117
Neo opical Ich hyology, 7(2):117-140, 2009
Copy igh © 2009 Sociedade B asilei a de Ic iologia
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
(Os a iophysi: Cha aci o mes: E y h inidae) wi h desc ip ions o wo
new species
Os aldo T. Oyakawa1 and Geo ge M. T. Ma ox2
Hoplias lace dae was o iginally desc ibed om he io Ribei a de Iguape, Ipo anga, São Paulo S a e. The Hoplias lace dae
g oup is de ined as con aining gene ally la ge ahi as wi h he medial ma gins o den a ies unning pa allel o each o he and
lacking ee h on he basihyal compa ed o he H. malaba icus g oup in which he medial ma gins o he den a ies con e ge
owa ds he mandibula symphysis and which ha e ee h on he basihyal. A axonomic e ision o he g oup based on me is ic
and mo phome ic da a iden i ied i e dis inc species: H. lace dae dis ibu ed in he io Ribei a de Iguape and io U uguai; H.
in e medius om he io São F ancisco, uppe io Pa aná basin, and io Doce; H. b asiliensis om i e s o he A lan ic Coas al
d ainage om he io Pa aguaçu o he io Jequi inhonha; H. aus alis new species, endemic o he io U uguai; and H.
cu upi a new species p esen in no he n Sou h Ame ica, including he ios Neg o, T ombe as, Tapajós, Xingu, Tocan ins and
Capim in he Amazon basin, uppe io O inoco nea he io Casiquia e (Venezuela), and coas al i e s o Guyana and Su iname.
A lec o ype o Hoplias in e medius and a neo ype o H. b asiliensis a e designa ed.
Hoplias lace dae oi desc i a o iginalmen e do io Ribei a de Iguape, Ipo anga, São Paulo. O g upo Hoplias lace dae oi
de inido como um conjun o de espécies pa a ab iga as aí as ge almen e de g ande po e, com as ma gens mediais dos
den á ios dispos as pa alelamen e e sem den es no basi-hial, em oposição às aí as do g upo Hoplias malaba icus, cujas
ma gens mediais dos den á ios con e gem em di eção à sín ise mandibula e possuem den es no basi-hial. A e isão axonômica
do g upo baseada em dados me ís icos e mo omé icos iden i icou cinco espécies dis in as: Hoplias lace dae dis ibuída nos
ios Ribei a de Iguape e U uguai; H. in e medius no io São F ancisco, al o io Pa aná e io Doce; H. b asiliensis nos ios
cos ei os da d enagem A lân ica, desde o io Pa aguaçu a é o io Jequi inhonha; H. aus alis espécie no a, endêmica ao io
U uguai; e H. cu upi a espécie no a no no e da Amé ica do Sul, incluindo os ios Neg o, T ombe as, Tapajós, Xingu,
Tocan ins e Capim na bacia Amazônica; al o io O inoco p óximo ao io Casiquia e (Venezuela) e ios cos ei os da Guiana e
Su iname. São designados um lec ó ipo pa a Hoplias in e medius e um neó ipo pa a H. b asiliensis.
Key wo ds: T ai ão, F eshwa e ish, Sou h Ame ica, Taxonomy, Dis ibu ion pa e ns.
1Museu de Zoologia da Uni e sidade de São Paulo, Caixa Pos al 42494, 04218-970 São Paulo, SP, B azil. [email p o ec ed]
2Depa amen o de Zoologia, Ins i u o de Biociências, Uni e sidade de São Paulo, Caixa Pos al 11461, 05422-970 São Paulo, SP, B azil.
[email p o ec ed]
In oduc ion
Species o he cha aci o m genus Hoplias a e dis ibu ed
h oughou o he many hyd og aphic sys ems o Sou h
Ame ica and a e popula ly known as ahi as, ai ões (= big
ahi as) o lobós. They inhabi a g ea numbe o habi a s,
anging om lo ic en i onmen s such as small o la ge i e s
and wa e alls, o s ic ly len ic wa e s such as lakes and
ese oi s. In all habi a s hey a e p eda o s o o he ish
(Tapho n, 1992; Planque e e al., 1996). Many species a e
impo an in subsis ence and comme cial ishe ies in a ious
egions o B azil and o he Sou h Ame ican coun ies
(Goulding, 1980). Species o Hoplias can achie e conside able
size, and H. lace dae and H. aima a a e among he gian s o
he g oup, eaching up o one me e in o al leng h.
The genus can be dis inguished om all o he
cha aci o ms by he ollowing de i ed cha ac e s: (1) con ac
be ween he ome and pa asphenoid ia in e digi a ions; (2)
a mese hmoid wi h an an e io p ojec ion ha sepa a es he
con ala e al p emaxillae; (3) he con ac o he an e io pa s
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
118
o in ao bi als 2 and 5 wi h consequence exclusion o
in ao bi als 3 and 4 om he o bi al im; (4) he pos e io end
o he maxilla wi h a la ge p ojec ion along i s do sal ma gin;
(5) he expansion o he median egion o haemal spine o he
i h e eb a; and (6) he absence o he en al p ocess o
he hi d e eb a (Oyakawa, 1998).
Oyakawa (1990) used mo phological cha ac e s o de ine
h ee species g oups in Hoplias, a p ac ice done in o he
cha aci o m gene a such as Aces o hynchus (Menezes, 1992;
Toledo-Piza, 1996) and As yanax (Ga u i & B i ski, 1997). One
o hese g oups wi hin Hoplias was in o mally called he H.
lace dae g oup, which can be de ined by he absence o oo h-
bea ing pla es in he leshy issue on he do sal su ace o he
basihyal and basib anchials (i.e. smoo h ongue) and by he
medial ma gins o con ala e al den a ies lying almos pa allel
o each o he , esul ing in U-shaped ma gin in en al iew
(Figs. 1a-c). The second assemblage is he H. malaba icus
g oup, ha has s ill no been e ised, bu which can be
ecognized by he p esence o oo h-bea ing pla es a ached
o leshy co e ing on do sal su ace o he basihyal and
basib anchials (i.e. ough ongue) and by ha ing he medial
ma gins o con ala e al den a ies ab up ly con e ging owa ds
he mandibula symphysis he eby o ming a V-shaped ma gin
in en al iew (Fig. 1d). The hi d is he H. mac oph halmus
g oup, which was ecen ly e ised (Ma ox e al., 2006) and
de e mined o con ain a single species, Hoplias aima a. I is
de ined by he absence o an accesso y ec op e ygoid and he
p esence o a da k o al spo on he ope cula memb ane.
Eigh species a e conside ed alid in Hoplias (Oyakawa,
2003; Ma ox e al., 2006), h ee o which we e included by
Oyakawa (1990) in he Hoplias lace dae g oup: H. b asiliensis
(Spix in Spix & Agassiz, 1829), H. mic ocephalus (Agassiz in
Spix & Agassiz, 1829) and H. lace dae Mi anda-Ribei o (1908).
A ou h nominal species, Mac odon in e medius Gün he
(1864), was conside ed a junio -synonym o H. mic ocephalus
(Oyakawa, 1990, 2003). The numbe o specimens o he H.
lace dae g oup in ich hyological collec ions has g ea ly
inc eased in ecen decades, e lec ing an expansion o i s
known geog aphic dis ibu ion h oughou Sou h Ame ica.
The pu pose o his pape is o e iew he species o he H.
lace dae g oup, edesc ibe he alid species and desc ibe
wo new species o he g oup.
Ma e ial and Me hods
This s udy was based on me is ic and mo phome ic da a
om 467 specimens. Me is ic da a was aken unde a
s e eomic oscope and he mo phome ic da a was aken poin -
o-poin wi h digi al calipe s o 0.1 mm. Measu emen s and
coun s we e made on le side o body, whene e possible,
and ollow Fink & Wei zman (1974) and Ma ox e al. (2006).
Unb anched and b anched in- ay coun s a e p esen ed as
Roman and A abic numbe s, espec i ely. Os eological names
ollow Wei zman (1962) and Robe s (1969). Desc ip ions o
colo a ion we e based on specimens p ese ed in e hanol.
An iden i ica ion key o species o he Hoplias lace dae g oup
is p esen ed wi h modal me is ic alues p esen ed in b acke s.
Ins i u ional abb e ia ions ollow Fe a is (2007) wi h addi ion
o UFRGS, Uni e sidade Fede al do Rio G ande do Sul.
Fig. 1. Ven al iew o head o (a) Hoplias cu upi a, MZUSP 24403, 55.5 mm SL; (b) H. cu upi a, MZUSP 24403, 92.2 mm SL;
(c) H. cu upi a, MZUSP 24403, 162.2 mm SL and (d) H. malaba icus, MZUSP 90416, 101.7 mm SL.
O. T. Oyakawa & G. M. T. Ma ox 119
Resul s
Key o species o Hoplias
1. Medial ma gins o con ala e al den a ies ab up ly
con e ging owa d mandibula symphysis, o ming a V-
shaped ma gin in en al iew (Fig. 1d); do sal su ace o
basihyal and basib anchials wi h oo h-bea ing pla es (i.e.
ough ongue) ............................ Hoplias malaba icus g oup
1’. Medial ma gins o con ala e al den a ies almos pa allel
o each o he , esul ing in a U-shaped ma gin in en al
iew (Figs. 1a-c); do sal su ace o basihyal and
basib anchials wi hou oo h-bea ing pla es (i.e. smoo h
ongue) ................................................................................... 2
2. Da k o al spo on ope cula memb ane p esen ; accesso y
ec op e ygoid absen .................................... Hoplias aima a
2’. Da k o al spo on ope cula memb ane absen ; accesso y
ec op e ygoid p esen ............... 3 (Hoplias lace dae g oup)
3. Specimens la ge han 60 mm SL wi h 4 well-de ined pa ches
o se e al small po es o la e osenso y sys em along he
en al su ace o den a y, specimens smalle han 60 mm
SL only wi h ou po es (Figs. 1a-c); la e al line scales 34-
39 [36] ............................................... Hoplias cu upi a n. sp.
3’. Specimens o any size always wi h 4-8 ela i ely la ge po es
o la e osenso y sys em along he en al su ace o
den a y; la e al line scales 38-48 .......................................... 4
4. La e osenso y sys em along en al su ace o den a y wi h
6-8 [7] po es; la e al line scales 43-48 [46]
.................................................................... Hoplias lace dae
4’. La e osenso y sys em along en al su ace o den a y
wi h 4-6 po es; la e al line scales 38-46 .............................. 5
5. An e io p o ile o head ounded in la e al iew;
la e osenso y sys em along en al su ace o den a y
always wi h 5 po es; g ound colo a ion o head and body
da k b own o da k g ey ................. Hoplias aus alis n. sp.
5’. An e io p o ile o head angula in la e al iew;
la e osenso y sys em along en al su ace o den a y wi h
4-6 po es; g ound colo a ion o head and body da k o
ligh b own ............................................................................ 6
6. La e al line scales 38-43 [41] ................... Hoplias b asiliensis
6’. La e al line scales 42-46 [44] ............... Hoplias in e medius
Hoplias aus alis, new species
Fig. 2
Holo ype. BRAZIL. San a Ca a ina S a e. U uguay basin: MCP
40175, 220.0 mm SL, io das An as, io do Ou o d ainage, on
Fo mosa do Sul-I a i oad, Fo mosa do Sul, 26º38’55’’S 52º48’05’’W,
25 Jan 2006, C. A. Lucena, E. Pe ei a, V. Be aco & J. P. Sil a.
Pa a ypes. ARGENTINA. Misiones. U uguay basin: MCP 13318,
2, 41.9-42.2 mm SL, a oyo I aca ua é, 27º53’S 55º17’W, 28 Jun
1989, E. Le ne ; MCP 13320, 1, 59.3 mm SL, a oyo San a Ma ia,
27º56’S 55º27’W, 28 Jun 1989, E. Le ne . BRAZIL. Rio G ande do
Sul S a e. U uguay basin. MCP 18561, 1, 200.6 mm SL, io
Dou ado be ween T ês Ba as and Ma iano Mo o, Ma iano Mo o,
21 Sep 1995, E. Filho, V. Schulz, S. Meu e & P. Iaczinski; MCP
19344, 1, 64.5 mm SL, io U uguay nea mou hs o ios Canoas and
Pelo as, 27º36’ 22”S 51º27’40”W, 13 No 1996, E. Pe ei a, J. Sil a
& R. Reis; MZUSP 99307, 1, 225.3 mm SL, io U uguay, nea ly 3
km om he Sal o do Yucumã, Pa que Es adual do Tu o, 27º08’13”S
55º31’55”W, 29 Oc 1991, L. Cama a, L. Hanh, T. Melo & E. Sil a;
MZUSP 39791, 1, 166.2 mm SL, a oio Lageado União in Linha
dos Lima, Palmi inho, 22 Dec 1985, L. Malaba ba, R. Reis & S.
Mallmann; UFRGS 4940, 2, 98.9-151.7 mm SL, io Ce quinha, São
José dos Ausen es, 7 Ap 2000, Equipe Ic io UFRGS; UFRGS
7064, 1, 59.1 mm SL, s eam on he oad o Pi apó, São Nicolau, 5
Ma 2005, M. Aze edo, J. Fe e , L. Malaba ba & C. Oli ei a.
San a Ca a ina S a e. U uguay basin: MCP 12090, 1, 243.8 mm
SL, io Jacu inga, on Seá a-Concó dia oad, Concó dia, 22 May
1988, R. Reis, E. Pe ei a, P. Aze edo & A. Be gmann; MCP 12410,
2, 217.1-234.6 mm SL, same locali y, 2 Oc 1988, E. Pe ei a, A.
Be gmann, P. Aze edo & A. Rami es; MCP 12741, 1, 211.7 mm
SL, io Canoas, on Abdon Ba is a-Ani a Ga ibaldi oad, Campos
No os, 11-12 No 1988, C. Lucena, E. Pe ei a, P. Aze edo & A.
Rami es; MCP 13146, 1, 253.6 mm SL, same locali y, 30-31 Ma
1989, A. Be gmann, E. Pe ei a, P. Aze edo, A. Rami es & M. Va gas;
MCP 18516, 2, 140.0-227.1 mm SL, io Riacho G ande, on Pi i iba-
BR 153 oad, Concó dia, 17 Sep 1995, E. Filho, V. Schulz, S. Meu e
& P. Iaczinski; MCP 18820, 1, 199.8 mm SL; MCP 18821, 1, 198.8
mm SL, same locali y, 29 Jul 1995, same collec o s; MCP 18923, 2,
114.3-176.8 mm SL, same locali y, 11 Jan 1996, same collec o s;
MCP 20847, 7, 25.2-37.2 mm SL, a oio Lageado G ande, 3 km
No hwes o Linha San ana, San a Helena, 26º53’35”S 53º36’4”W,
27 Jan 1998, C. Lucena, M. Lucena, E. Pe ei a & J. Pezzi; MCP
20860, 1, 45.1 mm SL, io Jundiá, on São Ped o-Tunápolis oad,
27º01' 17”S 53º38’58”W, same da e and collec o s; MCP 20866, 1,
120.9 mm SL, A oio da Taipa, on Mondaí-I api anga oad, Mondaí
27º07’46”S 53º28’27”, same da e and collec o s; MCP 40088, 1,
56.1 mm SL, io Lageado Se ão, unde b idge o BR 158 oad,
Cunha Po ã, 27º02’42”S 53º13’33”W, 25 Jan 2006, V. Be aco, C.
Lucena, J. Pezzi & E. Pe ei a; MCP 40093, 1, 32.9 mm SL, io
Bu o B anco, bo de be ween Se a Al a and Sul B asil, 26º41’12”S
52º49’2”W, same da e and collec o s; MCP 40117, 1, 34.9 mm SL,
io Pesquei o, on I a i-T ês Amigos oad, I a i, 26º39’39”S
52º55’34”W, same da e and collec o s; MZUSP 99306, 1, 210.4
mm SL, io Saudade, b idge a e own o Modelo owa ds BR 282,
Modelo, 26º47’10”S 53º2’43”W, same da e and collec o s; MCP
41287, 7, 32.9-87.5 mm SL, collec ed wi h he holo ype; UFRGS
7531, 2, 246.6-252.9 mm SL, io Jacu inga, downs eam om wa e
cap u e, Concó dia, 30 Oc 2004, J. Scalcon. URUGUAY. Ce o
La go. U uguay basin: MCP 9909, 1, 352.0 mm SL; MCP 9910, 1,
443.9 mm SL, io Neg o, A e ia, 31º50’S 54º28’W, 19 May 1984,
C. Lucena.
Diagnosis. The numbe o po es o he la e osenso y canal
along he en al su ace o den a y dis inguished Hoplias
aus alis om H. lace dae (always 5 s. 6-8 po es,
espec i ely). The possession o an an e io p o ile o he
head ounded in la e al iew dis inguished H. aus alis om
H. b asiliensis and H. in e medius, which ha e he an e io
p o ile o head angula . Hoplias aus alis can be u he
dis inguished om H. cu upi a in he numbe o scales along
la e al line (40-45 s. 34-39 espec i ely).
Desc ip ion. Mo phome ic da a p esen ed in Table 1. Body
cylind ical, deepe han wide. G ea es body dep h a e ical
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
120
h ough hi d scale an e io o do sal- in o igin in specimens
smalle han 90 mm SL, and h ough six h scale an e io o
do sal- in o igin in la ge specimens (ca. 220 mm SL). An e io
p o ile o head ounded in la e al iew, bu mo e angula in
specimens smalle han 40 mm SL. Do sal p o ile o head
a ying om sligh ly con ex in specimens smalle han 40
mm SL o almos s aigh in la ge indi iduals. Do sal ma gin
o o bi a ho izon al h ough do sal p o ile o head in
specimens smalle han 40 mm SL, bu no eaching do sal
p o ile o head in specimens la ge han 90 mm SL. Do sal
p o ile o body con ex om e ical h ough i s se ies o
scale o do sal- in o igin; sligh ly s aigh and
pos e o en ally inclined along do sal- in base; sligh ly
conca e om e ical h ough base o las do sal- in ay o
o igin o do sal mos p ocu en caudal- in ay. La e po ion
o p o ile sligh ly s aigh e in specimens smalle han 90 mm
SL. Ven al p o ile o lowe jaw dis inc ly angula in egion o
den a y symphysis, s aigh o sligh ly con ex om e ical
h ough an e io nos il o pos e io ma gin o lowe jaw. Medial
ma gins o con ala e al den a ies app oxima ely pa allel (Fig.
1a-c). Ven al p o ile o body sligh ly con ex o pel ic- in
o igin; app oxima ely s aigh om la e poin o anal- in
o igin; s aigh and pos e odo sally inclined along anal- in
base; sligh ly conca e o s aigh om base o las anal- in
ay o an e io mos en al p ocu en caudal- in ay.
Uppe jaw usually as long as lowe jaw, bu sligh ly sho e
in some specimens. Pos e io po ion o maxilla do sally
enla ged and ex ending medially o an e io ma gins o second
and hi d in ao bi als. Uppe and lowe lips leshy wi h sho
skin p ojec ions co e ing canines ex e nally. An e io nos il
ubula wi h an e io sli along i s dis al hal . An e io and
pos e io nos ils si ua ed along ho izon al h ough cen e o
o bi , an e io nos il loca ed a poin wo- hi ds o o bi al
diame e om an e io ma gin o o bi . Pos e io nos il si ua ed
close o an e io nos il han o an e io ma gin o o bi . Eye
p opo iona ely la ge in smalle specimens. In ao bi al bones
well de eloped and ho izon ally elonga e. In ao bi als 3, 4,
and en al po ion o 5 pa ially co e ing p eope cle. Ven al
ma gin o in ao bi al 3 con ex. Pos e io ma gin o in ao bi al
4 ela i ely s aigh and ma gins o in ao bi als 5 and 6 sligh ly
con ex. Small specimens (ca. 90 mm SL) wi h in ao bi al 3
ba ely eaching o bi al im and p oximal po ions o
in ao bi als 2 and 4 almos in con ac wi h each o he . La ge
specimens (ca. 220 mm SL) wi h in ao bi als 3 and 5 comple ely
excluded om o bi al im and in ao bi al 4 ba ely eaching
im.Tee h in bo h jaws conical o canine. P emaxilla y ee h in
Fig. 2. Hoplias aus alis, holo ype, MCP 40175, 220 mm SL, io das An as, Fo mosa do Sul, San a Ca a ina S a e, B azil.
Table 1. Mo phome ic da a o Hoplias aus alis. S anda d
leng h in mm; alues 1-14 a e pe cen s o s anda d leng h and
alues 15-22 a e pe cen s o head leng h. n = numbe o
examined specimens.
Holo ype n Mean Range
S anda d
de ia ion
S anda d leng h 220.0 43 - 25.2-443.9 -
1. Body dep h 21.4 34 21.5 18.3-25.5 2.0
2. Head leng h 30.0 36 31.3 26.2-33.7 1.4
3. Pec o al- in leng h 16.8 36 18.3 15.8-21.9 1.3
4. Pel ic- in leng h 16.4 36 19.3 16.0-22.7 1.6
5. Anal- in leng h 17.9 36 19.3 17.4-22.0 1.3
6. Do sal- in leng h 26.7 36 29.1 25.7-33.3 1.8
7. Do sal- in base 15.0 36 17.4 15.0-20.9 1.2
8. Anal- in base 9.6 36 10.0 8.1-12.0 0.9
9. P e-pec o al dis ance 26.8 36 28.9 24.5-32.5 1.8
10. P e-pel ic dis ance 51.8 36 54.5 50.2-58.3 1.8
11. P e-do sal dis ance 49.4 36 49.9 45.0-53.4 2.0
12. P e-anal dis ance 79.3 36 80.0 74.6-86.0 2.6
13. Caudal-peduncle dep h 14.5 36 14.5 12.6-17.5 1.1
14. Caudal-peduncle leng
h
14.1 36 14.3 11.6-16.6 1.1
15. Head dep h 47.9 36 44.6 37.9-49.5 3.1
16. Snou leng h 26.7 36 24.4 20.3-28.5 2.1
17. Snou wid h 21.2 36 19.2 14.2-22.7 2.3
18. Snou dep h 23.5 36 21.3 16.7-27.0 2.7
19. P e-nasal dis ance 16.4 36 15.8 9.9-19.8 2.0
20. O bi al diame e 18.3 36 20.6 13.3-29.0 4.2
21. In e o bi al wid h 24.8 35 22.1 14.9-26.8 2.9
22. Uppe jaw leng h 50.4 36 47.6 41.0-54.8 3.8
O. T. Oyakawa & G. M. T. Ma ox 121
single ow. Fi s p emaxilla y oo h la ge canine, second,
se en h and nin h ee h medium sized. Eigh h oo h canine
and almos as la ge as an e io mos p emaxilla y canine in
la ge specimens (ca. 220 mm SL). Thi d o six h p emaxilla y
ee h small. Maxilla wi h single ow o app oxima ely 30
ela i ely small ee h. Den a y wi h an e io ex e nal ow o
ee h and pos e io in e nal ow. Ex e nal se ies wi h h ee
small an e io ee h ollowed by wo well-de eloped canines,
wi h pos e io one la ge . Pos e io canine ollowed by 11
conical ee h, some hal size o an e io mos den a y canine.
In e nal se ies beginning p oxima e o small conical oo h
pos e io o las conical oo h o ex e nal se ies and composed
o app oxima ely 12 e y small ee h. Accesso y ec op e ygoid
and ec op e ygoid oo hed; la e wi h se ies o small conical
ee h along i s en ola e al ma gin and many smalle ili o m
ee h medially along i s en al su ace. Endop e ygoid
eden ulous.
Dis al ma gins o all ins ounded. To al do sal- in ays
12-14 (n = 22; ii,10-12; mode: ii,11, n = 11). Do sal in loca ed a
midbody, i s o igin a e ical h ough app oxima ely second
scale an e io on se ies along pel ic- in o igin. Leng h o
longes do sal- in ays wo- hi ds o body dep h. Anal- in
base sho . To al anal- in ays 9-11 (n = 21; ii,7-9; mode: ii,8, n
= 12). Tips o longes ays o dep essed do sal in no
ex ending beyond e ical h ough anal- in o igin. To al
pec o al- in ays 11-13 (n = 22; i,10-12; mode: i,11, n = 9).
Pec o al- in o igin loca ed a e ical h ough pos e io egion
o ope cle. Tip o pec o al in sepa a ed om pel ic- in o igin
by ou o six scales, and alling sho o e ical h ough do sal-
in o igin. Pec o al, pel ic and anal ins o simila sizes. Pel ic-
in ays i,7-8 (n = 22, i,8 in one specimen). Pel ic- in o igin
si ua ed a midbody and app oxima ely h ee scales pos e io
o e ical h ough do sal- in o igin. Tip o pel ic in sepa a ed
om e ical h ough anal- in o igin by h ee scales in specimens
smalle han 90 mm SL, and by up o eigh scales in la ge
specimens (ca. 220 mm SL). Caudal- in ays i,15,i (n = 9).
Well-de eloped, imb ica ed cycloid scales along body.
Do sal scales begin in se ies a pos e io ma gin o pa ie als
and o e lap sup aoccipi al spine. Las e ical se ies o scales
on caudal peduncle o ming sligh ly con ex a ch on caudal-
in base. An e io ma gin o scales wi h small ecess and
pos e io ma gin ounded. App oxima ely wel e adii
ex ending om cen e o scale o i s an e io ma gin and a ound
wen y- wo adii, some anas omosed, ex ending om cen e
o scale o i s do sal, pos e io and en al ma gins. La e al
line s aigh and comple e, ex ending om pos e o en al
ma gin o sup aclei h um o pos e io mos scale in body.
La e al line wi h 40-45 pe o a ed scales (n = 38, 44 in one
specimen and 45 in one specimen; mode: 41, n = 12) (Table 2).
La e al-line scales wi h single la e osenso y canal.
Longi udinal se ies o scales be ween la e al line and do sal-
in o igin 4.5-5.5 (n = 21; mode: 4.5, n = 15). Longi udinal
se ies o scales be ween la e al line and pel ic- in o igin 3.5-
5.5 (n = 20; mode: 4.5, n = 18). Longi udinal se ies o scales
a ound caudal peduncle 19-20 (n = 21; mode 20, n = 20).
App oxima ely 12 gill ake s on i s epib anchial, mos in
o m o small den icula ed pla es. Lowe b anch o i s
b anchial a ch wi h i e mo e elonga e ake s and
app oxima ely 10 pla e-like ake s (n = 9). La e osenso y canal
along en al su ace o den a y always wi h i e po es (n =
43, Table 3).
Colo in alcohol. G ound colo a ion o head and body da k
b own o da k g ey, da ke do sally and pale en ally, la e
egion whi e o da k yellow o ligh b own in mos specimens.
Da k b own midla e al s ipe along la e al line weakly e iden
in specimens smalle han 90 mm SL and almos absen in
la ge specimens (ca. 220 mm SL). App oxima ely se en da k
blo ches along la e al line wi h wid h o blo ches abou six
longi udinal se ies o scales; pa e n mo e conspicuous in
specimens smalle han 90 mm SL. Da k colo a ion o body
obscu ing da k midla e al s ipe and blo ches in la ge
specimens. Th ee o ou da k blo ches p esen along do sal
su ace o body. Some specimens wi h se e al small da k spo s
loca ed on do sola e al po ion o body. Do sal su ace o
head da k b own o g ey. Nume ous small black spo s on
do sal su ace o head o se e al specimens. Lips da k g ey,
wi h small e ical ligh bands in specimens smalle han 40
mm SL. Colo a ion o in ao bi al egion simila o ha o do sal
su ace o head. Some specimens wi h one o wo da k s ipes
adia ing pos e io ly om eye ac oss ope cula se ies. G ound
colo a ion o ope cula se ies simila o es o head. Ope cle
memb ane usually as da k as ope cle, bu sligh ly ligh e in
some specimens.
All ins ligh b own o g ey wi h da k spo s on ays and
in e adial memb ane o ming pa e n o i egula da k s ipes.
Da k and ligh s ipes o simila wid h in specimens smalle
han 90 mm SL, wi h da k s ipes wide han ligh ones in
la ge specimens. Ven al su aces o pec o al and pel ic ins
ei he ligh e o wi h pa e n simila o ha o do sal su ace
bu less conspicuous.
Po es 4 5 6 7 8
H. aus alis 43
H. b asiliensis 6 66 3
H. cu upi a 151
H. in e medius 17 74 6
H. lace dae 17 67 5
Table 2. F equency dis ibu ion o o al numbe o la e al line
scales in species o he Hoplias lace dae g oup.
Table 3. F equency dis ibu ion o o al numbe o po es o
cephalic la e al sys em along he en al su ace o he
den a y in species o he Hoplias lace dae g oup.
La e al Line
Scales 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48
H. aus alis 11 12 10 3 1 1
H. b asiliensis 2 5 23 24 15 5
H. cu upi a 6 17 60 38 18 3
H
. in e medius 5 32 36 9 4
H. lace dae 1 7 22 28 25 6
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
122
Dis ibu ion. Th oughou io U uguay basin, including i s
ibu a y io Neg o (Fig. 3).
E ymology. The name aus alis, an adjec i e, e e s o he
sou he n ange o his species, one o he sou he n mos in
he genus.
Rema ks. Valenciennes (1846) desc ibed Mac odon au i us
based on one specimen collec ed in Mon e ideo, U uguay,
om io U uguay basin. I is no clea , howe e , om he
o iginal desc ip ion whe he he species belongs o he
Hoplias lace dae g oup o o ei he o he wo axa o his
g oup documen ed om he io U uguay in his s udy.
Un o una ely, he ype specimen o M. au i us has no been
loca ed in MNHN, con a y o he o he ype specimens o
he E y h inidae desc ibed in Valenciennes (1846) [P. P u os ,
pe s. comm.]. Mac odon au i us is he e o e conside ed a
nomen dubium (ICZN, 1999).
Two species o Hoplias lace dae g oup, H. aus alis and
H. lace dae, occu in sympa y and syn opy in he io
U uguay basin, and we e collec ed oge he a some locali ies.
Hoplias lace dae also occu s in he io Ribei a de Iguape
(see below), and H. aus alis is, hus, he only Hoplias species
appa en ly endemic o he io U uguay basin. Judging om
he amoun o ma e ial in museum collec ions, H. aus alis is
much less common in he io U uguay han H. lace dae.
Hoplias b asiliensis (Spix, 1829)
Fig. 4
E y h inus b asiliensis Spix in Spix & Agassiz, 1829: 45, pl.
20 [o iginal desc ip ion, ype locali y: Pa aguaçú, Bahia
S a e, B azil]. -Gün he , 1864: 281 [species lis as synonym
o Mac odon ahi a]. -Lü ken, 2001: 78 [ oo no e,
Fig. 3. Map o Sou h Ame ica showing geog aphic dis ibu ion o species o he Hoplias lace dae g oup based on ma e ial
examined in his s udy. Some symbols may ep esen mo e han one lo . Numbe s indica e ype locali ies o (1) Hoplias
aus alis (squa es); (2) H. b asiliensis (losanges); (3) H. cu upi a ( iangles); (4) H. in e medius (do s); and (5) H. lace dae
(in e ed iangles).
O. T. Oyakawa & G. M. T. Ma ox 123
synonym o Mac odon ahi a]. -Eigenmann & Eigenmann,
1889: 103 [species lis as synonym o Mac odon
malaba icus]. -Eigenmann, 1910: 448 [species lis as
synonym o Hoplias malaba icus]. -Eigenmann, 1912: 416
[species lis as synonym o Hoplias malaba icus]. -Fowle ,
1950: 364 [species lis as synonym o Hoplias malaba icus
malaba icus]. -Godoy, 1975: 405 [as synonym o Hoplias
malaba icus malaba icus].
Mac odon b asiliensis: -Mülle , 1842: 309 [species lis ]. -
Mülle , 1843: 316 [species lis ]. -Mülle & T oschel, 1844:
82 [species lis ]. -Mülle & T oschel, 1845: 6 [species lis ].
-Godoy, 1975: 405 [as synonym o Hoplias malaba icus
malaba icus].
Hoplias b asiliensis: -Oyakawa, 2003: 239 [species lis ,
dis ibu ion]. -Oyakawa & Ne o-Fe ei a, 2007: 63 [species
lis , dis ibu ion]. -Menezes e al., 2007: 152 [ ype-locali y,
dis ibu ion].
Neo ype. BRAZIL. Bahia S a e. MZUSP 45483, 1, 162.9 mm SL,
io Pa aguaçú nea Iaçú, 25 Jul 1988, R. Cas o, S. Jewe & H.
San os. [designa ed he ein].
Ma e ial examined. BRAZIL. Bahia S a e. Pa aguaçú basin:
MZUSP 40171, 10, 39.3-122.8 mm SL; MZUSP 40172, 7, 41.3-
94.9 mm SL, io Pa aguaçú nea Iaçú; MZUSP 47939, 11, 23.7-66.2
mm SL; MZUSP 49234, 12, 23.1-65.1 mm SL, io do Una, io
Pa aguaçú d ainage, I ae ê; MZUSP 40173, 1, 114.5 mm SL, lagoa
Cha ca Na i idade, loodplain o io Pa aguaçú, Iaçú; MZUSP
86123, 1, 112.5 mm SL, io do Una, Rumo illage, I ae ê; MZUSP
87684, 2, 83.9-132.2 mm SL, io do Una, Fazenda No a Iguaçú,
I ae ê. Bahia, io de Con as basin: MZUSP 40174, 1, 82.7 mm SL,
io Gongoji, ibu a y o io de Con as, Fazenda I ama a i, 5 km
om Dá io Mei a; MZUSP 41500, 2, 32.1-35.1 mm SL; MZUSP
39797, 7, 19.2-54.0 mm SL, io de Con as, Ipiaú. Bahia. Pa do
basin: MZUSP 40170, 2, 193.9-205.7 mm SL, io Pa do, Fazenda
He mógenes, 5 km ups eam om e y-c ossing, I ape ininga.
Minas Ge ais S a e. Jequi inhonha basin: MZUSP 40269, 7, 95.5-
273.4 mm SL, io Jequi inhonha nea A açuaí, Vila I i a; MZUSP
40270, 7, 38.7-137.0 mm SL, io Jequi inhonha, Ilha Almena a,
Cu iango; MZUSP 74032, 3, 27.8-39.4 mm SL, ma ginal lagoon o
io Jequi inhonha, km 205 o Sal o da Di isa-Jacin o oad.
Diagnosis. The numbe o po es o he la e osenso y canal
along he en al su ace o he den a y dis inguished
Hoplias b asiliensis om H. lace dae (4-6 s. 6-8
espec i ely). I can be dis inguished om H. in e medius
by he numbe o he scales along he la e al line (38-43 s.
42-46 espec i ely). The an e io p o ile o head ha is angula
in la e al iew dis inguished H. b asiliensis om H. aus alis,
H. cu upi a and H. lace dae, which ha e he an e io p o ile
o head ounded. I can be u he dis inguished om H.
cu upi a by ha ing ligh b own g ound colo a ion o he
head and body.
Desc ip ion. Mo phome ic da a p esen ed in Table 4. Body
cylind ical, deepe han wide. G ea es body dep h a e ical
h ough i h scale an e io o do sal- in o igin in specimens
smalle han 35 mm SL, close o do sal- in o igin in la ge
specimens. An e io p o ile o head angula in la e al iew,
mo e ounded in specimens smalle han 100 mm SL. Do sal
p o ile o head a ying om sligh ly con ex in specimens
smalle han 100 mm SL o almos s aigh in specimens la ge
han 130 mm SL. Do sal ma gin o o bi loca ed a ho izon al o
h ough do sal p o ile o head in specimens smalle han 100
mm SL bu no eaching do sal p o ile o head in specimens
la ge han 130 mm SL. Do sal p o ile o body sligh ly con ex
om e ical h ough i s se ies o scale o body o do sal- in
o igin in specimens smalle han 130 mm SL and almos s aigh
in la ge specimens (ca. 280 mm SL); s aigh and
pos e o en ally inclined along do sal- in base; s aigh and
less inclined o sligh ly conca e om e ical h ough base o
las do sal- in ay o o igin o do sal mos p ocu en caudal-
in ay. La e po ion o p o ile sligh ly mo e conca e in
specimens smalle han 130 mm SL. Ven al p o ile o lowe jaw
dis inc ly angula in egion o den a y symphysis, s aigh
Fig. 4. Hoplias b asiliensis, neo ype, MZUSP 45483, 162.9 mm SL, io Pa aguaçú nea Iaçú, Bahia S a e, B azil.
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
124
and sligh ly inclined om e ical h ough an e io nos il o
pos e io ma gin o lowe jaw. Medial ma gins o con ala e al
den a ies unning in pa allel (Fig. 1a-c). Ven al p o ile o body
sligh ly con ex o pel ic- in o igin; app oxima ely s aigh
om la e poin o anal- in o igin; s aigh and pos e odo sally
inclined along anal- in base; sligh ly conca e om las anal-
in ay o an e io mos en al p ocu en caudal- in ay.
Uppe jaw sligh ly sho e han lowe jaw. Pos e io po ion
o maxilla do sally enla ged and ex ending medial o an e io
ma gins o second and hi d in ao bi als. Uppe and lowe
lips leshy wi h sho p ojec ions o skin co e ing canines
ex e nally. An e io nos il ubula wi h an e io sli along i s
dis al hal . An e io and pos e io nos ils si ua ed along
ho izon al h ough cen e o o bi , an e io nos il wo- hi ds
o o bi al diame e om an e io ma gin o o bi ; pos e io
nos il midway be ween an e io ma gin o o bi and an e io
nos il. Eye p opo iona ely la ge in smalle specimens.
In ao bi al bones well de eloped and ho izon ally elonga e.
In ao bi als 3, 4, and en al po ion o 5 pa ially co e ing
p eope cle. Ven al ma gin o in ao bi al 3 and pos e io
ma gins o in ao bi als 4, 5, and 6 con ex. Small specimens
(ca. 35 mm SL) wi h in ao bi al 3 excluded om o bi al im
and p oximal ends o in ao bi als 2 and 4 in con ac . La ge
specimens (ca. 130 mm SL) wi h in ao bi al 4 ba ely eaching
o bi al im and in ao bi al 5 g adually sepa a ed om im.
P oximal ends o in ao bi als 2 and 6 almos mee ing in some
o la ges specimens (ca. 280 mm SL).
Tee h in bo h jaws conical o canine. P emaxilla y ee h in
single ow. Fi s p emaxilla y oo h la ge canine, second, six h
and eigh h ee h medium sized. Se en h oo h canine almos
as la ge as an e io mos p emaxilla y canine. Thi d o i h
and nin h p emaxilla y ee h small. Maxilla wi h single ow o
app oxima ely 32 ela i ely small ee h, excep o e y well-
de eloped canine-like ou h oo h. Den a y wi h an e io
ex e nal ow o ee h and pos e io in e nal ow. Ex e nal se ies
wi h ou an e io small ee h, ollowed by wo well-de eloped
canines, wi h pos e io canine la ge han an e io canine and
ollowed by 8 conical ee h sligh ly smalle han an e io mos
den a y canine. In e nal se ies beginning a leng h o nea ly
one conical oo h pos e io o las conical oo h o ex e nal
ow and o med o app oxima ely 11 e y small ee h.
Accesso y ec op e ygoid and ec op e ygoid oo hed; la e
wi h se ies o small conical ee h along i s en ola e al ma gin
and wi h many smalle ili o m ee h on i s en omedial
su ace. Endop e ygoid eden ulous.
Dis al ma gins o all ins ounded. To al do sal- in ays
12-14 (n = 75; ii,11; ii,10 in wo specimens and ii,12 in i e
specimens). Do sal in loca ed a midbody, i s o igin a e ical
h ough app oxima ely hi d scale an e io on se ies along
pel ic- in o igin. Leng h o longes do sal- in ays wo- hi ds
o body dep h. Anal- in base sho . To al anal- in ays 9-10 (n
= 75; ii,8, ii,7 in i e specimens). Tip o dep essed do sal in
eaching e ical h ough anal- in o igin. Specimens smalle
han 35 mm SL wi h ip o anal in ( hi d o six h b anched
ays) eaching o igin o an e io mos en al p ocu en ay
o caudal in. To al pec o al- in ays 12-14 (n = 70; i,11-13;
mode: i,12, n = 40). Pec o al- in o igin loca ed a e ical
h ough cen al po ion o ope cle. Tip o pec o al in
sepa a ed om pel ic- in o igin by ou o six scales. Pec o al
and pel ic ins o simila size and sligh ly la ge han anal in.
Pel ic- in ays i,7 (n = 75, i,5 in one specimen). Pel ic- in o igin
si ua ed a midbody app oxima ely h ee scales pos e io o
e ical h ough do sal- in o igin. Tips o pel ic in sepa a ed
om e ical h ough anal- in o igin by ou o i e scales.
Caudal- in ays i,15,i (n = 71, 14 b anched ays in one
specimen).
Well de eloped cycloid scales imb ica ed along body.
Do sal scales begin in se ies a pos e io ma gin o pa ie als
and o e lap sup aoccipi al spine. Las e ical se ies o scales
on caudal peduncle o ms sligh ly con ex a ch on caudal- in.
An e io ma gin o scales wi h small ecess and pos e io
ma gin ounded. App oxima ely wel e adii ex ending om
cen e o scale o i s an e io ma gin and a ound wen y- i e
adii, some anas omosed, ex ending om cen e o scale o
do sal, pos e io and en al ma gins. La e al line s aigh and
comple e, ex ending om pos e o en al ma gin o
sup aclei h um o pos e io mos scale in body. La e al line
wi h 38-43 pe o a ed scales (n = 74, 38 in wo specimens;
mode: 41, n = 24) (Table 2). La e al-line scales wi h single
la e osenso y canal. Longi udinal se ies o scales be ween
la e al line and do sal- in o igin 5.5 (n = 75). Longi udinal
se ies o scales be ween la e al line and pel ic- in o igin 4.5-
5.5 (n = 75; mode: 5.5, n = 72). Longi udinal se ies o scales
a ound caudal peduncle 18-20 (n = 73; 18 scales in wo
specimens).
Table 4. Mo phome ic da a o Hoplias b asiliensis. S anda d
leng h in mm; alues 1-14 a e pe cen s o s anda d leng h and
alues 15-22 a e pe cen s o head leng h. n = numbe o
examined specimens.
Neo ype n Mean Range S anda d
de ia ion
S anda d leng h 162.9 75 - 19.2-273.8 -
1. Body dep h 23.8 75 22.0 17.7-25.9 1.8
2. Head leng h 35.2 75 33.7 30.9-37.4 1.2
3. Pec o al- in leng h 20.5 72 19.7 14.5-22.8 1.8
4. Pel ic- in leng h 21.2 75 20.4 17.4-23.3 1.2
5. Anal- in leng h 19.2 75 19.5 16.6-33.1 2.0
6. Do sal- in leng h 28.1 75 29.8 25.6-34.8 1.8
7. Do sal- in base 14.8 75 16.8 14.1-21.8 1.4
8. Anal- in base 10.3 75 9.7 7.7-20.1 1.5
9. P e-pec o al dis ance 32.5 75 31.6 28.4-35.6 1.6
10. P e-pel ic dis ance 58.7 75 56.2 53.9-58.7 1.1
11. P e-do sal dis ance 52.9 75 52.4 49.0-55.9 1.4
12. P e-anal dis ance 83.6 75 80.3 76.4-85.4 1.6
13. Caudal-peduncle dep h 14.3 75 13.9 12.3-15.6 0.7
14. Caudal-peduncle leng h 13.8 75 13.5 10.2-15.4 1.1
15. Head dep h 41.4 75 40.1 34.2-45.4 2.2
16. Snou leng h 26.1 75 23.7 18.8-28.4 1.9
17. Snou wid h 19.7 75 17.8 12.6-21.9 1.7
18. Snou dep h 21.4 75 17.9 9.8-22.4 2.4
19. P e-nasal dis ance 16.5 74 14.5 9.4-18.1 1.8
20. O bi al diame e 18.4 75 24.5 16.2-33.3 3.8
21. In e o bi al wid h 22.1 75 19.1 16.4-26.0 1.8
22. Uppe jaw leng h 48.7 75 46.2 36.6-52.5 3.0
O. T. Oyakawa & G. M. T. Ma ox 125
App oxima ely eigh gill ake s on i s epib anchial, mos
in o m o small den icula ed pla es. Lowe b anch o i s
b anchial a ch wi h i e o six mo e elonga e ake s and
app oxima ely 10 pla e-like ake s (n = 8). La e osenso y canal
along en al su ace o den a y wi h 4-6 po es (n = 75; mode:
5, n = 66) (Table 3).
Colo in alcohol. G ound colo a ion o head and body da k o
ligh b own, da ke do sally and pale en ally. Single da k
midla e al s ipe along la e al line sepa a ing da k do sal om
ligh e en al egion. S ipe mo e e iden in specimens smalle
han 130 mm SL, he only colo ea u e in e y small specimens
(ca. 35 mm SL). Specimens la ge han 100 mm SL wi h i e o
six da k blo ches along da k longi udinal s ipe, some o hem
connec ed wi h con ala e al blo ch by da k do sal saddle.
Do sal su ace o head da k b own. Lips wi h al e na ing da k
and ligh , hin, e ical bands. Ven al su ace o head whi e.
Colo a ion o in ao bi al egion and do sal su ace o head
simila . Many specimens wi h wo da k s ipes adia ing
pos e io ly om eye ac oss ope cula se ies and wi h hi d
da k s ipe ex ending pos e o en ally om eye along
in ao bi al 2. G ound colo a ion o ope cula se ies da k
b own. Ope cula memb ane usually ligh e han ope cle.
All ins ligh b own and ligh e han body, wi h da k spo s
on ays and in e adial memb ane o ming pa e n o i egula
da k s ipes. S ipes o anal and caudal ins wide han hose
o do sal in wi h s ipes mo e i egula in anal in. Pec o al,
pel ic, and anal ins wi h pale pa e n han do sal and caudal
ins. Ven al su aces o pec o al and pel ic ins ei he ligh e
han, o wi h same pa e n, as do sal su ace o in, bu wi h
pa e n less conspicuous.
Dis ibu ion. Coas al i e s o no heas e n B azil, om he
io Pa aguaçú in Bahia S a e o io Jequi inhonha in Minas
Ge ais and Espí i o San o S a es, including io de Con as and
io Pa do (Fig. 3).
Rema ks. Specimens used by Spix & Agassiz (1829) o
desc ibe E y h inus b asiliensis and deposi ed in he
Zoologische S aa ssammlung München, we e des oyed
du ing a bombing aid in he Second Wo ld Wa (Te o al,
1983; Ko ela , 1988; Neumann, 2006). The holo ype o
E y h inus b asiliensis is no among he specimens s udied
by Spix & Agassiz in he Musée d’His oi e Na u elle,
Neuchâ el, and is he e o e judge o be missing. I s o iginal
desc ip ion, howe e , men ioned he “ ongue […] eden a e”
and “b anches o he lowe jaw a he dis an om each o he
and somewha sp ead ou below […] pa allel o one ano he ”
[ou ansla ion]. These obse a ions con i m ha E.
b asiliensis is a membe o he Hoplias lace dae g oup. Da a
p esen ed in his s udy ma ch he emaining in o ma ion in
he o iginal desc ip ion. Since he holo ype o E. b asiliensis
is los , and in o de o cla i y he axonomic s a us o his
nominal species, a neo ype is designa ed in acco dance o
A icle 75 o he In e na ional Code o Zoological
Nomencla u e (ICZN, 1999). The axon is edesc ibed he ein,
wi h cha ac e s diagnosing i om o he membe s o he genus.
Cas elnau (1855) iden i ied some specimens collec ed in
he io Ca andaí, a ibu a y o io das Mo es, Minas Ge ais
S a e, as E y h inus b asiliensis and s a ed ha he specimens
a ained 50 cm SL. Acco ding o he esul s o he p esen
s udy, Hoplias b asiliensis a ains only 30 cm SL, being he
smalles among species o he H. lace dae g oup. The i e
men ioned by Cas elnau (1855) belongs o he io G ande
basin, uppe io Pa aná, and he a ailable name o he
specimens ci ed by ha au ho is H. in e medius, a o m ha
can each 50 cm SL.
Ri e s in which Hoplias b asiliensis occu s d ain he
A lan ic coas o B azil, om he io Pa aguaçu o he io
Jequi inhonha. The egion o occu ence o his species lies
sou h and eas o he io São F ancisco and no h o he io
Doce. Pelleg ino e al. (2005) p oposed ha he o igin o io
Doce, oge he wi h he o ma ion o cu en io São F ancisco,
we e esponsible o isola ing his egion. The a ea om
sou he n coas al Bahia o he no he n po ion o Espí i o
San o is inhabi ed by many endemic axa, and is conside ed a
“ho spo ” o biodi e si y (Bize il, 1994; Chomi z e al., 2005;
Galindo-Leal & Câma a, 2005; Rocha e al., 2005). Examples o
endemic ishes om ha egion can be ound in he s udies
o Menezes (1988), Wei zman e al. (1988), Zana a & Akama
(2004), B i o e al. (2005), Ga a ello (2005), Lucinda (2005),
Be aco & Lucena (2006), Sa men o-Soa es e al. (2006a, b),
Zana a & Camelie (2009), among o he s. Many o he
o ganisms a e also known o be endemic o his egion,
including plan s (Ha ley, 1988; Sode s om e al., 1988;
Thomas e al., 1998), amphibians (C uz & Pimen a, 2004), liza ds
(Rod igues e al., 2002; Pelleg ino e al., 2005), bi ds (Sil a e
al., 2004; Sil ei a e al., 2005), and mammals (Emmons &
Vuce ich, 1998; Ven u a e al., 2004).
Hoplias cu upi a, new species
Fig. 5
Hoplias a . lace dae: -Lima e al., 2005: 155 [species lis ,
occu ence in he io Tiquié, ibu a y o uppe io Neg o,
biological, ecological and e hnoich hyological no es].
Holo ype. BRAZIL. Pa á S a e. Tocan ins basin: MZUSP 45582,
157.9 mm SL, io I acaiúnas, Caldei ão, Se a dos Ca ajás, 05º45’S
50º30’W, Ma 1983, M. Goulding.
Pa a ypes. BRAZIL. Pa á S a e. Tocan ins basin: INPA 26893, 3,
156.4-219.3 mm SL io I acaiúnas, Caldei ão, 15 Oc 1983, M.
Goulding; MCP 41288, 4, 128.2-211.2 mm SL, same locali y, 14
Oc 1983; MNRJ 30395, 4, 136.1-168.7 mm SL, same locali y and
da e; MZUSP 33481, 1, 250.0 mm SL, same locali y and collec o ,
Jun-Jul 1983; MZUSP 22283, 4, 94.9-225.7 mm SL, Iga apé Pacuí
on km 97 o Belém-B asília oad, 15 Aug - 20 Oc 1959, Expedição
Depa amen o de Zoologia.
Ma e ial examined. BRAZIL. Amazonas S a e. Neg o basin:
MZUSP 34011, 2, 216.4-220.8 mm SL; MZUSP 34013, 1, 271.5
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
132
88, 13 b anched ays in one specimen and 14 b anched ays in
six specimens).
Well de eloped cycloid scales imb ica ed along body.
Do sal scales begin in se ies a pos e io ma gin o pa ie als
and o e lap sup aoccipi al spine. Las e ical se ies o
scales on caudal peduncle o ms sligh ly con ex a ch on
caudal- in base in la e al iew. An e io ma gin o scales
wi h small ecess and pos e io ma gin ounded.
App oxima ely eigh adii ex ending om cen e o scale o
i s an e io ma gin and a ound eigh een adii, some
anas omosed, ex ending om cen e o scale o i s do sal,
pos e io , and en al ma gins. La e al line s aigh and
comple e, ex ending om pos e o en al ma gin o
sup aclei h um o pos e io mos scale in body. La e al line
wi h 42-46 (n = 90, 42 in i e specimens and 46 in ou
specimens; mode: 44, n = 36) pe o a ed scales (Table 2).
La e al-line scales wi h single la e osenso y canal.
Longi udinal se ies o scales be ween la e al line and do sal-
in o igin 5.5-6.5 (n = 92; mode: 5.5, n = 86). Longi udinal
se ies o scales be ween la e al line and pel ic- in o igin 4.5-
5.5 (n = 93; mode: 5.5, n = 75). Longi udinal se ies o scales
a ound caudal peduncle 20 (n = 92).
App oxima ely 9 gill ake s on i s epib anchial, mos in
o m o small den icula ed pla es. Lowe b anch o i s
b anchial a ch wi h i e o six mo e elonga e ake s and
app oxima ely 10 pla e-like ake s (n = 96). La e osenso y canal
along en al su ace o den a y wi h 4-6 po es (n = 97; mode:
5, n = 74) (Table 3).
Colo in alcohol. G ound colo a ion o head and body da k
o ligh b own, da ke do sally and pale en ally. Ven al
egion whi e in se e al specimens. App oxima ely se en
da k pa allel diagonal ba s along do sal egion, ba s
ex ending an e o en ally o jus below la e al line.
Diagonal ba s in many specimens con inuing
pos e o en ally o app oxima ely wo longi udinal scale
se ies below la e al line, esul ing in che on-like pa e n.
Do sal po ions o ba s mo e e iden han en al egion.
Ba s mo e conspicuous in specimens wi h ligh b own
g ound colo a ion. Do sal su ace o head da k b own. Lips
wi h al e na ing da k and ligh e ical bands, bu
comple ely black in some o la ges specimens examined
(ca. 300 mm SL). Ven al su ace o den a ies a ying om
whi e in small specimens (ca. 40 mm SL), o al e na ing
da k and ligh ans e se s ipes in specimens a ound 130
mm SL, o comple ely da k in la ge specimens. Colo a ion
o in ao bi al egion and do sal su ace o head simila .
Many specimens wi h wo da k s ipes adia ing pos e io ly
om eye along in ao bi al 6 and do sal po ion o
in ao bi al 3. G ound colo a ion o ope cula se ies da k
b own. Ope cula memb ane usually ligh e han ope cle.
All ins ligh b own, ligh e han body in specimens smalle
han 130 mm SL, wi h da k spo s on ays and in e adial
memb anes o ming pa e n o i egula da k s ipes. S ipes
on anal and caudal ins wide and mo e egula han hose on
do sal in. Ven al su aces o pec o al and pel ic ins ei he
ligh e han, o wi h same pa e n as, do sal su ace, bu wi h
pa e n less conspicuous. Fins o la ges specimens (ca. 300
mm SL) usually da ke han es o body.
Dis ibu ion. In he io São F ancisco basin and uppe eaches
o io Pa aná basin, including ios G ande, Pa anaíba and
Piqui i. Hoplias in e medius also occu s in ibu a ies o io
Doce, Minas Ge ais S a e (Fig. 3).
Rema ks. Syn ypes o Mac odon in e medius a e
deposi ed in he BMNH collec ion, and pe mi i s clea
placemen in he Hoplias lace dae g oup. Da a p esen ed
he ein ma ches in o ma ion om he o iginal desc ip ion o
M. in e medius. Oyakawa (1990) included he nominal
species E y h inus mic ocephalus Agassiz, 1829 and
Mac odon in e medius Gün he , 1864 in he Hoplias
lace dae g oup, wi h la e being conside ed a junio
synonym o he o me (Oyakawa, 2003). These names a e
a ailable o he axon occu ing in he uppe io São
F ancisco d ainage, since E. mic ocephalus was desc ibed
om “ io São F ancisco” and M. in e medius om “ io Cipó”,
one o i s ibu a ies. As discussed abo e, many specimens
o species desc ibed by Spix and Agassiz (1829) we e los
du ing Wo ld Wa II, as well as he holo ype o E y h inus
mic ocephalus is also missing (Te o al, 1983; Ko ela , 1988).
Unlike he si ua ion wi h E y h inus b asiliensis (see abo e),
he o iginal desc ip ion o E. mic ocephalus, al hough
men ioning ha he ongue is eden a e, is unclea as o he
alignmen o he con ala e al den a ies no ing only ha
“b anches o he lowe jaw na ow and con ex h oughou
hei leng h, a he apa when he mou h is closed and
dis an om he o he by a g ea e space han hei
ans e se diame e ” [ou ansla ion]. This makes i di icul
o de e mine whe he Hoplias mic ocephalus belongs o
he H. lace dae o o he H. malaba icus species g oups.
Nei he Hoplias mic ocephalus no H. in e medius ha e
been ecen ly used, excep o a ca alog based on p elimina y
da a (Oyakawa, 2003), bu p e ious au ho s included bo h
epi he s as synonyms o H. malaba icus (e.g. S eindachne ,
1874; Lü ken, 2001). The younge name, H. in e medius is
ce ainly a membe o he H. lace dae g oup whe eas he
olde one, H. mic ocephalus canno be unambiguously
assigned o his g oup. Since nei he hese names has been
used ecen ly, H. in e medius is he ein chosen o designa e
his axon.
In desc ibing Mac odon in e medius, Gün he (1864) ci ed
wo specimens, bu made no e e ence o ei he being he
holo ype, and bo h a e hus syn ypes o he species.
Following A icle 74 o he In e na ional Code o Zoological
Nomencla u e (ICZN, 1999), one o he syn ypes is designa ed
a lec o ype (BMNH 1861.5.16.6) (Fig. 9), and he o he becomes
a pa alec o ype (BMHN 1861.5.16.7).
Hoplias in e medius has an in e es ing disjunc
dis ibu ion, occu ing in he uppe s e ches o io São
F ancisco, io Doce and io Pa aná. In he la e basin, he
species is no known o occu downs eam o he Se e
O. T. Oyakawa & G. M. T. Ma ox 133
Quedas. Simila dis ibu ions a e epo ed o o he ish
species (Menezes, 1988; Va i & Ha old, 2001). De gam e al.
(2002) demons a ed ha popula ions o H. malaba icus om
he uppe io Doce, uppe io Pa aná, and uppe io Pa aíba
do Sul a e gene ically close, indica ing a possible common
o igin o ecen exchanges be ween hese basins. Viei a e
al. (2005) also sugges ed a common ances y be ween he
ios São F ancisco and Doce based on hei ich hyo auna.
Cu en knowledge o he geological his o y o eas e n
Sou h Ame ica indica es ha he o igin o coas al i e s
in ol ed successi e e osion o he eas e n ma gin o he
pla o m associa ed wi h consequen upland i e cap u e,
gi ing ise o he a ious independen basins d aining in o
he A lan ic Ocean (Po e , 1997; Ribei o, 2006). Acco ding
o hese au ho s, six majo upli e en s c ea ed megadomes
ha wo ked syne gis ically wi h ma ginal e osion and upland
s eam cap u e along he A lan ic coas o Sou h Ame ica
when he la e and A ica we e sepa a ed du ing he opening
o he A lan ic Ocean. One o hese megadomes, he
“Man iquei a-Angola”, is ep esen ed in B azil by highlands
ex ending om São Paulo and Rio de Janei o s a es no h o
Minas Ge ais s a e, and was esponsible o isola ing he
uppe s e ches o he ios São F ancisco, Pa aná and Doce.
The e o e, i is possible ha he disjunc dis ibu ion pa e n
o Hoplias in e medius, as well as o o he examples ci ed
abo e, esul s om a common his o y o hese basins.
Goeldi (1898) ci ed Mac odon in e medius in he Ilha de
Ma ajó, in he mou h o io Amazonas. Unde cu en
unde s anding o he dis ibu ion o H. in e medius i is
clea ha hese eco ds o he species in he Amazon basin
we e based on misiden i ica ions. The only species o H.
lace dae g oup occu ing in he icini y o Ilha de Ma ajó is
H. cu upi a, desc ibed in he p esen s udy. Howe e , he
numbe o la e al line scales (43) epo ed by Goeldi (1898)
does no ma ch wi h he ange egis e ed o H. cu upi a
(34-39).
Al es & Pompeu (2001), Pompeu & Al es (2003), Viei a e
al. (2005) and Sil a e al. (2006a) s a ed ha Hoplias a .
lace dae (= H. in e medius) was in oduced in o io São
F ancisco because i was o iginally desc ibed om io Ribei a
de Iguape and was missing om ea lie ich hyological su eys
in he io São F ancisco. The absence o H. in e medius in
p e ious epo s o he São F ancisco ish auna is due o he
ac ha pas na u alis s ailed o ecognize wo g oups o
species in ha basin based on he o ien a ion o he
con ala e al den a ies (Lü ken, 2001).This esul ed in
axonomic con usion and he placemen o H. in e medius in
he synonym o H. malaba icus. A e hese wo g oups had
been o mally de ined by Oyakawa (1990), Hoplias a .
lace dae (= Hoplias in e medius) began o be included in
he ich hyological su eys o he io São F ancisco, bu was
e oneously ega ded as an exo ic species in oduced om
he io Ribei a de Iguape basin.
Al hough se e al ka yo ypes ha e been epo ed o he
Hoplias malaba icus g oup (e.g. Be ollo e al., 1983; De gam
& Be ollo, 1990; Be ollo e al., 2000), only one ka yo ype
has been desc ibed o specimens o he Hoplias lace dae
g oup collec ed in he uppe io Pa aná basin (Be ollo e al.,
1978). The diploid numbe ci ed by hese au ho s, 2n = 50, is
ega ded he ein as belonging o H. in e medius, as ha species
occu s in he uppe io Pa aná basin. Fu he ka yo ype and
molecula s udies in ol ing Hoplias in e medius om io São
F ancisco and io Doce basins would g ea ly aid in he
knowledge o his species ega ding i s disjunc dis ibu ion.
Hoplias in e medius inhabi s main i e s channels and
ma ginal lagoons (Viei a e al., 2005). Ju eniles we e cap u ed
in loodplain lakes (Meschia i e al., 2000). The species is
common in ese oi s (e.g. Cunico & Agos inho, 2006; Sil a
e al., 2006a) and has economic impo ance (Al es & Pompeu,
2001; Viei a e al., 2005). I s pisci o ous habi (Al im & Pe e ,
2004) makes i ulne able o he bioaccumula ion o me cu y.
In he io das Velhas, a ibu a y o he io São F ancisco, i s
lesh had he highes pe mi ed le el o me cu y (Al es &
Pompeu, 2001).
Fig. 9. Hoplias in e medius (lec o ype o Mac odon in e medius) BMNH 1861.5.16.6, 533.4 mm SL, io Cipó, Minas Ge ais
S a e, B azil.
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
134
Hoplias lace dae Mi anda-Ribei o, 1908
Fig. 10
Hoplias lace dae Mi anda-Ribei o, 1908: 3 [o iginal
desc ip ion, ype locali y: io Ribei a de Iguape, Ipo anga,
São Paulo s a e, B azil; Holo ype: MNRJ 211, d ied
specimen]. -Eigenmann, 1909: 374 [ e e ence, occu ence
in he io Ribei a de Iguape]. -Eigenmann, 1910: 448
[ e e ence, occu ence in he io Ribei a de Iguape]. -Fowle ,
1950: 364 [in species lis ]. -Aze edo e al., 1965: 103
[ edesc ip ion, di e ences om H. malaba icus]. -B i ski,
1972: 81-82 [in pa , species lis , di e ences om H.
malaba icus, occu ence in he io Ribei a de Iguape basin].
-Gé y e al., 1987: 368 [iden i ica ion key]. -Bize il, 1994: 56
[in pa , occu ence in eas e n i e basins]. -Bize il & Lima,
2000: 107 [occu ence in he io Ribei a de Iguape]. -
Oyakawa, 2003: 239 [species lis , dis ibu ion]. -Oyakawa
e al., 2006: 166 [species lis wi h pic u es, di e ences om
H. malaba icus]. -Oyakawa & Ne o-Fe ei a, 2007: 64
[species lis , dis ibu ion]. -Menezes e al., 2007: 153 [ ype-
locali y, dis ibu ion].
Hoplias a . lace dae: -Malaba ba, 1989: 127 [ ema ks on
occu ence in he io Jacuí, Rio G ande do Sul]. -Zaniboni
Filho e al., 2004: 104 [biological and ecological ea u es,
wi h pic u e].
Hoplias lace ae: -Gé y, 1977: 102 [una ailable name, lapsus
calami, ema ks on alidi y o name, iden i ica ion key].
Holo ype. BRAZIL. São Paulo S a e: MNRJ 211, 750.0 mm SL,
io Ribei a de Iguape, Ipo anga, no da e, R. K one. (Fig. 11).
Ma e ial examined. BRAZIL. São Paulo S a e. Ribei a de Iguape
basin: MNRJ 11217, 1, 443.0 mm SL, io Juquiá, ibu a y o io
Ribei a de Iguape, Regis o; MZUSP 7900, 1, 625.0 mm SL;
MZUSP 7901, 1, 380.3 mm SL, io Ribei a de Iguape; MZUSP
38569, 1, 157.2 mm SL, io Ribei a de Iguape, Regis o; MZUSP
7902, 1, 367.0 mm SL; MZUSP 37989, 2, 279.4-417.2 mm SL, io
Pa do on he oad Ipo anga-Ba a do Tu o, 14 km om Ba a do
Tu o; MZUSP 45433, 1, 234.9 mm SL, io Ribei a de Iguape by
io Pilões, Eldo ado; MZUSP 51104, 1, head leng h 244.5 mm SL,
io Juquiá, Regis o; MZUSP 61748, 1, 150.5 mm SL, io And é
Lopes, Ipo anga; MZUSP 72953, 1, 388.6 mm SL, io Jacupi anga,
Regis o. San a Ca a ina S a e. U uguay basin: MCP 12339, 1,
202.0 mm SL; MCP 12365, 1, 65.3 mm SL, io Canoas be ween
Va gem and São José do Ce i o, Campos No os; MCP 12731, 1,
295.5 mm SL; MCP 12732, 1, 224.53 mm SL; MCP 12879, 2,
130.7-245.0 mm SL, io Canoas a Passo do Canoas, SC-458,
Campos No os; MCP 12338, 1, 218.8 mm SL; MCP 12745, 2,
242-249.3 mm SL; MCP 13143, 1, 152.9 mm SL; MCP 12878, 3,
297.2-344.4 mm SL, io Canoas, on Abdon Ba is a-Ani a Ga ibaldi
oad, Campos No os; MCP 12884, 1, 387.8 mm SL; MCP 12898,
3, 63.4-147.1 mm SL; MCP 41003, 1, 115.3 mm SL, io Canoas,
on Abdon Ba is a-Ani a Ga ibaldi oad; MCP 13420, 1, 180.0
mm SL; MCP 18427, 1, 203.8 mm SL; MCP 18828, 1, 226.6 mm
SL; MCP 18925, 2, 242.6-301.0 mm SL; MCP 19151, 1, 402.1
mm SL; MCP 19035, 2, 243.1-325.6 mm SL, io do Peixe in Vol a
G ande, Concó dia; MCP 19146, 1, 411.5 mm SL; MCP 41004,
1, 253.3 mm SL, io Riacho G ande, on Pi i iba-BR 153 oad,
Concó dia; MCP 19147, 1, 415.1 mm SL, io U uguai in I á; UFRGS
7530, 1, 238.6 mm SL, io Jacu inga nea wa e cap u e, Concó dia.
Rio G ande do Sul S a e. U uguay basin: INPA 26892, 1, 144.1
mm SL; MCP 12719, 3, 130.5-185.3 mm SL, a oio Passo Al o,
São Nicolau; MCP 10896, 3, 110.0-207.7 mm SL, io Ga upá,
ibu a y o io Qua aí, on U uguaiana-Qua aí oad; MCP 11226,
1, 142.3 mm SL, ibu a y o io Qua aí-Mi im on Qua aí-Aleg e e
oad, Aleg e e; MCP 11231, 4, 120.0-179.9 mm SL; MNRJ 30393,
1, 145.7 mm SL, io Ga upá on Aleg e e-Qua aí bo de ; MCP
12089, 1, 288.8 mm SL; MCP 13421, 1, 301.1 mm SL; MNRJ
30394, 1, 302.1 mm SL, io Ligei o, on Ma celino Ramos-
Maximiliano de Almeida oad; MZUSP 37254, 1, 107.6 mm SL;
MZUSP 43683, 1, 196.3 mm SL, io Ga upá, ibu a y o io
Qua aí; MCP 12674, 1, 284.3 mm SL, a oio Canoin on Pi apó-
São Nicolau oad, São Nicolau, Rio G ande do Sul; MCP 13210, 1,
285.8 mm SL, mou h o io Ijuí-Mi im, Pi apó; MCP 14228, 1,
Fig. 10. Hoplias lace dae, MZUSP 61748, 150.5 mm SL, io Ribei a de Iguape, São Paulo S a e, B azil.
O. T. Oyakawa & G. M. T. Ma ox 135
28.4 mm SL, ibu a y o io Neg o on Bagé-Aceguá oad; MCP
14280, 1, 332.2 mm SL, a oio Ibicuí da Faxina, San ana do
Li amen o; MCP 16802, 2, 304.6-349.2 mm SL, io Ijuizinho
nea Ce missões dam; MCP 18875, 1, 250.3 mm SL, io U uguai
in Ma celino Ramos; MCP 16818, 276.8 mm SL, io Bu icá nea
CERTHIL dam, Independência; MCP 18924, 1, 251.4 mm SL, io
U uguai in Ma celino Ramos; MCP 19385, 2, 86.0-124.6 mm SL,
io Canoas, on Tupi inga-Celso Ramos oad, Tupi inga; MCP
19378, 2, 288.4-309.4 mm SL, io Pelo as a Passo da Ped a O ei a,
Esme alda; MCP 21650, 1, 170.8 mm SL, io Qua aí-Mi im, on
U uguaiana-Qua aí oad; MCP 26972, 2, 43.0-43.7 mm SL, io
Ibi apui ã, unde b idge ups eam om he Biological Rese e o
Ibi apui ã, Aleg e e; MCP 27452, 1, 360.5 mm SL, A oio Lagoão,
io Ibicuí d ainage, on San iago-Manoel de F ei as oad; MCP
27481, 1, 144.4 mm SL, io I u, ibu a y o io Ibicuí, San iago;
UFRGS 8394, 6, 68.8-94.9 mm SL, a oio Pi aí, ibu a y o io
Neg o be ween Bagé and Dom Ped i o, Bagé. Rio G ande do Sul
S a e. Jacuí basin: MCP 30714, 1, 230.7 mm SL, io Jacuí-Mi im,
Ca azinho; MZUSP 25026, 2, 215.9-225.4 mm SL, io Jacuí a
E nes ina dam nea Passo Fundo. URUGUAY. A igas. U uguay
basin: MCP 17616, 1, 79.2 mm SL, a oyo A umbeba, ci ca o 22
km Sou heas o A igas; UFRGS 8059, 1, 101.3 mm SL, a oyo
Mandiyú, ibu a y o ío U uguay, A igas. Ce o La go.
U uguay basin: MCP 9878, 1, 288.6 mm SL; MCP 9880, 1, 301.9
mm SL; MCP 9896, 1, 506.0 mm SL; MCP 9926, 1, 399.7 mm SL,
ío Neg o a A e ia. Tacua embó. U uguay basin: MZUSP 52006,
4, 289.6-321.9 mm SL, ío Neg o, Rincón del Bone e.
Diagnosis. Hoplias lace dae di e s om he o he species
o H. lace dae g oup in he numbe o po es o he
la e osenso y sys em along he en al su ace o den a y (6-
8 s. 4-6 in H. b asiliensis and H. in e medius, always 5 in H.
aus alis and 4 in H. cu upi a). I can be u he dis inguished
om H. b asiliensis and H. cu upi a, in he numbe o scales
along la e al line (43-48 s. 38-43 and 34-39 espec i ely).
Desc ip ion. Mo phome ic da a p esen ed in Table 7. Body
cylind ical, deepe han wide. G ea es body dep h a e ical
h ough i h scale an e io o do sal- in o igin in specimens
smalle han 70 mm SL, close o do sal- in o igin in la ge
specimens. An e io p o ile o head sligh ly ounded in la e al
iew, mo e angula in specimens smalle han 160 mm SL.
Do sal p o ile o head almos s aigh . Do sal ma gin o o bi
a ho izon al h ough o do sal p o ile o head in specimens
smalle han 100 mm SL bu no eaching do sal p o ile o
head in la ge specimens. Do sal p o ile o body sligh ly
con ex om e ical h ough i s se ies o scale o body o
do sal- in o igin; s aigh and pos e o en ally inclined along
do sal- in base; s aigh and less inclined o sligh ly conca e
om e ical h ough base o las do sal- in ay o o igin o
do sal mos p ocu en caudal- in ay. La e po ion sligh ly
mo e conca e in specimens smalle han 100 mm SL. Ven al
p o ile o lowe jaw dis inc ly angula in egion o mandibula
symphysis, s aigh o sligh ly inclined om e ical h ough
an e io nos il o pos e io ma gin o lowe jaw. Medial
ma gins o con ala e al den a ies unning in pa allel (Fig. 1a-
c). Ven al p o ile o body sligh ly con ex o pel ic- in o igin;
s aigh o app oxima ely con ex om la e poin o anal- in
o igin; s aigh and pos e odo sally inclined along anal- in
base; s aigh o sligh ly conca e om las anal- in ay o
an e io mos en al p ocu en caudal- in ay.
Uppe jaw sligh ly sho e han lowe jaw, mo e so in la ge
specimens (ca. 380 mm SL). Pos e io po ion o maxilla
do sally enla ged and ex ending medially o an e io ma gins
o second and hi d in ao bi als. Uppe and lowe lips leshy
wi h sho skin p ojec ions co e ing canines ex e nally.
An e io nos il ubula wi h an e io sli along i s dis al hal .
An e io and pos e io nos ils si ua ed along ho izon al
h ough cen e o o bi , an e io nos il loca ed a app oxima e
one o bi al diame e om an e io ma gin o o bi ; pos e io
nos il midway be ween an e io nos il and an e io ma gin
o o bi . Eye p opo iona ely la ge in smalle specimens.
In ao bi al bones well de eloped and ho izon ally elonga e.
P eope cle bea ing inume ous la e osenso y sys em po es
and pa ially co e ed by pos e io ma gin o in ao bi als 3,
4, and pos e o en al po ion o in ao bi al 5. An e o en al
ma gin o in ao bi al 3 ela i ely s aigh and pos e o en al
ma gin con ex. Pos e io ma gin o in ao bi al 4 ela i ely
s aigh and ma gins o in ao bi als 5 and 6 sligh ly con ex.
Small specimens (ca. 70 mm SL) wi h in ao bi al 3 ba ely
eaching o bi al im and p oximal ends o in ao bi als 2 and 4
in con ac . La ge specimens (ca. 160 mm SL) wi h in ao bi al
4 ba ely eaching o bi al im and p oximal ends o in ao bi als
2 and 5 almos in con ac . Specimens la ge han 380 mm SL
wi h in ao bi al 5 p og essi ely sepa a ed om im. P oximal
ends o in ao bi als 2 and 6 almos mee ing in some o la ges
specimens examined (ca. 750 mm SL).
Tee h in bo h jaws conical o canine. P emaxilla y ee h in
single ow. Fi s p emaxilla y oo h la ge canine, and i h
Table 7. Mo phome ic da a o Hoplias lace dae. S anda d
leng h in mm; alues 1-14 a e pe cen s o s anda d leng h and
alues 15-22 a e pe cen s o head leng h. n = numbe o
examined specimens.
Holo ype n Mean Range S anda d
de ia ion
S anda d leng h 750.0 93 - 28.4-750.0 -
1. Body dep h - 88 21.4 16.8-25.1 1.8
2. Head leng h 28.0 89 31.1 27.6-34.9 1.5
3. Pec o al- in leng h - 89 16.9 11.6-19.2 1.3
4. Pel ic- in leng h - 89 17.9 15.0-20.4 1.3
5. Anal- in leng h - 89 17.3 13.2-19.7 1.2
6. Do sal- in leng h - 89 26.2 23.1-30.6 1.6
7. Do sal- in base 13.9 89 15.5 13.6-18.0 0.9
8. Anal- in base 6.2 89 9.1 6.2-11.3 0.8
9. P e-pec o al dis ance - 88 28.5 24.6-34.4 1.7
10. P e-pel ic dis ance 56.7 88 56.0 50.6-60.8 2.1
11. P e-do sal dis ance 50.0 88 51.3 47.6-56.9 2.0
12. P e-anal dis ance 82.0 88 81.4 75.1-87.6 2.6
13. Caudal-peduncle dep h 11.0 88 13.1 10.8-15.1 0.7
14. Caudal-peduncle leng h 12.0 88 13.6 11.0-16.4 1.0
15. Head dep h 45.1 90 41.7 37.8-47.1 2.0
16. Snou leng h 26.3 90 25.1 22.5-29.0 1.1
17. Snou wid h - 90 18.9 13.6-22.9 1.3
18. Snou dep h - 89 19.1 16.2-21.8 1.3
19. P e-nasal dis ance - 90 15.0 11.4-19.1 1.2
20. O bi al diame e 8.0 90 18.5 8.0-27.8 3.7
21. In e o bi al wid h 31.0 90 21.1 15.6-30.1 2.9
22. Uppe jaw leng h 51.4 90 47.7 44.9-51.8 1.5
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
136
oo h medium sized. Six h oo h canine and almos as la ge as
an e io mos p emaxilla y canine. Second o ou h, se en h
and eigh h p emaxilla y ee h small. Maxilla wi h single ow
o app oxima ely 38 ela i ely small ee h, excep o e y well
de eloped canine-like ou h oo h. Den a y wi h an e io
ex e nal ow o ee h and pos e io in e nal ow. Ex e nal se ies
wi h h ee an e io small ee h, ollowed by wo well de eloped
canines, wi h pos e io canine la ge han an e io canine and
hen en conical ee h, sligh ly smalle han an e io mos
den a y canine. In e nal se ies beginning a le el o las conical
oo h o ex e nal ow and composed o app oxima ely 15 e y
small ee h. Accesso y ec op e ygoid and ec op e ygoid
oo hed. Ec op e ygoid wi h se ies o small conical ee h along
i s en ola e al ma gin and many smalle ili o m ee h
medially on i s en al su ace. Endop e ygoid eden ulous.
Dis al ma gins o all ins ounded. To al do sal- in ays
12-15 (n = 90; ii,10-13; mode: ii,11, n = 66). Do sal in loca ed a
midbody, i s o igin a e ical h ough app oxima ely ou h
scale an e io on se ies along pel ic- in o igin. Longes do sal-
in ay app oxima ely h ee-qua e s o body dep h in
specimens smalle han 160 mm SL and almos hal o body
dep h in la ge specimens. Anal- in base sho . To al anal-
in ays 9-12 (n = 89; ii,7-10, mode: ii,8, n = 56). Tip o
dep essed do sal in eaching e ical h ough anal- in o igin
in smalle specimens (< 160 mm SL), bu alling sho o
e ical h ough ha poin in la ge specimens. To al pec o al-
in ays 11-15 (n = 89; i,10-14; mode: i,12, n = 35). Pec o al- in
o igin loca ed a e ical h ough cen al po ion o ope cle.
Tip o pec o al in sepa a ed om pel ic- in o igin by i e o
nine scales. Pec o al and pel ic ins o simila size, sligh ly
la ge han anal in in specimens la ge han 380 mm SL.
Pel ic- in ays i,7 (n = 86). Pel ic- in o igin si ua ed a
midbody app oxima ely ou scales pos e io o e ical
h ough do sal- in o igin. Tips o pel ic in sepa a ed om
e ical h ough anal- in o igin by i e o se en scales.
Caudal- in ays i,15,i (n = 15).
Well de eloped cycloid scales imb ica ed along body.
Do sal scales begin in se ies a pos e io ma gin o pa ie als
and o e lap sup aoccipi al spine. Las e ical se ies o scales
on caudal peduncle o ms almos s aigh line on caudal- in
base in specimens smalle han 160 mm SL, and sligh ly con ex
a ch on caudal- in in la ge specimens. An e io ma gin o
scales wi h small ecess and pos e io ma gin ounded. Eigh
o ele en adii ex ending om cen e o scale o i s an e io
ma gin and a ound eigh een adii, some anas omosed,
ex ending om cen e o scale o i s do sal, pos e io and
en al ma gins. La e al line s aigh and comple e, ex ending
om pos e o en al ma gin o sup aclei h um o pos e io mos
scale in body. La e al line wi h 43-48 (n = 89, 43 in one specimen;
mode: 46, n = 28) pe o a ed scales (Table 2). La e al-line scales
wi h single la e osenso y canal. Longi udinal se ies o scales
be ween la e al line and do sal- in o igin 4.5-6.5 (n = 88; mode:
5.5, n = 61). Longi udinal se ies o scales be ween la e al line
and pel ic- in o igin 4.5-6.5 (n = 86; mode: 4.5, n = 63).
Longi udinal se ies o scales a ound caudal peduncle 18-22
(n = 88; mode: 20, n = 73).
App oxima ely 11 gill ake s on i s epib anchial, mos in
o m o small den icula ed pla es. Lowe b anch o i s
b anchial a ch wi h i e o se en mo e elonga e ake s and
app oxima ely 12 pla e-like ake s (n = 18). La e osenso y canal
along en al su ace o den a y wi h 6-8 po es (n = 90; mode:
7, n = 67) (Table 3).
Colo in alcohol. G ound colo a ion o head and body da k o
ligh b own, da ke do sally and pale en ally. Ven al egion
whi e o ligh yellow in mos specimens. Single da k midla e al
s ipe along la e al line s ipe mo e e iden in specimens smalle
han 100 mm SL. Midla e al s ipe c ossed by i e o se en
da k ba s, each h ee o i e scales wide. Diagonal ba s in
many specimens con inuing pos e o en ally app oxima ely
wo o h ee longi udinal se ies o scales en al o la e al
line, esul ing in che on-like pa e n mo e conspicuous in
specimens smalle han 160 mm SL. Do sal po ions o ba s
mo e e iden han en al egion. G ound colo a ion o do sal
su ace o head, in ao bi al egion and ope cula se ies da k
b own, wi h da k e icula ed pa e n on do sal su ace o head
in some specimens. Lips wi h al e na ing da k and ligh e ical
bands in specimens smalle han 160 mm SL and comple ely
da k in la ge specimens. Colo a ion o in ao bi al egion
simila o do sal su ace o head. Many specimens wi h one
o wo da k s ipes ex ending adially and pos e io ly om
eye ac oss ope cula se ies and hi d s ipe ex ending
pos e o en ally om eye ac oss pos e io egion o maxilla.
G ound colo a ion o ope cula se ies da k b own. Ope cula
memb ane usually ligh e han ope cle.
All ins ligh b own (ligh e han body) wi h da k spo s on
ays and in e adial memb anes o ming pa e n o i egula
da k s ipes. S ipes o do sal and caudal ins wide han hose
o anal in. S ipes less egula in do sal in. Ven al su aces
o pec o al and pel ic ins ei he ligh e han o wi h same
pa e n as do sal su ace, bu wi h pa e n less conspicuous.
Dis ibu ion. Known om he ype-locali y and i s icini y,
io Ribei a de Iguape basin, and om he io U uguay basin,
including he ío Neg o in U uguay (Fig. 3).
Rema ks. Hoplias lace dae has been ega ded as ha ing a
wide dis ibu ion, occu ing in he io Ribei a de Iguape as
well as in i e s o he Amazon, Pa aná, São F ancisco, U uguay
and o he basins. Resul s p esen ed he ein, howe e ,
demons a e ha specimens ound in he io Ribei a a e
mo phologically di e en om hose o emaining ci ed basins
wi h he excep ion o io U uguay. The dis ibu ion o Hoplias
lace dae is he e o e es ic ed o io Ribei a de Iguape and
io U uguay, being in sympa y wi h Hoplias aus alis in he
la e basin.
Al hough samples om he io Ribei a and io U uguay
did no di e signi ican ly in he me is ic and mo phome ic
ea u es examined in he p esen s udy, some geog aphical
dispa i ies should be poin ed ou . Mos specimens om he
io Ribei a ha e 6.5 and 5.5 scales abo e and below la e al
line, espec i ely, while he majo i y o specimens om he io
O. T. Oyakawa & G. M. T. Ma ox 137
U uguay ha e 5.5 and 4.5 scales abo e and below la e al line
espec i ely. Specimens om he io Ribei a end o ha e ewe
la e al line scales han do specimens om he io U uguay
(modes: 44 and 46, espec i ely) and mo e pec o al- in ays
han do samples om he io U uguay (i,13-14 s. i,10-13,
espec i ely). Ne e heless, hese sub le di e ences mus be
quali ied by he ac ha hey a e no absolu e and specimens
o H. lace dae om he io Ribei a a e a e in collec ions (only
en specimens we e a ailable o his s udy). Mo e specimens
om io Ribei a and he use o cy ogene ics and molecula
app oaches may de e mine whe he hese wo popula ions
cons i u e di e en species.
Be olle i e al. (1990) and Pessano e al. (2005) ci ed H.
lace dae om he io U uguay basin, he o me in he a ea o
Ga abi Hyd oelec ic S a ion and he la e in A oio Qua ai-
Chico. These au ho s, howe e , did no men ion he basis o
hose iden i ica ions, no did hey illus a e hei ma e ial.
The e o e, i is impossible o de e mine whe he hey we e
dealing wi h Hoplias lace dae o he new species, H. aus alis,
desc ibed he ein, bo h o which occu in he io U uguay basin.
A ew specimens o Hoplias lace dae om E nes ina
ese oi in he io Jacuí, a i e unning in o he A lan ic
Ocean in Rio G ande do Sul, sou he n B azil, we e examined.
Malaba ba (1989: 127) and Ca los Lucena (pe s. comm.)
men ioned ha he species was in oduced om io U uguay
o io Jacuí sys em in he 1980s. Na i e occu ence o Hoplias
lace dae is he e o e conside ed es ic ed o he io
U uguay and io Ribei a de Iguape d ainages. Zaniboni Filho
e al. (2004) epo ed some biological ea u es o his species
in he io U uguay including leng h, weigh , and g ow h
cu e. Acco ding o hem, he species has ac ional
spawning and an economic impo ance in local ishe ies and
aquacul u es.
In he io Ribei a de Iguape, he species is ega ded as
a e by local ishe man and a wo-yea su ey o i s
ich hyo auna ailed o yield specimens o his species
(Oyakawa e al., 2006). The small numbe o specimens o
Hoplias lace dae om io Ribei a in ich hyological collec ions
may be addi ional e idence ha his species is uncommon,
equi ing u he s udies on i s ecology and biology and
special a en ion o i s conse a ion s a us.
Acknowledgemen s
A p e ious e sion o his s udy was pa o a disse a ion
submi ed by he senio au ho in pa ial ul illmen o a
Mas e ’s deg ee in Zoology a he Uni e sidade de São Paulo.
This s udy was de eloped a he Museu de Zoologia,
Uni e sidade de São Paulo, which p o ided wo kspace and
access o acili ies. The au ho s a e deeply g a e ul o Luiz
Malaba ba (UFRGS), Pa ice P u os (MNHN), James
Maclaine and Ral B i z (BMNH) o sending pho og aphs o ,
and in o ma ion on, ype specimens. Ca los Lucena and
Robe o Reis (MCP), Luiz Malaba ba (UFRGS), Ma k Sabaj
Pé ez and John Lundbe g (ANSP) and Da id Tapho n
(MCNG) g ea ly con ibu ed o his s udy by p o iding
in o ma ion and pho og aphs, lending specimens, and
p o iding access o he collec ions. Va ious aspec s o his
s udy ecei ed aluable inpu om Mônica Toledo-Piza (IB-
USP), José Lima de Figuei edo, Naé cio Menezes, Flá io Lima
and José Bi indelli (MZUSP). Ma k Sabaj Pé ez (ANSP) and
Richa d P. Va i (USNM) g ea ly con ibu ed by kindly
e iewing an ea lie e sion o he manusc ip . The map o
Sou h Ame ica is based on one p epa ed by Ma ilyn
Wei zman. The pho og aph in Fig. 9 is copy igh ed by he
Na u al His o y Museum, London. Edua do Baena ook mos
o emaining pic u es and edi ed Fig. 1. Collec ing ips we e
suppo ed by CNPq - Conselho Nacional de
Desen ol imen o Cien í ico e Tecnológico/FINEP -
Financiado a de Es udos e P oje os/CAPES - Coo denação
de Ape eiçoamen o de Pessoal de Ní el Supe io , P ojec
No. 661058/1997-2, en i led “Conhecimen o, Conse ação e
U ilização Racional da Di e sidade da Fauna de Peixes do
B asil” sponso ed by Naé cio Menezes (MZUSP), and
Fundação de Ampa o à Pesquisa do Es ado de São Paulo
(FAPESP) wi hin he BIOTA/FAPESP - The Biodi e si y
Vi ual Ins i u e P og am (www.bio asp.o g.b ), P ojec No.
00/04300-9, en i led “Di e sidade de peixes de iachos e
cabecei as da Bacia do io Ribei a de Iguape no Es ado de
São Paulo”, sponso ed by he senio au ho . Pa o his
esea ch was unded by a Böhlke Awa d (ANSP) o he
second au ho , who also was inancially suppo ed by a
Doc o al ellowship om CNPq (P ojec No. 140141/2006-5).
Fig. 11. Hoplias lace dae, holo ype, MNRJ 211, 750.0 mm SL, io Ribei a de Iguape, São Paulo S a e, B azil.
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
138
Li e a u e Ci ed
Agos inho, A. A. & H. F. Júlio J . 1999. Peixes da bacia do Al o io
Pa aná. Pp. 374-400. In: Lowe-McConnell, R. H. Es udos ecológi-
cos de comunidades de peixes opicais. São Paulo, EDUSP, 534p.
Al es, C. B. M. & P. S. Pompeu (O gs.). 2001. Peixes do io das
Velhas: passado e p esen e. Belo Ho izon e, SEGRAC, 194p.
Al im, M. C. C. & A. C. Pe e . 2004. Food esou ces sus aining he
ish auna in a sec ion o he Uppe São F ancisco Ri e in T ês
Ma ias, MG, B azil. B azilian Jou nal o Biology, 64(2): 195-202.
Aze edo, P., J. O. Vaz & W. B. Pa ei a. 1965. Redesc ição do
ai ão, Hoplias lace dae (Ribei o). Pp. 101-106. In: Anais do
II Cong esso La ino-Ame icano de Zoologia, São Paulo, CNPq/
FAPESP, 379p.
Ba ella, W. & M. Pe e e J . 2003. Fish communi y al e a ions due
o pollu ion and damming in Tie ê and Pa anapanema Ri e s
(B azil). Ri e Resea ch and Applica ions, 19: 59-76.
Be aco, V. A. & C. A. S. Lucena. 2006. Two new species o As yanax
(Os a iophysi: Cha aci o mes: Cha acidae) om eas e n B azil,
wi h a synopsis o he As yanax scab ipinnis species complex.
Neo opical Ich hyology, 4(1): 53-60.
Be olle i, J. J. , C. A. S. Lucena, Z. M. S. Lucena, L. R. Malaba ba
& R. E. Reis. 1990. Es u u a e composição da u u a usina hid e-
lé ica de Ga abi, Rio G ande do Sul, B asil. Comunicações do
Museu de Ciências da PUCRS, sé ie Zoologia, 3(2): 33-97.
Be ollo, L. A. C., G. G. Bo n, J. A. De gam, A. S. Fenocchio & O.
Mo ei a-Filho. 2000. A biodi e si y app oach in he neo opical
E y h inidae ish, Hoplias malaba icus. Ka yo ypic su ey,
geog aphical dis ibu ion o cy o ypes and cy o axonomic
conside a ions. Ch omosome Resea ch, 8: 603-613.
Be ollo, L. A. C., C. S. Takahashi & O. Mo ei a-Filho. 1978.
Cy o axonomic conside a ions on Hoplias lace dae (Pisces,
E y h inidae). Re is a B asilei a de Gené ica, 1(2): 103-120.
Be ollo, L. A. C., C. S. Takahashi & O. Mo ei a-Filho. 1983.
Mul iple sex ch omosomes in he genus Hoplias (Pisces:
E y h inidae). Cy ologia, 48: 1-12.
Bize il, C. R. S. F. 1994. Análise axonômica e biogeog á ica da
ic io auna de água doce do les e b asilei o. Ac a Biológica
Leopoldensia, 16(1): 51-80.
Bize il, C. R. S. F. & N. R. W. Lima. 2000. Le an amen o da
ic io auna da bacia do Rio Ribei a, B asil. Ac a Biológica
Leopoldensia, 22(1): 103-110.
B i ski, H. A. 1972. Peixes de água doce do Es ado de São Paulo.
Sis emá ica. Pp. 79-108. In: Poluição e Piscicul u a. Faculdade
de Saúde Pública e Ins i u o de Pesca, São Paulo, 216p.
B i ski, H. A., Y. Sa o & A. B. S. Rosa. 1984. Manual de iden i ica-
ção de peixes da egião de T ês Ma ias: com cha es de iden i i-
cação pa a os peixes da Bacia do São F ancisco. B asília, Câma-
a dos Depu ados, Coo denação de Publicações, CODEVASF -
Di isão de Piscicul u a e Pesca, 115p.
B i o, M. R., F. C. T. Lima & A. C. A. San os. 2005. A new
Aspido as (Silu i o mes: Callich hyidae) om io Pa aguaçu
basin, Chapada Diaman ina, Bahia, B azil. Neo opical
Ich hyology, 3(4): 473-479.
Cas elnau, F. L. L. 1855. Poissons. Pp. 55-56. In: Animaux no eaux
ou a es ecueillis pendan l’Expedi ion dans les pa ies cen ales
de l’Ame ique du Sud, de Rio de Janei o a Lima, e de Lima au
Pa á; exécu ée pa o d e du Gou e nmen F ançais pendan les
années 1843 a 1847. Pa is, P. Be and. . 2, 112p.
Chomi z, K. M., K. Alge , T. S. Thomas, H. O lando & P. V. No a.
2005. Oppo uni y cos s o conse a ion in a biodi e si y
ho spo : he case o sou he n Bahia. En i onmen and
De elopmen Economics, 10: 293-312.
C uz, C. A. G. & B. V. S. Pimen a. 2004. New species o Physalaemus
Fi zinge , 1826 om sou he n Bahia, B azil (Anu a,
Lep odac ylidae). Jou nal o He pe ology, 38(4): 480-486.
Cunico, A. M. & A. A. Agos inho. 2006. Mo phological pa e ns o
ish and hei ela ionships wi h ese oi s hyd odynamics.
B azilian A chi es o Biology and Technology, 49(1): 125-134.
De gam, J. A. & L. A. C. Be ollo. 1990. Ka yo ypic di e si ica ion
in Hoplias malaba icus (Os eich hyes, E y h inidae) o he São
F ancisco and Al o Pa aná basins, B azil. Re is a B asilei a de
Gené ica, 13(4): 755-766.
De gam, J. A., S. R. Pai a, C. E. Schae e , A. L. Godinho & F. Viei a.
2002. Phylogeog aphy and RAPD-PCR a ia ion in Hoplias
malaba icus (Bloch, 1794) (Pisces, Teleos ei) in sou heas e n
B azil. Gene ics and Molecula Biology, 25(4): 379-387.
Eigenmann, C. H. 1909. The eshwa e ishes o Pa agonia and an
examina ion o he A chipla a A chelenis Theo y. Pp. 225-374.
In: Repo s on he P ince on Uni e si y expedi ions o
Pa agônia, 1896-1899, . 3, p . 1, 374p.
Eigenmann, C. H. 1910. Ca alogue o he eshwa e ishes o opi-
cal and Sou h empe a e Amé ica. Pp. 375-511. In: Repo s o
he P ince on Uni e si y expedi ions o Pa agonia, 1896-1899,
. 3, p . 2, 777p.
Eigenmann, C. H. 1912. The eshwa e ishes o B i ish Guiana,
including a s udy o he ecological g ouping o species and ela ion
o he auna o he pla eau o ha o he lowlands. Memo ies o
he Ca negie Museum, 5(1): 1-554.
Eigenmann, C. H. & R. S. Eigenmann. 1889. A e iew o he
E y h ininae. P oceedings o he Cali o nia Academy o Science,
2(2): 100-117.
Emmons, L. H. & M. G. Vuce ich. 1998. The iden i y o Winge’s
Lasiu omys illosus and he desc ip ion o a new genus o
Echimyid oden (Roden ia: Echimyidae). Ame ican Museum
No i a es, 3223: 1-12.
Fe a is, J . C. J. 2007. Checklis o ca ishes, ecen and ossil
(Os eich hyes: Silu i o mes), and ca alogue o silu i o m p ima y
ypes. Zoo axa, 1418:1-628.
Fink, W. L. & S. H. Wei zman. 1974. The so-called chei odon in ishes
o Cen al Ame ica wi h desc ip ions o wo new species (Pisces:
Cha acidae). Smi hsonian Con ibu ions o Zoology, 172: 1-46.
Fowle , H. W. 1950. Os peixes de água doce do B asil (2a en ega).
A qui os de Zoologia, 6: 205-404.
Galindo-Leal, C. & I. G. Câma a (Eds.). 2005. Ma a A lân ica:
biodi e sidade, ameaças e pe spec i as. Belo Ho izon e, Fun-
dação SOS Ma a A lân ica and Conse a ion In e na ional, 472p.
Ga a ello, J. C. 2005. Re ision o genus S eindachne idion (Silu i o mes:
Pimelodidae). Neo opical Ich hyology, 3(4): 607-623.
Ga u i, V. & H. A. B i ski. 1997. Desc ição de uma espécie no a de
As yanax (Teleos ei, Cha acidae), com mancha ume al o alada
ho izon almen e, p oceden e da bacia do Rio Guapo é, Amazô-
nia. Papéis A ulsos de Zoologia, 40(15): 217-229.
Gé y, J. 1977. Cha acoids o he Wo ld. New Je sey, T. F. H.
Publica ions, 672p.
Gé y, J., V. Mahne & C. Dlouhy. 1987. Poissons Cha acoïdes non
Cha acidae du Pa aguay (Pisces, Os a iophysi). Re ue Suisse
de Zoologie, 94(2): 357-464.
Godoy, M. P. de. 1975. Família E y h inidae. Pp. 399-444. In:
Peixes do B asil - subo dem Cha acoidei. Bacia do Rio Mogi
Guassú. Edi o a F ansciscana, São Paulo, ol. 3, 399-628 [229p].
Goeldi, E. A. 1898. P imei a con ibuição pa a o conhecimen o dos
peixes do alle do Amazonas e das Guyanas, es udos
ich hyologicos dos annos 1894-1898. Bole im do Museu
O. T. Oyakawa & G. M. T. Ma ox 139
Pa aense, 2: 443-488.
Goulding, M. 1980. The ishes and he o es . Explo a ions in
Amazonian Na u al His o y. Los Angeles, Uni e si y o
Cali o nia P ess, 280p.
Gün he , A. 1864. Cha acinidae. Pp. 278-380 In: Ca alogue o he
ishes in he B i ish Museum. London. V. 5, xxii, 455p.
Ha ley, R. M. 1988. E olu ion and dis ibu ion o E iope (Labia ae),
and i s ela i es, in B azil. Pp. 71-120. In: Vanzolini, P. E. & R.
H. Heye (Eds.). P oceedings o a wo kshop on neo opical
dis ibu ion pa e ns. Rio de Janei o, Academia B asilei a de
Ciências, 488p.
ICZN [In e na ional Commission on Zoological Nomencla u e].
1999. In e na ional Code o Zoological Nomencla u e. Fou h
Edi ion. London, In e na ional T us o Zoological
Nomencla u e, 306p.
Ko ela , M. 1988. Au ho ship, da es o publica ion, s a us and ypes
o Spix and Agassiz’s B azilian ishes. Spixiana, 11(1): 69-93.
Lima, F. C. T., L. Ramos, T. Ba e o, A. Cabalza , G. Tenó io, A.
Ba bosa, F. Tenó io & A. S. Resende. 2005. Peixes do Al o
Tiquié - Ic iologia e conhecimen os dos ukuya e ukano. Pp.
111-282. In: Cabalza , A. (O g.). Peixe e Gen e no Al o Rio
Tiquié. São Paulo, Ins i u o Socioambien al, 339p.
Lucinda, P. H. F. 2005. Sys ema ics and biogeog aphy o he genus
Phallop ychus Eigenmann, 1907 (Cyp inodon i o mes:
Poeciliidae: Poeciliinae). Neo opical Ich hyology, 3(3): 373-382.
Lü ken, C. F., 2001. Peixes do Rio das Velhas: Uma con ibuição
pa a a Ic iologia do B asil. Pp: 23-164. In: Al es, C. B. M. & P.
S. Pompeu (O g.) Peixes do Rio das Velhas: passado e p esen e.
Belo Ho izon e, SEGRAC, 194p.
Malaba ba, L. R. 1989. His ó ico sis emá ico e lis a comen ada das
espécies de peixes de água doce do sis ema da Laguna dos Pa-
os, Rio G ande do Sul, B asil. Comunicações do Museu de
Ciências da PUCRS, sé ie Zoologia, 2(8): 107-179.
Ma ox, G. M. T., M. Toledo-Piza & O. T. Oyakawa. 2006.
Taxonomic s udy o Hoplias aima a (Valenciennes, 1846) and
Hoplias mac oph halmus (Pelleg in, 1907) (Os a iophysi,
Cha aci o mes, E y h inidae). Copeia, 2006(3): 516-528.
Menezes, N. A. 1988. Implica ions o he dis ibu ion pa e ns o he
species o Oligosa cus (Teleos ei, Cha acidae) om cen al and
sou he n Sou h Ame ica. Pp. 295-304. In: Vanzolini, P. E. & R. H.
Heye (Eds.). P oceedings o a wo kshop on neo opical dis ibu ion
pa e ns. Academia B asilei a de Ciências, Rio de Janei o, 488p.
Menezes, N. A. 1992. Rede inição axonômica das espécies de
Aces o hynchus do g upo lacus is com a desc ição de uma
no a espécie (Os eich hyes, Os a iophysi, Cha aci o mes). Co-
municações do Museu de Ciências da PUCRS, sé ie Zoologia,
5: 39-54.
Menezes, N. A., S. H. Wei zman, O. T. Oyakawa, F. C. T. Lima, R.
M. C. Cas o & M. J. Wei zman. 2007. Peixes de Água Doce da
Ma a A lân ica - Lis a p elimina das espécies e comen á ios
sob e conse ação de peixes de água doce neo opicais. Museu
de Zoologia da Uni e sidade de São Paulo, 408p.
Meschia i, A. J., M. S. A ci a & N. Fene ich-Ve ani. 2000. Fish
communi ies associa ed wi h mac ophy es in B azilian loodplain
lakes. En i onmen al Biology o Fishes, 58: 133-143.
Mi anda-Ribei o, A. 1908. Peixes da Ribei a. Resul ados de excu -
são do S . Rica do K one, memb o co esponden e do Museu
Nacional do Rio de Janei o. Kosmos, 5: 1-5.
Mülle , J. 1842. Beobach ungen übe die Schwimmblase de Fische,
mi Bezug au einige neue Fischga ungen. A chi es ü
Ana omie, Physiologie und wissenscha liche Medicin (J.
Mülle ), 1842: 307-329.
Mülle , J. 1843. Bei äge zu Kenn niss de Na ü lichen Familien
de Fische. A chi es ü Na u geschich e, 9(1): 292-330.
Mülle , J. & F. H. T oschel. 1844. Synopsis gene um e specie um
amiliae Cha acino um (P odomus desc ip ionis no o um gene um
e specie um). A chi es ü Na u geschich e, 10(1): 81-99.
Mülle , J. & F. H. T oschel. 1845. Ho ae Ich hyologicae,
Besch eibung und Abbildung neue Fische. Die Familie
Cha acinen. Be lin, 40p.
Neumann, D. 2006. Type ca alogue o he Ich hyological collec ion
o he Zoologische S aa ssammlung München. Pa I: His o ic
ype ma e ial om he “Old Collec ion”, des oyed in he nigh
24/25 Ap il 1944. Spixiana, 29(3): 259-285.
Oyakawa, O. T. 1990. Re isão sis emá ica das espécies do gêne o
Hoplias (g upo lace dae) da Amazônia b asilei a e egião les e
do B asil (Teleos ei: E y h inidae). Unpublished Mas e
Disse a ion, Uni e sidade de São Paulo, São Paulo, 114p.
Oyakawa, O. T. 1998. Relações ilogené icas das amílias
Py hulinidae, Lebiasinidae e E y h inidae (Os eich hyes:
Cha aci o mes). Unpublished Ph.D. Disse a ion, Uni e sida-
de de São Paulo, São Paulo, 200p.
Oyakawa, O. T. 2003. Family E y h inidae. Pp. 238-240. In: Reis,
R. E., S. O. Kullande & C. J. Fe a is J . (O gs.). Check lis o
he eshwa e ishes o Sou h and Cen al Ame ica. Po o Ale-
g e, Edipuc s, 729p.
Oyakawa, O. T. & A. L. Ne o-Fe ei a. 2007. Família E y h inidae.
Pp. 63-64. In: Buckup, P. A., N. A. Menezes & M. S. Ghazzi
(O gs.). Ca álogo das espécies de peixes de água doce do B asil.
Rio de Janei o, Museu Nacional, Uni e sidade Fede al do Rio
de Janei o, 195p.
Oyakawa, O. T., A. Akama, K. C. Mau a i & J. C. Nolasco. 2006.
Peixes de iachos da Ma a A lân ica nas Unidades de Conse a-
ção do Vale do Rio Ribei a de Iguape no Es ado de São Paulo.
São Paulo, Edi o a Neo ópica, 201p.
Pelleg ino, K. C. M., M. T. Rod igues, A. N. Wai e, M. Mo ando,
Y. Yonenaga-Yassuda & J. W. Si es J . 2005. Phylogeog aphy
and species limi s in he Gymnodac ylus da winii complex
(Gekkonidae, Squama a): gene ic s uc u e coincides wi h i e
sys ems in he B azilian A lan ic Fo es . Biological Jou nal o
he Linnean Socie y, 85: 13-26.
Penczak, T., L. C. Gomes, L. M. Bini & A. A. Agos inho. 1998. The
impo ance o quali a i e in en o y sampling using elec ic ishing
and ne s in a la ge, opical i e (B azil). Hyd obiologia, 389:
89-100.
Pessano, E. F. C., C. L. O. Aze edo, M. V. M. Que ol, E. Que ol, L.
G. B asil, L. R. B. Cas o, T. B. Pin o & F. V. Co êa. 2005.
Ic io auna do a oio Qua ai-Chico, bacia do médio Rio U uguai,
no in e io do Pa que Es adual do Espinilho, Rio G ande do Sul,
B asil. Bio emas, 18(2): 143-153.
Planque e, P., P. Kei h & P.-Y. Le Bail. 1996. A las des poissons
d’eau douce de Guyane. Tome I. Pa is, Muséum Na ionale
d’His oi e Na u elle, 429p.
Pompeu, P. S. & C. B. M. Al es. 2003. Local ish ex inc ion in a small
opical lake in B azil. Neo opical Ich hyology, 1(2): 133-135.
Po e , P. E. 1997. The Mesozoic and Cenozoic paleod ainage o
Sou h Ame ica: a na u al his o y. Jou nal o Sou h Ame ican
Ea h Sciences, 10(5-6): 331-344.
Ribei o, A. C. 2006. Tec onic his o y and he biogeog aphy o he
eshwa e ishes om he coas al d ainages o eas e n B azil:
an example o aunal e olu ion associa ed wi h a di e gen con-
inen al ma gin. Neo opical Ich hyology, 4(2): 225-246.
Robe s, T. 1969. Os eology and ela ionships o cha acoid ishes,
pa icula ly he gene a Hepse us, Salminus, Hoplias,
Re ision o he Neo opical ahi as o he Hoplias lace dae species-g oup
140
C enolucius, and Aces o hynchus. P oceedings o he Cali o nia
Academy o Sciences, 35(15): 391-500.
Rocha, C. F. D, M. an Sluys, H. G. Be gallo & M. A. S. Al es.
2005. Endemic and h ea ened e apods in he es ingas o he
biodi e si y co ido s o Se a do Ma and o he Cen al da
Ma a A lân ica in eas e n B azil. B azilian Jou nal o Biology,
65(1): 159-168.
Rod igues, M. T., M. Dixo, D. Pa an & V. K. Ve dade. 2002. A new
species o Leposoma (Squama a, Gymnoph halmidae) om he
emnan A lan ic o es s o he s a e o Bahia, B azil. Papéis
A ulsos de Zoologia, 42(14): 335-350.
Sa men o-Soa es, L. M., R. F. Ma ins-Pinhei o, A. T. A anda & C.
C. Chamon. 2006a. Mic oglanis pa axo, a new ca ish om
sou he n Bahia coas al i e s, no heas e n B azil (Silu i o mes:
Pseudopimelodidae). Neo opical Ich hyology, 4(2): 157-166.
Sa men o-Soa es, L. M., R. F. Ma ins-Pinhei o, A. T. A anda & C.
C. Chamon. 2006b. I uglanis cahyensis, a new ca ish om
Bahia, B azil (Silu i o mes: T ichomyc e idae). Neo opical
Ich hyology, 4(3): 309-318.
Sa o, Y. & H. P. Godinho. 1999. Peixes da bacia do Rio São F ancisco.
Pp. 401-413. In: Lowe-McConnell, R. H. (Ed.). Es udos ecológi-
cos de comunidades de peixes opicais. São Paulo, EDUSP, 534p.
Sil a, A. R. M., G. B. San os & T. Ra on. 2006a. Fish communi y
s uc u e o Ju amen o ese oi , São F ancisco Ri e basin, Minas
Ge ais, B azil. Re is a B asilei a de Zoologia, 23(3): 832-840.
Sil a, F. S. D., J. R. M. Deus & A. W. S. Hilsdo . 2006b. The uppe
eaches ich hyo auna o he Tie ê Ri e , São Paulo, B azil:
aspec s o hei di e si y and conse a ion. Biodi e si y and
Conse a ion, 15: 3569-3577.
Sil a, J. M. C., M. C. Sousa & C. H. M. Cas elle i. 2004. A eas o
endemism o passe ine bi ds in he A lan ic o es , Sou h
Ame ica. Global Ecology and Biogeog aphy, 13: 85-92.
Sil ei a, L. F., P. F. De eley, J. F. Pacheco & B. M. Whi ney. 2005.
A i auna o he Se a das Lon as-Ja i mon an e complex, Bahia,
B azil. Co inga, 24: 45-54.
Sode s om, T. R., E. J. Judziewicz & L. G. Cla k. 1988. Dis ibu ion
pa e ns o neo opical bamboos. Pp. 121-157. In: Vanzolini, P.
E. & R. H. Heye (Eds.). P oceedings o a wo kshop on
neo opical dis ibu ion pa e ns. Rio de Janei o, Academia
B asilei a de Ciências, 488p.
Spix, J. B. on & L. Agassiz. 1829. Selec a gene a e species Piscium
quos in i ine e pe B azilian annis 1817-1820; Pe ac o e pigendus
cu a i D . J.B. de Spix. Digessi desc ipsi e obse a ionibus
ana omicis illus a i D . L. Agassiz. [Memo iae J.B. de Spix]
Munich: Typis C. Wol . x i, 1-6, 438p.
S eindachne , F. 1874. Die Süsswasse ische dês südös lichen
B asilien (I). Si zungsbe ich e de Kaise liche Akademie de
Wissenscha en, Wien, Ma hema isch-na u wissenscha lichen
Classe, 69: 1-40.
Tapho n, D. C. 1992. The cha aci o m ishes o he Apu e Ri e
d ainage, Venezuela. Biollania, edición especial, 4. Monog a ias
Cien í icas del Museo de Ciencias Na u ales, UNELLEZ,
Guana e, Venezuela, 537p.
Te o al, F. 1983. Die Fischausbeu e de B asilien-Expedi ion 1817-
1820 on J. B. . Spix und C. F. Ph. . Ma ius. Spixiana, 9
(supplemen ): 313-317.
Thomas, W. W., A. M. V. Ca alho, A. M. A. Amo im, J. Ga ison &
A. L. A beláez. 1998. Plan endemism in wo o es s in sou he n
Bahia, B azil. Biodi e si y and Conse a ion, 7: 311-322.
Toledo-Piza, M. 1996. Taxonomic ede ini ion o he species o
Aces o hynchus o he mic olepis g oup wi h he desc ip ion
o Aces o hynchus apu ensis, a new species om Venezuela
(Os a iophysi: Cha aci o mes: Cha acidae). Ame ican Museum
No i a es, 3160: 1-23.
Valenciennes, A. 1846. Le Mac odon aima a. In: Cu ie , G. L. C. F.
D. and A. Valenciennes. His oi e Na u elle des Poissons, . 19:
523-527. P. Be and, Pa is, 544p.
Va i, R. P. & A. S. Ha old. 2001. Phylogene ic s udy o he
neo opical ish gene a C eag u us Gün he and Piabina
Reinha d (Teleos ei: Os a iophysi: Cha aci o mes), wi h a
e ision o he cis-andean species. Smi hsonian Con ibu ions
o Zoology, 613, 239p.
Ven u a, K., M. J. J. Sil a, V. Fagundes, R. Pa dini & Y. Yonenaga-
Yassuda. 2004. An undesc ibed ka yo ype o Thap omys
(2n=50) and he mechanism o di e en ia ion om Thap omys
nig i a (2n=52) e idenced by FISH and Ag-NORs. Ca yologia,
57(1): 89-97.
Viei a, F., G. B. San os & B. M. Al es. 2005. A ic io auna do Pa que
Nacional da Se a do Cipó (Minas Ge ais, B asil) e á eas adja-
cen es. Lundiana, 6 (supplemen ): 77-87.
Wei zman, S. H. 1962. The os eology o B ycon meeki, a gene alized
cha acid ish, wi h an os eological de ini ion o he amily.
S an o d Ich hyological Bulle in, 8(1): 1-75.
Wei zman, S. H., N. A. Menezes & M. J. Wei zman. 1988.
Phylogene ic biogeog aphy o he Glandulocaudini (Teleos ei:
Cha aci o mes: Cha acidae) wi h commen s on he dis ibu ion
o o he eshwa e ishes in eas e n and sou heas e n B azil.
Pp. 379-427. In: Vanzolini, P. E. & R. H. Heye , (Eds.).
P oceedings o a wo kshop on neo opical dis ibu ion pa e ns.
Rio de Janei o, Academia B asilei a de Ciências, 488p.
Zana a, A. M. & A. Akama. 2004. Myxiops aphos, new cha acid
genus and species (Cha aci o mes: Cha acidae) om he io
Lençóis, Bahia, B azil. Neo opical Ich hyology, 2(2): 45-54.
Zana a, A. M. & P. Camelie . 2009. As yanax e milion and As yanax
buge ai: new cha acid ishes (Os a iophysi: Cha aci o mes)
om No heas e n Bahia, B azil. Neo opical Ich hyology, 7(2):
175-184.
Zaniboni Filho, E., S. Meu e , O. A. Shiba a & A. P. O. Nuñe .
2004. Ca álogo ilus ado de peixes do Al o Rio U uguai.
Flo ianópolis. Edi o a UFSC, T ac ebel Ene gia, 128p.
Accep ed Ap il 2009
Published June 17, 2009
