Geophagus pappaterra Heckel, 1840
Satanoperca: from the Ancient Greek Σατάν (Satán), meaning ‘Satan’ (the devil), and πέρκη (pérkē), meaning ‘perch’.
pappaterra: from a local vernacular name for the species, literally translating as ‘earth eater’.
Originally native to the rio Guaporé in Brazil and Bolivia (where it is known as the Río Iténez) and the upper rio Paraguai in Brazil and northern Paraguay, these two basins being connected during the wettest part of the year via the vast Pantanal wetland, where they both have their headwaters.
The Guaporé is an affluent of the rio Mamoré within the rio Madeira basin, a major branch of the western Amazon watershed, and S. pappaterra is also known from several of its tributaries including the rios San Martin, Alegre, and Branco. The Paraguai is part of the upper Río de la Plata basin thus it flows in the opposite direction, and the species has been recorded in the Cuibá and São Lourenço drainages which feed it.
It may also occur in the Madre de Dios system, the other main affluent of the rio Madeira, but reports are scarce. Specimens apparently collected from the rio Tambopata in the Madre de Dios basin were referred to as S. pappaterra in the phylogenetic analysis published by Willis et al. (2012).
S. pappaterra has also been introduced to the upper rio Paraná in southeastern Brazil with several records from the rio Tietê tributary system, but populations from the upper Tapajós, Tocantins-Araguaia, and Xingu rivers are not conspecific.
Type locality is ‘Rio Guaporé, Mato Grosso, Brazil’.
Has been observed in main river channels, smaller tributaries, oxbows, and floodplain lakes, typically over substrates of sand or mud with patches of leaf litter and submerged tree branches.
In some cases it can also be found in artificial reservoirs created by dams.
Maximum Standard Length
180 – 200 mm.
Aquarium SizeTop ↑
An aquarium with a base measuring at least 210 ∗ 60 cm or equivalent is required.
The most essential item of décor is a soft, sandy substrate so that the fish can browse naturally (see ‘Diet’).
Coarser materials such as gravel or small pebbles can inhibit feeding, damage gill filaments and even be ingested with the potential of internal damage or blockages.
Additional furnishings are as much a case of personal taste as anything else but the most favoured set-ups tend to feature relatively dim lighting plus some chunks of driftwood and scattered roots or branches.
Leaf litter is a typical feature of the natural environment but not really recommended in aquaria because the feeding behaviour of Satanoperca spp. tends to cause an excess of partially-decomposed material in suspension which not only looks unsightly but can block filter and pump mechanisms.
Water quality is of the utmost importance since these cichlids are extremely susceptible to deteriorating water quality and swings in chemical parameters so should never be introduced to a biologically immature aquarium. S. daemon also requires soft, acidic conditions and in most cases will not thrive in untreated tap water.
The best way to achieve the desired stability is to filter the tank using a combination of external canister filters and/or a sump system and perform minimum weekly water changes of 50-70%.
If the maintenance regime is insufficient health issues such as head and lateral line erosion or stunted growth can occur.
Mechanical filtration should also be tailored to trap small particles stirred up by the fish as sand can cause blockages and wearing issues with filter mechanisms if allowed to continually run through the system. High flow rates should be avoided so position filter returns accordingly.
Temperature: 20 – 28 °C
pH: 5.5 – 7.5
Hardness: 36 – 179 ppm
Satanoperca spp. are benthophagous by nature, employing a method of feeding whereby mouthfuls of substrate are taken and sifted for edible items with the remaining material expelled via the gill openings and mouth. For this reason they are commonly termed ‘eartheaters’ and the provision of a suitable substrate is essential to their long-term well-being. Once settled they readily rise into the water column when food is introduced but continue to browse normally at other times.
The stomach contents of wild specimens mostly comprise small aquatic and terrestrial invertebrates including cladocerans, ostracods, and insect larvae (particularly chironomids), plus plant material (seeds), organic detritus and sediment.
Even large individuals seem unable to properly ingest larger food items, meaning the diet should contain a variety of high quality, fine-grade prepared foods plus small live or frozen bloodworm, Tubifex, Artemia, mosquito larvae, etc. At least some of the dried products should contain a high proportion of vegetable matter such as Spirulina or similar.
Home-made, gelatine-bound recipes containing a mixture of dried fish food, puréed shellfish, fresh fruit and vegetables, for example, are proven to work well and can be cut into bite-sized discs using the end of a sharp pipette or small knife.
Rather than a single large meal offer 3-4 smaller portions daily to allow natural browsing behaviour as this seems to result in the best growth rate and condition.
Behaviour and CompatibilityTop ↑
Unless breeding this species is surprisingly peaceful and will not predate on fishes larger than a few millimetres in length. Suitable tankmates are numerous and include most peaceful species enjoying similar environmental conditions. Best avoided are aggressive or territorial substrate-dwelling species, or those requiring less acidic or mineral-rich water.
Some aquarists keep Satanoperca spp. alongside freshwater stingrays of the genus Potamotrygon which in many cases has proven successful but in some has resulted in the cichlids disappearing at night.
S. pappaterra is gregarious and tends to exist in loose aggregations unless spawning, with juveniles in particular displaying strong grouping instincts. A group of 5-8 individuals should be the minimum purchase and these will form a noticeable dominance hierarchy. When maintained in smaller numbers weaker specimens can become the target of excessive antagonism by dominant individuals or the group may fail to settle and behave nervously.
Adult females tend to grow slightly smaller than males and are deeper-bodied when gravid.
This species is an ovophilous maternal mouthbrooder that has been bred in aquaria. There does not appear to be any particular trigger for the spawning process with the main requirements being good diet and stringent maintenance regime involving relatively large weekly water changes.
Since accurate sexing is very difficult it is perhaps best to begin with a group of young fish and allow pairs to form naturally, while a degree of patience is also required since it can be at least a year, often two or more, until they become sexually mature.
Courtship is initiated by the male, with nuptial individuals forming territories which are typically centred around a piece of driftwood or other solid object. Gravid females are courted and when a pair forms they may defend the territory together for a few days prior to spawning. Other fishes, especially conspecifics, are not tolerated in the vicinity and in smaller aquaria may need to be removed.
Eggs are deposited in small batches and once fertilised immediately collected by the female, the process normally being repeated until she is carrying several hundred. In some cases the male may also pick up some eggs but does not usually hold them for long, and the female is normally ejected from the territory soon after spawning is complete although apparently some males do display extended brood care behaviour.
Once the fry are free swimming they are easily-fed, accepting good quality powdered dry foods, Artemia nauplii, microworm, etc.
If maintaining the adults in a community situation it is recommended to remove brooding females since the fry become easy prey for other fishes, including conspecifics, once released.
S. pappaterra is relatively rare in the ornamental trade and much sought after by enthusiasts.
It is easily distinguished from all known congeners by presence of a series of prominent black blotches beneath the dorsal-fin, plus a well-defined dark band extending along the side of the body. In recent genetic analyses its genetic distinctness was strongly supported, with S. jurupari and S. leucosticta its closest relatives.
Satanoperca was considered a synonym of Geophagus for a number of decades before being revalidated by Kullander (1986). It is separated from all other members of the putative cichlid subfamily Geophaginae by presence of a small ocellated spot at the upper caudal-fin base, and from Geophagus by a number of morphological characters as follows: equal numbers of abdominal and caudal vertebrae or one more abdominal vertebra (vs. more caudal than abdominal vertebrae in Geophagus); no ribs on caudal vertebrae and no postabdominal swimbladder extensions (vs. presence of both in Geophagus; cheek scaled rostrally to centre (vs. naked on anterior half; dorsal and anal fins naked (vs. scaled); external gill rakers on lower limb of first gill arch attached to the skin covering the gill filaments (vs. attached to ceratobranchial skin); jaw dentition reduced to one or two short series in each jaw (vs. pluriseriate with wide band of inner teeth); absence (vs. presence) of tooth plates on 4th ceratobranchial; presence (vs. absence) of an ocellus on caudal-fin base; both or only upper lip fold continuous (vs. both interrupted anteriorly); preoperculum and supracleithrum frequently serrated (vs. entire); absence (vs. presence) of median hypural cartilage; 3 (vs. 4) infraorbitals.
The genus is likely to contain additional species, and has been separated into three groups with distinctive morphology, namely the S. jurupari group, the S. daemon group, and the monotypic S. acuticeps group.
Among these, members of the S. jurupari group are characterised by a colour pattern comprising a dark lateral stripe or row of blotches along the side of the body, normally XV.10 (XIV-XVII.8-11) dorsal-fin rays, normally 27 (26-28) E1 scales (the longitudinal row immediately above the lower lateral line, excluding the pectoral girdle), and 12-18 rakers on the first gill arch.
S. daemon group members grow larger than other congeners, possess one or two large dark blotches on the side of the body and a prominent ocellus at the upper caudal-fin base, usually XIX.13 (XII-XIV.12-14) dorsal-fin rays, 29-31 E1 scales, and 18-22 gill rakers. S. acuticeps is unique in that it has a relatively more terminal mouth than all other genus members, normally XIII.12 (XIII.11-12) dorsal-fin rays, 27-28 E1 scales, and 17-20 gill rakers.
The groups also differ in reproductive strategy, with the S. daemon and S. acuticeps assemblages being substrate spawners, and the S. jurupari group mouthbrooders.
Satanoperca and a number of related genera are often included in the putative subfamily Geophaginae, and Kullander (1998) conducted a morphology-based phylogenetic study in which the neotropical Cichlidae was divided into six subfamilies of which the Geophaginae contained 16 genera divided among three ‘tribes’:
Acarichthyini – Acarichthys and Guianacara.
Crenicaratini – Biotoecus, Crenicara, Dicrossus and Mazarunia.
Geophagini – Geophagus, Mikrogeophagus, ‘Geophagus‘ brasiliensis group, ‘Geophagus‘ steindachneri group, Gymnogeophagus, Satanoperca, Biotodoma, Apistogramma, Apistogrammoides and Taeniacara.
Later molecular studies by Farias et al. (1999, 2000, 2001) resulted in the additions of Crenicichla and Teleocichla to the Geophaginae, a result supported by López-Fernández et al. (2005) who conducted the most detailed molecular analysis of the grouping to date including 16 of the 18 genera and 30 species.
However their conclusions regarding interrelationships between genera did vary somewhat from previous hypotheses and can be summarised by the following loosely-defined groups:
– a weakly-supported sister group relationship between Acarichthys and Guianacara.
– a well-supported ‘Satanoperca clade’ comprising Satanoperca, Apistogramma, Apistogrammoides and Taeniacara.
– a ‘big clade’ with Geophagus, Mikrogeophagus, ‘Geophagus‘ brasiliensis group, ‘Geophagus‘ steindachneri group, Gymnogeophagus, Biotodoma, Crenicara and Dicrossus.
– a ‘crenicarine clade’ with Biotoecus and Crenicichla.
No representatives of Teleocichla or Mazarunia were included in the study but the former is well-established as sister to Crenicichla while the latter has grouped closely with Dicrossus and Crenicara in earlier works.
The other main conclusions of the paper are confirmation that Geophaginae is a monophyletic group exhibiting strong signs of having undergone rapid adaptive radiation.
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