups eam
om b idge. Pe u: CAS 158882, 1 (90.8 mm SL) (R), Ampiyacu
i e , nea Pebas, Lo e o depa men ; CAS 158883, 1 (69.1 mm SL)
(R), Chancho i ule , nea Pebas, Lo e o depa men ; CAS 158884,
1 (49.2 mm SL) (R), Yaguas Yacu, nea Pebas, Lo e o depa men ;
INHS 43890, 2 (48.9-77.3 mm SL), Queb ada Shushuma, Nanay
i e d ainage, Amazon basin; USNM 124901, 1 (60.0 mm SL) (R),
Shansho c eek, Ampiyacu i e d ainage.
Ta ia galaxias Mees, 1974
Figs. 22-24
Cen omochlus aulopygius. Pelleg in, 1899: 188 [Apu é, Ven-
ezuela]. Schul z, 1944: 240 [Venezuela]. Puyo, 1949: 97
[s eams on egion o App ouage].
Ta ia aulopygia. Gosline, 1945: 10 [in pa , Apu é]. Mago-
Leccia, 1967: 225 [Venezuela]. Bu gess, 1989: 242, pl. 113,
114 [no heas e n Sou h Ame ica].
Ta ia galaxias Mees, 1974: 86-88, ig. 23. [ ype locali y:
Qui ibana i ule , O inoco Basin, Venezuela]. Sands, 1984:
36-37 [Venezuela]; Mees, 1988: 411-412 [Gua apiche i e ,
Venezuela]; Bu gess, 1989: 242 [Apu e, Venezuela]; F anke,
1990: 20-34 [no es on ep oduc ion]. Soa es-Po o, 1998:
331-350 [ci a ion]. Fe a is, 2003: 476 [checklis ]. Lasso e
al., 2004: 139 [Ca aniapo, Bi a, Me a, Suapu e, Apu e,
Cau a, Mo ichal La go, Del a, O inoco]. Fe a is, 2007: 77
[checklis ].
Ta ia in e media. Soa es-Po o, 1998: 333 [ci a ion]. Fe a is,
2003:477 [checklis ]. Lasso e al., 2004: 139 [Bi a, Apu e,
Ca oní, O inoco].
Diagnosis. Ta ia galaxias is uniquely dis inguished om
congene s by i s well de eloped pos clei h al p ocess, each-
ing almos o a e ical h ough he do sal- in o igin; a la ge
eye, 37.0-42.9% HL ( s. 17.4-31.6% in o he Ta ia); and by a
sho snou 23.1-28.9% HL ( s. 31.2-43.6% in o he Ta ia).
The species is also dis inguished by he ollowing combina-
ion o cha ac e is ics: na ow ellip ical c anial on anel; ibs
7 ( s. 8-11 in mos o he Ta ia, excep T. s iga a wi h 7 ibs
and T. gy ina wi h 5-6 ibs). Addi ional ea u es use ul o
dis inguishing T. galaxias include: nasal ossi ied wi h wide
medial langes pa ially su u ed o la e al ma gin o
mese hmoid; colo a ion a iable, sides o body usually da k
wi h ligh , ounded spo s o do s, o some imes uni o mly
pale b own; oo hed p e ome in examined la ge adul speci-
mens.
Desc ip ion. Measu ed adul specimens 54.2-125.6 mm SL;
mo phome ic da a p esen ed in Table 7. Body slim, head
sligh ly dep essed do so- en ally. Head la ge, obus , ou -
line o head in do sal iew ellip ic, b oade han long. Do sal
ou line o unk om do sal- in base o caudal peduncle g adu-
ally comp essed pos e io ly. La e al p o ile o head om snou
ip o abo e ope cula ma gin sligh ly con ex o pec o al- in
inse ion. Ven al p o ile o head and abdomen la . Ven al
p o ile o body conca e pos e io o anal in.
Head in egumen hin, c anial oo isible; well-de eloped
adipose eye lid; eye la e o-do sally loca ed in an e io po -
ion o head; mou h e minal, uppe lip ex ended pos e o-
la e ally as well-de eloped leshy ic al old; snou ma gin
ounded; an e io nos il ubula , loca ed on an e io bo de
o snou ; pos e io nos il la ge, ounded, limi ed by small
skin lap; ans e se dis ance be ween an e io nos ils la ge
han dis ance be ween pos e io ones. Maxilla y ba bel each-
ing e ical h ough end o do sal- in; men al ba bel sho ,
ips no eaching pec o al- in base, a anged in a c along en-
al su ace o jaw; inne men al ba bel abou 50.0-67.0% leng h
o ou e men als. Pos clei h al p ocess sho , eaching e i-
cal h ough o igin o do sal in. Caudal peduncle deep, i s
dep h abou 14.2-18.3% SL.
Ros al bo de o c anium wi h mese hmoid as long as
b oad; p emaxilla unde nea h wi h synchond al a icula ion;
c anial on anel ellip ical, bounded by mese hmoid and on-
al (Fig. 23); nasal ossi ied, wi h medial langes su u ed o
la e al ma gin o mese hmoid; au opala ine ubula , o ien ed
obliquely o longi udinal axis o body; maxilla small, sho e
Fig. 21. Male modi ied anal in o Ta ia dunni, holo ype,
ANSP 71705, 83.3 mm SL. Le side la e al iew. Abb e ia-
ions: ac, an o sely cu ed den icula ion; b1, b anched i s
ay; b6, b anched six h ay; dd, de e en duc ; c, e o sely
cu ed den icula ion; k, egumen a y keel; ui, unb anched
i s ay; uii, unb anched second ay; uiii, unb anched hi d
ay. Scale ba = 1 mm.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 517
han au opala ine. P e ome expanded an e io ly, wi h well
de eloped a ow-shaped la e al p ocesses, bea ing small ee h
a ached o p ocess in a la ge specimen wi h 85 mm SL (MCNG
25983). Jaws o equal size; p emaxilla and den a y wi h wo o
h ee ows o conical ee h. Fi s nuchal pla e pen agonal;
second nuchal pla e b oad, la e ally conca e; hi d nuchal
pla e p ojec ed la e ally, wi h cu ed dis al ip. Epioccipi al
p ocess e y small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion. Sup ap eope cle la ge.
Six b anchios egal ays a icula ed wi h hyoid a ch: ou wi h
an e io ce a ohyal and wo wi h pos e io ce a ohyal; las
one la ges and expanded.
Se en o eigh in ao bi al bones in usually comple e se-
ies. Ra ely incomple ely ossi ied wi h 5 canal bones. In ao -
bi al 1 hin, wi h sho en o-la e al p ocess limi ed o an e-
io bo de o eye; emaining in ao bi als hin, educed o
canalicula po ions. In ao bi al 2-4 small, close o in ao -
bi al 1. In ao bi als 5-6 long ossi ied canals, bo de ing eye
below and pos e io ly. Las in ao bi al much small, o ming
pos e io o bi al im, in con ac o spheno ic. La e al line on
body wi h ossi ied canal bones pos e io ly o e ical h ough
pel ic o igin.
Do sal in I,5 (n=42), do sal- in spine wi h 15-18 an o se
se a ions along en i e an e io ma gin; pos e io ma gin smoo h.
Pec o al in I,5 (n=42), pec o al- in spine wi h 19-25 an o se
se a ions along an e io ma gin; 13-17 e o se se a ions along
pos e io ma gin; se a ions along bo h ma gins p og essi ely
la ge owa ds spine ip. Pel ic- in i,5 (n=42), ma gin ounded.
Adipose in mode a e in size, o igin on e ical h ough anal-
in base o igin. Anal in iii, 6-7 (n=42); anal- in p e ygiopho es
in eigh od-like p oximal adials and se en ca ilaginous dis al
adials. Caudal in o ked, lobes wi h ounded ips, 8+9 p inci-
pal ays, 18-20 uppe p ocu en , 18-20 lowe p ocu en ays
(n=42). Se en pleu al ibs a ached o consecu i e e eb ae.
Pos -Webe ian e eb ae 32-33 (n=4), a ely 30.
Colo in alcohol. Body usually da k b own wi h e enly spaced
small whi e do s; chin wi h sca e ed pigmen ; unde pa s
om chin o anus and en als unpigmen ed; pec o al in
pigmen ed ligh ly (Mees, 1974: 86). Do sal in wi h da k base
and i egula spo s; caudal in usually wi h whi ish spo s
i egula ly dis ibu ed o e lobes (Fig. 22).
Colo a ia ion. Al hough he body usually is da k b own
wi h whi ish spo s, T. galaxias may be polymo phic o colo
pa e n, as a la ge a ay o a ia ion ega ding colo is ob-
se ed. Some specimens ha e a ligh b own body. Some ha e
small oundish spo s o e he body and caudal in, a common
pa e n in e y young specimens < 40 mm SL. Some ha e
Fig. 22. Ta ia galaxias.(a) ANSP 138105, male, 64.6 mm SL. (b) ANSP 138105, emale, 54.2 mm SL, El Vien o i ule , Colombia.
A sys ema ic e ision o Ta ia
518
caudal in wi h da k ips. All dis inc pa e ns may be ound
oge he in he same locali y.
Sexual dimo phism. Based on examina ion o gonads, T.
galaxias a ains sexual ma u i y abo e 54.2 mm SL. In ma u e
emales a geni al papilla is no e iden . A geni al papilla is
isible in ma u e male, wi h a long slende eme gen de e en
duc (Fig. 24, dd). The male anal in is s ongly modi ied, wi h
he an e io ays enla ged and hickened. The i s unb anched
anal- in ay ha e segmen s used and is p eceded by a egu-
men a y keel (Fig. 24, k). The second unb anched ay is elon-
ga ed, wi h an in e media e size be ween he neighbo ing i s
and hi d ays. Thi d unb anched plus he i s b anched a e
he longes ays, o ming a poin ing ip. The hi d unb anched
ay has he dis al segmen s an o sely cu ed (Fig. 24, ac).
Fi s and second b anched ays ha e e o sely cu ed dis al
segmen s (Fig. 24, c). The emaining b anched ays a e p o-
g essi ely sho e , wi h he las ay no educed (Fig. 24, b7).
Hemal spines 15-18 in e digi a e wi h he anal- in
p e ygiopho es, being he hemal spines 15-17 hickened in
ma u e males, bu undi e en ia ed in emales. The caudal- in
lobes ha e he same leng h in ma u e emales, whe eas in
males he uppe lobe is sligh ly elonga ed.
Dis ibu ion. Endemic o he O inoco basin in Venezuela, in-
cluding Me a d ainage, in Colombia (Fig. 6).
Rema ks. Mees (1974) desc ibed T. galaxias on he basis o
i s diagnos ic colo pa e n o small whi e spo s e enly spaced
o e a da k body (pa e n exempli ied in Fig. 22). Mees, (1974:
88) conside ed i s colo a ion eminiscen o ha o T.
in e media, excep in T. galaxias he body is da ke , he spo s
smalle , and he ail da k wi h whi e spo s ( s. pale wi h da k
spo s and c oss-ba s in T. in e media). We obse ed he da k
body o be geog aphically a iable in bo h T. galaxias and T.
in e media. In acidic black wa e i e s specimens o T.
galaxias end o ha e a da ke body, when compa ed o hose
om clea wa e i e s. Toge he wi h he pa e n o small
whi e spo s, o he pa e ns o colo a ion we e ecognized
among O inoco specimens, such as uni o mly da k and da k
wi h whi e ellipsoid spo s, he la e being a es . In T. galaxias
he spo s a e usually ound, whe eas in T. in e media he
spo s a e ypically mo e ellipsoidal. Ta ia galaxias also has a
la ge eye 37.0-42.9% HL ( s. 21.4-26.3% in T. in e media) and
ewe ibs, 7, and ewe e eb ae 32-33 ( s. 10 ibs and 35-36
e eb ae in T. in e media).
Ta ia galaxias and T. in e media a e he only species o
Fig. 23. Neu oc anium o Ta ia galaxias, RMNH 26493,
pa a ype, 55.6 mm SL. Do sal iew. Abb e ia ions: ep,
epioccipi al; o, single c anial on anel, , on al; le, la e al
e hmoid; me, mese hmoid; na, nasal; n2, second nuchal pla e;
n3, hi d nuchal pla e; pe, pos e io epioccipi al p ocess; ps,
pos empo al-sup aclei h um; p , p e o ic; so, sup aoccipi al;
sp, spheno ic. Scale ba = 1.0 mm.
Range
(n=42) Mean SD N
S anda d leng h 54.2-125.6 64.7 15
Pe cen s o s anda d leng h
Body dep h 16.1-22.9 19.2 1.98 15
Body wid h 18.8-24.8 22.3 1.35 15
Caudal peduncle dep h 14.2-18.3 15.6 1.96 15
Caudal peduncle leng h 21.5-28.0 24.6 1.99 15
P edo sal leng h 28.7-38.1 32.6 1.97 15
P eanal leng h 64.0-69.9 66.3 1.95 15
P epel ic leng h 45.6-53.2 49.5 1.87 15
Do sal o igin o pec o al o igin 20.8-28.9 23.9 1.98 15
Do sal o igin o pel ic o igin 25.0-29.9 27.7 1.88 15
Pec o al o igin o pel ic o igin 29.6-35.8 32.2 1.99 15
P epec o al leng h 18.8-25.3 21.8 1.64 15
Do sal- in base leng h 7.9-12.5 10.4 1.17 15
Adipose- in base leng h 5.7-14.9 11.8 1.98 15
Anal- in base leng h 4.6-12.9 9.6 1.99 15
Do sal- in spine leng h 15.3-22.1 18.6 1.96 15
Pec o al- in spine leng h 20.0-26.8 23.1 1.97 15
Pos clei h al p ocess leng h 16.1-21.3 18.4 1.48 15
Fi s b anched pel ic- in ay 11.5-15.3 13.8 1.00 15
Longes anal in ay 6.0-20.1 11.2 2.14 15
Maxilla y ba bel leng h 22.6-39.8 31.6 3.80 15
Ou e men al ba bel leng h 5.4-11.0 8.2 1.54 15
Men al ba bel leng h 3.7-13.0 5.9 1.84 15
Head leng h 21.7-28.2 24.4 1.63 15
Pe cen s o head leng h
Head wid h 78.0-83.6 80.7 1.85 15
Snou dep h 42.3-48.6 45.5 1.92 15
In e o bi al dis ance 54.1-59.9 57.9 1.89 15
Le in e na ial wid h 18.6-28.6 22.0 1.86 15
An e io in e na ial dis ance 40.8-48.1 44.2 1.87 15
Pos e io in e na ial dis ance 39.2-45.8 42.1 1.96 15
Snou leng h 23.1-28.9 25.3 1.95 15
O bi al diame e 37.0-42.9 39.3 1.94 15
Mou h wid h 50.9-55.9 53.2 1.87 15
Table 7. Mo phome ic da a o Ta ia galaxias. SD = s an-
da d de ia ion.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 519
he genus bea ing a oo hed p e ome . Wi hin he Aucheni-
p e idae p e ome ian ee h appea in As e ophysus ba achus,
in bo h adul s and young specimens. Fe a is (1988) consid-
e ed he p esence o p e ome ian ee h in Ta ia an abe a-
ion. In ou examina ion o T. galaxias only a single specimen
exhibi ed ee h. In T. in e media, howe e , p e ome ian ee h
we e ound in all la ge specimens > 74 mm SL (n= 4). In all
o he la ge size (>100 mm SL) cen omochlin species, such as
Glanidium albescens,G. ibei oi,G. melanop e um,G.
leopa dus,Cen omochlus schul zi,C. heckelii and C.
exis ima us, he p e ome is eden ulous.
Ma e ial examined. 57 specimens (27.0-125.6 mm SL). Pa a ypes.
CAS 6568, 4 (49.0-56.6 mm SL) and RMNH 26493, 2 (48.5-55.6
mm SL) (R), Qui ibana i ule , nea Caica a, O inoco basin
(pa a ypes o Ta ia galaxias). Non- ype specimens. AMNH 58320,
2 plus 1CS, no locali y, p obably Venezuela. Colombia: ANSP
138104, 1 (43.4 mm SL); ANSP 138106, 1 (38.0 mm SL), Humaci a
a m, small s eam, p obably belonging o he complex o lake
Mozambique, Me a i e d ainage; ANSP 138105, 2 (54.2-64.6
mm SL), El Vien o i ule , Finca El Vien o sou h o Ma azul. Ven-
ezuela. Amazonas: AMNH 91381, 1 (125.6 mm SL), Ma aca i e ;
ANSP 160648, 1 (44.0 mm SL), Sipapo i e abo e Penda e; CAS
158873, 1 (40.8 mm SL), bi u ca ion on O inoco i e in Tama ama
beach. FMNH 103487, 2 (30.0-36.4 mm SL), A abapo i e close
o Sapo island, abo e San Fe nando de A abapo. MCNG 25983, 1
(85.6 mm SL), Siapa i e . MCNG 27966, 1 (39.3 mm SL), O inoco
i e in El Tig e island. Apu e: FMNH 69928, 2 (27.0-39.6 mm
SL), Cina uco i e sou h o San Fe nando de Apu e; FMNH 103487,
2 (31.5-37.5 mm SL), A abapo i e ups eam om San Fe nando
de A abapo. INHS 60322, 2 (R), Gua i ico i ule , sou h o B uzual,
Apu e i e d ainage. MCNG 7080, 1 (32.9 mm SL); MCNG 15891,
1 (43.1 mm SL), Mapo al Ri ule . Boli a : ANSP 149691, 1 (39.2
mm SL), U bani i e on oad Ma ipas-Las T inche as. ANSP
149973, 17 (28.8-44.4 mm SL), Qui ibana i ule nea Caica a,
O inoco basin. ANSP 160571, 2 (34.1-35.5 mm SL), Los Pa ones
i ule , ibu a y o Sipao i e , nea Fa m Fundo Malama. CAS
158860, 2 (28.7-32.0 mm SL); CAS 158865, 3 (43.7-58.4 mm SL);
CAS 158879, 1 (54.8 mm SL), Qui ibana i ule , nea Caica a,
O inoco basin. MCNG 1028, 2 (33.1-34.0 mm SL), Tie a Blanca
i ule , Km 82 on oad San a Elena. MCNG 16632, 2 (43.3-64.0
mm SL), Las Cla i as i e Km 85 on oad San a Elena. MCNG
15595, 1 (57.0 mm SL), Sacuima i e , ia Caica a-Pijiguaos. MCNG
15970, 1 (39.0 mm SL), Cha i ipa i e . MCNG 23254, 1 (44.8 mm
SL), Queb ada, close o C. Se apia del Nicha e i e ; MZUSP 44072,
2 (68.5-72.3 mm SL), Cocniza i ule , ibu a y o O inoco i e ,
nea Caica a del O inoco. Gua ico: INHS 61870, 5 (20.5-29.5 mm
SL) Gua iqui o i e nea Aguas Mue es, Pa que Agua o-Gua iqui o.
INHS 61940, 3 (24.4-25.0 mm SL) San Ba olo i e . INHS 69420,
4 (22.8-30.0 mm SL) San Ba olo i e wes om La Espe anza.
MCNG 1914, 3 (31.4-32.5 mm SL) Agua o i e in Ha o San Jose.
MCNG 15001, 3 plus 1CS (25.3-29.4 mm SL) San Ba olo i e ,
Pa que Agua o. MCNG 17137, 1 (28.3 mm SL) i ule in Ha o
Ba olena. MCNG 17138, 2 (25.1-32.2 mm SL) San Ba olo i e ,
Pa que Agua o. Monagas: RMNH 30502, 1 (31.0 mm SL)
Gua apiche i e . USNM 265661, 1 (30.0 mm SL) Gua guapo i ule ,
abou 500 me e s ups eam om i s mou h in O inoco i e , up-
s eam om Ba ancas.
Ta ia gy ina (Eigenmann & Allen, 1942)
Figs. 2-5, 25-26
Cen omochlus aulopygius. Eigenmann & Eigenmann, 1888:
157 [Cudajas (Codajás)]. Eigenmann & Eigenmann, 1890:
270 [dis ibu ion]. Eigenmann & Eigenmann, 1891: 34 [lis -
ing].
Cen omochlus gy inus Eigenmann & Allen, 1942: 118, pl.5
ig. 4 [ ype locali y: Pe u, Iqui os, b ook nea I aya i e ].
Gosline, 1945: 10 [lis ing]. Fowle , 1945a: 62 [Iqui os];
Fowle , 1945b: 112 [lis ing]; Fowle , 1951: 462, ig. 491 [li -
e a u e compila ion].
Cen omochlus c eu zbe gi Boeseman, 1953: 7, ig. 1c [ ype
locali y: Djaic eek (Djao s eam)]. New synonym. -Hoede-
man, 1957:151, ig. 8 [Co opina s eam nea Republiek].-
Rössel, 1962: 30 [no locali y].-Hoedeman, 1968: 148
[Su iname].
Ta ia gy ina. Mees, 1974: 74-75 [no es on holo ype and dis-
ibu ion]. O ega & Va i, 1986:14 [ e e ence]. Bu gess,
1989:242 [ e e ence]. Soa es-Po o, 1995:205 [ci a ion].
Soa es-Po o, 1998: 333 [ci a ion]. Fe a is, 2003:477
[checklis ]. Fe a is, 2007: 77 [checklis ].
Ta ia c eu zbe gi. Mees, 1974: 77-80, ig. 18 [no es and dis i-
bu ion, no he n Su iname lowlands and Cudajas
(=Codajás), B azil]. Sands, 1984: 40 [lis ing]. Mees, 1985:
241 [no he n Su iname]. Mees, 1988: 410 [Pa a s eam and
Kaboe i s eam, Su iname]. Bu gess, 1989:242 [ e e ence].
Soa es-Po o, 1995:205 [ci a ion].Chang & O ega, 1995:4
[ e e ence]. Soa es-Po o, 1998: 333 [ci a ion]. Fe a is,
2003:476 [checklis ]. Fe a is, 2007: 77 [checklis ].
Diagnosis. Ta ia gy ina is dis inguished om all o he spe-
cies o Ta ia by i s educed mese hmoid, wide han long,
(mese hmoid leng h 12.1-15% HL; wid h 17.2-19% HL); lowe
jaw sligh ly p o uding beyond uppe ; nasal ossi ied, ubu-
la , no su u ed o mese hmoid; hi d nuchal pla e educed
(Fig. 3, n3); small p e ome , wi h sho os o-la e al p ocess;
elonga e pos clei h al p ocess, some imes as long as head
leng h; ibs 5-6 ( s. 7-11 in o he Ta ia); and e eb ae, 29-30
Fig. 24. Male modi ied anal in o Ta ia galaxias, MCNG
25983, 85.6 mm SL. Le side la e al iew. Abb e ia ions: ac,
an o sely cu ed den icula ion; b1, b anched i s ay; b7,
b anched se en h ay; dd, de e en duc ; c, e o sely cu ed
den icula ion; k, egumen a y keel; ui, unb anched i s ay;
uii, unb anched second ay; uiii, unb anched hi d ay. Scale
ba = 1.0 mm.
A sys ema ic e ision o Ta ia
526
nus. I occu s in he uppe eaches o he A aguaia, Tocan ins,
Xingu, and Capim i e s, ibu a ies o he lowe Amazon o
B azil, no hwa ds o he Su iname coas al i e s and he
Essequibo i e in Guyana (Fig. 6).
Rema ks. Recogni ion o T. in e media emained di icul o
a long ime. Mees (1974: Fig. 11) mis akenly associa ed he
specimens om Suia Missu i e in Ma o G osso, B azil, as T.
aulopygia. Mees’ specimens we e young, bea ing small spo s
o e he body, and hei colo a ion con o ms wi h ha ob-
se ed o o he T. in e media.Ta ia in e media and T.
aulopygia a e compa ed unde he edesc ip ion o T.
aulopygia.
The pec o al- in spine wi h ans e se bands was consid-
e ed dis inc i e o T. in e media by Soa es-Po o (1998).
He ein his ea u e is conside ed a iable, as in some popula-
ions he spine is ba ed, some no . Soa es-Po o (1998: 341)
conside ed a ba ed spine as a de i ed ea u e sha ed be-
ween T. in e media and T. b unnea, bu such a condi ion
a ies in aspeci ically in bo h species.
Some la ge T. in e media may be supe icially simila o
la ge T. dunni, as bo h species may ha e ain ligh spo s on
he sides o he body. In spi e o his simila i y, T. in e media
di e s by ha ing a sho pos clei h al p ocess, no eaching
he e ical h ough he do sal- in o igin ( s. long, eaching
e ical h ough do sal- in o igin in T. dunni); a oo hed
p e ome in la ge specimens ( s. eden ulous in la ge T. dunni);
and a male modi ied anal in wi h 3-5 elonga e an o se cu ed
segmen s ( s. 1-3 sho an o se cu ed segmen s in T. dunni).
Ma e ial examined. 57 specimens (13.1-91.2 mm SL). Syn ype.
NMW 47355, 1 (R), B azil: Pa á: Ma abi anos (syn ype o
Cen omochlus in e medius). B azil: Amapá: MNRJ 12132, 1 (66.0
mm SL), B azil: Amapá: Ma uanum i e , ibu a y o Ma api i e ,
Macapá. MNRJ 12133, 1 (31.9 mm SL) and MNRJ 12134, 9 (40.2-
65.1 mm SL), B azil: Amapá: Apo ema i e in Fazenda Modelo do
Apo ema, ibu a y o A agua i i e . B azil: Goiás: MNRJ 5896, 2
(28.0-66.6 mm SL), B azil: Goiás: A aguaia i e , A uanã (olde
Leopoldina). B azil: Ma o G osso: MZUSP 44256, 2 (29.5-35.2
mm SL), B azil: Ma o G osso: lake Ipa u, Pa que Nacional do
Xingu. RMNH 26491, 2 (20.3-31.6 mm SL), B azil: Ma o G osso:
Suia Missu i e (=suiá miçú). B azil: Pa á: INPA 2978, 1 (45.4 mm
SL), iga apé Tucu uizinho, Tocan ins i e . INPA 11076, 1 (84.7
mm SL), Tocan ins i e . INPA 21006, 2 (50.4-56.5 mm SL),
T ombe as i e . MZUSP 9332, 1 (37.3 mm SL), iga apé Ribei a,
nea Badajós. MZUSP 43250, 15 (19.0-40.0 mm SL), lake
Be na dino, nea Badajós, Capim i e ; Km 97. MZUSP 44125, 2
(24.1 mm SL); MZUSP 44255, 5 (29.6-65.3 mm SL), Vila San ana,
Capim i e . MZUSP 44257, 7 (33.0-48.4 mm SL) and MZUSP
47503, 1, CS, iga apé Ca anandena, Capim i e . MZUSP 47507,
20 (17.2-49.0 mm SL), lake Ma ia P e a, Capim i e ; MZUSP
47508, 1 (40.0 mm SL), iga apé Ja i-Mi im, Capim i e ; MZUSP
50860, 1 (17.2 mm SL), iga apé Can ei o, nea Vila San ana, Capim
i e . MZUSP 44254, 3 (13.1-16.4 mm SL), iga apé Oxipucu,
Mocajuba, Tocan ins i e . MCZ 46032, 1 (20.8 mm SL), A a i
i e . MZUSP 50861, 1 (24.2 mm SL), A a i i e , A a i wa e alls,
Ma ajó island. MCZ 46130, 3 (37.3-40.6 mm SL); MZUSP 43139,
5 (35.2-42.0 mm SL); MZUSP 47504, 1 CS, and RMNH 26492, 1
(40.0 mm SL) iga apé Pa acu i, nea Icoa aci, Belém. Tocan ins:
INPA 20042, 1 (58.2 mm SL) A agua i i e , locali y o A i anhas in
Casea a dis ic . MZUSP 41838, 1 (55.5 mm SL), có ego Jóia,
San a Ri a do A aguaia. MZUSP 41422, 1 (57.7 mm SL), Có ego
Jaca é, San a Ri a do A aguaia. Su iname. RMNH 26195, 2 (51.3-
55.1 mm SL); RMNH 26199, 12 (38.5-50.0 mm SL), Sipaliwini.
RMNH 26201, 1 (27.4 mm SL), s eam nea ae od ome
Käyse gebe g e. ZMA 105.791, 3 (51.2-74.2 mm SL) (R), s eam
on le ma gin o Su iname i e , 2,5 km no h om Bo opasi Village,
B okopondo dis ic . Guyana. AMNH 12947, 1 (72 mm SL),
Wisma . AMNH 17643, 1 (91.2 mm SL), Cuyuni i e . CAS 24798,
1 (49.3 mm SL) Wisma i e , Deme a a, Deme a a basin.
Ta ia meesi, new species
Fig. 30-32
Holo ype. INHS 99772 (44.3 mm SL), Guyana, Po a o-Sipa uni:
lowe Po a o i e (Essequibo i e D ainage), Wa a uk ca a ac ,
05º15’31.9”N 59º24’01.0”W, 26 Oc 1998, L. M. Page, M. H. Sabaj,
J. M. A mb us e , M. Ha dman, J. H. Knou & W. S. P ince.
Pa a ypes. Guyana, Po a o-Sipa uni: AUM 28140, 14 (27.3-43.1
mm SL); INHS 49549, 9, (1 CS, 43.3 mm SL) (30.3-46.6 mm SL),
MBML 2046, 3 (32.2-42.6 mm SL), and MNRJ 28720, 3, (1 CS,
34.0-39.5 mm SL) all collec ed wi h he holo ype.
Fig. 28. Neu oc anium o Ta ia in e media, MZUSP 44255,
65.5 mm SL. Do sal iew. Abb e ia ions: ep, epioccipi al; o,
single c anial on anel, , on al; le, la e al e hmoid; me,
mese hmoid; na, nasal; n1, i s nuchal pla e, n2; second
nuchal pla e; n3, hi d nuchal pla e; pe, pos e io epioccipi al
p ocess; ps, pos empo al-sup aclei h um; p , p e o ic; so,
sup aoccipi al; sp, spheno ic. Scale ba = 1 mm.

L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 527
Diagnosis. Ta ia meesi is uniquely dis inguished om con-
gene s by i s colo a ion: do sal su ace o body, back and
uppe sides da k-b own some imes wi h depigmen ed a ea
a ound adipose in. C anial on anel wi h wo na ow open-
ings: an e io one be ween mese hmoid and on al and pos-
e io one limi ed o on al (Fig. 31). The on anel ape u es
a e sepa a ed om each o he h ough an o bi osphenoid
su u e benea h on al. The new species also is dis inguished
om congene s by a combina ion o cha ac e is ics: nasal
ossi ied as ube, wi h na ow la e al lange no su u ed o
mese hmoid; pec o al in I,4; caudal- in lobes a e he same
leng h in bo h adul emales and males. An addi ional ea u e
use ul o dis inguishing T. meesi has he smalle adul size,
less han 50 mm SL.
Desc ip ion. Measu ed adul specimens 39.5-46.6 mm SL;
mo phome ic da a p esen ed in Table 10. Body slim, head
sligh ly dep essed do so- en ally. Head obus , ou line o
head in do sal iew somewha ellip ic, b oade han long.
T unk om do sal- in base o caudal peduncle becoming
g adually comp essed la e ally. La e al p o ile o head om
snou ip o ope cula ma gin sligh ly con ex un il pec o al-
in inse ion. Ven al p o ile o head and abdomen almos
s aigh . Ven al p o ile o body gen ly cu ed, conca e be-
hind anal- in o igin. Head in egumen hin, c anial oo is-
ible; well-de eloped adipose eye lid; eye la e o-do sally lo-
ca ed in an e io po ion o head; mou h e minal, uppe lip
ex ended pos e o-la e ally as well-de eloped leshy ic al old;
snou ma gin ounded, in do sal iew; an e io nos il ubu-
la , loca ed on an e io bo de o snou ; pos e io nos il la ge,
ounded, limi ed by small skin lap; ans e se dis ance be-
ween an e io nos ils p opo ionally same as dis ance be-
ween pos e io ones. Maxilla y ba bel sho , ex ending close
o pos e io ip o pos clei h al p ocess; men al ba bel sho ,
ips no eaching pec o al- in base, a anged in a c along en-
al su ace o jaw; inne men al ba bel abou 51.0-64.0% leng h
o ou e men als. Pos clei h al p ocess long almos eaching
e ical h ough middle o end o do sal in. Caudal peduncle
mode a ely deep, dep h abou 10.4-12.3 SL.
Ros al bo de o c anium wi h mese hmoid app oxima ely
as long as b oad; p emaxilla unde nea h wi h synchond al
a icula ion; ellip ical c anial on anel, wi h wo na ow open-
ings: an e io one be ween mese hmoid and on al and pos-
e io one limi ed o on al. Fon anel ape u es sepa a ed o
each o he by su u e benea h on al, along o bi osphenoid
(Fig. 31). Nasal ossi ied, ubula , wi h na ow medial langes,
no su u ed o mese hmoid; au opala ine ubula , o ien ed
obliquely o longi udinal axis o body; maxilla abou same
size o au opala ine; p e ome expanded an e io ly , wi h well
de eloped a ow-shaped la e al p ocesses; jaws o equal size;
p emaxilla and den a y na ow wi h wo o h ee ows o coni-
cal ee h. Fi s nuchal pla e e y sho , pen agonal; second
nuchal pla e sligh ly conca e along la e al ma gins; hi d
nuchal pla e cu ed, p ojec ed la e ally, wi h b oad ip.
Epioccipi al p ocess e y small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion. Sup ap eope cle p esen
as sho canal bone. Fi e slende b anchios egal ays a icu-
la ed wi h hyoid a ch: h ee o ou wi h an e io ce a ohyal
and wo wi h pos e io ce a ohyal; las wo la ened.
Basib anchial 2 o ming osseous od wi h a b oad ca ilagi-
nous an e io ip, sepa a ed om sho e basib anchial 3.
Fi e in ao bi al bones in incomple e se ies. In ao bi al 1
hin wi h sho en o-la e al p ocess; emaining in ao bi als
hin, educed o canalicula po ions. In ao bi al 2 smalles ,
close o in ao bi al 1, ollowed by non-ossi ied po ion o
canal below eye and wo pos e io ossicles much close o
spheno ic, o ming pos e io o bi al im. La e al line on body
wi h ossi ied canal bones only close o head.
Do sal in I,5, do sal- in spine wi h 9-11 an o se se a-
ions along en i e an e io ma gin, pos e io ma gin smoo h.
Pec o al in I,4, pec o al- in spine wi h 15-18 an o se se a-
ions along an e io ma gin; 10-11 e o se se a ions along
pos e io ma gin; se a ions along bo h ma gins p og essi ely
la ge owa d spine ip. Pel ic- in i,5, ma gin ounded. Adi-
pose in small, o igin on e ical h ough end o anal- in base.
Anal in iii,7; anal- in p e ygiopho es in eigh od-like p oxi-
mal adials and se en ca ilaginous dis al adials. Caudal in
deeply o ked, lobes wi h poin ed ips, 8+9 p incipal ays, 5-8
uppe p ocu en , 5-6 lowe p ocu en ays. Pleu al ibs 9
a ached o consecu i e e eb ae. Pos -Webe ian e eb ae
34 (n=2).
Colo in alcohol. Do sal su ace o body, back and uppe
sides da k-b own some imes wi h depigmen ed a ea a ound
adipose in. Lips and an e io nos il whi ish. Head pigmen a-
ion pale, wi h pos e io nuchal pla es ligh e . Lowe sides
and en al su aces whi ish. Do sal spine da k, subsequen
ays da ke only on i s bases. Pai ed, anal and adipose ins
unpigmen ed. Caudal- in base wi h ch oma opho es becom-
ing p og essi ely mo e widely spaced, wi h hyaline lobes
and ips.
Fig. 29. Male modi ied anal in o Ta ia in e media, MZUSP
44255, 65.5 mm SL. Le side la e al iew. Abb e ia ions: ac,
an o sely cu ed den icula ion; b1, b anched i s ay; b6,
b anched six h ay; dd, de e en duc ; c, e o sely cu ed
den icula ion; k, egumen a y keel; ui, unb anched i s ay;
uii, unb anched second ay; uiii, unb anched hi d ay. Scale
ba = 1.0 mm.
A sys ema ic e ision o Ta ia
528
Sexual dimo phism. Based on examina ion o gonads, T. meesi
a ains sexual ma u i y abo e 39.5 mm SL. A geni al papilla is
no e iden in emales. The geni al papilla o ma u e male is
isible, wi h a sho de e en duc . Adul male anal in (Fig.
32) is s ongly modi ied wi h a poin ed ip. Fi s unb anched
anal- in ay hickened. Second unb anched ay wi h a sha p
ip cu ed owa ds he hi d unb anched ay, bu no p o-
longed un il he in ip (Fig. 32, uii). Thi d unb anched ay is
he longes one, o ming a na ow in ip, wi h dis al seg-
men s gen ly cu ed (Fig. 32, uiii). Fi s , second and hi d
b anched anal- in ays a e p og essi ely sho e and cu ed
owa ds he in ip. Pos e io b anched ays no mally de el-
oped; las ay no educed (Fig. 32, b7). The caudal in lobes
ha e abou same leng h in bo h adul males and emales.
Dis ibu ion. Known only om ype locali y in he Essequibo
basin, Guyana (Fig. 6).
E ymology. The speci ic name hono s D . Ge lo F. Mees, o
his signi ican con ibu ions o ou knowledge o he genus
Ta ia, especially wi h ega ds o he species o he Guyana
shield.
Rema ks. The Wa a uk ca a ac , he ype locali y o T. meesi,
consis s o apidly lowing wa e o e huge boulde s. A he
base o he ca a ac is a la ge pool and backwa e a ea wi h a
subs a e o sand, g a el and ubble o e bed ock gi e shel-
e o some ca ish species, including he iny T. meesi (L.
Page, pe s. comm.).
Among Ta ia, subdi ided on anel ape u es, wi h wo
openings sepa a ed o each o he by o bi osphenoid su u e
is obse ed only in T. meesi. Dissec ion o alcoholic speci-
mens con i med he p esence o an an e io and pos e io
on anel. The o bi osphenoid is he place o o igin o he
acial mandibula muscle adduc o a cus pala ini.Ta ia meesi
has some ea u es ha a e uncommon wi hin he genus such
as a ubula nasal bone and caudal- in lobes o equal size in
bo h males and emales. I sha es wi h T. gy ina a low numbe
o canalicula in ao bi als and a low numbe o b anchios egal
ays. I is dis inc om T. in e media, he only congene in-
Fig. 30. Ta ia meesi.(a) holo ype, INHS 99772, male, 44.3 mm SL. (b) pa a ype, INHS 49549 , emale, 46.6 mm SL, Essequibo
i e , Guyana.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 529
habi ing he Essequibo basin, due o a c anial on anel wi h
wo openings ( s. single opening in T. in e media); nasal u-
bula , od-like, no su u ed o mese hmoid ( s. wi h la e al
bony langes, su u ed o mese hmoid in T. in e media); adul
specimens small sized, less han 50 mm SL ( s. la ge adul
size, mo e han 50 mm SL).
The colo pa e n, consis ing o small ch oma opho es con-
cen a ed mainly on do sola e al pa o body in T. meesi, e-
sembles supe icially he pa e n in T. boemia. In T. meesi, how-
e e , he caudal in is hyaline s. wi h small spo s in T. boemia.
Ta ia nei ai (Ihe ing, 1930)
Fig. 33-35
Ta ia in e media. Mi anda Ribei o, 1918: 734 [Pique e &
Pi acicaba, São Paulo]. Fowle , 1951: 470 [li e a u e compi-
la ion]. Mi anda Ribei o, 1962: 10 [ e e ence].
Glanidium nei ai Ihe ing, 1930: 99, pl. 13, ig. 1 [ ype locali y:
Pi acicaba i e , Pi acicaba, São Paulo]. Gosline, 1945: 11
[lis ing]. Mi anda Ribei o, 1962: 3 [ e e ence].
Ta ia nei ai. Mees, 1974: 71-74, ig. 16 [no es and dis ibu-
ion]. Sands, 1984: 42 [lis ing]. Bu gess, 1989: 242, pl. 108
[ e e ence]. Soa es-Po o, 1998: 333 [ci a ion]. B i ski e
al., 1999: 113 [Pan anal Ma o-G ossense]. Casa i e al.,
2001:5 [uppe Pa aná i e ]. Naka ani e al., 2001: 248 [la -
al de elopmen ]. Fe a is, 2003:477 [check lis ]. Ve íssimo
e al., 2005: 7 [Manso i e ]. Fe a is, 2007: 77 [check lis ].
Ta ia c . aulopygia. Mees, 1988: 409 ig. 1 [Pa aguay]. Bu -
gess, 1989:242 [ e e ence].
Diagnosis. Ta ia nei ai is dis inguished om o he species
o Ta ia by i s unique colo pa e n wi h sca e ed small whi e
spo s o e he sides o he body and he caudal in wi h da k
e ical ba s. The species is also dis inguished by he ollow-
ing combina ion o cha ac e s: hi d nuchal pla e b oad, wi h
sho la e al p ojec ion, and pos -Webe ian e eb ae
al e na ingly ibbed and un ibbed. Addi ional ea u es use ul
o dis inguishing he species include: caudal- in lobes o
equal leng h in ma u e emales, uppe lobe mo e elonga ed in
ma u e males; 31-33 pos -Webe ian e eb ae.
Desc ip ion. Measu ed adul specimens 48.1-82.1 mm SL;
mo phome ic da a p esen ed in Table 11. Body slim, head
sligh ly dep essed do so- en ally. Head la ge, obus , do -
sal p o ile o head ellip ic, b oade han long. Do sal ou line
o unk om do sal- in base o caudal peduncle g adually
Holo ype Range Mean SD N
INHS 99772 (n=10)
S anda d leng h 44.3 39.5-46.6 43.6 10
Pe cen s o s anda d leng h
Body dep h 19.4 19.4-22.5 20.7 1.01 10
Body wid h 19.2 18.3-20.2 19.2 0.58 10
Caudal peduncle dep h 12.0 10.4-12.3 11.4 0.66 10
Caudal peduncle leng h 28.0 24.5-28.1 27.0 1.41 10
P edo sal leng h 30.5 28.8-34.9 31.7 1.87 10
P eanal leng h 71.3 62.2-71.3 65.1 3.08 10
P epel ic leng h 49.9 48.1-51.5 49.7 1.12 10
Do sal o igin o pec o al o igin 24.4 22.3-25.5 23.9 1.25 10
Do sal o igin o pel ic o igin 28.2 28.1-32.6 30.4 1.69 10
Pec o al o igin o pel ic o igin 30.0 30.0-35.4 32.8 1.37 10
P epec o al leng h 19.9 18.4-22.1 19.8 1.04 10
Do sal- in base leng h 10.6 8.4-10.9 9.8 0.87 10
Adipose- in base leng h 8.1 8.1-10.4 9.1 0.84 10
Anal- in base leng h 4.7 4.7-10.5 8.4 1.88 10
Do sal- in spine leng h 19.0 14.0-19.0 16.4 1.52 10
Pec o al- in spine leng h 25.3 20.2-25.3 21.9 1.50 10
Pos clei h al p ocess lengh 19.2 18.7-22.0 20.0 1.01 10
Fi s b anched pel ic- in ay 13.8 11.4-14.5 13.0 1.13 10
Longes anal in ay 8.4 7.4-12.6 9.5 1.71 10
Maxilla y ba bel leng h 36.1 32.2-39.6 35.8 2.36 10
Ou e men al ba bel leng h 9.3 7.1-10.5 9.3 1.07 10
Men al ba bel leng h 4.5 4.3-5.8 5.0 0.54 10
Head leng h 24.6 23.0-27.4 24.6 1.14 10
Pe cen s o head leng h
Head wid h 70.6 67.3-74.7 71.0 2.44 10
Snou dep h 40.4 40.4-44.8 42.7 1.81 10
In e o bi al dis ance 55.0 51.8-56.6 54.7 1.51 10
Le in e na ial wid h 23.9 21.7-26.1 23.7 1.57 10
An e io in e na ial dis ance 40.4 36.9-40.4 38.7 1.20 10
Pos e io in e na ial dis ance 38.5 36.9-40.6 38.7 1.18 10
Snou leng h 36.7 34.6-39.6 37.4 1.96 10
O bi al diame e 23.9 19.7-27.4 23.9 1.87 10
Mou h wid h 51.4 47.2-53.3 49.5 1.54 10
Table 10. Mo phome ic da a o Ta ia meesi. SD = s anda d
de ia ion.
Fig. 31. Neu oc anium o Ta ia meesi, new species, INHS
49549, pa a ype, 43.3 mm SL. Do sal iew. Abb e ia ions: ep,
epioccipi al; o, single c anial on anel, , on al; le, la e al
e hmoid; me, mese hmoid; na, nasal; n1, i s nuchal pla e,
n2; second nuchal pla e; n3, hi d nuchal pla e; pe, pos e io
epioccipi al p ocess; ps, pos empo al-sup aclei h um; p ,
p e o ic; so, sup aoccipi al; sp, spheno ic. Scale ba = 1 mm.
A sys ema ic e ision o Ta ia
530
comp essed pos e io ly. La e al p o ile o head om snou ip
o abo e ope cula ma gin sligh ly con ex un il pec o al- in
inse ion. Ven al p o ile o head and abdomen la . Ven al
p o ile o body conca e pos e io o anal in.
Head in egumen hin, c anial oo isible; well-de eloped
adipose eye lid; eye la e o-do sally loca ed in an e io po -
ion o head; mou h e minal, uppe lip ex ended pos e o-
la e ally as well-de eloped leshy ic al old; snou ma gin
ounded; an e io nos il ubula , loca ed on an e io bo de
o snou ; pos e io nos il la ge, ounded, limi ed by small
skin lap; ans e se dis ance be ween an e io nos ils sligh ly
la ge han dis ance be ween pos e io ones. Maxilla y ba bel
mode a e in size, ex ending o pos e io ip o pos clei h al
p ocess, some imes longe ; men al ba bel a anged in a c
along en al su ace o jaw, ips no eaching pec o al- in
base; inne men al ba bel abou 60.0-70.0% leng h o ou e
men als. Pos clei h al p ocess well de eloped, almos each-
ing e ical h ough middle o do sal in. Caudal peduncle
mode a ely deep, i s dep h abou 13.6-14.8% SL.
Ros al bo de o c anium wi h mese hmoid as b oad as
long; p emaxilla unde nea h wi h synchond al a icula ion;
c anial on anel na ow, ellip ical, bounded by mese hmoid
and on al (Fig. 33); nasal ossi ied, wi h sho medial langes
pa ially su u ed o la e al ma gin o mese hmoid; au opala ine
ubula , o ien ed obliquely o longi udinal axis o body; max-
illa small, sho e han au opala ine; p e ome expanded an-
e io ly, wi h well de eloped a ow-shaped la e al p ocesses;
jaws o equal size; p emaxilla and den a y wi h h ee o ou
ows o conical ee h. Fi s nuchal pla e sho , somewha el-
lip ical; second nuchal pla e la e ally conca e, pa ially in
con ac wi h sup aoccipi al in some specimens; hi d nuchal
pla e ela i ely s aigh , p ojec ed la e ally. Epioccipi al p o-
cess small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion. Sup ap eope cle p esen
as a e y sho canal bone. Six b anchios egal ays a icu-
la ed wi h hyoid a ch: ou wi h an e io ce a ohyal and wo
wi h pos e io ce a ohyal; las one la ges and expanded.
Fi e in ao bi al bones in incomple e se ies. In ao bi al 1
b oad, wi h sho de eloped en o-la e al p ocess a ound
an e io bo de o eye; emaining in ao bi als hin, educed
o canalicula po ions. In ao bi al 2 smalles , close o in-
ao bi al 1, ollowed by non-ossi ied po ion o canal below
eye and by h ee pos e io canal bones o ming pos e io
o bi al im. La e al line on body wi h ossi ied canal bones
limi ed o head.
Do sal in I,4-5, a ely I,4 (n=25); do sal- in spine wi h 12-
15 an o se se a ions along en i e ma gin; pos e io ma gin
smoo h. Pec o al in I,4-5, a ely I,5 (n=25); pec o al- in spine
wi h 19-24 an o se se a ions along an e io ma gin, small
se a ions close o spine base; 13-17 e o se se a ions along
pos e io ma gin; se a ions along bo h ma gins p og essi ely
la ge owa ds spine ip. Pel ic- in i,5 (n=25), ma gin ounded.
Adipose in sho , o igin on e ical h ough end o anal- in
base. Anal in iii, 6-7, a ely iii,6 (n=25); anal- in p e ygiopho es
in eigh od-like p oximal adials and se en ca ilaginous dis-
al adials. Caudal in o ked, lobes wi h ounded ips, 8+9
p incipal ays, 19-21 uppe p ocu en , 19-20 lowe p ocu en
ays (n=25). Se en o nine pleu al ibs a ached o consecu-
i e e eb ae plus one a ached o 9 h o 11 h e eb ae. The
8 h o 10 h e eb ae co espond o gap, wi h no ibs a ached.
Ribs p og essi ely small an e opos e ioly. Pos -Webe ian
e eb ae 31-33 (n=4).
Colo in alcohol. Colo pa e n usually wi h sca e ed small
whi e spo s o e la e al sides o body mainly dis ibu ed o e
en ola e al pa s. Do sal spine usually wi h ans e se
bands; pec o al- in spine no pigmen ed. Caudal in wi h i -
egula da k e ical ba s. Pos e io nuchal pla es somewha
pale. Lips and nos ils whi ish. Ven al po ion o head a ound
chin ligh b own.
Colo a ia ion. Regional a ia ion in colo a ion was obse ed
ega ding specimens om he uppe Pa aná i e immedia ely
abo e he Se e Quedas wa e alls (MHNG 2355.92). These
specimens ha e a da k body colo a ion, wi h only a ew whi e
spo s en ally, and a caudal in wi h e ical ba s. Some
specimens om he uppe Pa aguay i e in he Pan anal o
Ma o G osso a e ligh b own o e do sal pa s o he body,
wi h sca e edspo s along en ola e al pa s o body, and
wi h a e ically ba ed caudal in.
Sexual dimo phism. Based on examina ion o gonads, T.
nei ai a ains sexual ma u i y abo e 48.1 mm SL. A geni al
papilla is no e iden in emales. The geni al papilla o ma u e
male is isible, wi h a sho de e en duc . Male anal in (Fig.
35) is modi ied wi h he h ee unb anched and i s b anched
ays enla ged and hickened. Fi s unb anched ay is non-
segmen ed, and immedia ely p eceded by a egumen a y keel
(Fig. 35, k). The second unb anched ay is elonga ed, coa-
lescing owa ds in ip. The hi d unb anched and i s
b anched ays a e he longes , o ming a sho in ip. The
hi d unb anched ay has wo an o sely cu ed dis al seg-
men s (Fig. 35, al). The i s b anched ay has ou o i e
Fig. 32. Male modi ied anal in o Ta ia meesi, INHS 49549,
pa a ype, 43.3 mm SL. Le side la e al iew. Abb e ia ions:
b7, b anched se en h ay; dd, de e en duc ; ui, unb anched
i s ay; uii, unb anched second ay; uiii, unb anched hi d
ay. Scale ba = 1 mm.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 531
e o sely cu ed dis al segmen s (Fig. 35, l).The pos e io
b anched ays become p og essi ely sho e , wi h he las
ay no educed (Fig. 35, b7).
Hemal spines 14-17 a e in e digi a ing wi h he anal- in
p e ygiopho es. The hemal spines 15-17 a e hick in ma u e
males, bu undi e en ia ed in emales. The male uppe cau-
dal- in lobe is sligh ly elonga e, abou 15.0% longe han he
lowe lobe, whe eas in ma u e emales he e a e equal lobes.
Dis ibu ion. Ta ia nei ai occu s in he uppe Pa aguay i e ,
in he Pan anal o Ma o G osso, B azil, uppe Pa aná i e ,
and uppe Pa aíba do Sul i e (Fig. 6). I is eco ded om
se e al ibu a ies o he uppe Pa aná i e , B azil, such as
he G ande i e in Minas Ge ais, he Tie ê and Pa anapanema
i e s in São Paulo, and he Tibagi i e in Pa aná. In he
uppe Pa aíba do Sul i e he species was eco ded om
Pique e, São Paulo S a e. The single eco d om he São F an-
cisco i e is unce ain (MZUSP 3801) because he locali y
da a a e dubious.
Rema ks. Ta ia nei ai was obse ed in semi-lo ic en i on-
men s. Some o hese ca ishes we e ound hidden wi hin
ma ginal ege a ion o we e cap u ed inside c e ices o sub-
me sed unks (O. Shiba a pe s. comm.).
Ta ia nei ai was desc ibed based on a single specimen
om he Pi acicaba i e . The holo ype is los , bu Ihe ing
(1930: pl. 13) illus a ed he specimen as ha ing i egula ly
sc a ched colo pa e n on he sides o he body and he
caudal in wi h da k e ical ba s. This colo pa e n dis in-
guishes T. nei ai om mos Ta ia species, excep some T.
aulopygia. Va ia ion in colo a ion may occu among popula-
ions, bu he caudal in is always ba ed, and some imes
esembles ha o T. aulopygia. In spi e o his simila i y, T.
nei ai and T. aulopygia can be dis inguished on he basis o
p opo ional measu emen s, he size o c anial on anel
(bounded by mese hmoid and on al in T. nei ai s. es ic ed
o on al in T. aulopygia); he modi ied anal in o adul males
(dis al ma gin con inuous, wi hou no ch in T. nei ai s.
no ched in T. aulopygia) and he pos -Webe ian e eb ae
(31-33 in T. nei ai s. 38-39 in T. aulopygia).
Ma e ial examined. 60 specimens (23.1-82.1mm SL). B azil: Goiás:
NUPELIA unnumbe ed, 1, (58.4 mm SL), Co umbá i e , nea Ipame i,
Pi es do Rio Dis ic . Ma o G osso do Sul: MZUSP 35882, 1 (26.6
mm SL), Piqui i i e , San o An ônio do Pa aíso a m, I iqui a;
MZUSP 36364, 2 (27.7-46.8 mm SL), Co ixão, Capão G ande,
Nhecolândia, Co umbá; MZUSP 42145, 2 (23.6-33.9 mm SL), lake
abou 25 Km om Poconé, on oad o Po o Ce cado. Pa aná: MZUEL
206, 1 (56.0 mm SL); MZUEL 207, 1 (53.7 mm SL); MZUEL 1098,
1 (63.5 mm SL), Tibagi i e , município de Se anópolis. Minas Ge ais:
MNRJ 28722, 2 (61.1-76.4 mm SL), G ande i e , ups eam om
Po o Colombia dam. São Paulo: MZUSP 1993, 1 (62.1 mm SL),
Pique e; MZUSP 2000, 1 (54.4 mm SL), Pi acicaba i e ; MZUSP
35360, 1 (62.3 mm SL), A aguá i e , Bo uca u; MZUSP 42569, 2
(58.2 mm SL), Pi acicaba i e , Pi acicaba; MZUSP 42570, 1 CS
Fig. 33. Ta ia nei ai.(a) MHNG 2355.92, male, 51.6 mm SL. (b) MHNG 2355.92, emale, 53.3 mm SL, Yuguy y s eam, Pa aguay.

A sys ema ic e ision o Ta ia
532
(60.5 mm SL), lake on A ibaia i e , nea Jagua iuna; MZUSP 45099,
1 (48.9 mm SL), Tie ê i e , I ápolis; MZUSP 46009, 1 (56.0 mm
SL), Co umba aí i e , Co umba aí; MZUSP 46855, 2 (23.1-41.0
mm SL), Pa do i e , nea Guaí a; MZUSP 47505, 7 (53.1-62.1 mm
SL), ilha Sol ei a, Pa aná i e , Ilha Sol ei a dam. Pa aguay: MHNG
2159.45, 1 (56.7 mm SL), I apua: Tembey i e . MHNG 2353.30, 1
(48.0 mm SL), sal o Pi a e a, La Co dille a. MHNG 2355.92, 12
(50.7-63.7 mm SL), Caaguazu: Yuguy y s eam, abou 17 Km sou h
o Yhu. MHNG 2430.19, 1 (R) (76.1 mm SL), lake on Iguazu i e ,
uppe Pa aná. MHNG 2480.48, 1 (82.1 mm SL). MHNG 2480.49, 1
(43.6 mm SL), Canindeyú: Sal o Guai a, I aipu lake. MHNG 2537.73,
8, 1 CS, (37.2-46.6 mm SL), Taga ija-mi i e , abou 30 Km eas om
Pue o Max; UMMZ 206602, 3 (29.6-30.4 mm SL), Agua ay i e ,
San Ped o; USNM 249826, 1 (54.5 mm SL), Canindeyú: Ca apa
i e , abo e wa e alls, old oad om S oessne o Guai a; ZMA
116.801, 1 (R) (66.2 mm SL), I apua: Tembey i e , 4 Km abo e
ca a ac s.
Ta ia nig a, new species
Fig. 36-39
Holo ype. INPA 12291 (116.8 mm SL), B azil, Amazonas: Ua umã
i e , app oxima e coo dena es 02o14’35”S 58 o49’58”W, Oc 1983,
Ich hyology INPA eam.
Pa a ypes. B azil, Amazonas: INPA 25501, 1 (107.3 mm SL), col-
lec ed wi h he holo ype; INPA 11073, 1 (93.7 mm SL); INPA
11075, 1 (113.5 mm SL); INPA 11081, 6 (86.6-107.8 mm SL);
MNRJ 32024, 1 plus1 CS (97.0-98.2 mm SL), Ua umã i e ,
Samaúma lake, P esiden e Figuei edo; INPA 12296, 1 (72.0 mm SL)
Ua umã i e , iga apé Mi i i. Pa á: MNRJ 12358, 3 (70.0-82.3 mm
SL) T ombe as i e , Po o T ombe as.
Diagnosis. Ta ia nig a is uniquely dis inguished om con-
gene s by ha ing da k b own body colo (Fig. 36) and sho
pos clei h al p ocess abou 60.0% HL ( s. mo e han 80.0%
HL in o he Ta ia). Addi ional ea u es use ul o dis inguish-
ing T. nig a include: pos clei h al spine no eaching e ical
h ough o igin o do sal in; nasal wi h well de eloped medial
langes; 32 e eb ae.
Desc ip ion. Measu ed specimens 70.0-116.8 mm SL; mo -
phome ic da a p esen ed in Table 12. Body deep, head de-
p essed do so- en ally. Ou line o head in do sal iew al-
mos ec angula , sligh ly b oade han long. T unk om do -
sal- in base o caudal peduncle becoming g adually com-
p essed la e ally. La e al p o ile o head om snou ip o
ope cula ma gin con ex un il pec o al- in inse ion. Ven al
p o ile o head and abdomen sligh ly con ex. Ven al p o ile
o body conca e behind anal- in o igin.
Head in egumen hin, c anial oo isible; well-de eloped
adipose eye lid; eye la e o-do sally loca ed in an e io po -
ion o head; mou h e minal, uppe lip ex ended pos e o-
la e ally as a well-de eloped leshy ic al old; an e io nos il
ubula , loca ed on an e io bo de o snou ; pos e io nos il
la ge, ounded, limi ed by small skin lap; ans e se dis ance
be ween an e io nos ils la ge han dis ance be ween pos-
e io ones. Maxilla y ba bel o mode a e size, ex ending well
beyond pos e io ip o pos clei h al p ocess, eaching e i-
cal line angen o o igin o do sal in; men al ba bel sho ,
ips no eaching pec o al- in base, a anged in a c along en-
al su ace o jaw; inne men al ba bel abou 54.0-65.0% leng h
o ou e men als. Pos clei h al p ocess sho , almos eaching
e ical h ough middle nuchal pla es. Caudal peduncle deep,
dep h abou 14.4-17.0% SL.
Fig. 34. Neu oc anium o Ta ia nei ai, MZUSP 42570, 60.5
mm SL. Do sal iew. Abb e ia ions: ep, epioccipi al; o, single
c anial on anel, , on al; le, la e al e hmoid; me, mese hmoid;
na, nasal; n1, i s nuchal pla e, n2; second nuchal pla e; n3,
hi d nuchal pla e; pe, pos e io epioccipi al p ocess; ps,
pos empo al-sup aclei h um; p , p e o ic; so, sup aoccipi al;
sp, spheno ic. Scale ba = 1 mm.
Fig. 35. Male modi ied anal in o Ta ia nei ai, MZUEL 206,
56.0 mm SL. Le side la e al iew. Abb e ia ions: ac, an o sely
cu ed den icula ion; b7, b anched se en h ay; dd, de e en
duc ; c, e o sely cu ed den icula ion; k, egumen a y keel;
ui, unb anched i s ay; uii, unb anched second ay; uiii,
unb anched hi d ay. Scale ba = 1.0 mm.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 533
C anium wi h mese hmoid as long as b oad; p emaxilla
unde nea h wi h synchond al a icula ion; ellip ical na ow
conca i y o c anial on anel, opening bounded by
mese hmoid and on al (Fig. 37); nasal ossi ied, wi h wide
medial langes pa ially su u ed o la e al ma gin o
mese hmoid; au opala ine ubula , o ien ed obliquely o lon-
gi udinal axis o body; maxilla abou same size o au opala ine;
p e ome expanded an e io ly wi h well de eloped a ow-
shaped la e al p ocesses; jaws o equal size; p emaxilla and
den a y wi h h ee o ou ows o conical ee h. Fi s nuchal
pla e somewha pen agonal; second nuchal pla e conca e
along la e al ma gins; hi d nuchal pla e cu ed, p ojec ed
la e ally, wi h small ip. Epioccipi al p ocess e y small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion. Sup ap eope cle p esen
as sho canal bone. Six b anchios egal ays a icula ed wi h
hyoid a ch: ou wi h an e io ce a ohyal and wo wi h pos e-
io ce a ohyal; las wo la ened and expanded. Basib anchial
2 o ming osseous od wi h a b oad ca ilaginous an e io ip,
sepa a ed om sho e basib anchial 3.
Fi e in ao bi al bones in incomple e se ies. In ao bi al 1
b oad, wi h sho en o-la e al p ocess; emaining
in ao bi als hin, educed o canalicula po ions. In ao bi al
2 smalles , close o in ao bi al 1; in ao bi al 3 elonga e, ol-
lowed by sho non-ossi ied po ion o canal. Pos e io h ee
canal bones o ming pos e io o bi al im. La e al line on body
wi h ossi ied canal bones only close o head.
Do sal in I,5 (n=12); do sal- in spine wi h 19-22 an o se
se a ions along en i e an e io ma gin; pos e io ma gin
smoo h. Pec o al in I,5 (n=12); pec o al- in spine wi h 28-31
an o se se a ions along an e io ma gin; 19-21 e o se se -
a ions along pos e io ma gin; se a ions along bo h ma -
gins p og essi ely la ge owa ds spine ip. Pel ic- in i,5
(n=12), ma gin ounded. Adipose in small, o igin on e ical
h ough end o anal- in base. Anal in iii,7 (n=12); anal- in
p e ygiopho es in eigh od-like p oximal adials and se en
ca ilaginous dis al adials. Caudal in o ked, lobes wi h
ounded ips, uppe lobe sligh ly elonga ed in compa ison o
lowe lobe, 8+9 p incipal ays, 17-19 uppe p ocu en , 17-20
lowe p ocu en ays (n=12). Pleu al ibs 10 a ached o con-
secu i e e eb ae. Pos -Webe ian e eb ae 32 (n=1).
Colo in alcohol. Ta ia nig a da k colo ed in la ge adul speci-
mens. Do sal mid-line and do sola e al pa s o body da k
b own, as well as do sal su ace o head, nuchal shield, do -
sal in and adipose in (Fig. 36). Body p og essi ely pale
owa ds en al pa s. Ven al po ion o head a ound chin,
lips and an e io nos ils ligh b own. Belly whi ish o o igin
o pel ic in. Pec o al, pel ic and anal ins ligh b own. Cau-
dal in da k b own.
Sexual dimo phism. Ta ia nig a a ains sexual ma u i y abo e
80.0 mm SL. The ma u e emales ha e a small in umescen
geni al papilla. Male geni al papilla is o med by a hick skin
lap a ound a slende eme gen de e en duc . Male modi ied
anal in (Fig. 38) has h ee unb anched and i s b anched
ays enla ged and hickened. The i s unb anched anal- in
ay is immedia ely p eceded by a egumen a y keel (Fig. 38,
k). The second unb anched ay has an in e media e size be-
ween he neighbo ing i s and hi d ays. Thi d unb anched
is he longes ay, sligh ly cu ed owa ds in ip; bea ing
smalle an o sely cu ed dis al segmen s (Fig. 38, ac). Fi s
b anched ay is sligh ly cu ed owa ds he in ip. Fi s and
second b anched ays ha e e o sely cu ed dis al segmen s
(Fig. 38, c). Pos e io b anched ays a e p og essi ely sho e ;
wi h he las ay no educed (Fig. 38, b7). Caudal- in lobes o
bo h analyzed ma u e emales and males ha e he uppe lobe
elonga ed.
Six ma u e males o T. nig a wi h a modi ied anal in we e
examined. In wo o hese males he modi ied anal in is la e -
ally cu ed and spoon shaped (Fig. 39), sugges ing ep o-
duc i e modi ica ion o in e nal insemina ing.
Dis ibu ion. Known om he cen al Amazon, in he Ua umã
and T ombe as i e d ainages, no he n ibu a ies o he
Amazon i e (Fig. 6).
E ymology. The speci ic name nig a, is a La in e e ence o
he da k colo pa e n o his species.
Range Mean SD N
S anda d leng h 48.8-82.1 58.4 9
Pe cen s o s anda d leng h
Body dep h 18.1-24.6 20.3 1.98 9
Body wid h 17.5-21.8 19.8 1.00 9
Caudal peduncle dep h 13.6-14.8 14.1 1.10 9
Caudal peduncle leng h 20.9-28.1 24.7 1.83 6
P edo sal leng h 26.8-36.4 32.1 1.68 9
P eanal leng h 64.9-74.8 69.3 2.84 9
P epel ic leng h 50.0-56.5 52.5 1.80 9
Do sal o igin o pec o al o igin 19.7-26.6 23.0 1.45 9
Do sal o igin o pel ic o igin 27.3-34.3 31.1 1.97 9
Pec o al o igin o pel ic o igin 30.6-38.2 35.0 1.97 9
P epec o al leng h 16.1-23.7 20.3 1.73 9
Do sal- in base leng h 8.0-11.8 10.2 0.93 9
Adipose- in base leng h 7.4-13.8 11.0 1.80 9
Anal- in base leng h 2.9-12.3 7.0 2.73 9
Do sal- in spine leng h 13.4-21.2 17.8 1.99 8
Pec o al- in spine leng h 22.0-27.1 23.6 1.66 9
Pos clei h al p ocess lengh 13.2-22.1 19.7 1.99 9
Fi s b anched pel ic- in ay 8.6-15.1 12.8 1.30 9
Longes anal in ay 5.7-10.3 8.3 1.31 9
Maxilla y ba bel leng h 30.0-34.9 32.1 1.94 9
Ou e men al ba bel leng h 6.9-11.1 8.9 1.26 9
Men al ba bel leng h 4.8-11.4 6.3 1.42 9
Head leng h 21.3-26.7 23.4 1.42 9
Pe cen s o head leng h
Head wid h 70.8-76.9 74.5 1.99 9
Snou dep h 52.1-57.7 54.8 1.96 6
In e o bi al dis ance 53.0-58.7 55.7 1.88 9
Le in e na ial wid h 20.8-28.5 24.3 1.94 6
An e io in e na ial dis ance 34.1-39.6 37.4 1.78 6
Pos e io in e na ial dis ance 33.5-39.7 36.7 1.98 9
Snou leng h 33.0-38.8 37.4 1.98 9
O bi al diame e 20.0-24.2 22.0 1.84 9
Mou h wid h 47.9-52.9 50.2 1.80 9
Table 11. Mo phome ic da a o Ta ia nei ai. SD = s anda d
de ia ion.
A sys ema ic e ision o Ta ia
534
Rema ks. Ta ia nig a has well-de eloped la e al o namen ed
medial langes o bone on he nasal, bu no as la ge as he
langes obse ed in T. aulopygia. The expanded nasal wi h
medial langes pa ially su u ed o mese hmoid was ecog-
nized as a de i ed ea u e o he genus (Soa es-Po o, 1998),
and was also obse ed in T. aulopygia, T. boemia, T. dunni,
T. nei ai, T. s iga a, and adul specimens o T. in e media.
Female geni al papilla in T. nig a is no a common condi-
ion wi hin he genus. The only o he Ta ia wi h emale geni-
al papilla is T. gy ina. The in umescen emale geni al papilla
may ep esen a ansi o y condi ion in species o Ta ia, and
pe haps is obse ed in specimens only close o ep oduc i e
phase. The elonga ed uppe caudal- in lobe in bo h ma u e
emales and males is a e wi hin he genus and may be also
associa ed o he ep oduc i e phase.
Ta ia nig a occu s in sympa y wi h T. b unnea in he
T ombe as i e , a lago do Ba a a, bu no syn opically. The
specimens o T. nig a we e cap u ed in he lake, while he T.
b unnea was eco ded om he iga apé Sa acazinho, a ibu-
a y o he Ba a a lake.
Ta ia s iga a Soa es-Po o, 1995
Figs. 40-42
Ta ia c . b unnea. Bu gess, 1989: 242, pl. 113. [ opical Sou h
Ame ica].
Ta ia s iga a Soa es-Po o, 1995: 202, ig. 1. [ ype locali y:
iga apé Limãozinho, Maués, Amazonas, B azil]. Bu gess
& Finley, 1996:166 [ e e ence]. Soa es-Po o, 1998: 333 [ci-
a ion]. Fe a is, 2003:477 [checklis ]. Lasso e al., 2005:
139 [Al o O inoco, O inoco]. Fe a is, 2007: 78 [checklis ].
Diagnosis. Ta ia s iga a is he only species o he genus
wi h a comple ely s iped colo pa e n, bea ing whi ish elon-
ga e longi udinal i egula s ipes along la e al sides o body.
A unique a angemen o adul male modi ied anal in is
p esen , wi h he second unb anched anal- in ay wi h an e-
io enla ged segmen s and a no ch o med by he educ ion
o he second and hi d b anched ays (Soa es-Po o, 1995).
Ta ia s iga a is also dis inguished om all congene s by a
combina ion o cha ac e is ics: ou b anched pec o al- in
ays; 7 ibs; 29-30 e eb ae; and ma u e males wi h uppe
caudal- in lobe sligh ly elonga ed, same leng h in emales.
Fig. 36. Ta ia nig a. (a) holo ype, INPA 12291, male, 116.8 mm SL. (b) pa a ype, INPA 25501, emale, 107.3 mm SL, Ua umã i e ,
Amazonas S a e, B azil.
L. M. Sa men o-Soa es & R. F. Ma ins-Pinhei o 535
Desc ip ion. Measu ed specimens 36.5-55.0 mm SL; mo pho-
me ic da a p esen ed in Table 13. Body slim, head sligh ly
dep essed do so- en ally. Head la ge, obus , ou line o head
in do sal iew somewha ellip ic, b oade han long. T unk
om do sal- in base o caudal peduncle g adually comp essed.
La e al p o ile o head om snou ip o ope cula ma gin
sligh ly con ex un il pec o al- in inse ion. Ven al p o ile o
head and abdomen la . Ven al p o ile o body gen ly cu ed,
conca e behind anal- in o igin. Head in egumen hin, c anial
oo isible; well-de eloped adipose eye lid; eye la e o-do -
sally loca ed in an e io po ion o head; mou h e minal, up-
pe lip ex ended pos e o-la e ally as well-de eloped leshy
ic al old; an e io nos il ubula , loca ed on an e io bo de
o snou ; pos e io nos il la ge, ounded, limi ed by small
skin lap; ans e se dis ance be ween an e io nos ils la ge
han dis ance be ween pos e io ones. Maxilla y ba bel sho ,
ex ending close o pos e io ip o pos clei h al p ocess, some-
imes sho e ; men al ba bel sho , ips no eaching pec o al-
in base, a anged in a c along en al su ace o jaw; inne
men al ba bel abou 50.0-60.0% leng h o ou e men als.
Pos clei h al p ocess almos eaching e ical h ough o igin
o do sal in. Caudal peduncle deep, dep h abou 13.6-14.5%
SL.
Ros al bo de o c anium wi h mese hmoid as la ge as
long; p emaxilla unde nea h wi h synchond al a icula ion;
la ge ellip ical c anial on anel bounded by mese hmoid and
on al (Fig. 41); nasal ossi ied wi h medial langes pa ially
su u ed o la e al ma gin o mese hmoid; au opala ine ubu-
la , o ien ed obliquely o longi udinal axis o body; maxilla
abou he same size as au opala ine; p e ome expanded an-
e io ly wi h well de eloped a ow-shaped la e al p ocesses;
jaws o equal size; p emaxilla and den a y wi h h ee ows o
conical ee h. Fi s nuchal pla e somewha pen agonal; sec-
ond nuchal pla e conca e along la e al ma gins; hi d nuchal
pla e cu ed, p ojec ed la e ally. Epioccipi al p ocess e y
small.
Suspenso ium, hyoid a ch, b anchial skele on and ope -
cula bones as in gene ic desc ip ion. Sup ap eope cle p esen
as sho canal bone. Six b anchios egal ays a icula ed wi h
hyoid a ch: ou wi h an e io ce a ohyal and wo wi h pos e-
io ce a ohyal; las one la ened.
Fou in ao bi al bones in incomple e se ies. In ao bi al 1
b oad, wi h sho en o-la e al p ocess; emaining
in ao bi als hin, educed o canalicula po ions. In ao bi al
2 smalles , close o in ao bi al 1, ollowed by non-ossi ied
po ion o canal below eye and by wo pos e io canal bones
much close o spheno ic, o ming pos e io o bi al im. La -
e al line on body wi h ossi ied canal bones limi ed o head.
Do sal in I,4-5 (n=10); do sal- in spine wi h 15-20 an o se
Table 12. Mo phome ic da a o Ta ia nig a. SD = s anda d
de ia ion.
Holo ype Range Mean SD N
INPA 12291
S anda d leng h 116.8 70.0-116.8 101.6 12
Pe cen s o s anda d leng h
Body dep h 19.9 17.0-21.9 19.5 1.47 12
Body wid h 18.7 18.7-21.6 19.6 0.81 12
Caudal peduncle dep h 15.4 14.4-17.0 15.8 0.77 12
Caudal peduncle leng h 28.6 22.2-28.6 26.4 1.99 12
P edo sal leng h 30.1 27.4-31.0 29.9 0.94 12
P eanal leng h 68.6 64.0-75.4 68.1 3.30 12
P epel ic leng h 47.5 47.3-51.9 49.5 1.47 12
Do sal o igin o pec o al o igin 21.0 18.5-23.3 21.0 1.36 12
Do sal o igin o pel ic o igin 27.4 27.2-32.4 29.9 1.83 12
Pec o al o igin o pel ic o igin 33.1 30.7-37.3 32.7 1.75 12
P epec o al leng h 17.8 17.8-22.1 19.4 1.26 12
Do sal- in base leng h 11.1 8.6-11.4 9.9 0.80 12
Adipose- in base leng h 10.4 10.3-12.5 11.0 0.68 12
Anal- in base leng h 4.2 3.0-11.9 7.9 3.50 12
Do sal- in spine leng h 19.7 16.3-22.0 19.7 1.54 9
Pec o al- in spine leng h 20.5 20.3-24.6 22.7 1.42 9
Pos clei h al p ocess lengh 12.3 12.0-14.4 13.1 0.81 12
Fi s b anched pel ic- in ay 13.8 11.8-15.1 13.6 1.19 12
Longes anal in ay 8.3 4.7-10.7 8.8 1.83 12
Maxilla y ba bel leng h 27.9 25.6-30.9 27.8 1.91 12
Ou e men al ba bel leng h 8.4 6.5-10.1 8.1 1.09 12
Men al ba bel leng h 4.3 3.6-5.8 4.8 0.65 12
Head leng h 19.2 19.2-22.9 21.3 1.08 12
Pe cen s o head leng h
Head wid h 82.1 77.5-82.2 80.0 1.97 12
Snou dep h 49.1 47.0-53.7 48.8 1.99 12
In e o bi al dis ance 66.5 61.3-66.7 63.5 1.99 12
Le in e na ial wid h 26.3 22.4-26.5 24.7 1.53 12
An e io in e na ial dis ance 44.6 39.8-44.6 42.7 1.98 12
Pos e io in e na ial dis ance 42.4 34.6-42.4 39.0 1.91 12
Snou leng h 40.6 38.5-43.1 40.6 1.95 12
O bi al diame e 24.1 23.0-27.8 25.8 1.92 12
Mou h wid h 53.6 48.5-53.6 51.2 1.96 12
Fig. 37. Neu oc anium o Ta ia nig a, MNRJ 32024, 97.0 mm
SL. Do sal iew. Abb e ia ions: ep, epioccipi al; o, single c a-
nial on anel, , on al; le, la e al e hmoid; me, mese hmoid;
na, nasal; n1, i s nuchal pla e, n2; second nuchal pla e; n3,
hi d nuchal pla e; pe, pos e io epioccipi al p ocess; ps,
pos empo al-sup aclei h um; p , p e o ic; so, sup aoccipi al;
sp, spheno ic. Scale ba = 1 mm.
A sys ema ic e ision o Ta ia
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