description of two new species ofanablepsoides ... · description of two new species...

KeywordsAnablepsoides limoncochae species group, Anablepsoides

urophthalmus species group, Rivulus, Anablepsoides luitali-mae, Anablepsoides vieirai.

AbstractAnablepsoides luitalimae, new species, from the rio

Madeira basin, southwestern Amazonia, Brazil, and An-ablepsoides vieirai, new species, from the rio Itapecurubasin, eastern Amazonia, Brazil, are described. A. luitali-mae belongs to the Anablepsoides limoncochae species groupand A. vieirai belongs to the Anablepsoides urophthalmusspecies group. Anablepsoides luitalimae is distinguishedfrom other species of the Anablepsoides limoncochae speciesgroup by having blue pelvic fins and one longitudinal linewith 6 minute contact organs on the sides of the body. An-ablepsoides vieirai is distinguished from other species of theAnablepsoides urophthalmus species group by the presenceof black spots in the head, opercular region and behind theeye and a higher number of scales in longitudinal series.

ZusammenfassungAnablepsoides luitalimae, neue Art aus dem Rio Madeira

Becken, südwestlichen Amazonien, in Brasilien und Ana -blepsoides vieirai, eine neue Art aus dem Rio ItapecuruBecken, Ost-Amazonien, in Brasilien, wird beschrieben. A.luitalimae gehört zur Anablepsoides-limoncochae-Arten-gruppe und A. vieirai gehört zur Anablepsoides-urophthal-mus-Artengruppe. Anablepsoides luitalimae unterscheidetsich von anderen Spezies der Anablepsoides-limoncochae-Artengruppe, indem sie blaue Beckenflossen und eineLängslinie mit 6 Minuten Kontaktorganen an den Seitendes Körpers haben. Anablepsoides vieirai unterscheidet sichvon anderen Spezies der Anablepsoides-urophthalmus-Artengruppe durch die Anwesenheit von schwarzen Fle -cken im Kopf, im opercularen Bereich und hinter demAuge und einer höheren Anzahl von Schuppen entlang derSeitenline.

ResumoAnablepsoides luitalimae, sp. nov., da área de drenagem da

bacia rio Madeira, sudoeste da Amazônia, Brasil, e An-ablepsoides vieirai, sp. nov., da área de drenagem da baciado rio Itapecuru, leste da Amazônia, Brasil, são descritas.Anablepsoides luitalimae pertence ao grupo de espécies An-ablepsoides limoncochae e Anablepsoides vieirai pertence aogrupo de espécies Anablepsoides urophthalmus. Anablep-soides luitalimae se distingue das demais espécies do grupoAnablepsoides limoncochae por possuir nadadeiras pélvicasazuis e uma linha com 6 diminutos órgãos de contatos nalateral do corpo. Anablepsoides vieirai se distingue das de-mais espécies do grupo Anablepsoides urophthalmus porpossuir machas pretas na cabeça, região opercular e atrásdo olho e um grande número de escamas na linha longitu-dinal.

RésuméAnablepsoides luitalimae, nouvelle espèce, du bassin du

rio Madeira, au sud-ouest de l’Amazonie, Brésil, et Ana -blepsoides vieirai, nouvelle espèce, du bassin du rio Itapecu-ru, à l’est de l’Amazonie, Brésil, sont décrits. A. luitalimaeappartient au groupe d’espèces d’Anablepsoides limoncochaeet A. vieirai, au groupe d’espèces Anablepsoides urophthal-mus. Anablepsoides luitalimae se distingue des autres espè -ces du groupe Anablepsoides limoncochae par des pelviennesbleues et une ligne longitudinale avec de minuscules or-ganes de contact sur les côtés du corps. Anablepsoidesvieirai se distingue des autres espèces du groupe Anablep-soides urophthalmus par la présence de taches noires dans larégion operculaire et derrière les yeux et au nombre plusélevé d’écailles en rangées longitudinales.

SommarioSono qui descritte due nuove specie: Anablepsoides luita -

limae, dal bacino del rio Madeira, sud-ovest Amazzonia, eAnablepsoides vieirai, dal bacino del rio Itapecuru, Amazzo-nia orientale, Brasile. Anablepsoides luitalimae appartiene al

aqua vol. 22 no. 4 - 22 October 2016165

aqua, International Journal of Ichthyology

Description of two new species of Anablepsoides (Cyprinodontiformes:Cynolebiidae) from Rio Madeira, Amazon drainage, Rondônia state and

from Rio Itapecurú basin, Maranhão state, Brazil

Dalton Tavares Bressane Nielsen

Laboratório de Zoologia, departamento de Biologia, Universidade de Taubaté,Pça Marcelino Monteiro 63, CEP: 12030-010, Taubaté, SP, Brazil. E-mail: [email protected]

Received: 11 July 2016 – Accepted 09 October 2016

AQUA-22(4).qxp_AQUA 24/10/16 12:29 Pagina 165

aqua vol. 22 no. 4 - 22 October 2016 166

Description of two n. sp. of Anablepsoides from Rio Madeira, Amazon drainage, Rondônia state and from Rio Parnaiba basin, Maranhão state, Brazil

gruppo di specie Anablepsoides limoncochae, mentre Ana -blepsoides vieirai a quello di Anablepsoides urophthalmus.Anablepsoides luitalimae si distingue dalle altre specie delgruppo per avere pinne pelviche blu e una linea longitudi-nale con 6 minuscoli organi di contatto sui lati del corpo.Anablepsoides vieirai si distingue dalle altre specie del grup-po per la presenza di macchie nere sul capo, sulla regioneopercolare e dietro l'occhio e un più alto numero di scagliein serie longitudinale.

INTRODUCTIONThe genus Anablepsoides is one of the most

species-rich genera in the Cynolebiidae family,with currently 49 valid described species. It has awide geographic distribution across South Americaand the Caribbean. At the southeastern boundaryof the distribution of the genus, Anablepsoides bahi-anus occur in the region surrounding the city ofSalvador, Bahia state, Brazil. At the southwesternboundary occurs A. beniensis, in the region aroundTrinidad City, Departamento Beni, Bolivia. At thenorthwestern boundary occurs A. monticola, at theProvincia Morona-Santiago, Ecuador and finally atthe northeastern boundary, A. cryptocallus on theCaribbean island of Martinique.The genus Anablepsoides was erected by Huber

(1992) as a subgenus of the genus Rivulus. Huber(1992) proposed two groups within this subgenus:the Anablepsoides urophthalmus species group, thatincluded species of central and eastern Amazon,Guianas and eastern Venezuela, and the Anablep-soides limoncochae species group, that includedspecies from central and western Amazon.Costa (2010) redefined both species groups, and

subsequently (Costa, 2011) raised Anablepsoides tothe generic status based on a combined phyloge-netic analysis including both mitochondrial DNAand morphological data. In this analysis the speciesof Rivulus are grouped into distinct paraphyleticlinages, following a pattern that was already knownin molecular analyses (e.g., Hrbek & Larson 1999;Murphy et al. 1999; Vermeulen & Hrbek 2005)The Anablepsoides limoncochae species group has

10 valid species: A. elongatus Fels & de Rham,1981 from the Río Ucayali drainage, Amazonbasin, Peru, A. christinae Huber, 1992 from the RíoMadre de drainage, Peru, A. iridescens Fels & deRham, 1981, from Río Ucayali drainage, Amazonbasin, Peru, A. limoncochae Hoedeman, 1962,from the Río Napo drainage, Amazon basin,Ecuador, A. parlettei Valdesalici & Schindler 2011from río Araza drainage, Amazon Basin, Peru, A.rubrolineatus Fels & de Rham, 1981 from Río

Ucayali drainage, Amazon basin, Peru , A. taeniatusFowler, 1945 from río Caquetá drainage, Colom-bia , A. urubuiensis Costa, 2013 from a creek adja-cent to the rio Urubuí, upper rio Urubu drainage,a tributary of the Rio Amazonas, Brazil, A. linea-soppilatae Valdesalici & Schindler 2013 from RioSan Gaban drainage, Amazon basin, Peru and A.hoetmeri Nielsen, Baptista Jr. & van den Berg,2016 from rio Purus drainage, Amazon basin,Brazil.The Anablepsoides urophthalmus species group has

10 valid species: A. urophthalmus Gunther, 1866from the region of Belém Pará, Amazon basin,Brazil, A. bahianus Huber 1990 from rio Catu,Bahia, Brazil, A. cearensis Costa & Vono, 2009from coastal river basins of Ceará state, Brazil, A.deltaphilus Seegers, 1983 from lower Río Orinoco,Venezuela, A. lungi Berkenkamp, 1984, fromsouthwest of Matoury, French Guiana, A.mazaruni Myers, 1924 from Mutusi hole near ríoMazaruni, Guyana, A. stagnatus Eigenmann,1909from coastal river basins of Guyana and Suriname,A. tocantinensis Costa, 2010 from floodplains ofthe left bank of the rio Tocantins, Brazil, A.xinguensis Costa, 2010 from a swamp close to theleft bank rio Xingu, Brazil, and A. jari Costa et al.,2013 from a stream tributary of rio Jari, lower rioAmazonas basin, Brazil.Although species of both groups being morpho-

logically similar, molecular evidence has indicatedthat species of the A. urophthalmus group are moreclosely related to other species of Anablepsoidesfrom the coastal rivers of northern South Americathan to species of the A. limoncochae group, where-as species of the latter group are more closely relat-ed to other species of Anablepsoides from the Ama-zon than to species of the A. urophthalmus group(Costa 2010).In the present contribution, two additional Ana -

blepsoides species from northern Brazil, the first be-longing to the A. limoncochae species group fromthe rio Madeira basin, Rondônia state, and the sec-ond belonging to the A. urophthalmus group fromthe rio Parnaiba basin, Maranhão state, are de-scribed.

MATERIALS AND METHODSMeasurements were taken point-to-point under a

stereomicroscope with a digital caliper to the near-est 0.1 mm on the left side of the specimen follow-ing Costa (1995). Measurements are expressed aspercents of standard length (SL), except subunits


of the head, which are recorded as percents of headlength (HL). Descriptions of color patterns were based on pho-

tographs of both sides of live individuals of eachpopulation taken in small aquaria one day aftercollection; color patterns derived from distributionof melanophores on fins were also observed in allpreserved specimens.In the description, counts of vertebrae and pleural

ribs were taken from cleared and stained (c&s) of 2males and 3 females paratypes A. luitalimae andone male and one female paratypes of A. vieirai,prepared according to Taylor & Van Dyke (1985).Frontal squamation followed Hoedeman (1958)and Costa (2006). For vertebral counts, the caudalcompounded centrum was counted as a single ele-ment. Osteological features included in the de-scription are those considered phylogenetically in-formative in recent studies of the species formerlyassigned to the genus Rivulus (Costa 2006, 2011).Institutional abbreviations are UNITAU (Univer-

sity of Taubaté, Taubaté, Brazil) and ZUEC(Museu de Zoologia da Universidade Estadual deCampinas, Campinas, Brazil). Comparisons withcongeners were based primarily on the literature(Huber 1992, Fels & De Rham 1981, Costa 2006,2007, 2008, 2010, 2013).

Anablepsoides luitalimae, n. sp. (Figs 1- 2)

Rivulus aff. taeniatus (not Fowler): Hrbek, 2013:294-295 (description, photo of preserved speci-men, middle rio Madeira basin, Brazil).

Holotype: ZUEC 13006, 1 male, 32.3 mm SL:Brazil, Rondônia, Porto Velho, stream tributary torio Madeira, near Porto Velho airport, 8°43’31.40”S 63°54’33.44” W, altitude 84 m; Gilson PontesLima, 23 August 2015. Paratypes: ZUEC 13007, 5 males, 30.4-37.2 mmSL, 8 females, 25.5-39.3 mm SL, 5 c&s, 2 males32.6-37.2 mm SL and 3 females 30.6-38.9 mmSL: all collected with the holotype. ZUEC 9943 3females, 33.2-38.8 mm SL, Porto Velho, streamtributary of rio Madeira, near Girau dam,9º21’21” S 64º40’30” W, Equipe UNIR, 19 July2004.Diagnosis: Distinguished from all other species of

the Anablepsoides limoncochae species group by sev-eral unique color pattern features: pelvic-fin light

blue (vs. orange, yellow or light yellow), caudal-finreticulated, light blue and light yellow at basis anddistal area hyaline, with its upper portion light yel-low (vs. orange), anal fin with reticulated basis,light blue with red brown dots at its basis, distalportion pale yellow (vs. reticulated basis, red dotswith distal portion golden yellow with a black barin A. elongatus and A. christinae, anal-fin withbright blue-green basis with rosy spots between therays, followed by a line of red dots also between therays and distal portion bright orange without blackborder in A. hoetmeri, blue basis with distal portionlight yellow with black bar in A. rubrolineatus, analfin pale yellowish, its basis whitish with red spotson posterior portion in A. parlletei, anal fin palegrey, with cream yellow basis and red spots in A.lineasoppilatae), dorsal fin origin posterior to thelast anal-fin ray (vs. at the level of the 13th ray in A.hoetmeri, A. rubrolineatus and A. elongatus), onelongitudinal line with 6 minute contact organs, be-ginning at the level of the trunk reached by tip ofthe pectoral fin and ending at the level of the analfin (vs. present at pectoral fin or 1-3 contact organson antero-ventral portions of flank or on each scaleof midventral portion of flank or two longitudinalrows of 18-20 contact organs in A. hoetmeri), dor-sal origin neural spines (26 vs. 21-23 in A.urubuiensis or 16-17 in A. hoetmeri), vomer teethnumber (4-6 vs. 1 in A. urubuiensis and A. hoetmerior 2-3 in A. parlettei and A. lineasoppilatae).Description: Measurement data presented in

Table I. Largest examined male 37.2 mm SL,largest examined female 38.9 mm SL. Dorsal andventral profiles slightly convex from snout to pos-terior end of dorsal and anal fins, nearly straight oncaudal peduncle. Body slender, subcylindrical,slightly depressed on head, compressed posteriorly.Greatest body depth at midlength between pec-toral-fin base and anal-fin base origin. Snout shortand rounded. Dorsal fin very small, rounded, with-out filaments in both sexes. Caudal-fin rounded.Pectoral-fin rounded, tip of pectoral fin reachingvertical at about 66% of the length between pec-toral fin and pelvic-fin base. Tip of pelvic-fin notreaching vertical through anus, pointed, base with-out interspace. Caudal-fin rounded. Dorsal-fin ori-gin at level of last anal-fin ray and neural spine of26th vertebrae. Anal fin origin between neuralspines of 19th-20th vertebrae. Dorsal-fin rays 7;anal fin rays 12; caudal-fin rays 25-26; pectoral-finrays 12; pelvic-fin rays 7. Scales large, cycloid.Body and head entirely scaled, except on jaws.






Fig. 2. Anablepsoides luitalimae, ZUEC 13007, paratype, female, 35.3 mm SL. Photo by D. Nielsen.

Fig. 1. Anablepsoides luitalimae, ZUEC 13006, holotype, male, 32.3 mm SL. Photo by D. Nielsen.


Scales on dorsal fin and anal-fin basis absent.Frontal squamation E-patterned. Longitudinal series of scales 43-44; transverse se-

ries of scales 10; scale rows around caudal-fin pe-duncle 16. One longitudinal line with 6 minutecontact organs, beginning at the level of the trunkreached by tip of the pectoral fin and ending at thelevel of the anal fin, absent in pectoral fin.Cephalic neuromasts: supraorbital 3 + 3, parietal

1,anterior rostral 1, posterior rostral 1, infraorbital1+14-15+1, preorbital 2, otic 1, post-otic 2,supratemporal 1, medianopercular 1, ventral oper-cular 1, preopercular 2 + 4, mandibular 4 + 1, lat-eral mandibular 2, paramandibular1. Two neuro-masts on caudal-fin base. Vomerine teeth 4-6. Sixbranchiostegal rays. Gill rakers on first and secondbrachial arch 8-9. Total vertebrae 34-35.C o l o r a t i o n i n l i f e : Males. Sides of body

metallic blue with six continuous, well defined lon-

gitudinal lines formed by red dots,. Six longitudi-nal lines present, five of which reaching caudal pe-duncle, one ending near anal-fin basis. Size of thered spots variable; red spots in the first (dor-salmost) longitudinal row larger in size than thosein the fifth and sixth (ventralmost) lines. Bluemetallic scales found between the longitudinal redlines. Dorsal region brown, from snout to dorsal-fin basis; irregular smoky pigmentation betweendorsal region and flanks. Abdominal region whitepinkish. Sides of head brown with dark spots, op-ercular region dark grey. Iris pale yellow. Dorsal-finhyaline , oblique transverse stripe on the middle ofthe dorsal fin, distal part light yellow. Anal-fin basepale blue with small red points, distal portion paleyellow, black border absent. Caudal fin with retic-ulated pattern, pale blue with yellowish pigmenta-tion, dorsal portion metallic gold. Pectoral fin lightorange. Pelvic fin light blue.



Fig. 3. Type locality of Anablepsoides luitalimae, flooded area at an urban forest fragment, middle rio Madeira basin, Rondônia, Brazil. Photo by G. Pontes Lima.


Females. Flanks dark, with four longitudinal linesformed by separated bright red dots, with inter-spersed dark and yellow blotches between the redlongitudinal lines from immediately behind headto region above anal fin. Interspersed large yellowblotches, from the anal fin region to caudal pedun-cle. Dorsal portion of body light brown. Top ofhead with a large metallic yellow scale. Sides ofhead brown, opercular region with smoky darkgrey pigmentation. Iris yellow. Dorsal-fin base witha white, transversal black line, distal portion hya-line. Caudal-fin hyaline, with reticulated dark pig-mentation. Large rounded black spot, preceded bysmaller whitish spot, on dorsal portion of caudalfin base. Anal fin with basis light blue, distal por-

tion hyaline, with interspersed small reddish spots.Pectoral-fin hyaline. Pelvic-fin hyaline with metal-lic blue tinge.Distribution: Known from flooded areas within

forest fragments along the right bank of the RioMadeira near Porto Velho, Rondônia state, Brazil(Fig. 7) .Habitat: The type locality is a flooded area with a

maximum depth of 50 cm lying at an urban frag-ment of very dense Amazonian rainforest,about 1km distance from the Rio Madeira. The water wasdark-colored, with pH 6.0 and low conductivity,with a dense leaf litter, and completely devoid ofaquatic vegetation (Fig. 3). Anablepsoides luitalimaewas the only fish species recorded at the site. Etymology: In honor of the discoverer’s aunt,

Luita Lima.

Anablepsoides vieirai, n. sp. (Figs 4-5)

Holotype: ZUEC 13008, 1 male, 36.3 mm SL: Brazil, Maranhão, Duque Bacelar, spring Riacho Joana, rio Parnaíba basin. 4°07’35.46” S42°58’48.76” W, altitude 61 m. Gilberto da SilvaVieira e Maria da Conceição Pereira da Silva, 01May 2015. Paratypes: ZUEC 10787, 3 males, 23.5-32.5 mmSL, 5 females, 20.9-36.9 mm SL, two c&s, onemale 32.5 mm SL and one female 31.5 mm SL: allcollected with holotype.Diagnosis: Distinguished from all other species of

the Anablepsoides urophthalmus species group bythe presence of black spots in the head and opercu-lar regions, behind the eye (vs. black spots absent),caudal fin hyaline with four centralmost rays withpale gray spots (vs. pale grays spots absent), lowercaudal peduncle depth (11.4-11.9 of SL vs. 12.5-15.9 of SL), greater number of scales in longitudi-nal series (44 vs. 42 in A. mazaruni), tip of pectoralfin reaching vertical at about 72% of length be-tween pectoral fin and pelvic-fin base (vs. 70% inA. bahianus and A. xinguensis), caudal fin withsmall distal filaments in (vs. distal filaments ab-sent), dorsal fin origin inserting between neuralspines of vertebrae 27-28 (vs. 31-32 A. xinguensis,32-35 A. urophthalmus), higher number of longitu-dinal series of scales, 43-44 (vs. 38-40 A. jari and42 in A. mazaruni), higher number of scale rowsaround caudal-fin peduncle (18 vs. 16), yellowstripe on the dorsal and ventral margins of the cau-



Table I. Morphometric and meristic data for the holotype(H) and paratypes of Anablepsoides luitalimae n. sp. SD = Standard Deviation.

H ParatypesMale Males n=5 Females n=8

Standard length (mm) 32.3 30.4-37.2 25.2-39.3

Percents of SL

Body depth 20.1 17.7-19.3 15.3-19.2

Caudal peduncle depth 13.6 11.3-12.2 11.1-12.4

Pre-dorsal length 77.7 74.1-77.7 71.2-78.7

Pre-pelvic length 51.7 50.0-52.4 48.3-54.6

Length of dorsal-fin base 6.5 6.7-7.6 6.2-7.2

Length of anal-fin base 15.4 15.6-17.9 13.3-15.8

Caudal-fin length 24.7 22.2-26.7 21.2-25.9

Pectoral-fin length 15.4 15.8-20.5 15.6-20.4

Pelvic-fin length 7.4 6.5-8.2 5.8-7.0

Head length 24.5 21.5-24.5 21.8-25.8

Percents of HL

Head depth 64.1 63.5-68.2 50.9-55.5

Head width 44.4 44.2-52.9 50.6-54.8

Snout length 14.5 14.5-17.0 14.9-16.5

Lower jaw length 17.3 14.3-17.8 13.7-15.3

Eye diameter 26.6 25.6-31.5 25.5-29.1

Counts

Dorsal fin 7 7 7

Caudal fin 25 25-26 25-26

Anal fin 12 12 12

Pelvic fin 7 7 7

Pectoral fin 12 12 12


Fig. 4. Anablepsoides vieirai, ZUEC 13008, holotype, male, 36.3 mm SL. Photo by D. Nielsen.



Fig. 5. Anablepsoides vieirai, ZUEC 10787, paratype, female, 36.9 mm SL. Photo by D. Nielsen.


dal fin in males (vs. inconspicuous in A. xinguensisand A. tocantinensis, yellowish white stripes on thedorsal and ventral margins or well-defined yellowpigmentation in other species). Distinguished from all members of the A. uroph-

thalmus specie group, except A. tocantinensis, bylower number of pelvic fin rays (5 vs. 6-7).Description: Measurement data presented in

Table II. Largest examined male 36.3 mm SL,largest examined female 36.9 mm SL. Dorsal andventral profiles slightly convex from snout to pos-terior end of dorsal and anal fins, nearly straightlyon caudal peduncle. Body slender, subcylindrical,slightly depressed on head, compressed posteriorly.

Greatest body depth at mid length between pec-toral-fin base and anal-fin base origin. Snout blunt.Dorsal-fin small, rounded without filaments inboth sexes. Caudal-fin rounded. Pectoral-finrounded, tip of pectoral fin reaching vertical atabout 72% of length between pectoral fin andpelvic-fin base. Tip of pelvic fin not reaching verti-cal through anus, small, pointed, base without in-terspace. Caudal-fin rounded with very small distalfilaments. Dorsal fin origin at level of base of the11th anal-fin ray and inserting between neuralspines of 27-28 vertebrae. Anal fin origin betweenneural spines of 17-18 vertebrae. Dorsal-fin rays 8;anal-fin rays 11-13; caudal-fin rays 29-30; pec-toral-fin rays 13; pelvic fin rays 5. Scales large, cy-cloid. Body and head entirely scaled, except onjaws. No scales on dorsal fin and anal-fin basis.Body squamation extending over anterior 30 % ofcaudal-fin base. Frontal squamation E-patterned. Longitudinal series of scales 43-44; transverse se-

ries of scales 10; scale rows around caudal-fin pe-duncle 18. Contact organs absent. Urogenitalpapilla small.Cephalic neuromasts: supraorbital 3 + 3, parietal

1, anterior rostral 1, posterior rostral 1, infraorbital1+13-15+1, preorbital 2, otic 1, post-otic 2,supratemporal 1, median opercular 1, ventral oper-cular 2, preopercular 2 + 4,mandibular 4 + 1, lat-eral mandibular 3, paramandibular 1. Two neuro-masts on caudal-fin base. Vomerine teeth absent.Four branchiostegal rays. Gill rakers on firstbrachial arch 8.Total vertebrae 33-34.C o l o r a t i o n i n l i f e : Males. Sides of the up-

per portion of the body metallic green, lower por-tion metallic purplish-blue with 6-7 longitudinalrows of conspicuous, continuous red dots, some-times interrupted and replaced by rows of blue orgreen metallic scales, the first (uppermost) line be-ginning immediately behind the head and finish-ing at the dorsal-fin level, the second to sixth lon-gitudinal lines beginning immediately behind thehead and finishing at caudal peducle, the seventh(lowermost) line, which is composed of small redspots, beginning behind pectoral-fin basis and fin-ishishing at the basis of the caudal fin. Dorsumbrown, with dark area between dorsum and sidesof body. Abdominal region whitish. Black spotspresent on the side of the head, opercular regionand below the eye. Iris pale yellow. Dorsal-fin lightyellow with small black dots. Anal-fin basis lightblue, distal portion light yellow. Caudal-fin hya-line, with four centralmost rays with pale gray



Table II. Morphometric and meristic data for the holotype(H) and paratypes of Anablepsoides vieirai n. sp. SD = Stan-dard Deviation.

H ParatypesMale Males n=3 Females n=5

Standard length (mm) 36.3 23.5-32.5 20.8-36.9

Percents of SL

Body depth 20.1 20.0-20.4 18.2-22.4

Caudal peduncle depth 11.8 11.4-11.9 11.9-12.2

Pre-dorsal length 74.4 77.2-78.2 78.1-80.7

Pre-pelvic length 50.0 49.3-54.4 54.1-57.2

Length of dorsal-fin base 9.2 6.3-10.7 8.6-10.1

Length of anal-fin base 20.6 20.4-24.6 18.4-19.2

Caudal-fin length 32.3 31.5-32.6 31.9-32.9

Pectoral-fin length 18.1 15.6-20.4 14.6-16.1

Pelvic-fin length 8.5 7.1-9.3 8.5-9.6

Head length 22.9 22.4-25.9 27.0-28.8

Percents of HL

Head depth 79.6 75.6-79.4 75.0-79.7

Head width 77.2 77.1-78.2 77.3-78.4

Snout length 13.9 13.4-15.8 13.8-15.9

Lower jaw length 19.7 19.7-24.6 19.7-21.8

Eye diameter 34.2 34.4-3 33.3-34.1

Counts

Dorsal fin 8 8 8

Caudal fin 29 29-30 29

Anal fin 13 13 11

Pelvic fin 5 5 5

Pectoral fin 13 13 13


Fig. 6. Type locality of Anablepsoides vieirai, spring running into the Riacho Joana, rio Parnaíba basin, Maranhão, Brazil. Photo by G. Vieira.

spots. Pectoral-fin light orange. Pelvic-fin yellowishat basis, bluish at distal portion. Females. Flanks light brown, without longitudi-

nal lines, with alternating light brown and darkbrown reticulated pattern. Sides of head brown.Opercular region with black spots. Iris yellow.Dorsal-fin base hyaline, with black spots. Caudal-fin hyaline with reticulated pattern formed byblack spots. A rounded, conspicuous black spot,preceded by yellow spot, on dorsal portion of cau-dal-fin base. Anal fin hyaline, basis with reticulatedpattern of black spots. Pectoral and pelvic-fins hya-line. Distribution: Anablepsoides vieirai is only known

from its type locality, near Duque Bacelar, rio Parnaíba basin, Maranhão state, northeasternBrazil. Habitat: The type-locality of Anablepsoides vieirai

is a spring that forms a Buritizal (a wetland domi-nated by the buriti palm, Mauritia flexuosa). Thespring forms a small stream running through sug-arcane field and is then dammed forming a smalllagoon of crystalline water. The pH varies from be-tween 5.0 and 6.5 at the source of the pond, tofrom 6.0 to 7.5 at the pond. Anablepsoides vieirai isfound in the flooded area below the Buritizal, in a

stream that flows directly into the Riacho Joana(Fig. 6).Etymology: In honor of the biologist and envi-

ronmentalist Gilberto da Silva Vieira, who discov-ered the species.

DISCUSSION The genus Anablepsoides as redefined by Costa

(2011) is a genus which comprises medium sizedspecies, ranging between about 30-60 mm SL,with short and rounded fins, except the caudal fin,that may be subtruncated, or elliptical and long ina few species. Longitudinal rows of red dots on theflanks are present in males, sometimes also in fe-males. Anablepsoides differs from all other Rivuli-nae genera by the possession of scales on the wholeventral surface of the head (vs. chin naked) (Costa2011).The two species groups recognized within An-

ablepsoides, Anablepsoides limoncochae species groupand Anablepsoides urophthalmus species group, havetheir geographical distributions well defined andseparated from each other (Fig. 7). The Anablep-soides limoncochae species group is distributedalong the piedmont of the Andes cordillera and ad-jacent foreland basins in western Amazonia (with




the exception of A. urubuiensis, which occurs atCentral Amazon) (Nielsen et al. 2016). Most of theknown species of the group (8 species) occur closeto the Andean cordillera, with A. lineasoppilataeand A. parlettei occurring at the Andean cordilleraitself.Anablepsoides luitalimae belongs to the Anablep-

soides limoncochae species group by sharing all thecharacteristics that define this group: males pre-senting contact organs on scales of the flanks, pres-ence of an oblique transverse stripe on the middleportion of the dorsal fin, and by the absence oflight yellow to orange areas on dorsal and ventralmargins of the caudal fin (Costa 2013). Anablep-soides luitalimae is recorded for the middle rioMadeira basin in Brazil, being the first species ofthe Anablepsoides limoncochae species group record-ed for this river basin. Although not as numerous as for A. hoetmeri (a

single longitudinal row with 6 minute contact or-

gans vs. two longitudinal rows of 18-20 minutecontact organs), the presence of contact organsalong the flanks of males of A. luitalimae indicatesthat its reproductive behavior is similar to that ofA. hoetmeri. Probably the reproductive behavior in-volves the female being positioned laterally to themale’s body, similar to what is observed in thegenus Fundulopanchax (Nothobranchiidae) (Belote2000).The Anablepsoides urophthalmus species group is

distributed across the northeastern portion ofSouth America: Venezuela, Guyana, FrenchGuyana , and in Brazil, at Amapá, Pará, Tocantins,Ceará and Bahia states (Fig. 7). Its distribution ex-tends southwards to the proximity of Salvador city,Bahia state, in Brazil (Costa 2010).Anablepsoides vieirai belongs to the Anablepsoides

urophthalmus species group by sharing the follow-ing combination of fetatures: longitudinal rows ofred or orangish brown spots on flanks in males, ab-



Fig. 7. Geographical distribution of the species of the Anablepsoides limoncochae species group and from the A. urophthalmusspecies group.


sence of contact organs on flanks, dark bluish orpurplish gray pigmentation over the side of thehead in males, presence of light yellow to orangepigmentation on the dorsal and ventral margins ofthe caudal fin, absence of an oblique transversestripe on the middle portion of the dorsal fin, analfin with 12-15 rays, and ventral surface of the headentirely scaled (vs. anterior portion of head entirelyscaled) (Costa 2008, 2010).Anablepsoides vieirai is found in the Rio Parnaíba

basin. This river is located in the transitional areabetween the Amazon biome and the Caatinga bio-me. The site of occurrence of Anablepsoides vieirailies in a straight line about 680 km from the areaof occurrence of A. urophthalmus and about 490km from the area of occurrence of A. cearensis, al-most exactly halfway between these geographicaldistributions. Anablepsoides vieirai appears to be closely related

to A. urophthalmus for sharing the following char-acteristics: body squamation extending over the an-terior 30 % of caudal fin base, color pattern ofmales with red dots forming lines almost uninter-rupted, and the absence of a broad dark red stripeon the dorsal and ventral margins of the caudal finin males. Huber (1992) considered Anablepsoidesbahianus a link between the Rivulus santensis group(currently the genus Atlantirivulus) with the R.urophthalmus group (currently the genus Anablep-soides). Costa (2008) redescribed Anablepsoidesbahianus and noticed that color patterns are similaracross the species belonging to the A. urophthalmusspecies group. Costa (2008) concluded that A.bahianus rather belongs to that geographically dis-tant species assemblage formerly assigned to thegenus Rivulus, which occurs at the eastern portionof the Amazon basin and adjacent parts of north-ern South America and Caribbean islands, insteadof being related to other Rivulus species endemic tothe Atlantic forest, currently assigned to the genusAtlantirivulus.The site of occurrence of Anablepsoides vieirai,

along with the site of occurrence of A. cearensis,demonstrates that there is continuity in the geo-graphical range of the A. urophthalmus species groupacross northeastern Brazil and decreases the geo-graphical gap between A. bahianus and the remain-ing species of the group. It is likely that the occur-rence of the genus Anablepsoides is more extensiveand includes many of smaller river basins along thecoastal Atlantic region of northeastern Brazil.

ACKNOWLEDGEMENTSThanks are due to Itamar Alves Martins from

Universidade de Taubaté (UNITAU) and RicardoBritzke (formerly at the Universidade EstadualPaulista, Botucatu) for laboratory support and toLuciano Araujo Medeiros for preparing the map.Dan Katz and Flávio C.T. Lima (ZUEC) reviewedthe manuscript and offered useful suggestions.

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description of two new species ofanablepsoides ... · description of two new species...

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