(Heckel 1840) Satanoperca pappaterra. Wayne S. Leibel 3574 Timberlane Dr. Easton, PA 18042

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Satanoperca pappaterra (Heckel 1840) Wayne S. Leibel 3574 Timberlane Dr. Easton, PA 18042 Some wonderful things happen by accident. This is an account of an accidental spawning of a wonderful Geophagine Eartheater, Satanoperca pappaterra with notes on its maintenance and taxonomic status. Geophagus pappaterra was described by Heckel in 1840 along with G. jurupari, G. daemon, and G. acuticeps with which it shares its juruparoid good looks. While G. jurupari was similar to G. daemon ( Geophago Daemoni similis ) and had a caudal peduncle blotch ( Macula parva ad basim pinnae caudalis ) with labial and head stripes ( fascia labia... and fasciis tribus transversis capitis ), G. pappaterra, among other characteristics, had a pronounced longitudinal black stripe running the length of its mid-flank ( vitta longitudinali media nigra ) just under the lateral line. Heckel further described five brown (indistinct) vertical bands which darkened conspicuously when they reached the uppermost two scale rows just below the dorsal fin insertion and, like G. jurupari, a dark caudal peduncle blotch. While G. jurupari was described from Natterer s Rio Negro (i.e. Amazon) material, G. pappaterra hailed from the Rio Gaupore, in the Mato Grosso region (southwestern Brazil). Pappaterra, in Portugeuse, means eartheater and is the local name given this fish! (according to Natterer, the original collector.) In a BB essay published in 1983 which attempted a resolution of the G. acuticeps identity problem, I suggested that the juruparoid eartheater bearing a dark longitudinal stripe and identified in various then authoritative cichlid compendia as G. acuticeps, was not (the real G. acuticeps, has 4 bloches), but rather might be G. pappaterra. Published along with the article was a picture of a pair of bronze-gold juruparoids of unknown provenance which I believed to be G. pappaterra sensu Heckel. However, without known locality data, my assertions were simply circumstantial at best a good guess. About one year ago (last January 1990), I was excited to find four medium-sized juruparoids (c. 4.5-6 inches TL) at George Fear s wholesale operation (Tropical Imports, Irvington, NJ). They were bright gold with stunning red-orange dorsals and were, at the same time, expressing a dark longitudinal band that is often part of the usual stress pattern of generic juruparoids. They had arrived a day or two before with a shipment of Mato Grosso fish (including the lime-green Buckelkopf (Laetacara sp. affin. curviceps) much the worse for wear. I thought: pappaterra! And I was right. There is no confusing this fish with any of the other animals that sell as G. jurupari in the hobby. Gosse (1976) in his monumental revision of the genus Geophagus, elected to lump several juruparoid species under the name G. jurupari despite some statistical differences in their meristics and morphometries. These Buntbarsche Bulletin 140 / 15

Above: Satanoperca pappaterra, the gold, striped jurupari from the Mato Grosso. Adult male 7" TL. Below: Satanoperca leucosticta, the common hobby jurupari. Adult female, 5" TL. W. S. Leibel photos. 14 / Buntbarsche Bulletin 140

included leucostictus Muller & Troschel 1848, macrolepis (Guenther 1862), mapiritensis Ferandez-Yepez 1950, and of course, G. pappaterra Heckel 1840. In arguing for the synonymization of G. pappaterra with G. jurupari, Gosse pointed out that Heckel s description was based on a single specimen which Haseman (1911) believed to be of extraordinary proportions (actually, it was 160 mm SL, 203 mm TL which is big, but not a monster for live material in my experience (eg. Agassiz s Thayer Expedition material that I viewed at Harvard; Leibel 1983). Following Haseman s lead, who suggested that this monster was simply one of the end variations of G., jurupari. All of the intervening stagey exist in the same locality, Gosse was inclined to lump the two, with G. jurupari having precedence since it was described 3 pages earlier in Heckel (1840). Gosse (1976) had confirmed the absolute accuracy of Heckel s original description upon reexamination of the 1840 holotype, but apparently did not think it sufficiently different from G. jurupari to retain the species as valid. Moreover, the name G. pappaterra had been applied infrequently but incorrectly in the literature to a number of non-mato Grosso specimens (eg. Schomburgk (1848) from Guyana, Eigenmann & Kennedy (1903) from Paraguay, (possibly correct), De Miranda Ribeira (1918) from Manaus) and thus it was poorly defined. Gosse (1976) recommended relegating pappaterra to synonymy with jurupari. With live fish in hand I am forced to reiterate the accuracy of Heckel s description of the Mato Grosso juruparoid (see photo) and I am forced to conclude, as did Kullander (1986), that Gosse (1976) was wrong or at least mislead by examining only the single Heckel G. pappaterra holotype in lumping the two fish. The Mato Grosso fish is juruparoid in overall shape, but it is iridescent gold, has a dark black longitudinal band extending from the operculum back to the peduncle that is 1-2 scale rows below the lateral line, and has five blotches on the dorsum just below the dorsal fin which, in certain behavioral states, extend vertically into light brown bars that continue below the longitudinal stripe (actually, the stripe sometimes fades except for five blotches which are part of the vertical bar series). Unlike what we in the hobby regard as G. jurupari (but see below), the stripe and the dorsal blotches are a regular, maintained feature of the coloration and not part of a transient stress pattern that jurupari expresses when you put it in a photo tank and threaten it, and lack any facial spotting. Apparently, Stawikowski & Werner (1988) would agree with me since the fish figured on page 194 and again on 196 are identical to mine, as are the 2-3 inch S. pappaterra imported from Germany by my friend Tony Orso (Water Life Imports, Fort Lee, NJ). Are they different and valid species? Before answering that, let's talk about what G. jurupari sensu stricto is. In his monumental Cichlid Fishes of the Amazon River Drainage of Peru, Kullander (1986) resurrected the genus Satanoperca Guenther 1862 to hold the juruparoid eartheaters. Guenther (1862) had originally divided Heckel s (1840) Geophagus into three genera: Mesops (now Biotodoma), Geophagus proper with a scaly dorsal fin, and Satanoperca with a naked dorsal fin. Although Gosse (1976) and others before him ignored or disagreed with Guenther s scheme, Kullander (1986) revalidated the Geophagus/Satanoperca split on 12 addi 16 / Buntbarsche Bulletin 140

S. pappaterra, 3" TL juvenile showing longitudinal stripe and sub-dorsal spots. W. S. Leibel photo. tional characters. In so redefining Satanoperca, Kullander (1986) redescribed Satanoperca jurupari from Peruvian Ucayali-Amazonas specimens. Kullander (1986) asserts that the real S. jurupari is found from eastern Peru east through the Amazon basin and its tributaries to the Rio Xingu. Kullander (1986) suggests that the juruparoid collected in the Orinoco drainage (Colombia, Venezuela) and upper Rio Negro is an as yet undescribed species, that S. pappaterra is a valid species and is found in the Guapore and Paraguay drainages, and that the juruparoid of the Guyanas (Essequibo and Corantijn drainages) is S. leucostica, a position he reiterates in his more recent Cichlids of Surinam (Kullander & Nijssen 1989) (see Leibel, 1990; BB 136). For aquarists, the best photo resource in identifying these fish (and all the Satanoperca species) is Stawikowski & Werner s (1988) Die Buntbarsche der Neuen Welt, Sudamerika (Reimar Hobbing, West Germany). Although the origin of the fish in question is very helpful in identifying the animals in question, some generalizations are possible, the jurupari" with the white iridescent dots on its muzzle and gill plate is S. leucosticta (Kullander 1989). The real S. jurupari apparently lacks all facial spotting (Kullander 1986) while the undescribed species from the Orinoco has spots on the face but not on the gill cover (Kullander 1989). Apparently the body color of all can vary from gun metal gray, to silver, to green-bronze, to gold depending on differing water/substrate conditions (Kullander 1986). Satanoperca pappaterra is distinguished from all of the above by its persistant longitudinal stripe and the 5 blotches on the dorsum just below the insertion of the dorsal fin. Most likely, there are other as yet undescribed juruparoid Satanoperca like the distinctive pappaterra - like fish I caught in Peru that expresses persistant black longitudinal blotches (see photo next page and BB 124). Buntbarsche Bulletin 1 4 0 /17

As expected, S. pappaterra present few surprises with regard to their general maintenance. As I have outlined previously (Leibel 1985a,b; 1987 a,b, 1988) the Satanoperca should be treated like large dwarf cichlids and kept warm (80-85 F), clean, and well-fed with a variety of pelleted and frozen foods, over a fine substrate (sand to #0 gravel) in a relatively large tank (at least 30 gallons long) with non-aggressive tankmates, preferably not other cichlids. I believe they take at least 1 ½ - 2 years to mature (my smallest spawning pairs, and I have spawned 4 different pairs of S. leucosticta, have been 4-5 inches) and will not spawn when kept with intimidating tankmates. They are apparently not sexually dimorphic, although in my largest pairs, including S. pappaterra, the male was the larger (1 + inch) of the two fish. Cichocki (1976) reported similar size differences in natural pairs observed in the wild. As has been reported by a number of breeders, the juruparoids that have been spawned to date (jurupari, pappaterra, leucosticta, and daemon) are all delayed (primitive) biparental mouthbrooders who lay their eggs on a substrate (a movable one in the wild!: Cichocki, 1976), fan and guard them for 24-48 hours, and then chew the yolksac wrigglers out of their shells for further buccal incubation until they are released, free-swimming 2-4 days later. The parents continue to provide buccal shelter for 3-4 weeks thereafter. In my case, I wasn t expecting a spawning, nor was I doing anything to provoke one. I lost two fish right off the bat and had left what I believed (correctly) to be a pair: one large one (c. 6 inches TL), the putative male, and a smaller, less elongate female (c. This undescribed Satanoperca sp. from Peru resembles S. pappaterra. Adult male, 7" TL. W. S. Leibel photo. 18 / Buntbarsche Bulletin 140

5 inches TL). I put them into a newly set-up 150 gallon community tank containing a shoal (15) of 6-inch elongate hatchets (Triportheus sp.-see recipe for eating these on page 10), a dozen banded silver dollars (4 inch Myleu s schomburgki), a shoal of Corydoras species, a dozen Clown Plecos (Pekoltia sp.) and a variety of other cichlids including large Biotodoma wavrini) and a few Satanoperca acuticeps. The hatchets and the silver dollars were in constant motion, but the S. pappaterra seemed unconcerned and ran the tank. There were two huge piles of Malaysian driftwood on either end of the tank (for the Pekoltia) and because the wood was new, the water (ph 7.0, 20 microsiemens conductivity) was quite tea brown with the leaching tannic acids. There was fine sand on the bottom, floating water sprite on the top, and the water heavily aerated with 3 Rena 301 pumps and filtered with a Hagen Fluval 403 canister filter. Fifty percent water changes were done routinely every two weeks. The water was maintained at 80 F, and the fish fed variously with dried krill, pelleted and floating dry foods, and a frozen, vegetable-based mixture (Lifeline, Herbivore, Bio-Tope). The S. pappaterra alternately got along terrifically for a few days and then didn t, the male chasing the disinterested female mercilessly; thank God for the driftwood piles. They continued like this from February until April. In April, I went away for a few days not long enough to ask a neighbor to feed the fish, but just long enough to worry about what they might do to each other without regular feedings. I adjusted the timers controlling the canopy lights to shut off at 7 P.M., rather than the usual midnight, to cut back on fighting time. When I returned, everyone was fine but hungry. I did a water change immediately, but since it was late at night, I simply switched one timer from auto to manual on then forgot to reset it. About 4 days later, I noticed the female S. pappaterra was not out front for feeding. She was sulking in the driftwood pile with what looked like a mouthful of eggs. I hadn t seen them spawn, and the male, rather than helping, would chase her every opportunity he got. When I fed them, the female refused even the choicest of sinking morsels. I knew she was carrying. But what to do? It was a miracle that the eggs had survived their 48 hour exposure on the driftwood (I presume they are delayed mouthbrooders but I have no direct proof) given the number of hungry catfish around. The mis-set timer had been their only salvation, providing the female with the light necessary to defend the clutch from the Pekoltia. Obviously she couldn t raise the fry in that tank, but my experience with other Satanoperca pairs suggested that attempting to move an ovigerous female would be disastrous (it works with Geophagus steindachneri but they re industrial strength). I thought I had nothing to lose because the male was making her life completely miserable and would soon do her in. Have you ever tried catching one fish out of a decorated 150 gallon tank? I put in an egg-crate divider to cut the six foot distance to two, carefully removed the sequestered driftwood pile where she was sulking and quickly but gently netted her out and into a 5 gallon pail. Shuddering all the way, I drove the pail 15 minutes to my office where the only other available running aquaria existed, and released her into a 20 gallon long after first building a driftwood cave. I feared the worst, but she held the eggs and released nearly 200 free-swimming fry Buntbarsche Bulletin 1 4 0 /19

Left: S. pappaterra, brooding female with young. W. S. Leibel photo. addition to any captive collection of juruparoid eartheaters. I m sure we haven t yet seen the last juruparoid surprise! References several days later. They were large enough to take newly hatched Artemia and gobbled it voraciously. She continued to care for them for 4 weeks post-release at which time they were too big for her to accommodate (she acted visibly annoyed by them) and I moved her to another tank to recondition her before returning her to the 150. The fry grew and continue to grow relatively slowly and the biggest are approaching one inch approximately 6 months post-spawn. The Guinacara (Acarichthys) cf. geayi which started out the same size as them when placed in the 30 gallon growout tank 3 months ago, are much larger than the S. pappaterra fry now. They begin to express the dark longitudinal stripe that characterizes this species at about one inch TL. I am continually amazed by the wealth of diversity in fishes that the Amazon Basin provides. It is estimated that perhaps only half of the existing fish species from that area have been formally described to the scientific (and aquaristic) community. Although formally named, Satanoperca pappaterra is another one of those surprises, and a worthy and beautiful Cichocki, F. P., 1976. Cladistic History of Cichlid Fishes and Reproductive Strategies of the American Genera Acarichthys, Biotodoma, and Geophagus. Dissertation, University of Michigan. Gosse, J-P, 1976. Revision du genre Geophagus (Pisces Cichlidae). Academie Royale des Sciences d Outre-Mer XIX-3, Bruxelles, Belgium (available in translated form in The Sifter, Journal of the Geophagus Aequidens Study Group, back issues available from author.) Heckel, J., 1840. Johann batterer s neue Flussfische Brasilien s. Annin. Wien. Mus. Natgesch. 2: 327-470. Kullander, S. O., 1986. Cichlids Fishes of the Amazon River Drainage of Peru. Monograph, Swedish Museum of Natural History. Kullander, S. O. & N. Nijssen, 1989. The Cichlids of Surinam (Teleostei: Labroideai), E. J. Brill, Leiden, The Netherlands. Leibel, W. S., 1983a,b. On the Identity of Geophagus acuticeps Heckel 1840. Buntbarsche Bulletin 94: 2-10; 95: 11-19. Leibel, W. S., 1985a,b. Maintaining Neotropical Geophagine Cichlids. Buntbarsche Bulletin 109: 3-12; 110: 2-11. Leibel, W. S., 1987 a,b, Demonfish, the Juruparoid Eartheaters. Freshwater and Marine Aquarist (FAMA) 10 (2): 12 et seq., 10 (3): et seq. Leibel, W. S., 1988. Satanoperca jurupari. Cichlid Index (ACA), 9 (6): 3-4. (in BB 128). Leibel, W. S., 1990. The Cichlids of Surinam - A Review of the New Monograph by Sven O. Kullander and Han Nijssen. Buntbarsche Bulletin 136: 8-25. Stawikowski, R. & U. Werner, 1988. Die Bunbarsche der Neuen Welt, Sudamerika. Reimar Hobbing, West Germany. 20 / Buntbarsche Bulletin 140