Zoologica Scripta, Vol. 11, No. 4, pp. 307-313,1982 0300-3256182104030747$03. 0010 Printed in Great Britain Pergamon Press Ltd. The Norwegian Academy of Science and Letters

Cichlid from the La Plata Basin. Part IV. Review of the , with Description of a New Species (Teleostei, Cichlidae)

WEN 0. KULLANDER

Section of Vertebrate , Swedish Museum of Natural History, Stockholm, Sweden

Received 28 September 1981

Kullander, S. 0.1982. fishes from the La Plata basin. Part IV. Review of the Apistogramma species, with description of a new species (Teleostei, Cichlidae).-Zool. Scr. 11: 307-313. Apistogramma inconspicua sp.n. is described from the Rio Candclaria in the Rio GuaporC drainage in Bolivia, and recorded also from Caceres on the Rio Paraguay in Brazil. The new species is closely related to the Paraguayan species A. commbrae (Regan) and an undescribed species in the GuaporC drainage system. These species share a specialized feature in the colour pattern, viz. an expanded vertical bar on the caudal peduncle which it confluent with the caudal spot. The now presumably complete record of Paraguayan Apistogramma species allows an analytical key to be given. Of the additional species, A. borellii (Regan) and A. rrifasciata (Eigenmann & Kennedy) represent two distinct lineages. Apistogramma pleurotaenia (Regan) is probably a La Plata basin species, although no localities are known, and its relationship remain obscure. The assemblage of Paraguayan Apistogramma species reflects the heterogeneity of the Paraguayan cichlid fauna as a whole. Sven 0. Kullander, Section of Vertebrate Zoology, Swedish Museum of Natural History, S-104 05 Stockholm, Sweden.

Introduction This paper features a review, chiefly of the distribution and diagnostic characters, of the Paraguayan Apisto- gramma species. Special attention is called to a species already announced (Kullander 1980a, 1982a), but not yet described. It is of particular interest because of being both parapatric with and obviously closely related to the Paraguayan A. commbrae (Regan), and found in both the GuaporC and Paraguay river systems. The methods employed have been described in Kullander (1980a,b). Museum acronyms used are: AMNH (American Museum of Natural History, New York), ANSP (Academy of Natural Sciences of Philadel- phia), BMNH (British Museum (Nat. Hist.), London), FMNH (Field Museum of Natural History, Chicago), IRSNB (Institut royal des Sciences naturelles de Belgique, Bruxelles), MZUSP (Museu de Zoologia da Universidade de Sio Paulo, SCo Paulo), NRM (Swedish Museum of Natural History, Stockholm), ZFMK (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn), and ZMB (Zoologisches Museum, Berlin, DDR).

Apistogramma inconspicua sp.n. (Figs. 1-2) Heterogramma corumbae Eigenmann & Ward; (part.) Haseman 1911, p. 359 (Sio Luiz de Caceres, Rio Paraguay, CM 2752,2755, now FMNH 54178). Heterogramma taeniatum (Gunther); (part.) Haseman 1911, p. 358 (SBo Luiz de Caceres, Rio Paraguay, CM 2737, now FMNH 54168; brief description). Aprsrogramma sp.; Kullander 1980a, pp. 20, 26, 49 (listed; undescribcd species of the commbrae group from Caceres, based on FMNH 54168,54178).-Kullander 1982a, pp. 36,43,45 (compared with A. commbrae). Material. IRSNB 637, holotype 13, 29.7 mm. IRSNB 638 + NRM A82/3405, 6,28.5 mm and 2P P, 23.5 and 24.4 mm. All from Bolivia, depto. Santa Cruz, R. Paraguay system, small pool of the R. Candelaria, Figs. 1-2.--1. Holotype of Apisfogramma inconspicua, a male, 29.7 mm above bridge on road Carmen-Santa Rosa (16"OO'S 61"40'W). 29 SL.-2. A female paratype of Apistogramma inconspicua, IRSNB 638, October 1977. Leg. J.-P. Gosse (sta. 24). 24.4mm SL.

307 Zoologica Scripta 11 308 S. 0. Kullander

Additional material. Notes, taken in 1975, on three more specimens, from Caceres, indicate that these belong to A. inconspicua. The largest specimen, a male, 37.4 mm SL, differs somewhat from the -material of A. inconspicua in having the lateral band extended to the tail spot. The soft dorsal and anal fins are produced, reaching backwards to about the end of the caudal fin. Two of the FMNH specimens have canals in the lower lateral line (two to four); all have 15 dorsal spines. (FMNH 54168. d,37.4 mm. Brasil, Est. Mato Grosso, R. Paraguaysystem, Caceres, 27 May 1909. Leg. J. D. Haseman. FMNH 54178. 8,28.2mm, P,24.9 mm. Same locality and leg. as preceding. Dates 24 and 27 May 1909.) Etymology. The species epithet is a Latin adjective meaning incon- spicuous. It is suggested by the near-absence of abdominal stripes, a feature distinguishing this species from A. commbrae, in which the abdominal stripes are very prominent. Fig. 3. Profiles of heads of a male (left, holotype) and a female (right, IRSNB 638,24.4 mm SL), to illustrate sex dimorphism in Apistogrumma Diagnosis. A species of the regani group, similar to A. inconspicua. Scale 1 mm. commbrae and to an undescribed species in the Bolivian Amazonas drainage, especially in having a dark tail spot to first anal soft ray in holotype, slightly shorter in the which includes the last vertical bar on the body and the smaller male, in females short, reaching to anal fin origin. caudal spot. The lateral band ends in the sixth vertical bar Caudal fin rounded in both sexes. rather than in the tail spot. Abdominal stripes appear as Teeth 17-21118-23 on one side in outer series of traces or are absent, rather than as conspicuous spots upper1lower jaw; about two short inner series, one in along the absominal sides or short vertical stripes below upper jaw continued a short distance along lateral margin the anterior part of the lateral band. All specimens of jaw. Lower pharyngeal tooth-plate examined in one examined have 15 rather than 16 dorsal spines. The specimen; anterior teeth simple, nearly conical, small, posteromedial teeth on the lower pharyngeal tooth-plate increasing in size posteriorly, base becoming elongate, bear two small projections on the rostral edge. with strong posterior cusp, except near middle in posterior series where the teeth have two very small projections on Description. Figures 1 and 2 illustrate general shape and the rostral edge, and towards edges where the teeth retain colour pattern; morphometric data are summarized in the narrow base, but develop a strong posterior cusp and Table I. The following data are chiefly from the holotype; an inconspicuous anterior elevation. Eight (3) or 10 (1) counts are from all specimens, and frequency is given in rakers on edge of lower pharyngeal tooth-plate. Verte- parenthesis. brae, including the urostylar element, 12 + 12 (4); one The body shape is similar to that of A. commbrae, supraneural. except that males are slightly deeper in appearance. Males with slightly more elevated head and slightly Coloration (alcohol). Ground colour brownish yellow, extended snout compared to females (Fig. 3). Ascending paler ventrally, increasingly brownish dorsally. Markings processes of premaxilla reaching about middle of orbit; dark brown; head stripes, lateral band, and tail spot most angle c. 48". Tip of maxilla exposed, reaching behind prominent. Gill-cover, snout, lips and forehead greyish. anterior margin of orbit; angle 55". Scales in lateral series Cheek brownish yellow. Dark spot on chin adjacent to (squ. long.) 22 (4), squ. caud. 3 (1) or 4 (1). Chest fully lower lip. Suborbital stripe about same width as pupil, scaled, except a naked area opposite tip of cleithrum in straight, to junction of sub- and interopercula. Supra- females; preventral and predorsal scale counts identical, orbital stripe not evident in general brownish colour of 10 (2) or 11 (1). Cheek fully scaled except for rostroventral nape. Moderately wide postorbital stripe, narrower than corner, scales in 2 (females) or 3 (males) series, cycloid pupil. Vertical bars diffuse above lateral band, indicated except dorsocaudally. All other head scales cycloid, by diffuse spots along dorsal fin base; slightly more except dorsal opercular scales; no satisfactory count of evident below lateral band. Lateral band about one scale opercular scales possible, but apparently two oblique wide, extending to just before Bar 6; not spotted, but series; subopercular scales 3 (1) or 4 (l), in one series; slightly intensified where crossing Bar 3. Bar 7, except interopercular scales 2 (1) or 3 (3). Lateral lines with 8-9 dorsal- and ventralmost portions much intensified, con- canal-bearing scales, total 13-15 scales in upper, 24 tinuous with squarish caudal spot which is margined subserial pores; only pore-bearing scales (6-7) in lower above and below by a whitish dot with narrow brownish line, which are continued on caudal fin (a pore, in two anterior and posterior edges. Dorsal pectoral spot dis- cases) or not (in one case). Proximal one-fourth of caudal tinct. Blackish close to ventral spine base laterally. fin scaled, outer scales cycloid. Abdominal stripes only as traces, one from upper, another D. XV.6 (2), XV.6.i (l), XV.7 (holotype). A. 111.6 (4). from lower edge of pectoral axilla, vanishing above anal P. 11 (2), 12 (2). Dorsal fin spines subequal from the sixth; fin origin (a third from below pectoral axilla to above anal lappets short, pointed in males, truncate in females; soft fin base distinguishable in two paratypes). Midventral fin pointed, third ray longest, reaching to about middle of stripe from vent or genital papilla faint and short in males, caudal fin in holotype; in females soft dorsal fin rounded, strong (blackish) and reaching further forwards, but not reaching to one-fourth of caudal fin. Soft anal fin similar to to ventral fin bases, in females (Fig. 4). A weak brownish soft dorsal fin; little beyond middle of caudal fin in spot on chest near ventral fin base in one female (Fig. 4). holotype. Pectoral fin asymmetrical, with rounded tip, Fins smoky. Dorsal fin with anterior two interspinous fourth ray longest, reaching to above genital papilla. membranes black; a dark spot on the base of each Ventral fin pointed, a short filament of the first ray reaches membrane and two or three series of clear spots on the last

Zoologicu Scripta I I Cichlidfishes from the La Plata basin. ZV 309

soft fin membranes. Anal fin with anterior edge and variation in the development of the tail spot, and is very margin dark, especially the anterior edge in females, different from A. inconspicua in other respects, e.g. in otherwise like dorsal fin. Caudal fin with six vertical series lacking a lateral band, but having a lateral spot. It has not of clear, dark-ringed dots across entire fin in males; in been possible to establish the relationships of A. nijsseni, females pale middle portion with about five vertical series but it is certainly distinct from all other Apistogramma of dark spots. Ventral fin with black base and outer edge; species (de Rham & Kullander, in preparation). The black area wider and more intense in females (Fig. 4). other two species, besides A. inconspicua, are A. comm- brae and an unnamed form in the Amazonas drainage Distributron. Known only from the type-locality, in the Bolivian system in Bolivia (see below). Amazonas drainage, and Caceres, in the Paraguay drainage (Fig. 6). Since these three last-mentioned forms also agree well with one another in other respects, I take the tail spot to Discussion indicate close phylogenetic relationship between these Relationships. Four species in the Apistogramma species. On the basis of the other character states, these are characterized by what I am calling a tail spot, i.e. a species must be referred to the regani group (Kullander spot formed by the confluence of the caudal base spot and 1980~).All other regani group species have a prominent the intensified middle portion of the last vertical bar caudal base spot, confined to the base of the caudal fin across the caudal peduncle. Apistogramma nijsseni Kul- (like most other Apistogramma species), but a tail spot is lander, in the Peruvian Amazonas system, presents some not developed or even indicated. The modification of the

Fig. 4. Abdominal and ventral fin sex dimorphism in melanophore pattern in Apistogramma inconspicua. From the hoiotype (male, to the right), and a 24.4 mm paratype (female, to the left). The ventral fin pigmentation of the male is chiefly on the external side. Scale I mm.

Zoologica Scripta 11 310 S. 0.Kullander

Bar 7 pigmentation thus represents an apomorph charac- undescribed Apistogramma species from the upper Rio ter state. Xingu system; all other Apistogramma examined have Apistogramma inconspicua is further characterized by teeth of the “bicuspid” type in the posteriormost series, its pharyngeal dentition, including “tricuspid” teeth on but many species remain to be studied. “Tricuspid” the posteromedial portion of the tooth plate. The other posterior pharyngeal teeth are also found in some species two tail spot forms have only simple teeth or a large of large geophagines (Gosse 1976), and I have noted the posterior cusp rises above an anterior shelf or bulge (cf. condition in non-geophagines. Fig. 9). I have found “tricuspid” teeth also in the three A. I am uncertain whether the “tricuspid” condition is steindachneri group species (Kullander 1982h), and in an derived relative to the “bicuspid” or vice versa. It may even be that tooth shape is easily modified, according to /-\ I- diet, in Apistogramma as in some other (cf. Greenwood 1965, for demonstration of plasticity in pharyngeal structures in an African cichlid species). Both conditions are found in unrelated groups of genera (smaller and larger geopagines, and non-geophagines). I do not think that it is possible to put any phylogenetic significance on the shape of the pharyngeal teeth on the basis of present knowledge only. Hence, there would be no contraindication of the close phylogenetic relationship of A. inconspicua with the other tail spot forms in this character. Distribution.The type-locality is a very small rivcr Fig. S. Occlusal view of the lower pharyngeal tooth plate and dentition in tributary to the Rio San Joaquin in the Rio Guapore Apistogrammu inconspicuu: a posteromedial tooth shown in lateral drainage system. It is not the better known Rio Candelaria view. From IRSNB 638,23.5 mm SL. Scale 1 mm. close to the Brazilian border east of the Serrania de San Lorenzo, where there are also two villages called Carmen and Santa Rosa (common village names in Bolivia). The geographical ranges of A. inconspicua and A. commbrae may overlap to some extent. Available collec- tions from the Paraguay system south of Caceres regularly feature A. commhrae, but not A. inconspicua, so it seems reasonable to assume that A. inconspicua is not present much further downstream in the Paraguay system. This circumstance suggests geographical replacement, but with only two localities for A. inconspicua there is not much ground for a characterization of the range of this species. The presumable northern and/or western range of A. inconspicuu relative to A. commbrue, nevertheless suggests that A. commhrae is a restricted endemic in the Paraguay drainage. The presence of A. inconspicua in two separate river systems must also be stressed, and it seems that it is not only the presence of a hydrological barrier that defines the northward extension of A. commhrae. The recognition of yet another tail spot species in the Guapore system makes it even more difficult to evaluate the distributional data for A. inconspicua. More collecting in the Paraguay and Guapore headwaters are needed before one can make a serious attempt at understanding distribution patterns.

The Paraguayan Apistogramma species It appears reasonable to assume that now all the Paraguayan species of Apistogramma have been described. The number of species recorded for the rather limited region is quite high, and the available collections are extensive in area coverage and represent over 80 years of collecting. Fig. 6. Map of the Paraguay system and nearby parts of adjacent Two of the species, A. horellii (Regan) and A. trifarciata drainages. Localities of Apisfogrummu inconspicuu: circles. Localities (Eigenmann & Kennedy), apparently represent distinct of A. commbme: black dots, except black triangle at Corrientes, representing southernmost locality area from literature records; based phyletic lineages, to judge from their characteristic mor- on data in Kullander (1981). phology and colour pattern (cf. Kullander 1980~).Apis-

Zoologica Scriptu 11 Cichlidfishes from the La Plata basin. IV 311 togrammapleurotaenia (Regan) has an uncertain position A. aequipinnis (ZMB 23409; ? Argentinien, ded. W. in the regani group (Kullander 1982~).Apistogramma Reitzig, 8 July 1938), is a large male, 34.7 mm SL, commbrae is closely related to A. inconspicua, and these apparently slightly dried and with a compressed ventral are taken to represent another species group in the region. region (due to postmortem pressure), giving the specimen The taxonomic diversity observed is analogous with a slightly atypical appearance. It has D. XVI.5 (not XV.5 that found in other Paraguayan cichlids, which form a as given by Ahl), and squ. long. 21 (not 22), otherwise it is taxonomically very diverse local group. So, even if the about as described by Ahl. genus Apistogramma is fairly well represented in the In the ZMB collection are two more males of A. Paraguay system, in contrast to other cichlid genera (with borellii, both aquarium specimens and identified by Ahl: one or two species), it conforms to the general picture. ZMB unreg., c. 36.5 mm SL, no locality, ded. Reitzig, 5 A study of the distribution of the Paraguayan Apisto- August 1938, identified as A. borellii, and ZMB unreg., c. gramma species faces several problems making it difficult 22 mm SL, Para, ded. Schreitmiiller, 19 February 1939. to reach any general conclusions. Uncertainty about the The latter specimen bears a manuscript name, which as range of A. inconspicua has already been indicated. For far as I can determine has not been published. The former A.plrurotaenia there is no precise locality at all available. specimen obviously represents the A. borellii material For the moment it is hardly possible to say whether the referred to by Ah1 (1938). species conform to a single general pattern of distribution The synonymy of A. reitzigi and A. borellii has already or, if not, how many. For instance, it may seem that A. been suggested by Schmettkamp (1978), who used a trifasciata has the same distribution as A. commbrae plus different data set. He also included the aquarium stock of A. inconspicua, but in the GuaporC A. trifasciata is known A. borellii, which is extensively treated in aquaristic and only from near the mainstream, not from far-away ethological literature as A. reitzigi. tributarieq. Ecological factors probably influence the Clear as this synonymy may appear, it should neverthe- dispersion a great deal. Greatest diversity is found to the less be recognized that a critical study of material from the north, and the number of species drops with increasing entire range of this inclusive A. borellii has still to be latitude. This could be due to climate shift from tropical to made. The type-localities of A. borellii (Carandazinho), temperate, but distributional limits are still too uncertain H. ritense (Rio Santa Rita, upper Corixa Grande system), to allow a search for environmental restraints on disper- and H. rondoni (Caceres) are, however, in the same sion, other than physical barriers. region. Those of A. reitzigi and A. aequipinnis can only be The following remarks on the species additional to A. guessed at, although Reitzig’s (1975, 1977) account of the commbrae (redescribed in Kullander 1982a) and A. history of the material suggests an imported stock from inconspicua summarize my findings from a study of the Parana medio region. mostly old material already described by Regan (1906) Apistogramma borellii may be distinguished from the and Haseman (1911). other La Plata basin species of the genus by its colour pattern, with immaculate caudal fin, a caudal base spot Apistogramma borellii (Regan,1906) terminating an interrupted or very uneven lateral band The synonymy of this species should be properly discussed which is strong or discernible only on the posterior sides of in the context of a revision, but since no material available the body, and the absence of any abdominal markings. to me would justify a redescription, and since the recogni- The low vertebral count given by Kullander (1980a, p. 39; tion of a number of probable junior synonyms as nominal 22, from ZFMK unreg.), appears to be due to difficulties species gives a misleading picture of the species richness in in discerning the penultimate vertebra on the indistinct the Paraguay system, the following comments appear radiograph, as suggested by comparison with an alizarin justified. stained aquarium specimen (NRM unreg.). I cannot (1 ) The type series of Heterogramma rondoni Ribeiro recognize any particularly close relationship to any other (1918) was not available for re-examination. The photo- Apistogramma species. graphs of three of the specimens (Ribeiro 1918, pl. XI), The distribution extends along the Paraguay system from the head- however, leave no doubt as to the identity of the species water region, southwards to the area of confluence with the Parana (Itati on the Alto Parana, ZFMK unreg., see also Liiling [1979,1980]; records with A. horellii. Meinken’s (1965) interpretation of from the Rio Corrientes. an affluent of the Parana medio, by Reitzig Ribeiro’s description and figures is grossly inaccurate. [1977], and the Argentinian Chaco near Corrientes, by Mayland [1980], (2) Haseman’s (1911) retouched photograph of the appear reliable although no preserved material exists). The localities on the map (Fig. 7) are based on BMNH 1900.4.14: holotype of his Heterogramma ritense suggests that this is 12-14 (Carandazinho), AMNH 1032, FMNH 54185, FMNH 52621 a spccimcn of A. borellii (cf. also Schmettkamp 1978), (Corumbi), ANSP 5391653922 (Descalvados), FMNH 54184 (Villa and there appears nothing in the description that would Hayes), FMNH 54186 (Puerto Suarez), ZFMK unreg. (Itati), BMNH 1895.1.30:24 (Colonia Risso), BMNH 1935.6.4;455460 (near Asun- justify the recognition of this form as a distinct species. cion), BMNH 1971.2.12;68-69 (Formosa), MZUSP unreg. (Ciceres), (3) Apistogramma aequipinnis Ah1 (1938) and A. MZUSP unreg. (internal lakes of the Piquiri-Itiquira system; approxi- reitzigi Ah1 (protologue in Mitsch [ 19381, formal descrip- mately located), MZUSP 4464pt. (Santo Antonio do Leverger), and tion in Ah1 [1939]), were based on aquarium material of Reitzig (1977; Rio Corrientes). A. borellii. The holotype of A. reitzigi (ZMB 23408; ded. Reitzig, 6 August 1938) is a male, 26.3 rnm., with the Apistogramma trifasciata (Eigenmann & Kennedy, 1903) characteristic facies of A. borellii. It differs somewhat What has been said about A. borellii as regards its from the description given by Ah1 (1939) in having D. synonymy and material available for a redescription, XVI.6 (not XVI.5) and A. 111.6 (not 111..5), and a lateral applies correspondingly to A. trifasciata. Two subspecies, line count of 7 + 6 pored6 pores (not 6/0). The holotype of Heterogramma trifasciatum maciliense Haseman, and A.

Zoologica Scripta 11 312 S. 0.Kullunder trifasciatu haruldschultzi Meinken, have been described (Kullander 1980a, p. 30), and indicates that the La Plata from the Rio GuaporC. I cannot distinguish the Guapo- basin origin is correct, although the vague locality data for rean material, but this inability may reflect the inadequate import material, especially that dating from the early material available. The relationships of the species are not years of the European aquarium hobby, are commonly clear to me. It can be distinguished from the other incorrect. Paraguayan species on the immaculate caudal fin, a conspicuous lateral band ending on the caudal fin base, an Key to the La Plata basin species of Apistogramma oblique dark stripe between the anal fin origin and the The key is based chiefly on colour pattern because it provides the most pectoral axilla. and the fin morphology of the male reliable and convenient characters. despite limitations imposed by faded specimens. Species descriptions should be used as a complement to the (produced anterior dorsal fin lappets, but rounded caudal key. Useful descriptions of A. borellii and A. trifasciarir may be found in fin). Regan (1906) and Haseman (1911), the latter paper also with photos. The only description of A. pleurotaenia is that of Regan (1909). The distribution extcnds along the Rio Paraguay from the headwaters 1. Abdominal stripes absent or very indistinct -7 south at least to Villa Hayes, perhaps to Santa Fe, on the Parand medio - Abdominal stripes prominent A. commbrae in Argentina, which would mark the extreme south of the distribution of 2. An oblique, narrow dark stripe from the anal fin origin forwards to the genus (cf. Bonetto et al. 1970). The Guapore material available is the pectoral axilla A. trifasciata from Sio Antonio dc Guapore, Bastos (cf. Haseman 1911). and - No oblique stripe across abdominal sides 3 Guajari-Mirim (IRSNB 18603). The localities on the map (Fig. 7) are 3. Tail spot present '4. inconspiciia based on FMNH 52616 (Corumba), FMNH 54172, FMNH 54183, - No tail spot 4 MZUSP unreg. (Cdccrcs), FMNH 54173 (Villa Hayes), FMNH 54174 4. Lateral band very uneven or repeatedly interrupted. strongest (Canipos Alegre. Rio Jauru), BMNH 1895.1.30: 5 (Colonia Risso), and posteriorly; three anal spines A. borellii Bonetto efa/. (1970; Santa FC). - Lateral band continuous; four anal spincs A. pleurotaeniu

Apistogramma pleurotaenia (Regan,1909) An undescribed species in the Guapore river system See Kullander (1982~)for comments. The precise distri- bution is unknown, the status and closer relationships The only specimen available of the unnamed species unclear. The reduced lateral line count (8 + 3 pored7 referred to above, was first thought to be an A. inconspicua pores, in the holotype) is a mark of Paraguayan species showing a different colour phase, but it more likely represents a different species, recognized by a commbrae- like pharyngeal dentition (Fig. 9) and wide interorbital

4

ze

"t Fig. 8. An undescribed Api.Ptograrnma species from the Rio Guapore system, IRSNB 19.975, female, 25.6 mm SL.

&

32L 64 68 +

Fig. 7.Map of tbc Paraguay system and nearby parts of adjacent drainages. Localities of Apistogramma borellii: black dots, except black Fig. Y. Occlusal view ot the lower pharyngeal tooth-plate in an un- triangle, which indicates southernmost locality, from Reitzig (1977). La dexribed Apistogrammrr specie3 from the RIO GuapoIC system; a Plata basin localities of A. trifasciata: circles, except Santa FC record posteriomedial tooth shown in lateral aspect. From IRSNB 19.975. (Bonetto c'tnl. 1970), an open triangle. 25.6 mm SL. Scale 1 mm.

Zoologica Scripta I1 Cichlidfishes from the La Plata basin. IV 313

Table I. Measurements of Apistogramma inconspicua (type-series) and an undescribed Apistogramma species in the Bolivian Amazonas system (IRSNB 19.975),in mm, except range and mean, which are in per cent of SL Apistogramma inconspicua IRSNB 19.975 Range x

SL 23.5 24.4 28.5 29.7 25.6 Head length 7.7 7.6 9.1 9.4 31.1-32.8 31.9 8.2 Head depth 6.5 6.5 8.1 8.5 26.6-28.6 27.8 7.0 Body depth 8.5 8.3 10.4 11.2 34.0-37.7 36.1 9.3 Predorsal length 8.7 8.6 10.3 11.2 35.2-37.7 36.5 9.7 Preventral length 9.3 9.4 10.7 11.9 37.5-40.1 38.9 10.4 Orbit diameter 3.0 3.0 3.6 3.7 12.3-12.8 12.6 3.2 Snout length 1.4 1.3 1.8 1.9 5.3- 6.4 6.0 1.6 Cheekdepth 1.4 1.4 1.9 2.1 5.7- 7.1 6.4 1.8 Head width 4.2 4.0 4.8 5.1 16.4-17.9 17.1 4.5 Interorbital width 1.8 1.9 2.3 2.3 7.7- 8.1 7.8 2.3 Preorbital depth 0.7 0.6 0.9 0.9 2.5- 3.2 2.9 0.7 Upper jaw length 2.2 2.5 3.1 3.1 9.4-10.9 10.2 2.5 Lower jaw length 3.3 3.5 4.3 4.3 14.0-15.1 14.5 3.7 Postorbital head length 3.3 3.4 4.0 4.1 13.8-14.0 13.9 3.5 CP depth 3.4 3.7 4.4 4.8 14.5-16.2 15.3 3.9 CP length 2.4 2.9 3.3 3.3 10.2-11.9 11.2 2.4 Dorsal base length 13.9 14.6 17.2 17.8 59.140.4 59.8 15.5 Anal base length 4.8 4.8 5.9 6.4 19.7-21.5 20.6 5.4 Pectoral fin length 7.0 7.2 8.9 9.3 29.5-3 1.3 30.5 7.1 Ventral spine length 3.3 3.5 4.2 4.2 14.0-14.7 14.3 3.7 Ventral fin length 7.2 7.8 9.2 11.2 30.637.7 33.2 8.3 Last D spine length 3.7 4.0 5.4 6.0 15.7-20.2 17.8 4.5 Last A spine length 4.4 4.3 5.1 5.5 17.6-18.7 18.2 5.1

(Table I). The lateral band ends in the tail spot, and the Haseman, J. D. 1911. An annotated catalog of the cichlid fishes collected by the Expedition of the Carnegie Museum to Central abdominal side markings are restricted to short narrow South America, 1907-10.-Ann. Carneg. Mus. 7: 329-373. vertical stripes, similar to those found in A. resticulosa Kullander, S. 0. 1980a. A taxonomical study of the genus Apisto- Kullander (1980~)(Fig. 8). The dorsal fin count is XVI.6, gramma Regan, with a revision of Brazilian and Peruvian species.- Bonn. zool. Monogr. 14: 1-152. and there is at least one lower lateral line canal. Kullander, S. 0. 19806. A redescription of the South American cichlid I refrain from naming this species until additional, and better pre- Papiliochromis ramirezi (Myers & Harry, 1948).-Stud. neotrop. served material, is available (IRSNB 19.975, P, 25.6 mm. Bolivia, Fauna Envir. 15: 91-108. depto. Santa Cruz, R. Guapore system, road from Ascension, 14 km Kullander, S. 0. 1980c. Description of a new species of Apistogramma north of Limon, R. Surucusi, tributary to the R. San Miguel. 9 from the Rio Madeira system in Brazil.-Bull. zool. Mus. Univ. November 1977. Leg. J.-P. Gosse (sta.no.31).) Amsterdam 7: 157-164. Kullander, S. 0. 1982a. Cichlid fishes from the La Plata basin. Part 11. Apistogramma commbrae (Regan, 1906).-Revue suisse Zool. 89: 33-48. Acknowledgements Kullander, S. 0. 19826. Beschreibung einer neuen Apistrogramma-Art I thank Dr Jean-Pierre Gosse (IRSNB) for the loan of the material upon aus Zentral-Amazonim-Dt. Cichliden Ges. Inf. in press. which the paper is founded. Additional material reported was placed at Luling, K. H. 1979. Biotope von Apistogrammu borellii (Regan, 1906) my disposal by Drs Hans-Joachim Paepke (ZMB), Herald0 Britski in Sudamerika.-Dt. Cichliden Ges. Inf. 10: 221-224. (MZUSP), and Karl Heinz Luling (ZFMK), Mrs Carol Hutchings Luling, K. H. 1980. Wissenschaftliche Ergebnisse des Forschungs- (AMNH), Messrs Garrett S. Glodek (FMNH), Gordon Howes aufenhaltes Dr. K. H. Liiling in Argentinien 1975176. 11. (BMNH), and William G.Saul (ANSP). Thank you for all the trouble. Ichthyologische und gewasserkundliche Beobachtungen und Unter- Visits to the ZMB, BMNH, and MZUSP were made possible by grants suchungen 9G-100 Km ostlich Corrientes (Rio Parana, Prov. Cor- from the British Council, the Swedish Institute, the University of rientes, Argentinien) .-Zool. Beitr. 26: 249-285. Stockholm (C. F. Liljevalch Jr travel grants), the Royal Swedish Mayland, H. J. 1980. Das Fischparadies Chaco.-Aquar. Mag. 14: Academy of Sciences (Yngve Sjostedt’s Travel Fund; J. A. Ahlstrand’s 128-1 35. Fund), and the Helge Axson Johnson Foundation. Thanks also go to my Meinken, H. 1965. Eine neue Apistogramma-Art aus Venezuela.- wife, Mrs Anita Hogeborn, for photography, and to Mrs Britt-Marie Senckenberg. biol. 46: 257-263. Lindkvist for typing assistance. Mitsch, H. 1938. Die Zwergcich1iden.-Aquarium, Berl. 1938: 180-181. Regan, C. T. 1906. A revision of the South-American cichlid genera Retroculus, Geophagus, Heterogramma, and Biotoecus.-Ann. Mag. nut. Hist. (7) 17: 49-66. References Regan, C. T. 1909. Description of a new cichlid fish of the genus Heterogramma from the La P1ata.-Ann. Mag. nut. Hist. (8) I: 270. Ahl, E. 1938. Uber einen neuen sudamerikanischen Fisch der Familie Reitzig, W. 1975. Zur Personalakte Apistogramma reitzigi.-Aquar. Cichlidae.-Zool. Anz. 123: 246-247. Mag. 9: 513-517. Ahl, F. 1939. Uber zwei neue Fische der Familie Cichlidae aus dem Reitzig, W. 1977. Der Schleier ist geluftet: Der “reitzigi” stammt aus Zoologischen Museum Berlin.-Zool. Anz. 127: 8CL82. dem Rio Parana.-Aquar. Mag. 11: 288-289. Bonetto, A. A. Cordiviola de Yuan, E. & Pignalberi, C. 1970. Nuevos Rham, P. de 8( Kullander, S. 0. In preparation. Apistrogramma datos sobre poblaciones de peces en ambientes leniticos per- nijsseni, un nouveau cichlidi nain pour 1’aquarium.-Revue fr. manentes del Parana medio.-Physis, B. Aires30: 141-154. Aquariol. (In preparation.) Gosse. J. P. 1976. Revision du genre Geophagus.-Mem. Acad r. Sci. Ribeiro, A. de Miranda. 1918. Cichlidae.-/‘ub[g5e.s Comm. Linh. d’oiitre-MerCI. Sci. nut. mid. 8“ N.S. 19 (3): 1-172. telegr. estrat. Matto Grosso Amazon. 46 (5): 1-18. Greenwood, P. H. 1965. Environmental effects on the pharyngeal mill Schmettkamp, W. 1978. Apistogramma reitzigi Ahl, 1939 ist ein of ‘I cichlid fish, Astatoreochrornis alluaudi, and their taxonomic zu Apistogramma borellii (Regan, 1906).-Dt. Cichliden implications.-/‘roc. Linn. Soc. Lond. 176: 1-10, Ges. Inf. 9: 144-147.

Zoologica Scripta I1