The Resurrection of

by Willem Heijns
On several occasions I have reported on the taxonomy of the genus Cichlasoma (Heijns, 1988, 1999, 2000) as it developed from a discarded name via a very speciose group, into a genus with only a dozen or so species. In these publications I discussed the history of the name Cichlasoma and then explained what would be the “correct” generic assignment of the species that once belonged to Cichlasoma, but had to be removed from that genus following the revision of Kullander (1983). As for Cichlasoma s.s., I restricted myself to naming the species Kullander (re)described and I commented on the difficulty of identifying them without locality data. After Kullander’s revision, Cichlasoma remained taxonomically untouched for almost thirty years. After Ottoni (2011) described a new species of Cichlasoma, I thought it might be a good idea to present an overview of this genus and its position within the family Cichlidae as it stands now.

Cichlasoma amazonarum in the aquarium. Photo Rainer Stawikowski.
Species of Cichlasoma

Cichlasoma was originally described by William Swainson (1839). For the turbulent history of the name see Heijns (1988). Of the species now considered to be part of this genus, Cichlasoma bimaculatum is the oldest one, being described by Linnaeus in 1758. The newest addition is Cichlasoma zarskei Ottoni 2011. In the following, I will give a summary of the taxonomic history of all thirteen Cichlasoma species.

Cichlasoma amazonarum Kullander 1983. As the name indicates this species occurs all along the mainstream of the Amazon river from Peru (Río Ucayali) to Gurupá at the mouth of the Amazon. Such a vast range almost inevitably contains different geographic variants. Kullander (1983) already recognized this, since he described several populations of this species. Before 1983, the cichlids now called Cichlasoma amazonarum were referred to as bimaculatum (with varying generic assignments), but for some reason Lüling thought fishes he found at Yarina Cocha, Peru belonged to Cichlasoma facetum (Lüling, 1975) or Aequidens portalegrensis (Lüling, 1978). I would not be surprised if Cichlasoma amazonarum were to be split into several new species someday.

Cichlasoma araguaiense with fry. Photo Uwe Werner.
Cichlasoma araguaiense Kullander 1983. This species was first reported on by Castelnau in 1855. He identified his specimen as Cichlasoma bimaculatum or, to be more precise, as “Chromys punctata?” with reference to Bloch’s (1792) drawing of the fish that later became known as Cichlasoma bimaculatum (see also Heijns, 2000). The question mark indicated that he wasn’t sure. Haseman (1911) published a catalog of the fishes collected by the expedition of the Carnegie Museum (1907-1910) in which he listed 13 localities (predominantly in the state of Bahia, Brazil) where a cichlid identified as Cichlasoma bimaculatum was found. One of these localities was the Rio do Sono, a tributary of the Rio Tocantins. The specimen found there (only 9 cm or 3.5” in size) was later determined as Cichlasoma araguaiense by Kullander (1983). One remark by Haseman is noteworthy here. In summarizing his records of Cichlasoma bimaculatum he stated “I cannot distinguish this species from Aequidens portalegrensis, except for its having as a rule four anal spines. One spine more or less does not always have either generic or specific significance”. Haseman was way ahead of his time, I would say.

Cichlasoma bimaculatum from Guyana. Photo Rainer Stawikowski.
Cichlasoma bimaculatum (Linnaeus 1758). This is the type species of the genus. It is also the very first species (now in Cichlasoma) to be described. In an earlier article (Heijns, 2000) I have discussed the history of the name and the problems that went with it. It was not until the year 2000 that these problems were finally solved by the International Commission on Zoological Nomenclature by placing Cichlasoma on the Official List of Generic Names with type species Labrus bimaculatus Linnaeus 1758 and by placing bimaculatus on the Official List of Specific Names with holotype NRM 7 (in the Swedish Museum of Natural History).

Since the name was already published in 1758, it is no great wonder that bimaculatus has a long and varied taxonomical history. Let’s look at synonyms first. Heckel (1840) believed that the specific epithets punctatus and bimaculatus applied to two separate species. Interestingly he did not refer to them as Labrus punctatus Linnaeus 1758 and Labrus bimaculatus Linnaeus 1758. Instead he used the generic name Sciaena for both species, a name used by Linnaeus in 1754. He assigned both species to his Acara (ignoring Swainson’s Cichlasoma) and thus called the first Acara punctatus, while for the second he created the new name Acara gronovii, to which species he also assigned Labrus bruneus Gronovius 1756. Today we know all of these names as synonyms of Cichlasoma bimaculatum (see Heijns, 2000). Another synonym is Sparus filamentosus Gray 1854.

Generic assignment (to which genus does bimaculatum belong) is quite another matter. Here’s a list of genera to which our species has been assigned over time. Labrus (Linnaeus 1758); Acara (Heckel, 1840); Cychlasoma (Gill, 1858); Heros (Cope, 1872); Acara (Heros) (Steindachner, 1875); Astronotus (Heros) (Eigenmann & Eigenmann, 1891); Astronotus (Cichlasoma) (Eigenmann & Bray, 1894); Cichlasoma (Eigenmann & Kennedy, 1903); Cichlasoma (Cichlasoma) (Pellegrin, 1904) and finally Cichlasoma (Kullander, 1983).

Fortunately the holotype of Cichlasoma bimaculatum has not been lost. It is in the Swedish Museum of Natural History in Stockholm and labelled as NRM 7. It dates from the collection of King Adolph Frederic back in the 18th century. Problem is that nobody knows where the specimen was collected. Linnaeus listed the Mediterranean, but that is of course wrong. So we don’t have a type locality for this species. A bit to my surprise, this is not seen as a major problem, whereas in other genera (i.e. Symphysodon, Australoheros) there has been a lot of discussion about correct type localities.

Head of Cichlasoma boliviense. Photo Uwe Werner.
Cichlasoma boliviense Kullander 1983. This species takes its name from the country in which the center of its distribution range lies. Earlier records (Haseman 1911, Fowler 1926) from this area were reported as Aequidens portalegrensis, a species now restricted to much more southern areas. Lüling appears to have found specimens in 1969 in the Río Chaparé (Bolivia) on two locations not far apart, which Kullander (1976) identified as Aequidens portalegrensis and Cichlasoma bimaculatum respectively. Both are now regarded to belong to Cichlasoma boliviense.

Cichlasoma dimerus with its spawn. The eggs are about to hatch. Photo Uwe Werner.
Cichlasoma dimerus (Heckel 1840). The only species described by Heckel which is currently assigned to Cichlasoma. As stated elsewhere (Heijns, 1988) Heckel did not recognize Swainson’s work and as a consequence did not use Cichlasoma as a valid generic name. Instead he described the genus Acara and proposed the name Acara dimerus for this species. He had only one fish at his disposal, found in the Rio Cuiabá, southern Mato Grosso (Brazil). Interestingly, Heckel (1840) also described Acara marginatus from the same river and also with just a single specimen at hand. His comment on these species was that they were very similar, but that Acara dimerus might be the smaller of the two. Both specimens Heckel used, were also studied by Kullander (1983). He found that they belonged to one and the same species, along with many other specimens from that area which he used in his revision of Cichlasoma. So he had to choose one of Heckel’s names to assign to this species. As first revisor Kullander selected dimerus as the specific epithet, the reason being that marginatus had never been used as a valid name after Heckel whereas dimerus had been recognized as valid by Günther (1862) and a few others.

Another interesting note, giving an insight into how nomenclature works, is the fact that Kullander considered “neither of the [Heckel] types as particularly representative for the species”. But by including these types into the species at hand, he had to choose between one of the two names Heckel (1840) had proposed, these two names being the oldest ones available. And by choosing dimerus, the fish used by Heckel for the description of Acara dimerus automatically becomes the holotype for what is now Cichlasoma dimerus. Thus the holotype of Cichlasoma dimerus is now considered “not particularly representative” for the species it stands for. Of course it doesn’t formally have to be representative as it is only the name bearer of Cichlasoma dimerus, but it would be much easier to identify other specimens as Cichlasoma dimerus if they would compare well with the holotype. I don’t think that is very satisfactory.

As with Cichlasoma bimaculatum this species has been assigned to a number of genera over time. Among them Cychlasoma (Gill, 1858), Astronotus (Aequidens) (Eigenmann & Bray, 1894) and Aequidens (Eigenmann & Kennedy, 1903). Heros centralis Holmberg 1891 appears to be a junior synonym, although this name was incompletely described and no types are available.

Cichlasoma orientale. Photo Sven Kullander.
Cichlasoma orientale Kullander 1983. This species has the most easterly distribution range of the genus, hence the name orientale (from the Latin orientalis, meaning eastern). Earlier records of it were all under the name Cichlasoma bimaculatum, as can be expected. Kullander’s description was based mainly on material from the Rio Curú, a tiny river flowing into the Atlantic Ocean just west of the city of Fortaleza (Ceará, Brazil), although other Ceará material was also considered. He was in doubt as to whether specimens from further east (Rio Grande del Norte, Pernambuco) could also be assigned to his new species, but these were later confirmed to be Cichlasoma orientale by Schindler (1995), who also found this species in the Rio Parnaiba, west of Ceará. Still further to the west, Schindler also collected cichlids in the Rio Itapecuru (Maranhão, Brazil), which seem to be closely related to Cichlasoma orientale, but show significant differences. These were later described as Cichlasoma zarskei (see below).

Cichlasoma orinocense with fry. Photo Rainer Stawikowski.
Cichlasoma orinocense Kullander 1983. This is a peculiar species. Although the name suggests a distribution range throughout the Orinoco drainage, Kullander’s description is based on a single specimen from a far corner of this great river system. The holotype was collected together with several other specimens in a lake in the upper part of the Río Metá, close to the Andes in Colombia. Three other specimens he identified as Cichlasoma orinocense came from the Río Guárico, near the city of Calabozo (Venezuela). As Kullander (1983) stated: “vast areas of the Orinoco basin remain to be sampled”. Near the mouth of the Rio Orinoco, in some coastal rivers and on the island of Trinidad another species of Cichlasoma can be found: Cichlasoma taenia (see below). This species can mainly be distinguished from Cichlasoma orinocense by its more elongated body and longer head. Schindler (1997) collected cichlids in the area between the ranges of Cichlasoma orinocense and Cichlasoma taenia (the lower part of the Orinoco) and found intermediate forms. He questions the significance of body-height and headlength as diagnostic characters to separate these two species, but suspects there may be other characters to be of equal value to warrant the specific status of both species.

Freshly collected pair of Cichlasoma paranaense. Photo Sven Kullander.
Cichlasoma paranaense Kullander 1983. As the name indicates, this species has its home range in the Rio Paraná, more precisely in the upper part of this great river drainage, locally known as Alto Paraná. In fact, it is the only Cichlasoma-species endemic to the Paraná drainage. The holotype was collected by Britski in 1966 in the Rio Sucuriú at Fazenda Santa Luzia. It measured only 68.1 mm (2.68 in). He reported on this fish in 1972, identifying it as either Aequidens portalegrensis or Aequidens paraguayensis. At first sight a bit strange, but Haseman (1911) had already used the same name (Aequidens portalegrensis) in his report on the fishes, collected during the Carnegie expedition (1908). De Miranda Ribeiro (1918) used the name Aequidens dimerus in his list of fishes in the Museu Paulista in Saõ Paulo.

Cichlasoma portalegrense, lectotype. Photo Sven Kullander.
Cichlasoma portalegrense (Hensel 1870). One of the older names for a species now included in Cichlasoma. It was described by Hensel as Acara portalegrensis from 11 fishes found in stagnant water near the town of Porto Alegre (hence the name). The largest specimen was 140 mm (5.5 in) long. This seems a bit strange, because none of the Cichlasoma species presently known reaches that size (SL). Kullander (1983) could only locate 10 fishes, the largest of which was 84.6 mm (3.33 in). He designated this as the lectotype of his Cichlasoma portalegrense without giving any specific reason.

Hensel assigned his new species to Acara Heckel 1840, following Günther who had classified species with 3 or 4 anal spines in this genus. Over time, our species has travelled quite a bit between different genera. Eigenmann & Eigenmann (1891) considered Acara to be a subgenus of Astronotus, an assignment which Eigenmann & Bray (1894) changed into Astronotus (Aequidens) after their description of this new subgenus. Ten years later Eigenmann & Kennedy (1903) raised Aequidens to generic rank changing the name into Aequidens portalegrensis, a name to be used for the next 80 years, right until Kullander assigned it to Cichlasoma. But even today some hobbyists still use the name Aequidens for this species, also known by its vernacular name “Port Cichlid”.

The Port Cichlid also has an impressive aquarium history. Problem with this history is that we are not really sure that it concerns Cichlasoma portalegrense. Schindler (1999) did some research on two different German aquarium populations of this cichlid and could only conclude that it might have been Cichlasoma portalegrense in one case and probably a hybrid in the other. In the USA (Leibel, 1984) many different cichlid species have been kept under the name Port Cichlid as well. Finally, the world famous ethologist G.P. Baerends (1984) studied cichlid behavior using Aequidens portalegrensis, at least that was what he called them…

Cichlasoma pusillum in the collection of NRM Stockholm. Photo Sven Kullander.
Cichlasoma pusillum Kullander 1983. According to Kullander (1983) the smallest species in the genus. The holotype measured only 59.2 mm (2.33 in) SL. Kullander chose the name pusillum because of its small size. He could not choose a name derived from a locality, as he had done for all other new species described in his revision of Cichlasoma because this species occurs in two drainages. Interestingly Kullander (2003) stated that the Uruguay material (3 fishes from Artigas) do not belong to this species after all, leaving only the material from the Paraná as Cichlasoma pusillum. Except one fish, collected by Haseman in 1909, of which the identity is uncertain. Recently Kullander & Santos De Lucena (2013) confirmed the identification of the Uruguay material as Cichlasoma dimerus.

Cichlasoma sanctifranciscense. Photo Sven Kullander.
Cichlasoma sanctifranciscense Kullander 1983. Again a species owing its name to its distribution area, the Rio São Francisco in Brazil. Kullander studied well over a hundred specimens from this river drainage collected by the expedition of the Natural History Museum in Vienna (NMW) in 1903 in which the famous ichthyologist Franz Steindachner participated. From these he also chose the holotype. Haseman (1911) also collected in this area, but Kullander did not review the specimens from the São Francisco, identified by Haseman as Cichlasoma bimaculatum.

Cichlasoma taenia, pair guarding fry. Photo Rainer Stawikowski.
Cichlasoma taenia Bennett 1831. The second oldest described species, currently in Cichlasoma. And with an interesting history. It was presented by a certain Mr. Bennett at the meeting of the Zoological Society of London held at 26 July 1831 as part of “a small collection of fishes formed during the voyage of the HMS Chanticleer”. How many specimens were presented is not known. Judging from the original description, which lacks any ranges of the given characters, it can fairly safely be assumed that there was only one specimen. The problem is that this specimen disappeared from sight for more than 21 years until it was registered in the British Museum of Natural History, now known as the Natural History Museum, on 13 September 1852. Günther (1862) subsequently included it in his famous catalog of the fishes in this museum and listed it as the “type of Chromis taenia”, notwithstanding the fact that he considered it to be a synonym of Acara bimaculata.

Over time, the name taenia has also travelled quite a bit between genera. After its initial assignment to Chromis, Heckel (1840) transferred it to Acara, followed by Gill (1858) who assigned it to Cychlasoma. Subsequent authors thought, as Günther did, that it was a synonym of bimaculatus and as such it followed the path of this name through the genera to which bimaculatus has been assigned (see above). Regan (1905), in his revision of the cichlids in and around Cichlasoma, also believed taenia to be a synonym of his Cichlosoma bimaculatum and listed one specimen from the “Zoological Society” collected in Trinidad. It is assumed that this specimen is the same as the one mentioned by Günther as the holotype of Chromis taenia. Finally, Kullander (1983) resurrected the name from synonymy and the species has since been known under its current name Cichlasoma taenia.

Cichlasoma zarskei. Photo Wolfgang Staeck.
Cichlasoma zarskei Ottoni 2011. The most recent addition to the genus. According to Kullander (1983) the distribution range of Cichlasoma orientale had its western limit in the state of Ceará with a possible extension into the state of Piauí from which only a literature record was known (Nomura & Barbosa 1980). Schindler (1995) found cichlids, belonging to the genus Cichlasoma in the state of Maranhão, west of Piauí. Due to the bad state the fishes were in, he was not able to distinguish them from Cichlasoma orientale, so he tentatively regarded them conspecific with orientale. Schindler’s suspicion that it might concern a separate species was confirmed by Ottoni (2011), who subsequently described the Maranhão population as Cichlasoma zarskei. After 28 years the genus Cichlasoma was expanded from 12 to 13 described species.

The precise locality where the type series were found is a bit of a mystery. The holotype and some paratypes supposedly were collected in a lake called “Lago Malhada Grande” in the Rio Maranhão basin. Extensive search yielded that the Rio Maranhão does not exsist and neither does the lake. Several other paratypes were collected at Igarapé Arari, a small municipality just south of Vitória do Mearim at the junction of the rivers Pindaré and Mearim. Cichlasoma zarskei was named in honour of Axel Zarske, the editor of the scientific magazine Vertebrate Zoology.

Part 2 will continue in the next issue.


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