(This article was originally published in Freshwater and Marine Aquarium Magazine, February & March, 1987;. It is here reproduced with the permission of author Wayne Leibel.
Satanoperca jurupari from the rio Mormon in Peru. Fish and Photo by Lee Newman.
When the sobriquet "Eartheater" is dropped in casual aquaristic conversation, the fish that comes most readily to mind is the neotropical cichlid Satanoperca jurupari known for its long-nosed metallic splendor and its curious habit of sifting substrate in search of food. In fact, the genus Satanoperca Günther 1862, as currently understood consists of six valid species sensu stricto. For the neotropical cichlid aficionado, few natural taxonomic groupings of species provide the range of aquaristic experience and challenge as the South American Geophagines. Some of the most easily maintained and readily spawned cichlids, as well as some of the most difficult, hail from these genera. Moreover, the species comprising this assemblage exhibit the full range of cichlid reproductive behaviors from simple substrate spawning to delayed (primitive) and immediate (advanced) mouthbrooding. Members of the jurupari species complex: jurupari, daemon, and acuticeps, offer the advanced aquarist more than adequate challenge in acquisition, maintenance, and captive propagation, and hence are the focus of this article. I hope, in the next few pages, to review the biology and captive maintenance of this suite of three closely-related and highly interesting neotropical cichlids.
Satanoperca jurupari Heckel 1840
Heckel described the flagship of this triumvirate, Satanoperca jurupari, in 1840 from collections made by Johann Natterer in Brazil. The specific nomen, jurupari, is a transliteration of the Tupi word for "Forest Demon" or "Devil" and derives from the native name for these fish as reported by Natterer: juruparipindi or "devil's fishhook" (lure). No further explanation is given by Heckel, but the South American Indians who feared the deep jungle after nightfall, and who believed the demon Jurupari, in various animal forms, lured unwitting hunters to their deaths, perhaps attributed similar demonic properties to this most bizarre looking, long-snouted fish.
Like all Satanoperca sensu stricto, jurupari possesses the characteristic lobed first gill arch and numerous setiform rakers on the free edges of all four paired gill arches that enable fish of this genus to effectively sift the substrate for food. The latter adaptation, coupled with the long nose and underslung, protrusible mouth of the juruparoids makes these three species extremely efficient sifting machines. This propensity for substrate sifting is reflected in the Portuguese name for these fish: pappaterra, which translates to eartheater and which is the source of Heckel's Latinized nomen Satanoperca, the specific name of S. pappaterra and the vernacular name of these fishes.
In his revision of the genus Geophagus, Gosse (1975) had occasion to inspect the type specimens of several jurupari look-alikes. jurupari is widely distributed throughout the Amazon drainage, the Orinoco drainage, and throughout the Guiana Highlands. As is true for most fishes with such cosmopolitan distributions, jurupari exhibits considerable geographically-based variation. Although Gosse (1975) found statistical bases on which to split the species along geographic lines, he conservatively elected to retain all populations under the species name jurupari. Moreover, examination of Heckel's (1840) S. pappaterra from the Mato Grosso, Meuller and Troschel's (1848) leucosticta from Guyana, and Fernandez-Yepez' (1950) mapiritensis from Venezuela, revealed that all fell within the normal range of S. jurupari and thus were synonymized with the latter by Gosse (1975). All jurupari morphs share the same basic morphology, differing in the relative height of head and body with respect to length. All living morphs that I have seen have in common the parallel metallic flank striping and the conspicuous black splotch in the upper corner of the caudal peduncle (Figure 1). The base color of the fish is variable from silvery to yellow-bronze and the color and extent of facial vermiculation and fin spotting varies as well. When stressed, all jurupari morphs express a dark horizontal stripe from peduncle to eye and several indistinct vertical bars (Figure 7). Satanoperca jurupari is sexually isomorphic with respect to size, finnage, and coloration. The largest specimens examined by Gosse (1975) reached nearly 20 cm. in standard length (7.8 inches), and I have seen several adults of that size and larger in Agassiz' preserved material from the Thayer Expedition to Brazil (1865-1866) at Harvard's Museum of Comparative Zoology (Leibel 1983).
Satanoperca daemon Heckel 1840
Satanoperca daemon was also described by Heckel from material in the Natterer collection. Though the Tupi likewise refer to this fish as "juruparipindi," Heckel astutely noticed differences in their coloration and described it as a second species; daemon, Latin for demon or devil.
Superficially, daemon and jurupari are similar in overall appearance (see Figure 3). There are several major differences however (see Table 1). In daemon, the last five rays of the soft dorsal are produced to magnificent black filamentous trailers that hang back past the caudal fin. In newly-imported specimens, the caudal and anal fins are blood red, fading to orange with prolonged captivity (presumably due to a diet insufficient in carotenes). The jurupari peduncle blotch is, in daemon, larger, darker, and ringed conspicuously in white. This peduncle ocellus is diagnostic for this species in both living and preserved animals. Unlike jurupari, daemon bears one or two mid-lateral splotches on its flanks; the actual number reflecting geographic origin. Most of the currently imported daemon are Colombian and these typically have two mid-lateral splotches in addition to the peduncle ocellus (Figure 3). Like jurupari, these daemon have parallel iridescent striping on every other scale row. Occasional Brazilian "daemon" have turned up this year as contaminants of jurupari shipments. Unlike their Colombian cognates, these have a single mid-lateral blotch, and are gun-metal gray in base coloration and lack the iridescent striping (Figure 4). Moreover, whereas the two mid-flank splotches in the Colombian fish occur below the upper lateral line, the single-splotch of the Amazonian morph is on and above the upper lateral line (see Axelrod l985, page 356 for an excellent photo of this animal). Heckel (1840) mentions having seen several one-spot daemon in the Natterer collection. Gosse (1975) examining over fifty examples, principally Amazonian in origin, notes that the two-spot (Colombian) daemon is rare in his sample, but speculates no further. Occasional four-spot daemon have turned up. Whether these are simply geographic variants, or distinct species, remains to be investigated.
Like jurupari, daemon is sexually isomorphic and reaches comparable sizes: Gosse (1975) records a maximum standard length of 29.6 cm. (11.6 inches) for this species. I have never personally seen a living specimen this large in aquaria, but this may be due to the relative fragility of this species in captivity (see below). S. daemon and acuticeps both occur sympatrically with jurupari and are often contaminants in shipments of the latter, but their range is restricted to the Amazon drainage with daemon crossing the Casquiare into the Orinoco). Neither daemon nor acuticeps are reported from the Guiana Highlands. This restricted range is reflected in their relative scarcity in the trade compared to the ubiquitous jurupari.
Satanoperca acuticeps Heckel 1840
Satanoperca acuticeps was the third geophagine cichlid described by Heckel (l840) in the Natterer collection and is distinctive for its sharp head profile (acuticeps = sharp head) and for its flank blotch pattern. Satanoperca acuticeps, prior to l983, was one of the most mis-identified cichlids in the aquarium literature. Innes' (1938) assertion that acuticeps was essentially a long, ventral-finned version of jurupari has been taken as gospel for over forty years and a great number of photographs depicting various jurupari variants have been committed to print bearing this false identification. Even Loiselle (1980) in his excellent and comprehensive review of the Satanoperca species assemblage in an earlier edition of Fresh Water and Marine Aquarium Magazine, erred in this regard by pronouncing the correctly labeled figures of acuticeps in the early German literature (e.g., Sterba 1966, Holly et. al. 1934) as wrongly-labelled daemon. In an earlier essay (Leibel 1983a) I reviewed the data that supports beyond a shadow of a doubt, the identification of the fish in Figure 5 as the real acuticeps. For those skeptics, I recommend that dense but conclusive article.
Satanoperca acuticeps is separable visually from daemon by the absence of a distinct peduncle ocellus (there is a black splotch, but it is not ringed in white) and the presence of three additional mid-lateral blotches evenly spaced from eye to peduncle blotch. Like daemon, the caudal and anal fins are blood-red in wild-caught individuals and fade to orange in captivity; the last 5-6 rays of the soft dorsal are produced to long black filaments, and the flanks bear parallel rows of iridescent scales - every other row - lending conspicuous metallic striping to the fish. Unlike both daemon and jurupari, the head profile is straight and steep, not convex, the ventrals are dramatically produced (Figure 6) as suggested by Innes (1938) and, the morphology of the lower lip; "detached" (continuous) in acuticeps and "attached" (discontinuous) in jurupari and daemon, is diagnostically different (see Leibel 1983a). A visual key to all three species is offered in Table 1. This, along with the accompanying photographs of normal (Figures 1-6) and stressed individuals (Figures 7-9) should allow correct identification of all three species in your dealer's tanks. Gosse (1975) reports a maximum standard length of 16 cm. (6.2 inches) for acuticeps, but I have seen examples in the Thayer Collection (MCZ/Harvard) every bit as large as adult daemon and jurupari. Like the other two species, acuticeps is sexually isomorphic.
Top to bottom, Left to Right: (1-L Figure 2) Satanoperca leucosticta female. (1-R Figure 3) Satanoperca daemon is easily identified by the pronounced caudal penduncle ocellus. As the fish matures, the dorsal fin extensions and long ventrals will continue to lengthen. 12 cm. SL specimen of indeterminant sex from Colombia. (2-L Figure 4) Two-spot Satanoperca daemon from Amazonia. (2-R Figure 5) Satanoperca acuticeps exhibiting diagnostic four-blotch pattern. Note the absence of penduncle ocellus. 14 cm. SL. specimen of indeterminant sex. (3-L Figure 6) Satanoperca acuticeps . The blotch pattern is behaviorally expressed. It is apparent in this picture how Innes and others described acuticeps (inaccurately) as simply a jurupari with elongate ventral fins. Photo by John O'Malley. (3-R Figure 7) Fright pattern of Satanoperca jurupari (4-L Figure 8) Fright pattern of Satanoperca daemon. The black lattice-work flank pattern is characteristic of this species. (4-R Figure 9). Fright pattern of Satanoperca acuticeps. The three mid-lateral blotches are transformed to complete vertical bars separated by half bars. Photos by Wayne S. Leibel unless otherwise stated.
A Visual Key To The Juruparoids
Maintenance of the Juruparoid Eartheaters
Reasonable strategies for maintaining Geophagine cichlids have been presented by Loiselle (1980) and by Leibel (1985b) and interested readers are directed to these sources for more information. The three juruparoids are best approached as the giant Apistogramma that Satanoperca are: the genera Geophagus, Gymnogeophagus, and Apistogramma share lobed gill arches and are believed to have arisen monophyletically (i.e., same branch in the evolutionary tree) from an Aequidens ancestor. In fact, I must confess that my initial interest in Geophagine cichlids was a retreat from the demands of this oft-finicky group of minute cichlids! Apistos, I'm afraid, don't eat trout chow with much relish, nor do they tolerate laxity in water quality maintenance. As I was later to find out, other than their slightly less demanding palate and expanded requirement for tank space, maintenance of these primitive mouthbrooding Geophagines is nearly identical to that of the dwarf Cichlidae: clean and hot are the two cardinal rules in the care of juruparoids.
My preference is to maintain mixed colonies with non-cichlid dithers in relatively large (surface area: long and wide, rather than high) tanks over fine sand for their sifting pleasure. And sift they do! Incessantly. They just do not seem (and probably are not) happy in bare tanks or over larger sized gravel. They still attempt to sift the latter, and in my early days, I performed many a pebble-ectomy on a sadly inconvenienced jurupari just trying to do its thing over an inappropriate substrate. Needless to say, plants don't stay rooted for long. One obvious solution - potting and protecting with a layer of coarse gravel - usually doesn't work: the fish somehow uproot them anyway. I favor a tangle of well-cured driftwood and clumps of Java Fern (Microsorium sp.) for cover. Floating duckweed and water sprite cut down on the top illumination and, in the case of acuticeps, the duckweed is sifted and eaten. The driftwood also leaches a modest amount of humic acids which these fish seem to prefer. I prefer to keep my juruparoids in soft acid water in the 78°-85°F range hailing as they do from the Equatorial regions of South America.
Water must be kept clean! These fish react to lax water maintenance by neuromast pitting (head hole) and, in the case of daemon and acuticeps, by bloating. Cephalic erosion, when caught in its earliest stages is usually reversible by massive, frequent water changes coupled with several appropriate drugs (see below). Neotropical bloat is the single greatest killer of daemon that I know of. And it may not be directly related to water condition. The course of the affliction is predictable and has been corroborated by a number of Geophagine enthusiasts (Rambo, Eckstein, 1984; Sifter 2:1). An apparently healthy group of daemon will break down seemingly at random and with little warning, one by one. Heavy ventilation and white stringy feces are the proximal symptoms - the fish refuses food and often hangs listlessly near the power filter return. If untreated (and often despite treatment) the fish will slowly bloat, its abdomen filling with fluid, and die a slow death several weeks later that is no doubt as agonizing for the fish as it is for the owner of that fish (Figure 10). I have had large colonies of daemon from several geographic origins and importations go (swelled) belly-up one by one while co-tanked with similarly heterogeneous jurupari that remained fit. I'd cite lax water maintenance as the proximal cause (I'm not the saint I'd like to be in this regard) if it weren't for shared experiences by aquarists of considerably more meticulous bent. Naladixic acid (see below), if administered early in the game, may sometimes reverse incipient bloat in these fish.
My prescription for water management? Frequent and massive water changes coupled with slow power filtration. The former is easily understood (50%-75% straight out of the tap), the latter, however, seems paradoxical. The juruparoids favor the slow-moving waters of the flooded varzea to the Amazon River proper. A Power 600 filter on a 210 liters (55-gallon) tank just doesn't make sense. We want efficient mechanical filtration, not the Iquacu Falls! I prefer to use small power filters (c. 450 lph (120 gph): Aquatronics Power 1 or Supreme PME) which I hang at opposite ends of the 210-265 liters (55-70 gallon) tanks I favor (and can afford) with outputs oriented complementarily. This positioning promotes slow but steady circulation without blowing the fish around. Although I use only floss and achieve chemical filtration simply by changing water; charcoal, PolyFilter, or Chemi-Pure would, I'm sure, provide useful adjuncts. However, given the incessant substrate sifting of these fishes, the floss does (and filter additives would) clog quickly with particulates and feces, and must be changed weekly. I also aerate heavily, which helps turn over oxygenated surface water. This is fairly important given the high temperatures and somewhat crowded conditions I favor. If I had the room and money I'd go to 1.80 meters 510 liters (six-foot 135 gallon) tanks with larger power filters at either end as above.
The palate of these fishes is rather catholic and they are heavy eaters once purged of parasites by initial prophylaxis. Since the fish do sift, I prefer pelleted foods. In my leaner years (employment, not corporeal), trout chow was my staple; it was nutritious and it sank! It also clouded up the water. With steady employment, both my sifting friends and I have enjoyed a dramatic rise in our collective standard of living: no more trout chow for either of us! I now offer a variety of pelleted foods including shrimp pellets and most recently Hikari wheat germ and Spirulina pellets with great success. Krill and high quality flake foods are also excellent and will be taken from the surface, but again, I believe it is important to encourage these fish to do what they have been evolutionarily engineered for: sifting. Another little known food: parboiled zucchini slices which most Geophagus and many cichlids will eat the centers out of (originally suggested by Eckstein, pers. commun.). I don't feed frozen brine shrimp because exoskeleton rupture during freezing results in loss of most of the digestible mass when the shrimp are thawed (Loiselle, pers. commun.) thus making this relatively expensive food generally a waste of money. I spend my frozen food dollars on bloodworms (Chironomus larvae), which do not rupture upon freezing, and use these as a conditioning food. When fattening the more exotic and demanding Geophagines, including the juruparoids, I also rely on live red earthworms which I culture for this purpose (see Piparo 1984 for details on their husbandry).
Since most of the commercially available juruparoids are wild-caught, it seems wise to make some general remarks concerning acclimatization of wild fish. Newly imported Saatanoperca are often in terrible condition: emaciated, usually parasitized, and in need of some immediate (and often pharmacologically supported) TLC. Coming as they do from rather invariably hot environments, they should be indulged accordingly in their new quarters: temperatures of 29°-32°C (85°-90°F) will often turn a marginal fish into a survivor in a matter of a few days. Discus afficionados often affect their first prophylaxis of wild specimens with temperatures in the over 32°C (mid-90s °F): many wild parasites seem less thermally tolerant than their piscine hosts and will succumb to a week of such treatment. I am not a purist, thus I have no hesitation in combining heat with drugs in the first few days post-dealer (see below).
In my experience, the juruparoids, particularly daemon and acuticeps, are usual victims of export-associated breakdown. They are often emaciated, listless, pick at all offered food (live or otherwise) and produce white stringy feces. Whether these symptoms are the result of Hexamitas, Spironucleus, or other insidious protozoans or bacteria in various permutation, is, practically speaking, irrevelant to me. My blockheaded approach: broad-spectrum drugs used calculatingly and judiciously. I have found two commercially available drugs, Naladixic Acid (Aquatronic's Naladin or Tetrid, or Aquarium Pharmaceutical's Nalagram) and/or Dylox (in combination with nitrofurazone and chloramphenical and sold as Paragon (Aquatronics) to be particularly useful prophylactic agents for newly-acquired neotropical cichlids. (Note: these should not be combined.)
Naladixic acid is proving to be an incredibly useful aquarium pharmaceutical for the treatment of several characteristically Geophagine diseases including Neotropical Bloat. It is a gram negative antibiotic which is readily and rapidly absorbed via the gills into the bloodstream and is the prophylactic of choice for wild Satanoperca. It is particularly useful for turning thin, white, stringy feces-producing animals around. A 24 hour dip as per instructions is usually sufficient.
Dylox is a powerful anti-helminthic which kills flukes and other parasitic worms. Dylox is believed by Discus afficionados to be cumulatively hepatotoxic (liver-damaging), but I have never had any obvious problems with it. Accomplished Discus breeder, Walter Cecil of Cambridge, Massachusetts, however, avoids this drug like the plague and uses, instead, Piperazine Hydrochloride. This latter drug is available from veterinarians as a solid (500 mg tablets), is dissolved in minimal water and food (dry or non-worm live) soaked in it. The drug acts as a local anesthetic for parasitic worms in the intestinal tract: the worms detach and are passed in the feces. Deworming is best accomplished in a bare aquarium and the fish removed soon after the worms are passed (i.e., a fecal smear is helpful here). My most recent experience with intestinal flukes have shown that Tetra's Medicated Food, which packs several antihelminthics in palatable form, may be useful in eliminating these parasites. The key here is palatable: most fish will eat this food readily (one problem with do-it-yourself piperazine-soaked foods, above), and should be fed it exclusively for ten days as per instructions.
One juruparoid "disease" best left untreated, are the black, pimple-like ectoparasites that often decorate wild-caught fish. These are flatworms that use cichlids and other fishes as intermediate hosts in their life cycle. They generally do not harm the fish host, infecting rather their primary host; fish-eating birds, who acquire these parasites with their food. Unless one concurrently maintains a harem of herons in the fish room, the "disease" will not spread to other, uninfected cichlids. Nothing short of sewing needle cautery will effectively remove them, and they are best ignored. Better: if these blemishes offend your aesthetic sensibility, avoid such pimpled fishes in the first place.
Breeding the Juruparoids
The juruparoids are the most difficult and frustrating of the Geophagines to spawn. Satanoperca jurupari has been spawned with some regularity and is being turned out in moderate numbers in the Far East. The trick with this fish, aside from clean, hot and well-fed, is patience. The fish apparently reach maturity at 2.5 to 3 years of age. It is also a rather shy, non-aggressive fish (at least with heterospecifics) and may be reproductively inhibited if co-tanked with other cichlids. I have spawned two pairs of Satanoperca jurupari in the past several years, and have had the good fortune to witness - and photograph - the event. Whereas S. jurupari can be somewhat rough with conspecifics, each of the two pairs were quite compatible amongst themselves. Courtship commenced with head-to-tail circling and slapping, opercular flaring, and genital butting usually initiated by the ripened female. In one of these pairs, courtship progressed to the point where the female, with tube down, swam alongside and at right angles to her consort, apparently attempting dry runs on his flanks! The male, though generally patient, responded with genital nudging that seemed to spur the female on.
Soon, both parents began scrubbing various prospective egg receptacles, finally focusing their attention on a piece of slate (Figure 11). In the wild, Cichocki (1976) reports that these fish favor movable platforms to hold their egg clutches. In one case; the sole of a shoe, in a second: a waterlogged block of wood, and in a third; a large leaf, were chosen as egg receptacles. In all three cases, the pairs tugged the spawn-bearing platforms to safety when the creature in mask and fins was noticed (see Leibel, 1985a, for a further discussion of movable platform spawning in neotropical cichlids). In the aquarium, immovable substrates; often smooth stones or even vertical slates, are acceptable.
Following several hours of meticulous scrubbing, the first eggs were laid. The female would express a line of 12-20 eggs at a time with the male following closely behind, presumably fertilizing them (Figures 12 & 13). Each succeeding track of eggs intersected, eventually forming a tight plaque of 200-300 eggs (Figure 14). These were moderately large (2mm x 1mm), amber colored, and attached via their long axes to the substrate. Egg laying was completed within one hour of its commencement. Within one hour after spawning, the eggs were covered with a fine layer of sand by both parents. Interestingly enough, even when laid on a near vertical slate, the parents still attempt to cover them - in vain - with sand! (Vince Edmondson, pers. commun.)
Both parents guard the clutch for roughly 36-48 hours before chewing them free of their egg shells and gently uptaking them into their mouths for further incubation (Figure 15). Satanoperca jurupari are biparental delayed (primitive) mouthbrooders who cooperate fully in the incubation and rearing of fry. It is not uncommon for extra eggs to remain on the substrate - apparently throat space is limited. These will fail to hatch unless removed for incubation in a special modified sponge-filter hatcher (Leibel 1984a) devised for this purpose. Unlike the eggs of mouthbrooding African cichlids, Geophagine eggs can not simply be reared in drum bowls with aeration owing to the delicate nature of the embryo after hatching from its protective shell (chorion) at 36-48 hours post-spawn, just prior to parental uptake.
Fry are released c. 48 hours 26.7°C (80°F) after their uptake, or 84-96 hours post-spawn. They resorb their yolk sacs for an additional four days at which time they feed voraciously on newly-hatched Artemia and finely-ground prepared foods. The adults continue to shelter the fry for 2-3 weeks post-release, offering them buccal shelter when threatened. When alarmed, parents assume a head-down position, sculling slowly backwards with mouth open. The fry respond to this parental behavior by schooling rapidly towards the mouth and diving deep into the throat. The behavior is apparently an innate (genetically-programmed) aspect of their behavior (see Leibel 1983b) which is not unexpected for any behavior with such important survival value. When danger is passed, the fry are released by spitting, or they are expelled gently through the gills to resume foraging under the parent's watchful eyes. Growth is moderate and lengths of 38-53 mm (1.5-2 inches) are attained within the first year. Fry should be removed within three weeks of release otherwise they will be eaten by the pair as they ripen and prepare for another spawning.
Satanoperca daemon and acuticeps have not, to the best of my knowledge, been spawned yet in captivity, but it is expected that these will prove, like jurupari, to be delayed, biparental mouthbrooders. Here, then, is a challenge for the rabid Cichlidophile. S. daemon is available regularly but, as mentioned previously, it is particularly delicate and prone to bloat. S. acuticeps is only infrequently available, seems a bit hardier than daemon, but less so than jurupari. Both are less shy and more aggressive than jurupari. Hot, clean, and well-fed go without saying as prerequisites for captive spawning of these two juruparoids. Probably patience as well: they may reach sexual maturity as slowly or more slowly than jurupari. It may well be that spawning may be elicited only by high temperatures (c. 35°C - 95°F) as is true for several difficult wild cichlids (e.g., Acarichthys heckelii, Leibel 1984b). Hopefully some of you will take up the challenge and be the first to spawn either of these highly desirable Geophagine cichlids. And even if you don't succeed, the aesthetic and aquaristic pleasures to be had in successfully raising a shoal of any one of the juruparoid triumvirate to healthy adulthood are considerable and truly a test of one's aquaristic virtuosity.
Top to bottom, Left to Right: (1-L Figure 10) Neotropical bloat is the single greatest killer of captive Satanoperca daemon and it is apparently stress related. The heavely pigmented "Half black" face of this unfortunate, results presumably from secondary bacterial damage to the nerves controlling melanocyte expansion. A very sick fish! Photo by Vince Edmondson. (1-R Figure 11) The male Satanoperca leucosticta cleans the intended spawn receptacle prior to spawning as the female looks on. (2-L Figure 12) The female lays down a track of 12-20 eggs egsg per spawning pass, The eggs are moderately large and attached by their long axes to the substrate. (2-R Figure 13) The male waits to fertilize each egg track as they are laid and alternates with the female. (3-L Figure 14) The female guards her tight plaque of 200-300 eggs for 36-48 hours prior to buccally uptaking them. Several hours after this photo was snapped, the parents covered the eggs, as is normal, with a fine layer of gravel. (3-R Figure 15) Here the male chews the last of the hatching fry out of their shells. Both parents cooperate in teh buccal incubation of the fry for c. 48 hours more prior to their release and will continue to give them buccal shelter when danger threatens for several weeks post-release. Photos by Wayne S. Leibel unless otherwise stated.
- Axelrod, H., et. al, 1985. Dr. Axelrod's Atlas of Freshwater Fishes, TFH, Neptune, N.J.
- Cichocki, F., 1976. Cladistic History of Cichlid Fishes and Reproductive Strategies of the American Genera Acarichthys, Biotodoma, and Geophagus. Dissertation, Univ. Michigan.
- Gosse, J.P., 1975. Revision du Genre Geophagus (Pisces: Cichlidae). Mem. Acad. Roy. Sci. Outre Mer (Bruxelles) 19(3): 1-172.
- Heckel, J., 1840. Johann Natterers Flussfische Brasiliense. Ann. Mus. Wien 2: 325-471.
- HoHy, M., H. Meinken, and A. Rachow, 1934. Die Aquarienfische im Wort und Bild. Alhed Kernen Verlag, Stuttgart West Germany.
- Innes, W.T., 1938. Exotic Aquarium Fishes. Innes Publ. Co., Philadelphia.
- Leibel, W.S., 1983a. On the Identity of Geophagus acuticeps. Part 1, Buntbarsche Bulletin 94: 2-10, Part 2. Buntbarsche Bulletin 95: ll-19.
- Leibel, W.S., 1983b. Fry Escape and Buccal Sheltering Behaviors in Geophagine Mouthbrooders. Buntbarsche Bulletin 99: 2-7.
- Leibel, W.S., 1984a. A Sponge Filter Hatcher for the Artipcial Rearing of Geophagus. The Sifter 1(1): 12.
- Leibel, W.S., 1984b. Heckel's Threadfinned Acara. FAMA (Jan.), p. 15 et seq.
- Leibel, W.S., 1985a. Movable Plaiform Spawning in an 1ncreasing Number of Neotropical Cichlids. Buntbarsche Bulletin 107: 2-11.
- Leibel W.S., 1985b. Sifiings, Tips for the Maintenance of Neotropical Geophagine Cichlids. Part 1 Buntbarsche Bulletin 109: 3-8, Part 2 Buntbarsche Bulletin 110: 2-11.
- Loiselle, P.V., 1980. South American Eartheaters. Geophagus, the Genus and its Allies. FAMA (June), p. 23 et seq.
- Piparo, A.J., 1984. Culturing and Feeding Red Worms. FAMA (March), p. 5 et seq.
- Rambo, C. and G. Eckstein, 1984. Disease Roundtable: Neotropical Bloat. The Sifter 2(1): 15.