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The Australoheros (Teleostei: Cichlidae) species of the Uruguay and Paraná River drainages

OLDŘICH ŘÍČAN1,2 & SVEN O. KULLANDER3

1 Department of Zoology, Faculty of Sciences, University of South Bohemia, Branišovská 31, CZ-37005 České Budějovice, Czech Republic2 Institute of Animal Physiology and Genetics of the Academy of Sciences of the Czech Republic, Rumburská 89, CZ-277 21 Liběchov, Czech Republic3 Department of Vertebrate Zoology, Swedish Museum of Natural History, P.O. BOX 50007, SE-104 05 Stockholm, SwedenCorresponding author. 1E-mail: [email protected]

Table of contents

Abstract ............................................................................................................................................................................... 1Introduction ......................................................................................................................................................................... 2Methods ............................................................................................................................................................................... 3Results ................................................................................................................................................................................. 3

Australoheros facetus (Jenyns, 1842) .......................................................................................................................... 4Australoheros scitulus (Říčan & Kullander, 2003) .................................................................................................... 12Australoheros tembe (Casciotta, Gómez & Toresanni, 1995) .................................................................................... 13Australoheros forquilha, sp. nov. ............................................................................................................................... 14Australoheros charrua, sp. nov. ................................................................................................................................ 23Australoheros kaaygua Casciotta, Almirón & Gómez, 2006 ..................................................................................... 26Australoheros minuano, sp. nov. ............................................................................................................................... 32Australoheros guarani, sp. nov. ................................................................................................................................. 36

Systematics and character diversity .................................................................................................................................. 39Discussion ......................................................................................................................................................................... 47Acknowledgements ........................................................................................................................................................... 49References ......................................................................................................................................................................... 49

Abstract

Australoheros Říčan and Kullander, 2006 includes four described species—Australoheros facetus (Jenyns, 1842), Aus-traloheros tembe (Casciotta, Gómez & Toresani, 1995), Australoheros scitulus (Říčan & Kullander, 2003) and Australo-heros kaaygua Casciotta, Almirón & Gómez, 2006. Four additional species are newly described in this paper based onresults presented in Říčan and Kullander (2006): Australoheros forquilha sp. nov., from the tributaries of the Upper RíoUruguay in Brazil; Australoheros charrua sp. nov. and Australoheros minuano sp. nov., from the tributaries of the Mid-dle Río Uruguay in Brazil and Uruguay, and Australoheros guarani sp. nov. from the Río Paraná tributaries in Paraguay.Sympatric species are not found closely related and all sister species have allopatric distributions. Four species groupsare recognized based on phylogenetic hypotheses generated from morphological and cyt b characters; the forquilhagroup—A. forquilha, A. tembe; the scitulus group—A. scitulus, A. charrua; the kaaygua group—A. kaaygua, A. minu-ano; and the facetus group—A. facetus, A. guarani.

Key words: ´Cichlasoma´ facetum group, Australoheros, new species, Cichlidae, South America, biogeography, RíoUruguay, Río Paraná

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ŘÍČAN & KULLANDER2 · Zootaxa 1724 © 2008 Magnolia Press

Introduction

Australoheros is the southernmost genus of cichlasomatine cichlids (tribe Heroini), distributed in the La Platabasin in the tributaries of the Río Paraná—Río Paraguay system and in the Río Uruguay system. Its distribu-tion extends in the west to the foothills of the Andes in Argentina and to the east in the Atlantic coastal drain-ages of Argentina (Buenos Aires province), Uruguay and Brazil up to the state Bahia, and also includes theSão Francisco drainage. Australoheros is thus one of the cichlid groups with the southernmost distributions onthe South American continent and the widest distribution compared to its closest relatives, the heroinecichlids.

Říčan and Kullander (2003, 2006) have shown that Australoheros is part of the tribe Heroini of the Neo-tropical cichlid subfamily Cichlasomatinae. Despite being the southernmost South American heroine genusAustraloheros is more closely related to Mesoamerican and Antillean heroines, than to most South Americanheroines (except Heroina and Caquetaia; Říčan, 2005; Říčan & Kullander, 2006; Concheiro Pérez et al.,2007).

Four Australoheros species are considered valid at present: Australoheros facetus (Jenyns 1842), A. tembe(Casciotta, Gómez & Toresani, 1995), A. scitulus (Říčan & Kullander, 2003) and A. kaaygua Casciotta, Alm-irón & Gómez, 2006. Several other nominal taxa are included in Australoheros, all considered junior syn-onyms of Australoheros facetus (Jenyns, 1842). These nominal taxa are: Chromys oblonga Castelnau, 1855,from the Rio Tocantins in Goiás, Brazil; Heros autochton Günther, 1862 from Brazil without any preciselocality; H. jenynsii Steindachner, 1869 from Montevideo, Uruguay; and H. acaroides Hensel, 1870 fromPorto Alegre, Brazil. In the present paper we have examined material of Australoheros from the drainages ofthe Middle Río Paraná and Río Uruguay and adjacent areas in Uruguay, Argentina, Paraguay and Brazil (RioGrande do Sul, Santa Catarina); for this area Říčan and Kullander (2006) have demonstrated that Australo-heros includes several undescribed species. In this paper we formally describe four new species and provide areview of the species diversity of the genus Australoheros in the drainages of the Uruguay and Paraná riverdrainages.

The Río Uruguay (2,262 km long) is one of the three rivers that form the Plate watershed, together withthe Paraná and Paraguay. Between the mouth of the Uruguay and the Atlantic Ocean lies an area of approxi-

mately 18,000 km2 that includes the Río de la Plata, a saline estuary with a depth of 4 to 18 m. The Río Uru-guay rises in the Serra Geral Mountains as the Rio Pelotas, near the southern coast of Brazil, at an altitude ofapproximately 1,800 m, bearing the name Río Uruguay down from the confluence of the Pelotas with theCanoas 1,816 km from the mouth. It runs inland along the border between Santa Catarina and Rio Grande doSul states for 938 km to the mouth of the Rio Pepirí. From the Rio Pepirí Guazú confluence the Uruguay flowssouth for 1,324 km, marking the borders between Brazil and Argentina, and Uruguay and Argentina, until itmeets the Río Paraná to form the estuary of the Río de la Plata, which flows into the Atlantic Ocean.

The river is a series of pools and rapids which divide it into natural sections and the river was thus evenbefore daming less navigable than other rivers of the La Plata basin. The Salto Grande Falls (353 km from themouth) dropped 9 m in 3 km, and are considered the border between the Middle and Lower Uruguay. In 1979these rapids were flooded after the construction of the Salto Grande Dam. The Yucumã (or Moconá) Falls,below the mouth of the Rio Pepirí Guazú (1,324 km from the mouth, just where the Río Uruguay leaves Braziland forms the border with Argentina), have a drop of 12 m, forming rapids approximately 1,800 m long.These rapids form the border between the Middle and Upper Río Uruguay (Zaniboni Filho & Schulz, 2003).

The Río Paraná (4,695 km long) originates at the confluence of the Rios Grande and Paranaíba in southernBrazil, and then runs generally southwest for 3,998 km before draining into the Río de la Plata estuary. East-ern tributaries in the upper part, such as the Rios Tietê, Paranapanema, and Iguaçu, originate similarly to theupper tributaries of the Río Uruguay in the coastal mountains a short distance from the Atlantic. The RíoParaná can be divided into upper, high, middle, and lower sections, each with distinctive geographic and bio-logical characteristics (Resende, 2003).


Zootaxa 1724 © 2008 Magnolia Press · 3AUSTRALOHEROS OF THE URUGUAY AND PARANÁ RIVERS

The Upper Paraná has been separated from the rest of the basin by the falls of Sete Quedas and is recog-nized as a distinct ichthyofaunal “province” (Bonetto, 1998). The whole Upper Paraná is heavily dammed,and also the falls of Sete Quedas are now within a dam, the large Itaipú Reservoir. The Upper Paraná is com-pletely within Brazilian territory except for the southern part including the Itaipú, where it forms the borderwith Paraguay. The High Paraná traces in a SW loop the contour of the Sierra de Misiones and forms the bor-der between Paraguay and the Argentinian province of Misiones. All tributaries of the High Paraná in this sec-tion are characterized by falls close to their confluence with the Paraná limiting fish passage upriver frommainstream. At the confluence with the Río Paraguay it becomes the Middle Paraná, turns again to the southand runs through Argentina. In this stretch it is a typical plains river, banked by its own alluvial deposits andhaving an extensive floodplain. At Santa Fé, the Paraná receives the last tributary, Río the Salado, andbecomes the Lower Paraná (Bonetto, 1986; Resende, 2003).

Methods

Most characters are based on alcohol-preserved specimens with notes on live coloration where available.Measurements and counts were taken as described by Kullander (1986). Measurements were taken with digi-tal calipers to 0.1 mm and are made point to point. Scale rows are numbered as described by Kullander (1990),i.e., the horizontal row including the lower lateral line is designated as row E0, and the rows are counted asE1, E2 etc. dorsally, and H1, H2 etc. ventrally. Scale counts between anterior insertion of the dorsal fin and theupper lateral line (L1) and between the posterior end of the upper lateral line and the dorsal fin are designatedas L1 and L2. Dorsal and anal fin rays and vertebrae were counted on radiographs. Vertebral counts includethe last halfcentrum. Color-marking terminology follows Kullander (1983, 1986) and Říčan and Kullander(2006). The number of specimens for each given count is indicated in parentheses. Drawings were made usinga drawing tube fitted to an Olympus SZX9 microscope.

Institutional abbreviations are as listed in Leviton et al. (1985) and Leviton and Gibbs (1988).Character data sets and methods of phylogenetic inference were used as described in Říčan and Kullander

(2006). Character states have been mapped onto cladograms using Mesquite (Maddison & Maddison, 2004).

Results

Key to the Australoheros species from the Rio Uruguay drainage and adjacent areas

We have chosen to use only readily accessible external characters, even though some characters visible only on x-rays, asthe numbers of caudal vertebrae or the number of vertebrae included in the caudal peduncle, are good diagnostic charac-ters. Some of the characters used in the key are color and coloration pattern characters, which will work only for adultspecimens (at least 50 mm SL). For this reason we have based the key mostly around meristic characters which are morestable in respect to size/age.

1a. 8–9 anal fin spines, 17 dorsal fin spines, dark blotches on operculum, suboperculum, posterior part ofhead, a row of spots in the anterior part of the E4 scale row (above the anterior part of the upper lateralline), spotted unpaired fins, three abdominal bars, 6 ceratobranchial gill rakers, body rounded, groundcolor yellowish ................................................................................. A. scitulus (Říčan & Kullander, 2003)

1b. 6–7 anal fin spines, 16 dorsal fin spines ..................................................................................................... 22a. 6 ceratobranchial gill rakers, mouth isognathous, midlateral blotch not well circumscribed and hidden in a

midlateral stripe (but in smaller individuals very distinct), well developed caudal spot, four abdominalbars, all bars very distinct, modally 12 pectoral fin rays, ground color yellowish, breeding males pinkish



or red ............................................................................................................................ A. minuano sp. nov.2b.(1c) . 6 ceratobranchial gill rakers, mouth small and isognathous, dominant large midlateral blotch, midlat-

eral stripe continuous and more intensive between the head and the midlateral blotch than posterior fromit, caudal spot absent or very inconspicuous, only three wide faint abdominal bars, very short caudalpeduncle ...................................................................................................................................................... 3

2c. 7–8 ceratobranchial gill rakers, mouth large .............................................................................................. 43a.(2b)Mouth positioned low on head, anterior upper margin of premaxilla well below the horizontal from the

ventral margin of the eye, unpaired fins and flank scales without spots, modally 13 pectoral fin rays ............................................................................................................................................... A. charrua sp. nov.

3b. Mouth situated higher on the head, anterior of premaxilla aligned with the horizontal from the ventralmargin of the eye, unpaired fins and flank scales spotted, modally 12 pectoral fin rays, rounded bodyshape, ground color yellow to orange, red corners of caudal fin ............................................................................................................................................................. A. kaaygua Casciotta, Almirón & Gómez, 2006

4a.(2c)25–26 E0 scales, 6 anal fin spines, long caudal peduncle covered with 4 to 5 scale rows, dorsal fin scale cover covering last 10 or more spines, no distinct midlateral stripe .......................................................... 5

4b. 24 E0 scales, 6 or 7 anal fin spines, short caudal peduncle covered with 3 scale rows, distinct caudal spot,fin scale cover covering 7–8 spines or fewer .............................................................................................. 6

5a.(4a). Dark markings forming a line below the orbit along the postero-lateral border of the suborbital series, unpaired fins with a checkerboard spotting pattern, downturned mouth with the upper jaw projecting infront of the lower jaw, membranes of the soft portions of the dorsal and anal fins heavily scaled ..................................................................................................................................................... A. forquilha sp. nov.

5b. Without spots on unpaired fins, no markings on head, some specimens with very thickened lips and termi-nal mouth, membranes of the soft portions dorsal and anal fins with very few scales, wild specimens withdistinctly green body color ....................................................................... A. tembe (Casciotta et al., 1995)

6a.(4b) Adult specimens with only three abdominal bars, only 2 ½ scales between anterior end of dorsal finand upper lateral line, preorbital distance about 25% HL, jaws subequal, mouth slightly inclined ventrally ........................................................................................................................................ A. guarani sp. nov.

6b. Adult specimens with four (sporadically three) abdominal bars, the shortest dorsal fin scale cover, cover-ing only 2–3 last spines (vs. 7–8 last spines), 3 ½ scales between anterior end of dorsal fin and upper lat-eral line, preorbital distance modally 20% HL or less, lower jaw projecting in front of upper jaw (i.e.mouth slightly upturned) ............................................................................................................................. 7

7a. Modally 6 anal fin spines, about 80% of specimens with four abdominal bars, but 80–90% of specimenswith the abdominal bars incompletely separated in their dorsal portion, caudal peduncle length modallybelow 40% of depth, ground color grey ............................................................... A. facetus (Jenyns 1842)

7b. Modally 7 anal fin spines, always four completely separated abdominal bars, caudal peduncle length mod-ally above 48% of depth, ground color yellowish to yellow ................................................... A. cf. facetus

Australoheros facetus (Jenyns, 1842)

Chromis facetus Jenyns, 1842: 104. Type locality: Maldonado, Rio la Plata, Uruguay. Types in Cambridge UniversityMuseum (Darwin no. 660).

Chromys oblonga Castelnau, 1855: 14. Type locality: Rio Tocantins, Goiás, Brazil. Holotype: MNHN A-9485.Heros autochthon Günther, 1862: 299. Type locality: Brazil. Syntypes: (4) BMNH uncat. (1).Heros jenynsii Steindachner, 1869: 149. Type locality: Montevideo, Uruguay. Syntypes: NMW 17324-27 (4), 58722 (1).Heros acaroides Hensel, 1870: 54. Type locality: “Bei Porto Alegre in stagnirenden Gewässern, Brazil”. Syntypes: ZMB

7455(2).



Material examined. NRM 33035 (2, 87.7–106.7 mm SL) Argentina, Buenos Aires, Moreno/Merlo, Río de laReconquista, 100 m downstream from Represa La Reja (3km S of the village La Reja). NRM 33050 (1, 40.6mm SL) Argentina, Entre Ríos, Gualeguaychú, Arroyo Ñancay, where crossing RN14. NRM 37036 (1, CS),NRM 37037 (1, CS), NRM 37038 (1, CS), NRM 37039 (1, CS), NRM 37040 (1, CS) Uruguay, Canelones,Laguna del Tronco at Salinas, 37 km from Montevideo on Ruta Nacional 1 to Atlántida, 500 m off roadtoward Pando. NRM 36495, (2, 45.2–63.5 mm SL), NRM 36774 (2, 53.2–56.0 mm SL), and NRM 36848 (3,30.6–35.8 mm SL) Uruguay, Treinta y Tres, Arroyo Local at margin of Valentines. NRM 37035 (1, CS), NRM39527 (13, 42.9–72.0 mm SL), NRM 37037 (1, CS) Uruguay, Treinta y Tres, Laguna Merín drainage, ArroyoLocal at Valentines. NRM 39546 (2, CS) Uruguay, Treinta y Tres, Laguna Merín drainage, Arroyo Averías atLas Pavas, 30 km from Valentines and 8 km from Ruta Nacional 19. NRM 39552 (1, 103.5 mm SL) Uruguay,Canelones, Laguna Arenera de Carrasco at San José de Carrasco, 18 km on road Montevideo-Atlántida, 200m off Ruta Nacional 1. NRM 48078 (39, 30.1–50.5 mm SL), NRM 47999 (20, 31.3–74.5 mm SL), and NRM48074 (16, 41.4–114.2 mm SL) Uruguay, Canelones, El Pinar.

Character diversity in Australoheros facetus. The type specimens of A. facetus are from Uruguay, Río dela Plata drainage. The description given by Jenyns (1842) agrees well with populations form both the Uru-guayan and Argentinian sides of the Río de la Plata, and these populations are fairly uniform. Říčan and Kul-lander (2006) distinguished a separate group of populations (A. cf. facetus) within the nominal Australoherosfacetus. We provide below a more detailed analysis of the variation between the two forms from Uruguay.Tackling this variation will be crucial in order to classify Australoheros populations from the coastal drain-ages of Brazil. The coastal populations from Southern Brasil are most certainly different species from both A.facetus and A. cf. facetus. We have not examined enough material from these areas, but preliminary resultsshow that these populations have a low proportion of four abdominal bars (only in 12% of the specimens) andalso have a different combination of meristic characters. In this paper we use as comparative material fishesfrom the Río Jacui coastal drainage of Río Grande do Sul (A. sp. “Jacui”).

The two forms from Uruguay are distinguished mainly by meristic differences. We provide below a sepa-rate description of the material that we refer to as A. cf. facetus. Steindachner (1869) describes Heros facetus(Jenyns) by the following characters: the mouth is upwards directed, lower projects in front of upper jaw,modal count of 10 dorsal rays, 6 anal spines in all 6 specimens, dorsal fin scale cover only covering the lasttwo spine bases and the base of the soft part of the fin (i.e. the shortest dorsal fin cover observed among Aus-traloheros species), L1 scale counts 17 (18–19). The 6 anal spines and 10 dorsal rays distinguish the twoforms according to the original description. A. cf. facetus is slightly different in some meristic characters fromthe description by Steindachner.

Description of A. cf. facetus from coastal Uruguay and southern Rio Grande do Sul, Brazil. Based onspecimens over 60 mm SL with notes on smaller specimens. Meristic data are summarized in Table 1. Mor-phometric data are summarized in Table 2. NRM 47999 (20); NRM 48074 (15); NRM 48078 (39); NRM36495 (2); NRM 36774 (2); NRM 36848 (1); NRM 39527 (13); NRM 43943 (2); NRM 37035 (1 C&S);NRM 37037 (1 C&S); NRM 37039 (1 C&S).

High proportion of four abdominal bars (88.0% of the specimens), with only 7.4 % individuals withoutfull separation of the four abdominal bars.

Scales in E0 row 23(1), 24(25), 25(18), 26(1). Upper lateral line scales 13(3), 14(2), 15(9), 16(17), 17(11),18(2). Lower lateral line scales 6(2), 7(7), 8(19), 9(14), 10(3). Scales between lateral lines 2. Circumpeduncu-lar scales 16 (7 dorsally + lateral line scale + 7 ventrally + lateral line scale).

D. XV,10 (3), XVI,8 (1), XVI,9 (30), XVI,10 (12), XVI,11 (1), XVII,8 (3), XVII,9 (13), XVII,10 (1). A.VI,8 (5), VI,9 (1), VII,7 (14), VII,8 (32), VII,9 (2), VIII,7(5), VIII,8(5). Anal fin pterygiophores 12(20),13(40), 14(5). Modally two anal pterygiophores in front of the first haemal spine [1(11), 2(51), 3(3)]. Dorsalpterygiophores posteriorly of the first caudal vertebra 10(35), 11(27), 12(3). Pectoral fin with a rounded tip,extending to about third anal spine. P. 12(5), 13(34), 14(5).



TABLE 1. Meristics of Australoheros.

continued.

continued.

Dorsal fin count frequences XIV XV XV XV XVI XVI XVI XVI XVII XVII XVII XVII XVIII

12 9 10 11 8 9 10 11 7 8 9 10 9

A. charrua 1 4 1 2

A. facetus Argentina-Uruguay 2 2 20 3 1

A. cf. facetus Uruguay 3 1 30 12 1 3 13 1

A. forquilha 2 4 3 16 6

A. sp. jacui 1 1 12 1

A. kaaygua upper RRo Uruguay 16 13 2

RRo Iguazu 3 1

A. guarani 1 1 3 1

A. scitulus 1 1 1 5 32 4 1

A. tembe 5

A. minuano 1 7 1

Anal fin count frequences V V VI VI VI VII VII VII VIII VIII VIII IX IX IX

8 9 7 8 9 7 8 9 6 7 8 6 7 8

A. charrua 2 5 1

A. facetus Argentina-Uruguay 2 9 13 1 3

A. cf. facetus Uruguay 5 1 14 32 2 5 5


A. guarani 5 1

A. kaaygua upper RRo Uruguay 2 3 17 8 1

RRo Iguazu 1 3

A. minuano 2 4 3

A. scitulus 14 9 6 18 1

A. sp. jacui 8 3 1 2

A. tembe 4 1

vertebrae pectoral fin rays C1 gill rakers

12 12 12 13 13 13 13 14 14 14

13 14 15 12 13 14 15 12 13 14 12 13 14 15 5 6 7 8 9

A. charrua 1 6 1 1 7 1 5 2

A. facetus Argentina-Uruguay 23 5 5 11 1 2 15

A. cf. facetus Uruguay 59 6 5 34 5 13 14

A. forquilha 1 23 6 1 19 9 7 15 4

A. guarani 6 7 6

A. kaaygua upper RRo Uruguay 29 2 11 5 4 11 1

RRo Iguazu 1 3 4 1 2 1

A. minuano 9 6 1 1 6 1

A. scitulus 4 5 2 36 2 25 9 3 25 6

A. sp. jacui 1 14 6 2 3 5

A. tembe 6



continued.

continued.

continued.

caudal peduncle vertebrae

-2 -1.5 -1 -0.5 0 0.5 1 1.5 2 2.5 3 3.5

A. charrua 1 3 2 2

A. facetus Argentina-Uruguay 3 9 7 6

A. cf. facetus Uruguay 6 37 17 4


A. guarani 3 3

A. kaaygua upper RRo Uruguay 3 1 4 3 10 1

RRo Iguazu 2 2

A. minuano 2 1 3 2 1

A. scitulus 1 27 7 6

A. sp. jacui 3 10 2

A. tembe 1 4 1

anal pterygiophores dorsal pterygiophores

11 11 12 12 13 13 13 14 14 14 15

1 2 1 2 1 2 3 1 2 3 2 9 10 11 12

A. charrua 1 3 3 1 2 5 1

A. facetus Argentina-Uruguay 3 14 1 7 1 2 19 5

A. cf. facetus Uruguay 7 13 3 34 3 1 4 35 27 3

A. forquilha 9 4 9 6 2 6 19 6

A. guarani 5 1 1 5

A. kaaygua upper RRo Uruguay 1 1 7 8 3 3 1 13 9

RRo Iguazu 1 1 2 3 1

A. minuano 2 5 2 1 8

A. scitulus 8 11 2 2 18 1 1 27 16

A. sp. jacui 5 5 3 2 1 12 2

A. tembe 3 1 2 6

E0 scale counts L1 scale counts L2 scale counts

23 24 25 26 13 14 15 16 17 18 19 6 7 8 9 10 11

A. charrua 6 2 6 1 1 1 3 3 1

A. facetus Argentina-Uruguay 1 9 7 15 2 1 4 10 2

A. cf. facetus Uruguay 1 25 18 1 3 2 9 17 11 2 2 7 19 14 3

A. forquilha 1 16 11 1 10 12 4 6 17 4 1

A. guarani 7 2 4 1 1 2 2 1

A. kaaygua upper RRo Uruguay 3 16 4 1 6 8 4 7 4

RRo Iguazu

A. minuano 7 1 5 2 2 6

A. scitulus 6 27 3 1 5 15 13 1 3 3 12 12 6

A. sp. jacui 7 1 5 2 8



Gill rakers externally on first gill arch, 2 epibranchial, 1 in angle, 7(13), 8(14) ceratobranchial. Vertebrae 13+13=26(59), 13+14=27(6). Caudal peduncle containing 0.5(6), 1(37), 1.5(17), 2(4) vertebrae. Revised diagnosis of Australoheros facetus. Australoheros facetus is the only prognathous species (i.e.

lower jaw longer than upper and mouth pointing upwards) in the genus. It also has the shortest dorsal scalecover of all Australoheros species and the least scaled dorsal and anal fins among Australoheros (togetherwith A. tembe). A. facetus is the only species of Australoheros with four abdominal bars, which are present inmore than 80% of individuals as compared to 50% or less in all other species.

Revised description. Based on specimens over 60 mm SL with notes on smaller specimens. Meristic dataare summarized in Table 1. Morphometric data are summarized in Table 2.

TABLE 2. Morphometry of Australoheros.

continued.

A. forquilha A. sp. "Jacui"

N Min-Max Mean±SD N Min-Max Mean±SD

Head length 30 31.5-39.1 35.3±2 17 32.1-35.5 33.8±1

Snout length 30 7.6-17.7 13.1±2.9 17 10.1-14.3 12.3±1.3

Body depth 30 40.9-47.8 45.1±1.7 17 44.2-48.4 45.8±1.2

Orbital diameter 30 9.3-13.8 11.3±1 17 9.0-12.7 10.1±1

Head width 30 15.6-19.1 17.5±0.9 17 16.5-18.2 17.4±0.4

Interorbital width 30 8.7-11.5 9.7±0.7 17 10.4-12.4 11.3±0.6

Preorbital distance 30 6.4-10.8 8.8±1.1 17 5.3-8.3 7.4±0.7

Caudal peduncle depth 30 15.8-18.8 17.4±0.8 17 16.3-18.6 17.2±0.6

Caudal peduncle length 30 8.4-11.1 9.9±0.7 17 7.9-10.1 9.0±0.7

Pectoral fin length 30 25.7-32.6 28.9±2 17 25.1-29.0 26.5±1.1

Ventral fin length 30 22.1-34.1 28.7±2.6 17 28.2-32.2 30.0±1.1

Last D spine length 30 10.4-17.6 14.0±1.7 13 11.3-18.0 14.6±1.6

A. guarani A. minuano


Head length 5 31.8-33.4 32.4±0.7 9 34.6-37.8 35.9±1.1

Snout length 5 8.9-10.5 9.8±0.8 9 9.2-11.7 10.6±0.8

Body depth 5 46.2-49.9 48.1±1.6 9 45.6-48.7 46.9±1.2

Orbital diameter 5 9.5-11.2 10.5±0.8 9 11.0-13.8 12.6±0.8

Head width 5 17.6-18.5 18.1±0.4 9 18.4-19.5 18.8±0.4





Pectoral fin length 5 28.2-30.7 29.4±1.0 9 28.4-33.2 29.8±1.7


Last D spine length 5 16.6-19.5 18.1±1.1



continued.

continued.

Scales on head and chest not distinctly smaller than on flanks. Scales in E0 row 23(1), 24(9), 25(7). Upperlateral line scales 17(15), 18(2). Lower lateral line scales 7(1), 8(4), 9(10), 10(2). Scales between lateral lines2. Circumpeduncular scales 16 (7 dorsally + lateral line scale + 7 ventrally + lateral line scale). Cheek scalerows 2(1), 3 (12), 4(4).

D. XV,11 (2), XVI,9 (2), XVI,10 (20), XVI,11 (3), XVII,10 (1). A. VI,7 (2), VI,8 (9), VI,9 (13), VII,7 (1),VII,8 (3). Anal fin pterygiophores 12(15), 13(8). Modally one anal pterygiophore in front of the first haemalspine [1(21), 2(2)]. Dorsal pterygiophores posteriorly of the first caudal vertebra 10(2), 11(19), 12(5). P. 13(5),14(11), 15(1).

Gill rakers externally on first gill arch, 2 epibranchial, 1 in angle, 7(2), 8(15) ceratobranchial. Vertebrae 13+13=26(23), 13+14=27(5). Caudal peduncle containing -1(3), 0(9), 0.5(7), 1(6) vertebrae.Color pattern in alcohol. Color pattern composed of vertical bars, a midlateral blotch in a longitudinal

stripe and a caudal fin blotch, as in all other Australoheros species. A. facetus (together with A. cf. facetus; see

A. facetus A. cf. facetus


Head length 21 33.8-37.1 35.3±1.1 59 33.5-38.4 35.8±1.0

Snout length 21 9.3-12.6 11.2±0.7 59 7.2-12.2 9.4±1.3

Body depth 21 43.9-52.8 49.1±2.1 59 42.1-49.2 44.5±1.5


Head width 21 18.1-19.9 19.0±0.5 59 18.3-20.7 19.4±0.5








A. kaaygua Brazil A. charrua


Head length 16 31.7-36.2 33.3±1.5 8 31.5-34.0 32.4±1

Snout length 16 7.8-11.4 9.5±0.9 8 8.0-9.5 8.5±0.6

Body depth 16 46.2-51.5 49.6±1.2 8 44.5-49.6 46.9±1.5


Head width 16 16.4-20.5 17.7±1.2 8 17.0-18.3 17.7±0.5










below) is the only Australoheros species known to have as a rule four abdominal bars. Four abdominal barspresent in more than 80% of individuals (84.2% based on our material), but 87% of specimens do not have thebars completely separated, producing interconnected bars unique to A. facetus. Most other species have onlythree, or four in less than 50% of individuals. The midlateral blotch is small, usually not prominent outside theborders of the longitudinal midlateral stripe.

Color in life. Ground color is greyish, not yellow as in A. cf. facetus, A. kaaygua, A. minuano, or A. scitu-lus. Breeding coloration with the typical Australoheros autapomorphy of interrupted dorsal portions ofabdominal bars (Říčan & Kullander, 2006).

Distribution. Australoheros facetus as currently conceived is a catch-all taxon, including several unde-scribed species apart from those described in this work (see below). The distribution limits thus currentlyinclude the large area of the Atlantic coast drainages of Brazil and Uruguay plus populations from Argentinawest from the Uruguay River drainage. Geographical distribution of A. facetus s. str. is shown on Fig. 1.

FIGURE 1. Collecting localities of Australoheros guarani and A. facetus (the facetus group). A symbol may cover morethan one collecting site.



Notes. A. facetus is most similar to A. cf. facetus, A. guarani and A. minuano. It can be distinguished fromA. cf. facetus by 6 anal fin spines (vs. 7), 10 dorsal fin rays, by being more deep bodied, with a shorter caudalpeduncle and a longer snout and preorbital distance (refer to Table 3 for exact values and additional separatingcharacters).

TABLE 3. Significantly different values in characters in A. facetus and A. cf. facetus (P < 0.05; decreasing order of sig-

nificance; mean values ±SD, proportional measurements in percent).

From the very similar A. guarani and A. minuano, it can be distinguished by a shorter dorsal fin scalecover, covering only the bases of the last 2–3 spines (vs. 7–8), in being slightly more deep-bodied with anupturned mouth. Additionally distinguished from A. guarani in the proportion of four abdominal bars (in ca84% vs. none), by a longer head, shorter preorbital and interorbital distance, more scales between the anteriorend of dorsal fin and upper lateral line (3 ½ scales vs. 2 ½), and more C1 gill rakers and L2 scales (refer toTable 4 for exact values).

TABLE 4. Significantly different values in characters in A. facetus and A. guarani (P < 0.05; decreasing order of signifi-

cance; mean values±SD, proportional measurements in percent).

Additionally distinguished from A. minuano by more C1 gill rakers, more pectoral fin rays and by asmaller orbit (refer to Table 5 for exact values).

Australoheros cf. facetus is unique among all Australoheros species in having always four fully developedabdominal bars. It is most similar to A. facetus, A. guarani and A. minuano, from which it can be distinguishedin having more than 7 anal spines (vs. 6) and 16–17 dorsal fin spines. Further distinguished from A. facetus by10 dorsal fin rays, in being less deep-bodied, with a longer caudal peduncle and a shorter snout and preorbital

facetus cf. facetus

Body depth 49.1±2.1 44.5±1.5

Caudal peduncle depth 39.1±3.7 48.1±3.5

Caudal peduncle vertebrae 0.26±0.6 1.16±0.4

Anal fin spines 6.1±0.3 7.1±0.5

Ventral fin length 33.3±5.1 28.2±1.4

Dorsal fin rays 10.1±0.5 9.2±0.6

snout length in head length 30.6±3.1 26.3±3.9

Pectoral fin rays 13.5±0.7 13.0±0.5

Interorbital width in head length 36.5±3.8 31.1±2.5

Anal fin rays 8.4±0.6 7.8±0.5

Ceratobranchial 1 gill rakers 7.9±0.7 7.5±0.5

L1scales 16.8±0.6 15.8±1.2

Preorbital distance in head length 20.3±2.4 16.0±2.5

facetus guarani

Head length 34.5±1.5 32.4±0.7




L2 scales 8.6±0.7 7.5±1.0



distance (refer to Table 3 for exact values and additional separating characters). Additionally distinguished from both A. guarani and A. minuano by a shorter dorsal fin scale cover, cov-

ering only 2–3 last spines (vs. 7–8). Additionally distinguished from A. guarani by more scales between theanterior end of dorsal fin and upper lateral line (3 ½ scales vs. 2 ½), by a much shorter preorbital distance, bya slightly longer head and narrower interorbital distance and in being less deep-bodied (refer to Table 7 forexact values).

TABLE 5. Significantly different values in characters in A. facetus and A. minuano (P < 0.05; decreasing order of signif-icance; mean values±SD, proportional measurements in percent).

TABLE 6. Significantly different values in characters in A. cf. facetus and A. minuano (P < 0.05; decreasing order ofsignificance; mean values±SD, proportional measurements in percent).

Additionally distinguished from A. minuano in being less deep-bodied with a longer caudal peduncle, 6C1 gill rakers (vs. more than 7), 12 pectoral fin rays (vs. 13)(refer to Table 6 for exact values and additionalseparating characters).

Říčan and Kullander (2006) further distinguished a population from the Yacui drainage (A. sp. “Jacui”),Atlantic Ocean drainage, Brazil, Rio Grande do Sul. A. sp. “Jacui” is distinct by several characters, as alsoexpressed in the RDA and phylogenetic analyses (Říčan & Kullander, 2006), and clearly represents a separatespecies. Pending an analysis of Australoheros material from the coastal drainages of Brazil, we currently treatit as undescribed.

Australoheros scitulus (Říčan & Kullander, 2003)

Cichlasoma scitulum Říčan and Kullander, 2003: 795, fig. 1 (Type locality: Uruguay, arroyo Colla, Río Rosa-rio drainage, Río de la Plata basin, upstream Paso Arballo (34°19'7"S 59°20'13"W). Holotype NRM 36647).

Říčan and Kullander (2003) provide a detailed description of Australoheros scitulus and compared it withA. facetus from Uruguay. Only additional observation on life coloration and distribution are given here.

Color in life. Ground color is yellowish as in e.g. A. cf. facetus, A. kaaygua or A. minuano. Breeding col-

facetus minuano



Obrit diameter in head length 30.3±2.4 35.0±1.6

Body depth 49.1±2.1 46.9±1.2

cf. facetus minuano

Caudal peduncle vertebrae 1.16±0.4 -0.06±0.7




Body depth 44.5±1.5 46.9±1.2


Anal fin rays 7.8±0.5 8.4¡À0.5

Head width in head length 54.2±2.3 52.4±1.3




oration with the typical Australoheros autapomorphy of interrupted dorsal portions of abdominal bars (Říčan& Kullander, 2006). Life specimens have been photographed by Körber and Stawikowski (1999) and Staeck(2003, p. 63 upper left). Some northern populations do not distinctly develop the A. scitulus apomorphic spot-ted patterns (e.g. Staeck, 2003 p. 63 upper left), but are still distinguishable as A. scitulus using meristic char-acters, some coloration characters as the posterior part of the midlateral stripe and mouth/head shape.Breeding animals are deep yellow in dorsal interbar spaces, but ventral portion of the body is almost evenlyblack. The iris is red. This is in contrast to A. kaaygua, where breeding animals have yellow interbar spacesalso on the belly (i.e. bars clearly demarcated all the way from dorsal fin base to anal fin base).

Distribution. Australoheros scitulus is distributed in the tributaries of the Lower and Middle Río Uruguayin Uruguay, Argentina, and the State of Rio Grande do Sul, Brazil (Fig. 2).

TABLE 7. Significantly different values in characters in A. cf. facetus and A. guarani (P < 0.05; decreasing order of sig-

nificance; mean values±SD, proportional measurements in percent).

Australoheros tembe (Casciotta, Gómez & Toresanni, 1995)

Cichlasoma tembe Casciotta, Gómez & Toresanni, 1995: 194, fig. 1 (Type locality: Argentina, Arroyo Urugua-í, aboveSalto del Urugua-í, at `Alto Paraná' company fields, Rio Paraná basin. Holotype: MLP 9059).

Material examined: 6 specimens, 50.9–117.8 mm SL. Argentina, Misiones, Arroyo Urugua-í drainage: STRI2517, 1, 71.4 mm SL; STRI 2518, 1, 82.6 mm SL; STRI 2524, 1, 117.8 mm SL; STRI 2467, 3, 50.9–110.5mm SL.

Casciotta et al. (1995) provide a detailed description of Australoheros tembe and its comparison with A.facetus from Argentina. Only additional observation on life coloration and distribution are given here.

Color in life. Life specimens collected in the wild have a beautiful green ground color, which they how-ever fail to develop under aquarium conditions (Casciotta et al., 2003). Such green ground color is not knownfrom any other species of Australoheros. Casciotta et al. (2003) further describe the breeding of A. tembeunder aquarium conditions.

Australoheros tembe in breeding coloration have a yellowish–whitish ground color, with distinct blackvertical bars as in all Australoheros species. Breeding females have the ventral area and lower portion of headblack, while breeding males have the same area pinkish, similar in color to A. minuano, where the whole bodyof the breeding male is pinkish to red. Australoheros tembe is the only Australoheros species that does notdevelop or only develops in very limited degree the Australoheros autapomorphic breeding color pattern(Říčan & Kullander, 2006) of interrupted dorsal portions of abdominal bars. With hatching and free-swim-ming of fry both sexes guard the free-swimming offspring and also the male now develops the black colora-tion on the belly and lower portion of head (Casciotta et al., 2003).

Distribution. Australoheros tembe is endemic to the Arroyo Urugua-í, Río Paraná drainage, Misiones,Argentina (Casciotta et al., 1995; Fig. 3). Casciotta et al. (2003) mention a record of A. cf. tembe (MLP 8574)from the Arroyo Fortaleza, Río Uruguay drainage, Misiones, Argentina. Říčan and Kullander (2006) have

cf. facetus guarani


Head length 35.8±1.0 32.4±0.7


Body depth 44.5±1.5 48.1± 1.6




demonstrated the presence of A. forquilha, a similar species, in the Río Soberbio, Río Uruguay drainage, Mis-iones, Argentina (Fig. 3). A. cf. tembe could thus also be A. forquilha, or the two species may occur together inthese NW Argentinian tributaries of the Río Uruguay.

FIGURE 2. Collecting localities of Australoheros charrua and A. scitulus (the scitulus group). A symbol may covermore than one collecting site.

Australoheros forquilha, sp. nov.(Fig. 4)

Australoheros sp. ”Forquilha” (Říčan & Kullander, 2006).

Holotype. MCP 13936, male, 110.4 mm SL, Brazil, Rio Grande do Sul. Rio Forquilha, Rio Uruguai drainage,road from Maximiliano de Almeida to Machadinho. 4 October 1988. E. Pereira, L. Bergmann, P. Azevedo,and A. Ramírez.



FIGURE 3. Collecting localities of Australoheros forquilha and A. tembe (the forquilha group). A symbol may covermore than one collecting site.

Paratypes. 18 specimens, 15.6–129.7 mm SL. Brazil, Rio Grande do Sul: MCP 12123, 1, 42.1 mm SL,Rio Forquilha, Rio Uruguai drainage, road from Maximiliano de Almeida to Machadinho, 6 km from Maxi-miliano de Almeida, 25 May 1988. E. P. Lerner; MCP 13389, 1, 15.6 mm SL, Rio Forquilha, Rio Uruguaidrainage, road from Maximiliano de Almeida to Machadinho. 27 February 1989. E. Pereira, L. Bergmann, P.Azevedo, and A. Ramírez; NRM 13389, 1, 92.3 mm SL, Rio Forquilha, Rio Uruguai drainage, road fromMaximiliano de Almeida to Machadinho. 27 February 1989. E. Pereira, L. Bergmann, P. Azevedo, and A.Ramírez; MCP 12525, 1, 107.4 mm SL, collected with the holotype; MCP 6262, 9/35, C, D, N, P, Q, R, T, U,V, S, Sanga das Aguas Frias, Rio Uruguai drainage, Irai. 1985. L. R. Malabarba, R. E. Reis, and S. B. Mal-man. Brazil, Santa Catarina: MCP 12777, 1, 85.2 mm SL, Rio Canoas, Rio Uruguai drainage, road from Tupi-tinga to Celso Ramos. 10 November 1988. C. Lucena, E. Pereira, and P. Azevedo; MCP 18743, 4, 61.4–129.7mm SL, Rio Rancho Grande where crossing road BR-153 to Piritiba, Rio Uruguai drainage, 11 January 1996.



E. Filho, V. Schulz, S. Meurer, and P. Iaczinski. Additional non-type material: 21 specimens, Argentina, Misiones: ZSM 23482, 15/16, A, B, C, D (C&S),

E, F, G, H, I, J, K, L, M, N, O, 65.8–99.5 mm SL, Río Soberbio, Río Uruguay drainage, Soberbio. 1966. J.Foerster; ZSM 23060, 6/12, D (C&S), I, J, L, C, G, 47.1–77.0 mm SL, Rio Soberbio, Río Uruguay drainage,Soberbio. 1966. J. Foerster. Fractions denote numbers of specimens in mixed lots including more than onespecies.

FIGURE 4. Australoheros forquilha. Holotype, MCP 13936, male, 110.4 mm SL, Rio Forquilha, Rio Uruguai drainage,Brazil.

FIGURE 5. Diagnostic head markings of A. scitulus (A) and A. forquilha (B, C).

Diagnosis. Australoheros forquilha is unique among all Australoheros species in having dark markingsbelow the orbit along the postero-lateral border of the suborbital series (Fig. 5 B, C), in having checkerboard-spotted dorsal, anal and caudal fins and in having more than 14 caudal vertebrae in some specimens, in having25–26 E0 scales (vs. 24–25), a downward pointing subterminal mouth (all other Australoheros species), andin having the shortest interorbital and longest preorbital distances (Figs. 6 and 7). Also distinguished from allspecies in the unique scale cover of the base of dorsal fin with two cover scales corresponding to one trans-verse scale row.



FIGURE 6. Interorbital distance plotted against standard length, showing clear separation of A. forquilha from theremaining species.

FIGURE 7. Preorbital distance plotted against standard length, showing clear separation of A. forquilha from theremaining species.

Description. Based on specimens over 60 mm SL with notes on smaller specimens. Meristic data are sum-marized in Table 1. Morphometric data are summarized in Table 2.

Australoheros forquilha is similar to A. tembe in having a more slender body with a narrower interorbitalregion of the head, shorter pectoral and ventral fins and a long caudal peduncle including two or more verte-brae, in having slightly thickened lips and a proportionally large mouth. It is hypothesized to have affinitieswith A. tembe (see results).

The head profile is more round than in the other species and the mouth is subterminal. Lachrymal bonedeeper than wide. Lips comparatively thick. Caudal peduncle considerably deeper than long (depth 45–70%of length; mean 56%).

Scales on chest about half the size than the biggest scales in the E0 row above the pectoral fin (thussmaller than in A. facetus with the largest chest scales). About 8 scale rows between the opercular flap and the



anterior insertion of the pelvic fin in the holotype. Scales in E0 row 24(1), 25(16), 26(11). Upper lateral linescales 16(1), 17(10), 18(12), 19 (4). Lower lateral line scales 8(6), 9(17), 10(4), 11(1). Scales between upperlateral line and dorsal fin scale cover 3 posteriorly, 4 plus two small parallel scales anteriorly, forming a sheathof smaller scales arranged in pairs per scale row, along the insertion of the dorsal fin (Fig. 8A, 9A). Scales

between lateral lines 2. Cheek scale rows 3 (1), 4(15), 5(4). Interradial scales appear from 15th (rarely 14th)spine membrane, in single rows. Two or three last interradial membranes without scales. Anal fin with one

basal scale row; interradial scales in single rows, from the 6th membrane (i.e. between the last spine and thefirst branched ray), lacking on two last interradial membranes. Caudal fin densely scaled, scales ctenoid; inter-radial scales in single rows; posterior margin of scaly area concave, extending to between one-third and mid-dle of caudal fin.

Soft dorsal fin pointed, extending to the middle of the caudal fin (up to the end of caudal fin in the holo-type). D. XV, 10 (2), XV, 11 (4), XVI, 9 (3), XVI, 10 (16), XVI, 11 (6). Soft anal fin pointed, of about thesame length as dorsal fin. A. V,8 (1), VI, 7 (12), VI, 8 (15), VI, 9 (1), VII, 8 (2). Anal fin pterygiophores 11(13), 12 (16), 13 (2). Pelvic fin base slightly posterior of pectoral fin base; first ray longest. Pelvic fin compar-atively short, not extending to the genital papilla in most specimens (reaching to the first anal spine in theholotype). Pectoral fin comparatively short, with a rounded tip, posteriorly reaching to the same level as thepelvic fin. P. 13(19), 14(9). Caudal fin with rounded corners, subtruncate with a smooth indentation due toslightly shorter midline rays.

All teeth caniniform, slightly curved. Outer row teeth increasing in size symphysiad, upper jaw anteriorteeth longest and more robust, lower jaw anterior teeth subequal, midline two pairs more robust.

Lower pharyngeal tooth plate in a dissected specimen about one quarter wider than long (length 75% ofwidth) and very shallow compared to other Australoheros species (Fig. 10; cf. Fig. 17 and fig. 2 in Říčan &Kullander, 2006). Dentigerous area wider than long. 8–9 teeth along midline, 26 teeth along posterior margin.Posterior teeth tend to be progressively more compressed, except for medial teeth. Larger teeth medially andposteriorly, gradually smaller anteriad and laterad. Posterior teeth with forwards curved posterior cusp andsubapical anterior shelf. Large laterally compressed teeth with a second cusp projecting anteriorly from shelf.

Gill rakers externally on first gill arch: 2 epibranchial, 1 in angle, 7(7), 8 (15), 9 (4) ceratobranchial. Vertebrae 13+13=26(1), 13+14=27(23), 13+15=28(6), 14+13=27(1). Caudal peduncle contains 1.5(6),

2(13), 2.5(6), 3(6) vertebrae. Color pattern in alcohol. Six or seven vertical flank bars, a midlateral blotch in the third flank bar from

posterior, an inconspicuous caudal fin base spot and the caudal peduncle bar make up the principal markings.Vertical bars are relatively wide, faint, indistinct in their ventral parts. The posteriormost bar bears the caudalspot, which is very faint or even absent in some specimens. The caudal peduncle bar covers the posterior lessthan half of the peduncle. The third bar, being the first flank bar, runs between the ends of dorsal and anal fins.The third flank bar, bearing the midlateral blotch is centered above the anteriormost portion of the anal fin.There are three or four abdominal bars. The number varies in adults, whereas all juveniles have four bars:compared to the situation in A. scitulus, the bar anterior from the midlateral blotch bar is divided into two. Thelast flank bar runs from anterior of the dorsal fin insertion, along the posterodorsal edge of the opercular cleftdown to the insertion of the pectoral fin or slightly anterior from it.

There are only weak traces of a midlateral stripe in adult specimens. Dorsal, anal and caudal fins with darkspots in a checker-board spotting pattern on interradial membranes. A line of black longitudinal blotchesbelow the orbit along the postero-lateral border of the suborbital series.

Color in life. Only one photograph of a life A. forquilha is known to us and it was presented in the Appen-dix to Stawikowski and Werner (2004: p. 455) as “Cichlasoma” cf. tembe. The fish in the photograph clearlyshows characters distinctive for A. forquilha (checkerboard spotting on unpaired fins, opalescent markings onhead and also on body scales, head shape with a low position of mouth). It was collected in an unspecifiedtributary of the Upper Uruguay in Brazil, and it thus also in distribution agrees with A. forquilha. The speci-



men is photographed in a field aquarium and is very colorful, with a red lower portion of body and head andgreen scales on body. The green color is similar to A. tembe.

Distribution. Australoheros forquilha is endemic to the tributaries of the Upper Uruguay in Brazil and itssouthernmost locality is in the Río Soberbio, an Argentinian tributary of the Middle Río Uruguay just belowthe Moconá falls separating the Middle and Upper Rio Uruguay (Fig. 3). It shares its distribution with A.kaaygua in the Río Uruguay basin (Fig. 14).

Etymology. The species is named after the Rio Forquilha, Rio Grande do Sul, Brazil, from where the spe-cies is described.

FIGURE 8. Differences in squamation among Australoheros species, showing two extreme situations. A, A. forquilha,B, A. facetus. Note the shorter basal scale row at the base of the dorsal fin, less scaled dorsal and anal fins, and less scalerows between the upper lateral line and the dorsal fin (both anteriorly and posteriorly) in A. facetus.



FIGURE 9. Differences in squamation among Australoheros species. A, A. forquilha, B, A. sp. “Jacui”, C, A. tembe, D,A. kaaygua, E, A. charrua, F, A. scitulus, G, A. guarani, H, A. minuano, I, A. facetus.



FIGURE 10. Lower pharyngeal jaw of a dissected specimen of Australoheros cf. forquilha, ZSM 23060 G, in (top tobottom) occlusal, caudal and left lateral aspect.

Notes. Australoheros forquilha can be easily distinguished from A. tembe by the diagnostic colorationmarkings lacking altogether in A. tembe (see above), a subterminal mouth and a more rounded head profile, 10or 11 dorsal fin rays (vs. 9 in A. tembe), 8 anal fin rays (vs. 7), and also by the mentioned shorter preorbital



and interorbital distance and a shorter caudal peduncle (including 2 vs. 3 vertebrae; modal values) (see Table8).

Distinguished from the A. scitulus group (A. scitulus, A. charrua) and A. kaaygua by body shape (beingmore elongated with shorter pectoral and ventral fins), larger mouth, higher C1 gill raker counts (7–8 vs. 6),longer caudal peduncle (modally containing 2–3 vertebrae vs. 1 or none) and coloration. It lacks the longitudi-nal stripe dominant in these three species and also has the bar in front of the midlateral blotch (in about half ofthe adult specimens and all juveniles) divided into two bars contrary to a single bar present in both juvenilesand adults of the three latter species.

Distinguished additionally from A. scitulus (and juvenile A. kaaygua) in lacking any dominant rows ofspots on flank scales (above anterior part of upper lateral line), in lacking the opercular and head spots of A.scitulus, also no prominent black spot in the axil of pectoral fin. Also distinguished from A. scitulus by having16 dorsal fin spines (vs. 17), 6 to 7 anal fin spines (vs. 8–9), and narrower interorbital distance (23.3–35.3%HL, mean 27.6% vs. 29.7–40.0%, mean 35.2% in specimens above 50mm SL).

Additionally distinguished from both A. kaaygua and A. charrua in having spotted unpaired fins, 6 analfin spines (vs. 7), 4–5 cheek scale rows (vs. 3), and narrower interorbital distance (23.3–35.3% HL, mean27.6% vs. 31.7–38.5%, mean 35.5% vs. 34.5–37.7%, mean 36.2% in specimens above 50mm SL). Addition-ally distinguished from A. kaaygua in having 14–15 caudal vertebrae (vs. 13) and 13–14 pectoral fin rays (vs.12–13).

In addition, distinguished from the A. facetus group (A. facetus, A. cf. facetus and A. minuano) and A. gua-rani by higher meristics, especially caudal vertebrae (14 to 15 vs. 13), caudal peduncle vertebrae (more than 2vs. fewer than 2), E0 scale row count (25 or more vs. 24), and from A. minuano additionally by fewer C1 gillrakers (7 to 8 vs. 6) and pectoral fin rays (13 vs.12).

Additionally distinguished from the A. facetus group species in mouth shape and position (inferior mouthvs. slightly up-turned mouth in A. facetus and A. cf. facetus , terminal in A. minuano), in coloration by nothaving a well-circumscribed midlateral stripe and a prominent caudal fin spot. The A. facetus group speciesalso have a much higher percentage of four well-developed abdominal bars.

TABLE 8. Significantly different values in characters in A. forquilha and A. tembe (P < 0.05; decreasing order of signif-icance; mean values±SD, proportional measurements in percent)..

The material from Soberbio (Misiones, Argentina) is in bad condition. This excludes most colorationcharacters, including the diagnostic suborbital markings and the checkerboard-spotting of unpaired fins frombeing studied. Soberbio specimens have slightly lower counts of caudal vertebrae (13–14 vs. 14–15), slightlyless caudal peduncle vertebrae (modally 2 vs. modally 3), 5–6 anal fin spines (vs. 6–7), slightly lower totaldorsal fin counts (25–26 vs. 26–27), 25 vs. 26 E0 scales and 3–4 vs. 4–5 cheek scale rows. Soberbio popula-tions possess the diagnostic shortest interorbital and longest preorbital distances. The inclusion of the Sober-bio populations into A. forquilha is in agreement with the RDA analysis (Říčan & Kullander, 2006), but inconflict with the populations as terminal taxa phylogenetic analyses (which include all informative charactersincluding the above mentioned proportions; op. cit.). Untill better-preserved specimens from the area are stud-

forquilha tembe


Caudal peduncle vertebrae 2.16±0.5 3.04±0.2)







ied, or until a phylogeographic study sheds more light on the matter, we prefer to treat the Soberbio populationas A. forquilha.

TABLE 9. Significantly different values in characters in A. scitulus and A. charrua (P < 0.05; decreasing order of signif-

icance; mean values±SD, proportional measurements in percent).

Australoheros charrua, sp. nov.(Fig. 11)

Australoheros sp. ”Pirapo” (Říčan & Kullander, 2006).

Holotype. MCP 13938, 77.9 mm SL, Brazil, Rio Grande do Sul, Arroyo Canoin, Rio Uruguai drainage, roadfrom Pirapo to São Nicolau. 2 November 1988. C. Lucena, L. Bergmann, E. Pereira, and P. Azevedo.

Paratypes. 7 specimens, 36.2–70.5 mm SL, all from Brazil, Rio Grande do Sul, collected with the holo-type. MCP 12667, 5, 36.2–69.7 mm SL; NRM 12667, 2, 61.0–70.5 mm SL.

Diagnosis. Most similar to A. scitulus, from which it is readily distinguished in lower dorsal (16 vs. 17)and anal (7 vs. 8–9) fin spine counts, in higher dorsal ray counts (10 vs. 9), and shorter head and larger inter-orbital distance (refer to Table 9 for exact values). It lacks the diagnostic blotches on head and body usuallypresent in A. scitulus.


In body form, proportions and most other features this species is very similar to A. scitulus (Říčan & Kul-lander, 2003). Mouth also small, but lower on the head with jaws of equal length. Lips narrow. Caudal pedun-cle considerably deeper than long (depth 30–49% of length; mean 40.5%).

Scales on head and chest not distinctly smaller than on flanks. Scales in E0 row 25(6), 26(2). Upper lateralline scales 17(6), 18(1), 19(1). Lower lateral line scales 7(1), 8(3), 9(3), 10 (1). Scales between upper lateralline and dorsal fin: 4 large anteriorly (or 3 large and one small), 2 large and one small posteriorly. Cheek scalerows 3(3), 4(5). About 8 scale rows between the opercular flap and the anterior insertion of the pelvic fin (asin A. scitulus).

Dorsal fin with one basal scale row, starting from the seventh or eighth spine and running posteriad; inter-

radial scales appear from 14th or 15th spine membrane, in single rows. Two or three last interradial membraneswithout scales. Anal fin with one basal scale row; interradial scales in single rows, starting between penulti-mate spine membrane and first ray membrane. Caudal fin densely scaled, scales ctenoid; interradial scales insingle rows; hind margin of scaly area concave, extending to between one-third and middle of caudal fin.

Soft dorsal fin pointed, extending to middle or three quarters of caudal fin. D. XVI,9 (1), XVI,10 (4),XVII,9 (1), XVII,10 (2). Soft anal fin pointed, of about the same length as dorsal fin. A. VII,7 (2), VII,8 (5),VIII,7 (1). Anal fin pterygiophores 12(4), 13(4). Pelvic fin extending to second or third anal spine. Pectoral finwith third and fourth rays longest, extending to about the level of anus and just to the midlateral blotch. P.12(1), 13(7) Caudal fin rounded to subtruncate.

scitulus charrua

Dorsal fin spines 16.9±0.4 16.3±0.5

Head length 34.6±1.4 32.4±1.0



E0 scales 24.9±0.5 25.3±0.5




All teeth caniniform, slightly curved. Outer row teeth increasing in size symphysiad, upper jaw anteriorteeth longest, lower jaw anterior teeth subequal.

Lower pharyngeal tooth plate not studied.

FIGURE 11. Australoheros charrua. Holotype, MCP 13938, 77.9 mm SL, Arroyo Canoin, Rio Uruguai drainage, Bra-zil.

Gill rakers externally on first gill arch, 2 epibranchial, 1 in angle, 5(1), 6(5), 7(2) ceratobranchial. Vertebrae 13+13=26(1), 13+14=27(6), 14+14=28(1). Caudal peduncle containing 2(1), 0(3), 0.5(2), 1(2)

vertebrae. Color pattern in alcohol. Basic coloration markings include six flank bars, i.e. three abdominal bars. The

caudal spot bar is very narrow, with only a very small or completely missing caudal spot. The caudal pedunclebar is due to the very short caudal peduncle confluent with the darker caudal spot bar. Juvenile and subadultspecimens do have a clearly developed caudal base spot above the lateral line. Vertical bars are relativelywide, faint, indistinct in their ventral parts. Other general aspects of the coloration are given above in the com-parison of coloration patterns of A. charrua, A. scitulus and A. kaaygua. The arrangement of the bars on thebody is essentially the same as described in A. scitulus (Říčan & Kullander, 2003). Slight differences arefound in the midlateral stripe. Compared to A. scitulus, the midlateral stripe is even better developed and morecontinuous anteriorly from the midlateral blotch. Posterior from the midlateral blotch, the stripe is betterdescribed as a series of two blotches, the anterior one being centered in the E2 scale row, the posterior one inthe E3 scale row.

Unpaired fins without spots. Flank and head scales also without spots. Color in life. No photographs of live specimens are known to us. Live fish probably look very similar to a

fish photographed by Staeck (2003: p. 63 upper left). This photograph shows A. scitulus of the form referredto as `C.` sp. ”Quarai” (Lucena & Kullander, 1992), where some northern populations do not distinctlydevelop the A. scitulus apomorphic spotting patterns, but are still distinguishable as A. scitulus using meristiccharacters and some coloration characters as the posterior part of the midlateral stripe and mouth/head shape.

Distribution. Known only from the type locality in Brazil, Rio Grande do Sul, Río Uruguay drainage (Fig.2), overlapping in distribution with A. minuano (Fig. 14). Also Crenicichla gaucho has the same limited areaof distribution.

Etymology. The species is named after the indigenous people of the Charrua, which inhabited an areaincluding the Rio Grande do Sul State (Brazil).



TABLE 10. Significantly different values in characters in A. charrua and A. facetus (P < 0.05; decreasing order of signif-

icance; mean values±SD, proportional measurements in percent).

Notes. Australoheros charrua is most similar to A. scitulus and also A. kaaygua (see phylogenetic analy-ses) in general appearance, body shape (relatively deep-bodied), small isognathous snout, few C1 gill rakers(6), short caudal peduncle and coloration. Similarities in coloration (in preserved specimens) to A. scitulus andA. kaaygua include a dominant longitudinal stripe and a prominent midlateral blotch and are described inmore detail below. In general, A. charrua is more similar to A. scitulus, both in higher meristic counts, detailsof coloration patterns and head shape (mouth positioned low on head with the anterior upper margin of pre-maxilla well below the horizontal from the ventral margin of the eye). The mouth is positioned even lower onthe head in A. charrua than in A. scitulus making the straight ventral outline of the head aligned even morehorizontally; Fig. 12). A. kaaygua on the other hand has similarities with A. scitulus in being spotted onunpaired fins and flank scales, but it can be easily distinguished from both, A. scitulus and A. charrua, in mer-istic counts. Both, A. charrua and A. kaaygua, lack spots on the operculum, suboperculum and posterior partof head, diagnostic for A. scitulus.

The similarities in coloration between A. scitulus, A. charrua, and the below described A. kaaygua includethe following:

1.The midlateral stripe is more distinctly developed anterior from the midlateral blotch than posteriorfrom it in adult specimens.

2.The midlateral stripe turns upwards posterior from the midlateral spot, whereas it follows the body axisin all the other species (except A. minuano, see below; i.e. it goes all the way along the body in scale rows E0and E1). This bending is most pronounced in A. charrua where the midlateral blotch is centered in the E1scale row, while the next posterior blotch is centered in the E2 scale row and the blotch in the last body bar iscentered in the E3 scale row. This successive shift in the position of the body bars one scale row up per blotchmakes the midlateral stripe look like turning up dorsally posteriad from the midlateral stripe. In A. kaayguaand A. scitulus the blotch posterior to the midlateral blotch is centered in the same scale row as the midlateralblotch (i.e. E1 scale row), and only the last blotch is very high on the body. However, all three share the fea-ture that the midlateral stripe does not continue in the E0 scale row posterior to the midlateral blotch (which isalso shared with A. minuano).

3.The midlateral blotch is very large, spanning from the E0 scales to the E3 scale row scales or at leastinto the dorsal half of the E2 row scales

4.There is no clearly visible caudal base spot, mostly only a more pigmented narrow bar at the caudalbase, which is fused with the caudal peduncle bar.

5.All species develop always only three abdominal bars and there are also no traces of four developingbars in developmental series.

charrua facetus




Head length 32.4±1.0 34.5±1.5

caudal vertebrae 13.9±0.4 13.2±0.4

Snout length in head length 26.3±2.0 30.6±3.1

E0 scales 25.3±0.5 24.5±0.6


Body depth 46.9±1.5 49.1±2.1



Additionally distinguished from A. kaaygua by having more vertebrae (14 vs. 13), more E0 scales (25 vs.24), more cheek scale rows (3–4 vs. 3) and pectoral fin rays (13 vs. 12), and in being less deep-bodied andwith a shorter snout (see A. charrua).

TABLE 11. Significantly different values in characters in A. charrua and A. cf. facetus (P < 0.05; decreasing order of

significance; mean values±SD, proportional measurements in percent).

Except for the coloration characters, distinguished from the A. facetus-like species also by a different headshape, with the mouth very low on the head, small terminal isognathous mouth and except A. guarani also inhaving only three (indistinct) abdominal bars (vs. 4). Additionally distinguished from all in having more cau-dal vertebrae (14 vs. 13) and more E0 scales (25 vs. 24), from all except A. minuano in having 6 C1 gill rakers(vs. 7–8), and from all except A. cf. facetus in having 7 anal spines (vs. 6). For all meristic and morphometricdifferences between A. charrua and the A. facetus-like species see Tables 10–13.

Also distinguished from A. forquilha and A. tembe by having fewer caudal peduncle vertebrae (none, orup to one vs. modally 2), and 7 anal fin spines (vs. 6 in A. forquilha and A. tembe). From A. forquilha addition-ally by 6 C1 gill rakers (vs. 8).

charrua cf. facetus

Head length 32.4±1.0 35.8±1.0



Caudal peduncle vertebrae 0.1±0.9 1.16±0.4



E0 scales 25.3±0.5 24.4±0.6


Body depth 46.9±1.5 44.5±1.5

Pectoral fin length 30.3± 1.4 28.7± 1.0



FIGURE 12. Head and snout shape differences among species of Australoheros The mouth has to be completely closedin the preserved fish to observe the sometimes slight differences.

Australoheros kaaygua Casciotta, Almirón & Gómez, 2006(Fig. 13)

Australoheros kaaygua Casciotta, Almirón & Gómez, 2006: 78, fig. 1 (Type locality: arroyo Ñandú, Río Iguazú basin,Misiones, Argentina. Holotype: MACN-ict 8917).

Australoheros sp. “Jacutinga” (Říčan & Kullander, 2006).



TABLE 12. Significantly different values in characters in A. charrua and A. guarani (P < 0.05; decreasing order of sig-


TABLE 13. Significantly different values in characters in A. charrua and A. minuano (P < 0.05; decreasing order of sig-


Material examined. 40 specimens, 24.6–77.0 mm SL. Brazil, Paraná: Rio Iguaçu drainage: NUP 3913, 2, fieldnumbers 3683 and 3967, Rio São Pedro, tributary to Rio Iguaçu, Pinhão county, Paraná State (26ºS/51º45'W),28.iii.1993, collectors: NUPELIA staff; NUP 3914, 1, field number 7743, Rio Iratim (Linígrafo), tributary toRio Iguaçu, Palmas county, boundary with Pìnhão-PR, Paraná State (26º05´S/51º45´W), 27.iv.1993, collec-tors: Nupélia staff; NUP 3915, 1, field number 8861, Rio São Pedro, tributary to rio Iguaçu, Pinhão county,Paraná State (26ºS/51º45'W), 28.iii.1993, collectors: Nupélia staff. Brazil, Santa Catarina: MCP 13937, 1,73.2 mm SL, Rio Jacutinga, Rio Uruguai drainage, road BR 283 from Ceará to Concordia. October 1988. L.Bergmann, E. Pereira, P. Azevedo, and A. Ramírez; MCP 13383, 6, 24.6–77.0 mm SL, Rio Jacutinga, RioUruguai drainage, road BR 283 from Ceará to Concordia. 16 February 1989. R. E. Reis, L. Bergmann, E.Pereira, and P. Azevedo; MCP 12509, 1, 75.0 mm SL, Rio Jacutinga, Rio Uruguai drainage, road BR 283from Ceará to Concordia. October 1988. E. Pereira, L. Bergmann, P. Azevedo, and A. Ramírez; MCP 13011,6, 44.2–61.4 mm SL, Rio Jacutinga, Rio Uruguai drainage, road BR 283 from Ceará to Concordia. 8 Decem-ber 1988. R. E. Reis, L. Bergmann, E. Pereira, and P. Azevedo. Brazil, Rio Grande do Sul: MCP 6262, 13/35:A, B, E, F, G, H, I, J, K, L, M, O, S. Sanga das Aguas Frias, Rio Uruguai drainage, Irai. 1985. L. R. Malabarba,R. E. Reis, and S. B. Malman; MCP 12710, J-K, 2, Arroyo Canoin, Rio Uruguai drainage, road from Pirapo toSão Nicolau. 2 November 1988. C. Lucena, L. Bergmann, E. Pereira, and P. Azevedo. Argentina, Misiones:ZSM 23482, 1/16: P, 63.1 mm SL, Río Soberbio, Río Uruguay drainage, Soberbio. 1966. J. Foerster; ZSM

charrua guarani

E0 scales 25.3±0.5 24.0±0.0







charrua minuano


Head length 32.4±1.0 35.9±1.1

E0 scales 25.3±0.5 24.1±0.4



Anal fin rays 7.6±0.5 8.4±0.5

L1 scales 17.4±0.7 16.3±0.5







23060, 6/12, A (C&S), B (C&S), E, H, K, F, 50.3–66.2 mm SL, Rio Soberbio, Río Uruguay drainage, Sober-bio. 1966. J. Foerster. Fractions denote numbers of specimens in mixed lots including more than one species.

Description of Australoheros kaaygua material from Brazil. Based on specimens over 60 mm SL withnotes on smaller specimens. Meristic data are summarized in Table 1. Morphometric data are summarized inTable 2.

The description of A. kaaygua from Misiones, Río Iguazú drainage, Argentina, by Casciotta et al. (2006)agrees well with our material from Brazil. Specimens from the Iguaçu drainage of Brazil have only an evenshorter caudal peduncle (-1 or fewer vertebrae; the shortest caudal peduncle found in Australoheros) andslightly larger scales, visible as only 3 scales anteriorly between the upper lateral line and origin of D fin, only1 and 1 small posteriorly. We treat these two differences as intraspecific variation and do not believe that theywould credit a separate species status. The analysis by Říčan and Kullander (2006) also supports the conspeci-fity of the Río Iguazú and Río Uruguay specimens.

Shape. See Fig. 13 for general aspect. Comparatively deep bodied (mean body depth 49.6% SL). Snoutshort, strait in lateral view. Jaws isognathous. Mouth small. In other respects it fits the description of A. scitu-lus (Říčan & Kullander, 2003).

Scales on head and chest not distinctly smaller than on flanks. Scales in E0 row 23(3), 24(16), 25(4).Upper lateral line scales 16(1), 17(6), 18(8). Lower lateral line scales 7(4), 8(7), 9(4). Scales between upperlateral line and dorsal fin 4 anteriorly, 1 big plus 1 small posteriorly. Cheek scale rows 3(14), 4(2). About 8scale rows between the opercular flap and the anterior insertion of the pelvic fin (as in A. scitulus and A. char-rua).

Dorsal fin with one basal scale row, starting from the seventh or eighth spine and running posteriad; inter-radial scales appear from 14th or 15th spine membrane, in single rows. Anal fin with one basal scale row; inter-radial scales in single rows, from penultimate spine. Caudal fin densely scaled, scales ctenoid; interradialscales in single rows; hind margin of scaly area concave, extending to between one-third and middle of caudalfin.

FIGURE 13. Australoheros kaaygua. MCP 13937, 73.2 mm SL, Rio Jacutinga, Rio Uruguai drainage, Brazil.

Soft dorsal fin pointed, extending beyond middle of caudal fin. D. XVI,9 (16), XVI,10 (13), XVII,8 (2).Soft anal fin pointed, of about the same length as dorsal fin. A. VI,7 (2), VI,8 (3), VII,7 (17), VII,8 (8), VIII,6(1). Anal fin pterygiophores 11(2), 12(22), 13(7). First pelvic fin ray longest, extending up to the second analfin spine. Pectoral fin with a rounded tip, third and fourth rays longest, extending just to the midlateral blotch.



P. 12(11), 13(5). Caudal fin rounded to subtruncate.All teeth caniniform, slightly curved. Outer row teeth increasing in size symphysiad, upper jaw anterior

teeth longest, lower jaw anterior teeth subequal. Number of lower jaw teeth up to 16 in one outer hemiseries,upper jaw tooth row much shorter, with about 7 or 8 teeth in one outer hemiseries.

Lower pharyngeal tooth plate not studied. Gill rakers externally on first gill arch, 2 epibranchial, 1 in angle, 5(4), 6(11), 7(1) ceratobranchial. Vertebrae 13+13=26(29), 13+14=27(2). Caudal peduncle containing -1(4), -0.5(1), 0(5), 0.5(4), 1(14),

1.5(1) vertebrae.Color pattern in alcohol. Six vertical flank bars, a caudal peduncle bar confluent with the caudal bar base

bar, and a midlateral stripe bearing the midlateral blotch in the third flank bar make up the principal markings.All fins and body are without conspicuous dark spots or blotches. The midlateral stripe is more distinct anteri-orly from the midlateral blotch posteriorly, and the midlateral blotch itself is a dominant coloration element.Vertical bars are relatively wide, faint, indistinct in their ventral parts. The midlateral stripe posteriorly fromthe midlateral blotch does not align with the lower lateral line and aligns with the E1 scale row and does notcontinue in the E0 scale row. Posteriorly from the midlateral blotch, the stripe is slightly decomposed into twoblotches in the respective vertical flank bars. The blotch posterior from the midlateral blotch is centered in thesame scale row as the midlateral blotch (i.e. E1 scale row), whereas the second blotch is more elongate alongthe vertical axis and centered in the E2 scale row, making the impression that the midlateral stripe makes adorsally directed turn at its posterior end. The arrangement of the bars on the body in essentially the same asdescribed for A. scitulus (Říčan & Kullander, 2003).

Dorsal, anal and caudal fins with very small dark spots on interradial membranes. Very small dark spotspresent also on the bases of some body scales in adult specimens. In juveniles the spotted pattern of the bodyis much more pronounced, with virtually every scale on the body having a dark spot at its base, includingthose in the anterior part of the E4 scale row (i.e. as in adult A. scitulus). Juveniles, on the contrary, withoutspots in unpaired fins.

Color in life. Life fishes have been photographed by Staeck (1998a, 1998b, 2003: p. 64). The ground colorof neutral fishes is yellowish to yellow or orange. Many other species of Australoheros have a yellowishground color, but it is best developed in A. kaaygua. The caudal fin has red dorsal and ventral margins andcorners. This character is not unique for A. kaaygua, and can also be seen in populations of A. facetus fromUruguay for example. Also specimens from the Rio Iguaçu tributaries clearly show the red markings on thecaudal fin. Breeding animals have the typical Australoheros breeding coloration with the horizontal interrup-tion of the black vertical bars in their dorsal portion between the opercle and the midlateral blotch (Říčan &Kullander, 2003; Staeck 1998a p. 82, 1998b: p. 62, 2003: p. 65). The yellow interbar spaces reach all the waydown to the belly, even in the breeding coloration (i.e. bars clearly demarcated all the way from D fin base toA fin base). Females in breeding coloration develop a black blotch in the dorsal fin. The yellowish groundcolor lasts till the female starts to guard free-swimming fry when they lose it and develop a pale, whitish-grey-ish ground color. This color change is typical for most other cichlasomatine cichlids. Males usually do notchange their ground color into white when helping with guarding fry and juveniles, which is also typical formost other cichlasomatine cichlids where the female takes most of the guarding of fry. Staeck (1998b, 2003)describes behavior and spawning under aquarium conditions.

Distribution. Australoheros kaaygua has a disjunct distribution, including tributaries of the Río Iguazú, atributary of Río Paraná in Brazil and Argentina (Casciotta, Almirón and Gómez, 2006), and tributaries of theUpper Rio Uruguay in Brazil (Fig. 14).



FIGURE 14. Collecting localities of Australoheros kaaygua and A. minuano (the kaaygua group). A symbol may covermore than one collecting site.

Notes. Juvenile A. kaaygua are similar to juvenile A. scitulus (of about 20 mm SL) in having a characteris-tic pattern made of rows of spots most marked anteriorly in the E4 scale row (above anterior part of upper lat-eral line) and also in the two scale rows below the upper lateral line (see Říčan & Kullander, 2006). It isunknown whether juvenile A. charrua have these spots as well.

Australoheros kaaygua can be distinguished from both A. scitulus and A. charrua in having 13 caudal ver-tebrae (vs. 14), and by modal counts of 12 pectoral fin rays (vs. 13), 24 E0 scales (vs. 25), 3 cheek scale rows(vs. 3–4) and in being more deep-bodied.

Additionally distinguished from A. scitulus in having 7 anal spines (vs. 8–9) and 16 dorsal spines (vs. 17),in having a larger interorbital distance, and longer ventral fins (refer to Table 14 for exact values). The differ-



ence in the anal fin spine counts is also evident even without counting the spines, since both adult as well asjuvenile A. kaaygua can be distinguished from A. scitulus by the origin of the anal fin, which in A. kaayguaoriginates in the middle of the spot-bearing vertical bar, whereas in A. scitulus the anal fin originates well infront of the spot-bearing vertical bar. The differences in coloration include retaining the spotted flank scales inA. scitulus into adult sizes in contrast to A. kaaygua, where they are mostly present only in juveniles, and adultA. scitulus also develop distinct spots on the head, opercular region, and in the anterior part of the E4 scalerow. Spotted unpaired fins are also less common and less pronounced in A. kaaygua than in A. scitulus speci-mens.

Additionally distinguished from A. charrua in having a shorter snout (refer to Table 15 for exact values)and by having the dorsal margin of the upper lip horizontally aligned with the ventral border of the eye (vs.mouth situated much lower on the head). Also distinguished by having the midlateral stripe posteriorly fromthe midlateral blotch not as distinctly decomposed into blotches, and also not as distinctly bent upwards. Themidlateral stripe posterior from the midlateral blotch is centered in the same scale row as the midlateral blotch(i.e. E1 scale row), and only the last blotch is very high on the body. The midlateral stripe does not continue inthe E0 scale row posterior to the midlateral blotch (see description of coloration patterns uniting A. charruawith A. scitulus and A. kaaygua in the description of A. charrua). Also distinguished from A. charrua by hav-ing fewer caudal vertebrae (13 vs. 14), fewer E0 scales (24 vs. 25), fewer cheek scale rows (3 vs. 3–4) andpectoral fin rays (12 vs.13), and in being more deep-bodied and with a longer snout (refer to Table 15 forexact values).

TABLE 14. Significantly different values in characters in A. scitulus and A. kaaygua (P < 0.05; decreasing order of sig-


TABLE 15. Significantly different values in characters in A. charrua and A. kaaygua (P < 0.05; decreasing order of sig-


scitulus kaaygua


Dorsal fin spines 16.9±0.4 16.1±0.3


Cheek scales 3.6±0.5 3.1±0.3


E0 scales 24.9±0.5 24.0±0.5

Body depth 46.8±2.0 49.5±1.2



charrua kaaygua

Caudal vertebrae 13.9±0.4 13.1±2.5

Body depth 46.9±1.5 49.5±1.2

E0 scales 25.3±0.5 24.0±0.5

Cheek scales 3.6±0.5 3.1±0.3





TABLE 16. Significantly different values in characters in A. kaaygua and A. facetus (P < 0.05; decreasing order of sig-


TABLE 17. Significantly different values in characters in A. kaaygua and A. cf. facetus (P < 0.05; decreasing order of

significance; mean values±SD, proportional measurements in percent).

kaaygua facetus





Anal fin rays 7.4±0.6 8.4±0.6


Head length 33.3±1.5 34.5±1.5


L2 scales 8.0±0.8 8.6±0.7


kaaygua cf. facetus

Body depth 49.5±1.2 44.5±1.5



Caudal peduncle vertebrae 0.4±0.7 1.16±0.4)



Head length 33.3±1.5 35.8± 1.0



Pectoral fin length 30.3±1.4 28.7± 1.0

L1scales 17.5±0.6 15.8±1.2


Number of anal fin pterygiophores 12.2±0.4 12.8 0.6

Anal fin spines plus rays 14.3±0.4 14.8±0.6

Supranumerary anal fin pterygiophores 1.5±0.5 1.9±0.4

Anal fin rays 7.4±0.6 7.8±0.5

E0 scales 24.0±0.5 24.4±0.6



TABLE 18. Significantly different values in characters in A. kaaygua and A. guarani (P < 0.05; decreasing order of sig-


Among the A. facetus-like species, this species is most similar to the more slender bodied A. minuano (inhaving a small terminal mouth, 6 C1 gill rakers, 12 pectoral fin rays). Distinguished from A. facetus, A. cf.facetus and A. guarani by a different head shape, and a smaller, terminal mouth (vs. larger, upturned mouth inA. facetus and A. cf. facetus and a larger and subterminal mouth in A. guarani), in having fewer C1 gill rakers(5–6 vs. 7–8), and modally 12 pectoral fin rays (vs. 13). Distinguished from the A. facetus-like species in col-oration characters, as explained in the description of A. charrua. Additionally distinguished from all, exceptA. guarani, by only three abdominal bars (vs. four), from all except A. cf. facetus by 7 anal spines (vs. 6), andfrom all except A. facetus in being more deep-bodied. For all meristic and morphometric differences betweenA. kaaygua and the A. facetus-like species see Tables 16–19.

Distinguished from A. forquilha and A. tembe by having 13 caudal vertebrae (vs. 14 or 15), by a shortercaudal peduncle including 1 or none vertebra (vs. modally 2 or 3), 7 anal fin spines (vs. 6), modally 12 pecto-ral fin rays (vs. 13–14), 24 E0 scales (vs. 26), 3 cheek scale series (vs. 4 to 5) and 5–6 C1 gill rakers (vs. 7–8in A. forquilha and A. tembe).

TABLE 19. Significantly different values in characters in A. kaaygua and A. minuano (P < 0.05; decreasing order of sig-


kaaygua guarani




Cheek scales 3.1±0.3 3.9±0.4




Pectoral fin rays 12.3±0.5 13±0.0

Supranumerary anal fin pterygiophores 1.5±0.5 1.0±0.0

Body depth 49.5±1.2 48.1± 1.6

kaaygua minuano

Body depth 49.5±1.2 46.9±1.2

Anal fin rays 7.4±0.6 8.4¡À0.5

Cheek scales 3.1±0.3 4.0±0.0



Head length 33.3±1.5 35.9±1.1

L2 scales 8.0±0.8 8.8±0.4



Australoheros minuano, sp. nov.(Fig. 15)

Australoheros sp. ”Uruguai” (Říčan & Kullander, 2006).

Holotype. MCP 12710 C, 70.8 mm SL, Brazil, Rio Grande do Sul, Arroyo Canoin, Rio Uruguai drainage,road from Pirapo to São Nicolau. 2 November 1988. C. Lucena, L. Bergmann, E. Pereira, and P. Azevedo.

Paratypes. 85 specimens, all from Brazil, Rio Grande do Sul: MCP 12710 A-I, 7, 34.7–83.9 mm SL, col-lected with the holotype; MCP 12724, 42, Brazil, Rio Grande do Sul, Arroyo Paso do Alto, Rio Uruguaidrainage, 5 November 1988. C. Lucena, L. Bergmann, E. Pereira, P. Azevedo and A. Ramírez; MCP 11227,24, Brazil, Rio Grande do Sul, Arroyo Garupa, divisa Quarai / Alegrete, Rio Uruguai drainage, 12 November1988. C. Lucena, L. Bergmann, and P. Azevedo; MCP 11216, 12, Brazil, Rio Grande do Sul, Arroyo Quarai-Mirim, estrada Quarai-Alegrete, Rio Uruguai drainage, 12 November 1988.

Diagnosis. A relatively elongated species, reaching more than 120 mm TL. Most similar to A. facetus, A.cf. facetus, A. kaaygua and A. guarani. Distinguished from all species except A. kaaygua in having a modalcount of 12 pectoral fin rays. Distinguished from A. facetus, A. cf. facetus and A. guarani by having modally 6C1 gill rakers. Also distinguished from A. facetus and A. cf. facetus in not having an upwards directed mouthand a longer dorsal fin scale cover, covering bases of 7–8 last spines (vs. only 2–3) (Fig. 9). Similar to A.kaaygua in having yellowish ground color, but without the red corners of caudal fin typical for A. kaaygua,and also distinguished by being much less deep-bodied. Distinguished from all Australoheros species by pinkto red dominant males.


Mouth is isognathous, somewhat downwards directed. The outline of the head when mouth closed dis-tinctly rounded (see Fig. 12).

Scales in E0 row 24(7), 25(1). Upper lateral line scales 16(5), 17(2). Lower lateral line scales 8(2), 9(6).Scales between upper lateral line and dorsal fin 3 large and one small anteriorly, two large and one small pos-

teriorly up to the 12th spine, than one large and one small posteriorly. Scales between lateral lines 2. Circumpe-duncular scales 16 (7 dorsally + lateral line scale + 7 ventrally + lateral line scale). Cheek scale rows 4 (3).Lower lateral line continued on caudal fin by 1 or 2 scales.

Dorsal fin with one basal scale row appearing from about tenth spine; interradial scales appear from mem-

brane between 16th spine and the first branched ray, along middle of soft portion running partially in doublerows. Two last interradial membranes without scales; i.e. 7–8 membranes with interradial scales. Anal fin withone basal scale row; interradial scales in single rows, from posterior of sixth spine, lacking on two last interra-dial membranes. Dorsal fin with one basal scale row appearing from about ninth spine; interradial scalesappear from fourteenth spine membrane, running in single rows. Two last interradial membranes withoutscales. Anal fin with one basal scale row; interradial scales in single rows, from penultimate or seventh spine,lacking on two last interradial membranes.

D. XVI,9 (1), XVI,10 (7). A. VI,8 (2), VI,9 (4), VII,8 (3). Anal fin pterygiophores 12(2), 13(7). One (7) ortwo (2) pterygiophores anterior of the first haemal spine. Pelvic fin base below pectoral fin base; first raylongest, extending to the first anal fin spine. Pectoral fin with a rounded tip, extending to about first analspine. P. 12(6), 13(1). Caudal fin rounded.


Lower pharyngeal tooth plate not studied. Gill rakers externally on first gill arch, 2 epibranchial, 1 in angle, 5(1), 6(6), 7(1) ceratobranchial.



FIGURE 15. Australoheros minuano. Holotype, MCP 12710 C, 70.8 mm SL, Arroyo Canoin, Rio Uruguai drainage,Brazil.

Vertebrae 13+13=26(9). First caudal vertebra located between 14th and 15th anal spines. Caudal pedunclecontaining -1(2), -0.5(1), 0(3), 0.5(2), 1(1) vertebrae.

Color pattern in alcohol. Australoheros minuano is most similar to the A. scitulus group, from which it isdistinguished by the following color pattern characters.

1.The midlateral blotch is much smaller in all developmental stages, never reaching above the dorsal halfof the E2 row scales (vs. midlateral blotch optically much larger, spans from the E0 scales to the E3 scale rowscales or at least into the dorsal half of the E2 row scales). When looking at the fish, in the case of A. charrua,the midlateral blotch clearly projects dorsally from the borders of the midlateral stripe, whereas it stays withinthe limits of the stripe in A. minuano. It is very easy to distinguish even juveniles, especially combined withpoints 2 and 3.

2.Develops a distinct rounded caudal base spot located above the lateral line (vs. no clearly defined caudalbase spot, at most a more pigmented narrow bar at the caudal base, which is centered along the peduncle axisand does not reach the borders of the peduncle).

3.The midlateral stripe follows the axis of the body and caudal peduncle. It runs in the E0 and E1 scalerows anterior of the midlateral stripe and in scale rows E0, E1 and E2 posterior to it, i.e. the midlateral stripegets wider posterior of the midlateral blotch (vs. a midlateral stripe also running in scale rows E0 and E1 ante-riorly to the midlateral blotch, but posteriorly from the midlateral blotch turning upwards and being of thesame width - i.e. the midlateral blotch is centered in the E1 scale row, while the next posterior blotch is cen-tered in the E2 scale row and the blotch in the last body bar is centered in the E3 scale row. This successiveshift in the position of the body bars one scale row up per blotch makes the midlateral stripe look like turningup dorsally posteriad from the midlateral stripe.

4.The midlateral stripe is of the same intensity all the way along the body and peduncle vs. the midlateralstripe is distinct mostly only in the area anterior from the midlateral blotch, posteriorly of it is much fainter,decomposed into the blotches as described in point 3 above.

5.Four abdominal bars (vs. three very indistinct abdominal bars). Color in life. The life coloration and breeding in captivity of A. minuano were described by Litz et al.

(2006; referred to as Cichlasoma sp. “Tacuarembó”). The coloration is similar to A. kaaygua, the ground coloris also yellowish to orange, dominant males are pink or reddish, unique among Australoheros. Breeding color-



ation is typical for Australoheros (Říčan & Kullander, 2006) as described for A. kaaygua (see above). Fishattributable to A. minuano have also been pictured by Staeck (2003: p. 63 upper right) and are usually referredto as ´Cichlasoma´ sp. ”Salto”.

Distribution. A. minuano is distributed in tributaries of the Middle and Lower Rio Uruguay in Brazil (RioGrande do Sul) and Uruguay (Fig. 14). Whether it also occurs in Argentinian tributaries is unknown. In thesouthern half of Uruguay and in the Atlantic drainages of Uruguay it is replaced by A. facetus. It is sympatricin distribution with A. scitulus in the northern half of its distribution area and with A. charrua. Litz et al.(2006) give a description of a locality in the Departamento Tacuarembó, Uruguay.

Etymology. The species is named after the native Minuano people, who formerly lived in the area of theBrazilian state of Rio Grande do Sul.

Notes. The difference from A. facetus, A. cf. facetus and A. guarani in coloration is subtle, with A. minu-ano being more similar to the A. scitulus group in having the midlateral stripe running also in the E2 scale rowposteriorly from the midlateral blotch, unlike the other similar species. The midlateral stripe is also more dis-tinctly developed. The mouth is slightly pointing down as in A. guarani, but unlike in A. facetus and A. cf.facetus, which have a slightly upturned mouth. Unlike A. guarani, the mouth of A. minuano is less sharp andmore rounded and the jaws also appear proportionally shorter (see Fig. 12).

Additionally distinguished from A. facetus and A. guarani by a larger orbit and from A. cf. facetus and A.guarani by a smaller head width. Additionally distinguished from A. facetus and A. cf. facetus by an interme-diate body depth and from A. cf. facetus also by fewer caudal peduncle vertebrae, 6 anal spines, and more analand dorsal rays.

Additionally distinguished from A. guarani by a longer head with a shorter preorbital and interorbital dis-tance and more L2 scales (refer to Tables 4, 5, 20 for exact values), and in the position of the caudal pedunclebar, which is in A. guarani in the posterior half of the caudal peduncle running through the last two perforatedscales of the lower lateral line. In A. minuano the bar runs through the central portion of the caudal peduncleand is bent anteriorad, so that at least the last scale of the lower lateral line lies outside the bar. Also distin-guished from A. guarani in featuring divisions of the abdominal bars in adult specimens (4 incompletely sep-arated abdominal bars in 7 of the 9 specimens), in having more scales between the anterior insertion of thedorsal fin and the upper lateral line (3 ½ vs. 2 ½).

FIGURE 16. Australoheros guarani. Holotype, MHNG 2237.58, 129.1 mm SL, Río Guyrau-gua, Río Paraná drainage,Paraguay.



Despite being superficially most similar to the A. facetus-like species (i.e. A. facetus, A. guarani and A.minuano), it has some character states resembling the A. scitulus group (A. scitulus, A. charrua and A.kaaygua), especially in color pattern (midlateral stripe running also in the E2 scale row posteriorly from themidlateral blotch), head shape, in having only 6 C1 gill rakers, and also by the more numerous scales betweenthe upper lateral line and the dorsal fin. Among the A. scitulus group most similar to A. kaaygua in having 13caudal vertebrae and 12 pectoral fin rays. Distinguished from the A. scitulus group by having 13 caudal verte-brae (vs. 14; except A. kaaygua), 6 anal fin spines (vs. 7 or more). Additionally distinguished from A. charruaand A. scitulus by modally 24 E0 scales (vs. 25), and from A. scitulus by 16 dorsal spines (vs. 17). Some col-oration characters still distinguish A. minuano from the A. scitulus group (see Color pattern in alcohol) .

Distinguished from A. forquilha and A. tembe by lower meristics, especially caudal vertebrae (13 vs. 14 ormore), E0 scale row count (24 vs. 25 or more), C1 gill rakers (6 vs. 7 to 8), pectoral fin rays (12 vs. 13).

Additionally distinguished from A. forquilha and A. tembe in coloration characters by having a well cir-cumscribed midlateral stripe and a prominent caudal fin spot. A. minuano specimens also have in a muchhigher percentage four well abdominal bars (5 of 9 specimens—two more with not completely divided poste-rior bar).

Australoheros guarani, sp. nov.(Fig. 16)

Australoheros sp. ”Paraguay” (Říčan & Kullander, 2006).

Holotype. MHNG 2237.58, 129.1 mm SL, Paraguay, Caaguazú, Río Guyrau-gua, Río Paraná drainage, km197 on road from Asunción to Stroessner, 12–14. 4. 1985.

Paratypes. 12 specimens, 29.3–100.1 mm SL, all from Paraguay, Río Paraná drainage: NRM 42215, 2,Caaguazú, arroyo crossing at about km 8 on road Caaguazú—Yhú (25°23'15" S, 56°0'35" W). Åhlander et al,20. 3. 1998; NRM 33498, 2, Itapúa, Arroyo Tembey, 4 km below the falls, Mahnert et al., 1. 11. 1982; MHNG2582.98, 1, Itapúa, Arroyo Tembey, 4 km below the falls, Mahnert et al., 1. 11. 1982. MHNG 2237.56, 7,29.3–100.1 mm SL, collected with the holotype.

Diagnosis. A species with unclear affinities among Australoheros species, with the largest scales betweenthe dorsal fin and upper lateral line (only 2 ½ scales vs. 3 ½ or more anteriorly, only one large scale posteri-orly). It is most similar to A. facetus, A. minuano and A. cf. facetus, and in coloration patterns also to A.forquilha and A. tembe, all being species with a small and inconspicuous midlateral blotch and a straight mid-lateral stripe (if present), which runs all along the flank in the limits of E0 and E1 scale rows. It can be clearlyseparated from A. forquilha and A. tembe in lower meristics, such as 13 caudal vertebrae (vs. 14 or more), ashorter caudal peduncle, shorter scale cover of the dorsal fin, larger (and fewer) scales and a narrower interor-bital space.


Scales ctenoid. Scales on head and chest not distinctly smaller than on flanks. Scales in E0 row 24(7).Upper lateral line scales 16(2), 17(4), 18(1). Lower lateral line scales 6(1), 7(2), 8(2), 9(1). Scales betweenupper lateral line and dorsal fin 3 anteriorly (or 2 large and 1 smaller), 1 large and one small posteriorly.Scales between lateral lines 2. Circumpeduncular scales 16 (7 dorsally + lateral line scale + 7 ventrally + lat-eral line scale). Cheek scale rows 3 (1), 4(6). Lower lateral line continued on caudal fin by 1 or 2 scales.

Dorsal fin with one basal scale row appearing from about tenth spine; interradial scales appear from mem-brane between 16th spine and the first branched ray, along middle of soft portion running partially in doublerows. Two last interradial membranes without scales; i.e. 7–8 membranes with interradial scales. Anal fin withone basal scale row; interradial scales in single rows, from posterior of sixth spine, lacking on two last interra-dial membranes.



TABLE 20. Significantly different values in characters in A. guarani and A. minuano (P < 0.05; decreasing order of sig-


Soft dorsal fin pointed, extending slightly beyond middle of caudal fin. D. XV,10 (1), XVI,9 (1), XVI,10(3), XVII,8 (1). Soft anal fin pointed, of about the same length as dorsal fin. A. VI,8 (5), VII,7 (1). Anal finpterygiophores 12(5), 13(1). Only one pterygiophor anterior of the first haemal spine. Pelvic fin base belowpectoral fin base; first ray longest, extending to first or second anal spine. Pectoral fin with a rounded tip,extending to the first anal spine only. P. 13(7). Caudal fin rounded.


Lower pharyngeal tooth plate in a dissected specimen about one quarter wider than long (length 76% ofwidth) and relatively robust (Fig. 17). Dentigerous area wider than long. 7 teeth along midline, 23 teeth alongposterior margin. Posterior teeth tend to be progressively more compressed, except for medial teeth. Largerteeth medially and posteriorly, gradually smaller anteriad and laterad. Posterior teeth with forwards curvedposterior cusp and subapical anterior shelf. Large laterally compressed teeth with a second cusp raising anteri-orly from shelf.

Gill rakers externally on first gill arch, 2 epibranchial, 1 in angle, 7(6) ceratobranchial.

Vertebrae 13+13=26(6). First caudal vertebra located between 14th and 15th dorsal spines. Caudal pedunclecontaining only the last halfcentrum (3) or only a half of it (3).

Color pattern in alcohol. All specimens examined show only three abdominal bars without any signs ofdivisions into four bars.

Color in life. Life colours are unknown. Breeding coloration is also unknown.Distribution. Australoheros guarani is so far the only Australoheros species known from Paraguay, where

it is sparsely distributed in the tributaries of the High Río Paraná (Fig. 1). Etymology. The species is named after the indigenous Guaraní people. Notes. Australoheros guarani is most similar to the Argentinian form of A. facetus, much more than to A.

cf. facetus. Except for the larger scales between the anterior end of dorsal fin and upper lateral line, it can bedistinguished from both these species by a longer dorsal fin scale cover covering the bases of more than sevenlast dorsal spines (vs. 2–3), by a terminal, slightly down-turned mouth. From both these species as well asfrom A. minuano, it is best distinguished by a larger preorbital and interorbital distance and in being slightlymore deep bodied and having a shorter head. From A. cf. facetus it also differs in having 6 vs 7 (8) anal finspines (refer to Tables 4, 7 and 20 for exact values).

From both A. facetus and A. cf. facetus , and also A. minuano, it can be distinguished by having only threeabdominal bars in all adult specimens and by a different head shape (see paragraph above).

Additionally distinguished from A. facetus in fewer C1 gill rakers and L2 scales (refer to Table 4 for exactvalues).

guarani minuano


Head length 32.4±0.7 35.9±1.1




Pectoral fin rays 13±0.0 12.1±0.4


L2 scales 7.5±1.0 8.8±0.4



It can additionally be distinguished from A. minuano by a larger head width and a smaller orbit, more C1gill rakers, pectoral fin rays, and L2 scales (refer to Table 20 for exact values) and also in the position of thecaudal peduncle bar, which in A. guarani is in the posterior half of the caudal peduncle running through thelast two perforated scales of the lower lateral line. In A. minuano, the bar runs through the central portion ofthe caudal peduncle and is bent anteriorad, so that at least the last scale of the lower lateral line lies outside thebar. The midlateral bar is developed as two pigmented lines running through the central portions of the 0 andE1 scale rows. The posterior portion of the stripe is different from the situation in A. minuano, where posteri-orly from the midlateral blotch the stripe is also distinctly developed in the E2 scale row (i.e. a pattern moresimilar to the A. scitulus group; the pattern of the posterior part of the stripe as observed in A. minuano can beviewed as an intermediate stage between A. guarani and the A. scitulus group). It can also be distinguishedfrom A. minuano in the head shape, which resembles more A. facetus in being sharper, but the mouth isinclined downward like in A. minuano.

Distinguished from the A. scitulus group in lacking the apomorphic color pattern (see description of A.charrua), by more C1 gill rakers, in head and mouth shape (much larger mouth, differently positioned, espe-cially compared to A. charrua). In addition distinguished from A. kaaygua by modally 13 pectoral fin rays (vs.12) and from A. scitulus and A. charrua by 13 caudal vertebrae (vs. 14), fewer anal fin spines (6 vs. 7 ormore), by larger scales (fewer scales in both the E1 scale row and between the upper lateral line and the dorsalfin) and additionally from A. scitulus by lacking the characteristic spotted pattern of that species.

Systematics and character diversity

Říčan and Kullander (2006) presented a morphological phylogeny of the three described and seven putativespecies. In the meantime, Casciotta et al. (2006) have described Australoheros kaaygua, a new species synon-ymous with A. sp. “Jacutinga” of Říčan and Kullander (2006). Říčan and Kullander (op. cit.) have found thattheir morphological phylogeny is incongruent with a phylogeny based on the cytb gene. We further explorethis incongruence. When the morphological matrix is forced to the cytb topology (Říčan & Kullander 2006),the resulting tree (N=1) is 6 steps longer (61 vs. 55; CI 0.60; RI 0.63 vs. CI 0.67; RI 0.72) but the two topolo-gies are not significantly different (Templeton test P=0.1336; Kishino-Hasegawa test P=0.1370; except for thecompare-2 test P=0.01). Since forcing the cyt b data set to the morphological topology produces a signifi-cantly longer topology and vice versa does not, the cytb topology has to be preferred. This result influencesthe interpretation of evolution and synapomorphy status of several characters. Two of the most important areshown in Figs. 18 and 19, where also the mapping of other characters can be observed.

The number of abdominal bars shows variation among the species of Australoheros, which is informativeat several levels. According to the morphological phylogeny (Říčan & Kullander, 2006) Australoheros is sup-ported by having four abdominal bars, at least in late stages on ontogeny and in a proportion of adults (charac-ter 13, state 1; Fig. 18A) and the situation of three abdominal bars in all developmental stages and all adults inthe A. scitulus group is explained as a secondary reversal to the ancestral situation (state 0; Fig. 18). The cytbphylogeny (op. cit.) and this constrained topology applied to the morphological matrix (Fig. 18B) places state1 one level down as a synapomorphy of part of Australoheros excluding A. scitulus and A. charrua. Reversalto the ancestral situation according to this topology occurred in A. kaaygua. Both topologies imply fivechanges to the character.



FIGURE 17. Lower pharyngeal jaw of a dissected specimen of Australoheros guarani, NRM 33498, in (top to bottom)occlusal, caudal and left lateral aspect.



Australoheros is also diagnosed by having the lowest numbers of caudal vertebrae among heroine cichlids(Říčan & Kullander, 2006). The alternative topology (Fig. 19B) does not change this result and both the MPas well as the alternative topologies imply three changes to the character (Fig. 19). Another diagnostic charac-ter of Australoheros, the lowest number of E0 scales among heroines, also has three changes in both topolo-gies (char 6; Figs. 18, 19). Variation in diagnostic characters of Australoheros is thus equally parsimoniouslydistributed on both the MP as well as on the constrained cladograms.

Characters found less parsimoniously distributed are those grouping A. scitulus / A. charrua with A.kaaygua, which is strongly rejected by the cytb data (Říčan & Kullander 2006). These characters are mainlycolor pattern characters, but also others (see descriptions). For some of these characters we do not know theoutgroup state, but the alternative topology implies that these aspects of the morphology of A. scitulus / A.charrua are ancestral states rather than secondary reversals.

The crucial point in deciphering the relationships between Australoheros species is that we do not knowits sister group. According to Říčan and Kullander (2006) and Říčan (2005), Australoheros is either sistergroup to a large clade of Mesoamerican heroines or is the sister group of all other Middle American heroinesplus their South American and Antillean sister groups, which gives a too wide range of possibilities for thechoice of outgroup. Říčan and Kullander (2006) have used Heroina as outgroup in their analyses, with somecharacters scored as unknown in the outgoup. Mapping characters onto the constrained topology (Figs. 18B,19B) produces an outgroup with a different ancestral character state suite than that used by Říčan and Kul-lander (2006) based on Heroina. According to the enforced cytb topology, the morphological character statesof the ancestral Australoheros were as follows (see Table 21):

0. 14 caudal vertebrae [1]; 1. very short caudal peduncle including less than 1 vertebral centrum [0]; 2.11–12 pterygiophores with 1–2 pterygiophore anteriorly from the first haemal spines [0]; 3. 11 dorsal finpterygiophores [1]; 4. VI anal fin spines and 7–9 anal fin rays [1]; 5. 13 or more pectoral fin rays [0]; 6. 25 E0scales [1]; 7. An intermediate long scale cover of the base of the dorsal fin, covering the bases of the middleportion of the hard part of the dorsal fin [1]; 8. 7 or more C1 gill rakers [1]; 9. Scale pattern along the anteriordorsal fin border with scale rows terminating with 1 small scale [0]; 10. 2 large 1 small or more scale rowsbetween posterior end of upper lateral line and the dorsal fin [0]; 11. 4 [1] scale rows between anterior inser-tion of the dorsal fin and the upper lateral line; 12. narrow interorbital distance [2]; 13. only three abdominalbars in all developmental steps and also in adults [0]; 16. Large and dominant and well circumscribed midlat-eral blotch in juveniles and adults [0]; 18. Midlateral stripe posterior from the midlateral blotch running inscale rows E0, E1 and E2 posterior to the midlateral blotch—i.e. the midlateral stripe gets wider posterior ofthe midlateral blotch [1]; 19. Midlateral stripe: clearly bordered [1]; 20. Spots in scales arranged into stripesalso ventral from the 0 scale row (at least into one): no [1];

Phylogenetic analysis rooted with this hypothetical outgroup results in two different trees, whose consen-sus includes A. kaaygua with A. scitulus and A. charrua, and A. minuano with A. facetus and A. cf. facetus. Inthese two clades it is more similar to the original MP tree (Figs. 18A, 19A). On the other hand, the consensustree also includes a clade containing A. tembe and A. forquilha with A. sp. “Jacui”, which is identical to theenforced tree (Figs. 18B, 19B). The tree length is 55, as in the original MP analysis. The most conservativeestimate for the moment is thus that Australoheros includes the following clades: A. scitulus and A. charrua;A. tembe, A. forquilha and A. sp. “Jacui”; A. minuano and A. kaaygua; A. guarani and A. facetus s. lat.



Variation in squamation patterns among Australoheros species is shown in Figs. 8 and 9. This variation isexpressed in characters 6, 7, 9, 10 and 11. The long scale-cover of the base of dorsal fin (character 7, state 0;Figs. 18B, 19B) is a synapomorphy of A. tembe, A. forquilha and A. sp. “Jacui” (Fig. 9A–C). The very short,almost non-existing scale cover, is a synapomorphy of A. facetus s. lat (state 2; Fig. 9I). The unique scalecover of the base of dorsal fin with two cover scales corresponding to one transverse scale row (character 9,state 1) is a synapomorphy of A. forquilha and A. sp. “Jacui” (Figs. 18B, 19B, 9A,B). A. tembe is in this char-acter intermediate. Large body scales, expressed as very few scale rows (basically only one scale) betweenposterior end of upper lateral line and the dorsal fin (character 10, state 2; Figs. 18B, 19B, 9D, G–I), are a syn-apomorphy of A. kaaygua, A. minuano, A. guarani and A. facetus s. lat. These species (and A. sp. “Jacui”) alsohave the lowest counts of scales in E0 row (only 24; character 6, state 2). On the other hand, scale counts can-not be attributed solely to scale size, as demonstrated by the number of scale rows between the anterior inser-tion of the dorsal fin and the upper lateral line (character 11). A. sp. “Jacui”, among the species with just onescale row between posterior end of upper lateral line and the dorsal fin, is in this character among the specieswith the highest number of scale rows. This character state (character 11, state 0) is a synapomorphy of A.tembe, A. forquilha and A. sp. “Jacui” (Fig. 9A–C). The likely source of variation is either different size ofscales in different parts of body or interspecific changes in the position of the upper lateral line.

Differences in head and mouth shapes among the species are also significant (Fig. 12). The most differentis A. tembe, showing very thick lips in combination with a very long head. A. kaaygua, A. charrua and A. scit-ulus can be separated by a very small mouth, making the head look very sharp. These features are synapomor-phic for A. scitulus / A. charrua (Figs 18, 19; not included in the character matrix). A forquilha and A. sp.“Jacui” have similar heads, with the mouth slightly inclined downwards (as in A. guarani and A. minuano),but the lower lip is much thicker than the upper lip (which is to a large extent hidden beneath the lachrymalbone and thick skin above the upper jaw) and the lower jaw is also shorter than the upper. These features aresynapomorphic for the two species (character 14, state 1). A. facetus s. lat. is the only species which has anupturned mouth, with the lower jaw projecting in front of the upper.

Phylogenetic hypotheses support the recognition of four species groups among the described species ofAustraloheros (see Results). The facetus group includes A. facetus, A. cf. facetus and A. guarani. Theforquilha group includes A. forquilha, A. sp. “Jacui” and A. tembe. The scitulus group is formed by A. scitulusand A. charrua. The kaaygua group includes A. kaaygua and A. minuano. The forquilha group is confined toUpper Río Uruguay (Santa Catarina and Rio Grande do Sul) and the Argentinian affluents of the Río Paraná(Misiones), and is probably also found in the coastal drainages of Rio Grande do Sul (A. sp. “Jacui”). Thefacetus group is found in the Río Paraná drainages in Paraguay (A. guaraní) and in the La Plata basin ofArgentina and Uruguay (A. facetus). The group is probably connected through the Río Paraná rather thanthrough Río Uruguay, from which the similar but distinct A. minuano is described. The kaaygua group isfound in Upper and Middle Río Uruguay drainages in Brazil and Argentina (Río Iguazú), while the scitulusgroup is found in the Middle and Lower Río Uruguay drainages in Brazil, Argentina and Uruguay. Since out-group character states are to a large extent unknown, we discuss the individual species groups as diagnosedwith combinations of character states.



18A

18B



FIGURE 18. Relationships among species of Australoheros (see Říčan and Kullander, 2006) with optimization of thenumber of abdominal bars. Drawings show juveniles with developing abdominal bars. A. Maximum parsimony topol-ogy, where state 1 is a synapomorphy of Australoheros, shown as the division of the abdominal bar in front of the midlat-eral blotch to produce two adult bars. The three abdominal bars in the A. kaaygua – A. charrua – A. scitulus group areoptimized as a character reversal, and these three species do show only three abdominal bars also in juvenile stages(character state 0, indistinguishable from the ancestral condition). B. Enforced topology produced by forcing the mor-phological data set to produce a tree in agreement with the cytb tree of Říčan and Kullander (2006). C. Photos show adultspecimens demonstrating character states 2–4, describing the variation in the character (partial tree dissected from theunconstrained MP topology in Fig. 198). 13–2 A. minuano, 13–3 A. facetus, 13–4 A. cf. facetus.

18C



FIGURE 19. Relationships among species of Australoheros (from Říčan and Kullander, 2006) with optimization of thenumber of caudal vertebrae. A. Maximum parsimony topology. B. Constrained topology produced by forcing the mor-phological data set to produce a tree in agreement with the cytb tree of Říčan and Kullander (2006).

19A

19B



FIGURE 20. Combined distributions of Australoheros and Crenicichla in the Rio Uruguay drainage. Two shaded areasrepresent the whole known distribution areas of Crenicichla species marked with 1 (C. celidochilus) and 2 (C. jurubi, C.igara and C. prenda).

The forquilha group (A. forquilha, A. sp. “Jacui”, A. tembe). The forquilha group is diagnosed by: 1) along scale cover of the dorsal fin; 2) highest number of scale rows between anterior insertion of dorsal fin andupper lateral line; 3) a long caudal peduncle including modally more than two vertebral centra; 4) modallymore than 25 E0 scales (except A. sp. “Jacui”) and 14 caudal vertebrae (except A. sp. “Jacui”); 5) 7–8 gill rak-ers on the first ceratobranchial; 6) midlateral stripe only very weakly developed, diffuse or almost missing; 7)small midlateral blotch; 8) no caudal fin base spot. In addition A. forquilha and A. sp. “Jacui” uniquely sharethe specific dorsal fin scale cover where two small scales correspond to one large scale in the anterior portionof the scale cover and the characteristic head and mouth shape.

The scitulus group (A. scitulus, A. charrua). Color pattern characters strongly support inclusion of A.kaaygua into the A. scitulus group, while cytb phylogeny (Říčan & Kullander 2006; see Figs. 18B, 19B herein) places it as the sister group of A. minuano, which is the next most similar species in coloration patterns tothe A. scitulus group.



Diagnosed by: 1) no clearly developed caudal base spot, mostly only a more pigmented narrow bar at thecaudal base, which is fused with the caudal peduncle bar; 2) midlateral stripe well developed and sharplydemarcated, more distinctly developed anterior from the midlateral blotch than posterior from it in adult spec-imens; 3) midlateral stripe turns upwards posterior from the midlateral spot, whereas it follows the body axisin all the other species (except A. minuano). This bending is most pronounced in A. charrua where the midlat-eral blotch is centered in the E1 scale row, while the next posterior blotch is centered in the E2 scale row andthe blotch in the last body bar is centered in the E3 scale row. In A. kaaygua and A. scitulus the blotch poste-rior to the midlateral blotch is centered in the same scale row as the midlateral blotch (i.e. E1 scale row), andonly the last blotch is very high on the body. However, all three share the feature that the midlateral stripe doesnot continue in the E0 scale row posterior to the midlateral blotch (which is also shared with A. minuano); (4)midlateral blotch is very large, it spans from the E0 scales to the E3 scale row scales or at least into the dorsalhalf of the E2 row scales; 5) only three abdominal bars and there are also no traces of four developing bars indevelopmental series; 6) 6 gill rakers on the first ceratobranchial; 7) 14 caudal vertebrae. 8) modally morethan 25 E0 scales.

The kaaygua group (A. kaaygua, A. minuano). A. kaaygua is very similar to the scitulus group in colorpattern characters (see above), but more similar to A. minuano in meristic characters (few pectoral fin rays,few E0 scales, few caudal vertebrae, few scales between upper lateral line and dorsal fin).

Diagnosed by: 1) modally only 12 pectoral fin rays, which is unique among heroines (pers. obs.), 2) mod-ally 13 caudal vertebrae; 3) 6 gill rakers on the first ceratobranchial; 4) modally only 24 E0 scales, 5) only onelarge and one small scale between the end of the upper lateral line and the dorsal fin.

The facetus group (A. facetus, A. cf. facetus, A. guarani). Diagnosed by: 1) well developed caudal finbase spot (also in A. minuano); 2) four abdominal bars in more than 80% of specimens (except A. guarani,where four bars mostly only in juveniles). (3) small midlateral blotch; 4) midlateral stripe weakly developed,running all the way along body in E1 scale row; 5) 7–8 gill rakers on the first ceratobranchial; 6) 13 caudalvertebrae; 7) modally only 24 E0 scales; 8) only one large one small scale between the end of the upper lateralline and the dorsal fin; 9) only 3 (to 4) scales between the anterior origin of the dorsal fin and the upper lateralline.

TABLE 21. Alternative outgroup character states produced by mapping morphological characters onto the enforced phy-logenetic hypothesis (Figs. 18B, 19B) derived from analysis of the cytb gene (Říčan & Kullander, 2006; compare with

character matrix in op. cit.).

Discussion

Říčan and Kullander (2006) recently described the genus Australoheros and thus provided a generic allocationfor ´Cichlasoma´ facetum and three recently described species. The type species of Australoheros, Australo-heros facetus, as understood at the onset of our study was a species of quite variable morphology with a sur-prisingly large distribution area. The data that we present here (and in Říčan & Kullander, 2003) and theiranalyses presented by Říčan and Kullander (2006) demonstrate that the large variability is not randomly dis-tributed, but well-clustered. Říčan and Kullander (2003, 2006) thus proposed that Australoheros facetus isbetter understood as a species complex. The limited amount of available sequence data (Říčan & Kullander,2006) agrees well with this view. In this paper we have described four new species of Australoheros, raising

character 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

state 1 0 0 1 1 0 1 1 1 0 0 1 2 0 0 ? 0 ? 1 1 1



the number of species to eight. As presented by Říčan and Kullander (2006), closely related species of Australoheros are allopatric, while

species groups are sympatric. This is the general pattern among South American fishes, where biogeographi-cal patterns are discernible in species relationships, while genera are mostly broadly overlapping in distribu-tion.

Australoheros scitulus is sympatric with A. facetus in Uruguay and the adjacent part of Argentina in drain-ages of the Lower Río Uruguay basin. Australoheros minuano is sympatric with A. charrua in the Braziliantributaries of the Middle Rio Uruguai in the state of Rio Grande do Sul. The type material of both speciescomes from the same locality in the Arroyo Canoin. A. minuano is most probably also sympatric with A. scit-ulus in Uruguay and adjacent part of Argentina in drainages of the Lower Río Uruguay basin. A. kaaygua issympatric with A. forquilha in the tributaries of the Upper Rio Uruguay in Brazil (see Figs. 1, 2, 3, 14). Whilethe relationships between Australoheros species are to some extent complex (see results), no analysis (seeFigs. 18 and 19) recovers any of the three sympatric species-pairs as monophyletic. Sympatric species are thusnot sister species among Australoheros. Speciation has thus obviously been largely allopatric, with secondarysympatry of already differentiated lineages.

The biogeography of Australoheros is very interesting and in great agreement with another cichlid genus,Crenicichla (Lucena & Kullander, 1992). The division of the Río Uruguay by rapids mentioned in the intro-duction, is well reflected in the diversity of its ichthyofauna. Among Australoheros, A. forquilha and A.kaaygua are found in the Upper Uruguay. Australoheros charrua and A. minuano are endemic to the MiddleUruguay and A. scitulus and A. facetus are endemic to the Lower Uruguay and the Río de la Plata. The samepattern is discernible in the Crenicichla species (Lucena & Kullander, 1992). Crenicichla jurubi, C. igara, C.prenda, C. tendybaguassu, C. missioneira and C. minuano are species of the Upper Uruguay. The latter threespecies are also found in the Middle Uruguay. C. gaucho and C. scottii are Middle-Lower Uruguay species.The widespread C. vittata and C. lepidota are also found in the Lower and Middle Río Uruguay, but not inUpper Uruguay. As noted by Bertoletti et al. (1989a–b, 1990) and Lucena and Kullander (1992), past drainagepatterns, specially in the Upper Río Uruguay were probably different. The Upper Río Uruguay is markedlydifferent from the Lower Río Uruguay in fish species composition (Bertoletti et al., op. cit.). This faunal dif-ference between Upper and Middle Río Uruguay is well evident in Australoheros. The forquilha groupincludes two species, which belong to two isolated drainages, the Upper and Middle Río Uruguay and a tribu-tary of the Río Paraná (Arroyo Urugua-í) (Fig. 3). A similar distribution is also found on the intraspecific levelin A. kaaygua (Fig. 14). The forquilha group and A. kaaygua distribution patterns may suggest that once theUpper Río Uruguay and Río Iguazú were connected.

The second interesting biogeographic aspect is the faunal turnover along the Middle Río Uruguay (Fig.20). This fact is again supportive for the independent histories of the upper and middle-lower parts of the RíoUruguay. The same faunal turnover is even better demonstrated for Crenicichla (Lucena and Kullander,1992). The Upper Uruguay is a center of endemism for Crenicichla, with three endemic species (C. igara, C.jurubi and C. prenda) and three additional species distributed in both Upper and Middle Río Uruguay (C. mis-sioneira, C. minuano and C. tendybaguassu; not including C. celidochilus, with a questionable distribution),but not in Lower Uruguay. Widely distributed species of Crenicichla (C. vittata, C. lepidota, C. scottii), on theother hand, ascend only as far as the Middle Uruguay, and overlap there in distribution with the three speciesof the Upper Uruguay Crenicichla. In the area of overlap is the small distribution area of Crenicichla gaucho.In Australoheros, A. forquilha and A. kaaygua are endemic species of the Upper Uruguay, reaching down intoMiddle Uruguay (but see above). Lower Uruguay species (A. scitulus, A. minuano, A. facetus) reach into theMiddle Uruguay (A. scitulus, A. minuano) and A. charrua is found in the zone of overlap between Upper andMiddle-Lower Uruguay Australoheros. The zone of overlap is the same area in both Australoheros andCrenicichla. The barrier prohibiting upstream migration of species from the Middle Uruguay are most proba-bly the Moconá falls.



This interesting biogeography of Australoheros and Crenicichla and the history of the Rio Uruguay prob-ably has its origin in the rise of Sierra de Misiones (Argentina). As this mountain range rose, it might haveseparated an once continuous drainage of the Paraná which probably also included the Upper Uruguay.Details of cichlid distribution and relationships also suggest that the northern portion of Misiones (Río Iguazúand Arroyo Urugua-í; affluents of the High Paraná) and the Upper Río Uruguay were once connected. It isquite possible, that most of the waterfalls forming significant barriers to fish dispersal in Misiones (includingthe famous Iguazú falls) in both the Upper Uruguay and High Paraná originated at the same time and are con-nected to this geological event. Studies further focusing on the phylogenetic relationships and historical bio-geography of freshwater species inhabiting this very interesting area are clearly needed to test this hypothesis.

The biogeography of Australoheros along the Lower Paraná and in most of its western area in Argentina isstill little known, but our results show that Australoheros facetus is most likely the sister group of A. guarani.A. facetus is found only in the coastal drainages of Río la Plata, but not along the Río Uruguay (where the sim-ilar A. minuano is found). A. guarani is found in the drainages of the High Río Paraná. We thus speculate thatthese two species are probably connected through the Río Paraná, rather than through the Río Uruguay. Wehave so far not been able to examine specimens from this area and set the distribution limits of A. facetusalong the Río Paraná.

The Río Uruguay drainages and adjacent areas from which we have been able to examine material is onlya very small fraction of the putative natural distribution area of Australoheros. Australoheros is known tooccur all the way west to the foothills of the Andes as well as to the northeast in the coastal drainages of theBrazilian highlands, in the Upper Paraná and in the São Francisco drainage. These areas are several timeslarger than the studied area and Australoheros is thus among the genera of Heroines with the largest distribu-tion. Australoheros may well prove to be one of the species-richest-genera of heroine cichlids.

Acknowledgements

We wish to thank the museum staff of MCP for enabling us to study specimens in their care. Important sam-ples and material of Australoheros were kindly provided by Stefan Körber and Thomas Litz (Germany), whoshared their invaluable knowledge of Argentinian and Uruguayan fishes and also read and improved thispaper. We would also thank three anonymous reviewers whose comments significantly improved the paper.Financial support was provided by grants to OŘ from the HIGH LAT program for Access to the SwedishMuseum of Natural History within the European Community program ‘‘Improving the Human ResearchPotential and the Socio-Economic Knowledge Base’’, by a scholarship from Stockholm University (Sweden)and by the MSM6007665801 grant (Czech Republic).

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