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'Aequidens' sapayensis: The Gold Acara
by Dean Hougen
Acquisition
I wasn't paying much attention to the auction, like most cichlid auctions, this one was dominated by overpriced African, rift-lake cichlids. The few "South American" cichlids being sold were mostly Discus and Angels, which had been selectively bred for many generations so that they would not resemble their beautiful ancestors. So, I was talking quietly with a friend sitting beside me while the auctioneer ran through the fish.
"Going once."
The friend on my other side nudged me.
"Going twice"
Some sort of rare Aequidens, he told me.
"And ..."
"Yo! I yelled, shooting my arm in the air."
The auctioneer took my bid, looked for another, and sold me the fish. I don't remember exactly how much I paid, but I thought they were quite the bargain. They could have been $200.00, for all I knew when I bid on them, but fortunately they weren't. They were more along the lines of what you might pay for Blue Acaras in a store.
Interestingly enough, that's just what they looked like: three young Blue Acaras. (Blue Acaras are supposedly 'Aequidens' pulcher, but they have been bred commercially for so long that it is difficult to ascertain what wild stock they might have originated from.)
Hmmn.
![Fin Organism Fish Marine biology Underwater]()
"Aequidens sapayensis. Probable trio," the bag read. Well, one was larger than the other two, but it seemed a bit premature to be sexing them at 1" long. Still, if these really were what they were said to be, I hoped that the size difference really did reflect a sex difference as well. The auction was in Chicago, so it was late the next evening before they were placed in a tank at my home in Minneapolis. I had changed some of their water at the hotel the evening of the auction and again the next morning, but the water looked rather foul in their bag when I got them home. The seemed to adjust well to their new tank, however.
Two days later I found one of the smaller fish dead. I couldn't tell if the stress of the trip or one of its siblings killed it, but my probable trio was now a probable pair. I kept the label from the bag, which listed the seller's name and phone number, in case my probable pair turned into a definite loner.
Identification
I hadn't heard of 'Ae.' sapayensis before, so my first step was to pull out my copy of Die Buntbarsche der Neuen Welt: Südamerika. Sure enough, there they were, in the chapter entitled "Der Buntbarsche der ..Aequidens" -pulcher-Gruppe." There was a page of text, and pair of pictures, no less. All this told me three things: (1) sapayensis is valid species, (2) they are 'Aequidens' sapayensis until they are assigned to some genus (as Aequidens has been restricted to exclude this group of species), and (3) my fish resembling Blue Acaras as they did, stood a chance of being 'Ae.' sapayensis.
That was pretty much where I let matters sit until I decided to write this article. Two years had past, the fish had bred successfully (see Keeping and Breeding, below), and I had accepted that my fish were, indeed. 'Ae.' sapayensis. As adults they still resembled Blue Acaras quite strongly, although they had horizontal rows of golden dots instead of the bluish dots of Blue Acaras. I had taken to referring to them as my "Gold Acaras."
As I prepared to write this article, however, I became curiousas to how the seller in Chicago had identified his fish. While there are a few small differences between these Gold Acaras and the Blue Acaras of the aquarium hobby, these differences are assuredly not striking. I doubt that I would notice anything unusual if I came across these fish while perusing sellers' tanks for rare fish. I doubted even more that most importers, wholesalers, or retailers would have noticed anything either.
Fortunately. I had kept the seller's tag from the auction, although I hadn't had to call him for more fish. Mike Brousil, the seller, told me that he got his fish from Steve Covolo. He also told me that Steve had either received his fish from, or at least had them identified by Dr. Wayne Leibel.
Wayne was unsure of the details (it had been four or five years now since his role in this saga), but was able to confirm that he had probably identified the ancestors of my fish somewhere along the route. He told me that 'Ae.' sapayensis were sometimes found in pet shops at small sizes being sold as Green Terrors, having been brought in from the wild misidentified. He said that based on the fact that they were found in Green Terror territory and that they matched closely with good color photos published in the German aquarium literature, he was convinced they were 'Ae.' sapayensis.
Good photos and collecting data probably provide as accurate an identification as can be reasonably had with these fish. The description of 'Ae.' sapayensis was done ninety years ago and the descriptions of the species most likely to be confused with 'Ae.' sapayensis (which are 'Ae.' pulcher. 'Ae.' coeruleopunctatas, and 'Ae.' latifrons) were made twenty-five to fifty years before that. As was the general case for descriptions of the time, these descriptions are brief and are made from a small number of specimens (one in the case of 'Ae.' sapayensis). All but one of them lacks drawings and, of course, all of them lack photographs.
Further, the published meristics for the various species are quite similar; the dorsal and anal fm spine/ray counts of 'Ae.'; coeruleopunctatus and 'Ae.' sapayensis overlap entirely. (See Table.) The case is probably even worse than suggested by the numbers in the original descriptions. Now that many more specimens are generally examined when describing a species, it is recognized that spine and ray counts can vary widely within a species. In the recently described 'Ae.' patricki, to chose an example from the true Aequidens, specimens are recorded with dorsal counts of 14-18 hard spines and 10-12 soft rays.2 If these sorts of variances are found in species of 'Aequidens' as well, it might well be impossible to assign an individual fish to any of these species on the basis of spine and ray counts alone.
Nonetheless, I examined photos of two specimens of Gold Acaras (a female from the original purchase and a male from two generations later) to obtain dorsal and anal counts. The counts of these two fish agreed with each other (not surprisingly) but do not match those from the descriptions of any similar 'Aequidens' species. (See Table.) They are closest to those of 'Ae.' latifrons but (as per the discussion above) this may well be insignificant.
The Blue Acara Group
One might tend to wonder given the similarity of meristics and general overall appearance, whether or not the species mentioned above are all valid species. This is a question that I am not qualified to answer and will not attempt to. Perhaps the best that we aquarists can do is wait for a modern review of these species by an ichthyologist.
It seems at least possible, however, that separate species of 'Aequidens' might be found where these species are. The type specimen of 'Aequidens' pulcher is from the island of Trinidad and the species is known to come from adjacent Venezuela as well. 'Aequidens' latifronsis is described from the Rio Magdalena in Columbia. The original description of 'Aequidens' coeruleopunctatus says that they originate from the Rio Chagres (in the canal zone) and from the western slope of Panama. Finally, 'Aequidens' sapayensis is named for its type locality, the tiny Rio Sapayo in Ecuador. (See map.) Quite notably, these localities fall into separate major ichthyofaunal provinces. (An ichthyofaunal province is a region characterized by a distinctive assemblage of fishes; ) It is much more common that a species will be wholly contained within such a province than that it will cross province boundaries. In fact, genera often are found completely within a single major province, although that is apparently not the case with these fish.
![Leaf Nature Organism Beak Fish]()
Keeping and Breeding
Gold Acaras resemble Blue Acaras not only in appearance, but also in behavior and in their requirements for successful keeping and breeding. Like most Acaras (but notably unlike the sympatric Green Terrors), Gold Acaras have a moderate temperament for a cichlid.
Mike Brousil reports (pers comm.) that his pair were quite timid and would not spawn if there were other fish in the tank. If isolated, however, they would spawn, at which point they would become very aggressive towards (ie: kill) any new fish introduced into their aquarium.
My own pair held their own in a tank with South American cichlids such as Acarichthys heckelii and Cichlasomataenia which were at the time of similar size. When they matured they were placed alone in a thirty gallon aquarium with an undergravel filter which had been cichlid proofed" by placing a piece of plastic window screen on top of the bottom two inches of gravel. Several rocks, pieces of driftwood and bark, and some homemade, plastic plants were present in the tank for hiding places and/ or spawning sites. The tank was lit only by the ambient room light. A rather defective heater varied the water temperature between 70° and 85° F. The water was very soft (approx. 3 DH), and the pH ranged from about 7 down to nearly 4. No attempt was made to record the temperature and pH fluctuations and correlate them with behavior.
![Map World Font Slope Rectangle]()
The larger of the fish staked out a territory at one end of the tank and the smaller fish spent most of its time hiding near the other end. The fish were fed primarily Doro-Min with a supplement of frozen blood worms. The larger fish ate greedily, but the smaller only caught an occasional food stick that floated near its hiding place or ventured cautiously forth after blood
worms. Both fish grew at about the same pace, but the larger fish grew much more robust, even fat. Then one day the larger fish showed a female breeding tube.
I was a bit surprised to see that the probable male of my probable pair was actually a female. I began to worry that I either did not have a pair (as I thought that the smaller fish was unlikely to be a male) or if Idid have a pair, that they wouldn't successfully spawn (as I doubted that such a timid male would leave hiding long enough to fertilize the eggs). So, I wasn't all that excited when I finally saw eggs. Many captive female cichlids will lay eggs by themselves if a suitable mate is not available.
The eggs were laid on a nearly vertical piece of rock at the female's end of the tank. The spawning was not observed, but the behavior of the other fish on the day the eggs were found did not differ from its previous behavior. It continued to hide on the far end of the tank from the larger fish; this helped to convince me that I did not have a breeding pair of fish.
I was surprised then when the eggs, roughly 100 in number, did not fungus on the following days. After 3 days they hatched, but the fry soon disappeared, never to be seen again. Still, I now knew I had a pair and I began to condition them with more feedings of blood worms in hopes of another spawn.
On the next spawning, there were approximately 300 eggs laid in the same location as before. This time, after the eggs hatched, the number of fry began to dwindle and the male became increasingly beaten-up. I suspected that the male was eating the fry and being attacked by the female as she attempted to defend them. I siphoned a couple of dozen fry out of the tank and set them up in a ten gallon aquarium with a sponge filter in order to raise them away from the parents. I also separated the male from the female using a piece of "eggcrate" light grating placed in the middle of the tank.
The fry in the thirty gallon with the parents grew much more quickly than those in the ten gallon tank, but their numbers continued to dwindle, although at a much slower rate than before. My guess is that now and then one of the fry in the parents' tank would find its way to the male's side of the tank and be eaten by him, but I never actually witnessed this. The presence of the divider did not significantly change the general behavior of either adult fish; the male continued to spend most of his time in hiding even though he was separated from the more outgoing female.
I decided to remove the remaining fry (now down to about a dozen) from the parents' tank. They were too big to place in with the fry previously removed from the thirty gallon, so I put them in a ten gallon tank of their own. A couple of days later I observed that there was still one young fish in the parents' tank, but I decided to leave it there.
In the next couple of weeks the growth of the single juvenile remaining in the parents' tank greatly outpaced those of its siblings in the fry tanks. I decided that it could probably fend for itself with the barrier removed and I was anxious to have the adults spawn again, so I took out the egg-crate divider.
No change in behavior was noted when the divider was removed, but a couple of weeks later the male was found dead. I imagine that he was killed by the female, but this is only speculation. In any case, this ended my attempts at spawning 'Aequidens' sapayensis, for the time being. I still have the female and the one of her offspring that grew up beside her. The remaining fry were all sold or given away to friends, local aquarists, or those attending the 1994 ACA convention. I know that at least one aquarist has bred the Gold Acaras he got from me (Vinny Kutty, pers. comm.) and now has fry from them. Also, considering the information provided by Dr. Wayne Leibel (above), it might be worthwhile to look for "Green Terrors" that really aren't.
![Eye Plant Petal Tints and shades Font]()
Bibliography
Conkel, D. 1990. "Fishes of the Balboan Jungle." Tropical Fish Hobbyist, 38 (8): 74ff.
Gill, T. 1858. "Synopsis of the Fresh Water Fishes of the Western Portion of the Island of Trinidad. W. r." Annals of the Lycewn of Natural History of New York. 6: 363-430.
Kner. R. and F. Steindachner. 1866. "Neue Gattungen und Arten von Fischen aus Central Amertka." Abhandlungen der Mathematisch - Phy sikalischen Classe der Koniglich Bayerischen Akademie der Wissenschaften. 10: 1-61.
Regan. C. 1903. "Descrtptions of new South Amertcan Fishes in the Collection of the British Museum." Annals and Magazine of Natural History. Ser. 7. Vol. 12: 621-630.
Stawikowski. R. and U. Werner, 1988. Die Buntbarsche der Neuen Welt: Siidamertka. Essen, West Germany.
Steindachner. F. 1878. "Zur Fisch-Fauna des MagdalenenStromes... Denkschriften der Kaiserlichen Akademie der Wissenschaften. Mathematisch Naturwissenschajtliche Classe. Wien. 39: 19-78
U.S. Office of Geography, Department of Interior. 1957 Gazetteer No. 36: Ecuador. Washington, D.C.
by Dean Hougen
Acquisition
I wasn't paying much attention to the auction, like most cichlid auctions, this one was dominated by overpriced African, rift-lake cichlids. The few "South American" cichlids being sold were mostly Discus and Angels, which had been selectively bred for many generations so that they would not resemble their beautiful ancestors. So, I was talking quietly with a friend sitting beside me while the auctioneer ran through the fish.
"Going once."
The friend on my other side nudged me.
"Going twice"
Some sort of rare Aequidens, he told me.
"And ..."
"Yo! I yelled, shooting my arm in the air."
The auctioneer took my bid, looked for another, and sold me the fish. I don't remember exactly how much I paid, but I thought they were quite the bargain. They could have been $200.00, for all I knew when I bid on them, but fortunately they weren't. They were more along the lines of what you might pay for Blue Acaras in a store.
Interestingly enough, that's just what they looked like: three young Blue Acaras. (Blue Acaras are supposedly 'Aequidens' pulcher, but they have been bred commercially for so long that it is difficult to ascertain what wild stock they might have originated from.)
Hmmn.
"Aequidens sapayensis. Probable trio," the bag read. Well, one was larger than the other two, but it seemed a bit premature to be sexing them at 1" long. Still, if these really were what they were said to be, I hoped that the size difference really did reflect a sex difference as well. The auction was in Chicago, so it was late the next evening before they were placed in a tank at my home in Minneapolis. I had changed some of their water at the hotel the evening of the auction and again the next morning, but the water looked rather foul in their bag when I got them home. The seemed to adjust well to their new tank, however.
Two days later I found one of the smaller fish dead. I couldn't tell if the stress of the trip or one of its siblings killed it, but my probable trio was now a probable pair. I kept the label from the bag, which listed the seller's name and phone number, in case my probable pair turned into a definite loner.
Identification
I hadn't heard of 'Ae.' sapayensis before, so my first step was to pull out my copy of Die Buntbarsche der Neuen Welt: Südamerika. Sure enough, there they were, in the chapter entitled "Der Buntbarsche der ..Aequidens" -pulcher-Gruppe." There was a page of text, and pair of pictures, no less. All this told me three things: (1) sapayensis is valid species, (2) they are 'Aequidens' sapayensis until they are assigned to some genus (as Aequidens has been restricted to exclude this group of species), and (3) my fish resembling Blue Acaras as they did, stood a chance of being 'Ae.' sapayensis.
That was pretty much where I let matters sit until I decided to write this article. Two years had past, the fish had bred successfully (see Keeping and Breeding, below), and I had accepted that my fish were, indeed. 'Ae.' sapayensis. As adults they still resembled Blue Acaras quite strongly, although they had horizontal rows of golden dots instead of the bluish dots of Blue Acaras. I had taken to referring to them as my "Gold Acaras."
As I prepared to write this article, however, I became curiousas to how the seller in Chicago had identified his fish. While there are a few small differences between these Gold Acaras and the Blue Acaras of the aquarium hobby, these differences are assuredly not striking. I doubt that I would notice anything unusual if I came across these fish while perusing sellers' tanks for rare fish. I doubted even more that most importers, wholesalers, or retailers would have noticed anything either.
Fortunately. I had kept the seller's tag from the auction, although I hadn't had to call him for more fish. Mike Brousil, the seller, told me that he got his fish from Steve Covolo. He also told me that Steve had either received his fish from, or at least had them identified by Dr. Wayne Leibel.
Wayne was unsure of the details (it had been four or five years now since his role in this saga), but was able to confirm that he had probably identified the ancestors of my fish somewhere along the route. He told me that 'Ae.' sapayensis were sometimes found in pet shops at small sizes being sold as Green Terrors, having been brought in from the wild misidentified. He said that based on the fact that they were found in Green Terror territory and that they matched closely with good color photos published in the German aquarium literature, he was convinced they were 'Ae.' sapayensis.
Good photos and collecting data probably provide as accurate an identification as can be reasonably had with these fish. The description of 'Ae.' sapayensis was done ninety years ago and the descriptions of the species most likely to be confused with 'Ae.' sapayensis (which are 'Ae.' pulcher. 'Ae.' coeruleopunctatas, and 'Ae.' latifrons) were made twenty-five to fifty years before that. As was the general case for descriptions of the time, these descriptions are brief and are made from a small number of specimens (one in the case of 'Ae.' sapayensis). All but one of them lacks drawings and, of course, all of them lack photographs.
| Species | Dorsal spines/rays | Anal spines/rays |
| 'Aequidens' pulcher | XIII/11 | III/7 |
| 'Aequidens' coeruleopunctatus | XV/10 | III/8-9 |
| 'Aequidens' latifrons | XI/9 | III/8 |
| 'Aequidens' sapayensis | XV/10 | III/8 |
| Gold Acaras | XIV/9 | III/7 |
| Table. Spine and ray counts from the original descriptions of 'Ae.' pulcher, 'Ae.' coeruleopunctatus. 'Ae.' latifrons, and 'Ae.' sapayensis and from two "Gold Acaras." |
Further, the published meristics for the various species are quite similar; the dorsal and anal fm spine/ray counts of 'Ae.'; coeruleopunctatus and 'Ae.' sapayensis overlap entirely. (See Table.) The case is probably even worse than suggested by the numbers in the original descriptions. Now that many more specimens are generally examined when describing a species, it is recognized that spine and ray counts can vary widely within a species. In the recently described 'Ae.' patricki, to chose an example from the true Aequidens, specimens are recorded with dorsal counts of 14-18 hard spines and 10-12 soft rays.2 If these sorts of variances are found in species of 'Aequidens' as well, it might well be impossible to assign an individual fish to any of these species on the basis of spine and ray counts alone.
Nonetheless, I examined photos of two specimens of Gold Acaras (a female from the original purchase and a male from two generations later) to obtain dorsal and anal counts. The counts of these two fish agreed with each other (not surprisingly) but do not match those from the descriptions of any similar 'Aequidens' species. (See Table.) They are closest to those of 'Ae.' latifrons but (as per the discussion above) this may well be insignificant.
The Blue Acara Group
One might tend to wonder given the similarity of meristics and general overall appearance, whether or not the species mentioned above are all valid species. This is a question that I am not qualified to answer and will not attempt to. Perhaps the best that we aquarists can do is wait for a modern review of these species by an ichthyologist.
It seems at least possible, however, that separate species of 'Aequidens' might be found where these species are. The type specimen of 'Aequidens' pulcher is from the island of Trinidad and the species is known to come from adjacent Venezuela as well. 'Aequidens' latifronsis is described from the Rio Magdalena in Columbia. The original description of 'Aequidens' coeruleopunctatus says that they originate from the Rio Chagres (in the canal zone) and from the western slope of Panama. Finally, 'Aequidens' sapayensis is named for its type locality, the tiny Rio Sapayo in Ecuador. (See map.) Quite notably, these localities fall into separate major ichthyofaunal provinces. (An ichthyofaunal province is a region characterized by a distinctive assemblage of fishes; ) It is much more common that a species will be wholly contained within such a province than that it will cross province boundaries. In fact, genera often are found completely within a single major province, although that is apparently not the case with these fish.
Keeping and Breeding
Gold Acaras resemble Blue Acaras not only in appearance, but also in behavior and in their requirements for successful keeping and breeding. Like most Acaras (but notably unlike the sympatric Green Terrors), Gold Acaras have a moderate temperament for a cichlid.
Mike Brousil reports (pers comm.) that his pair were quite timid and would not spawn if there were other fish in the tank. If isolated, however, they would spawn, at which point they would become very aggressive towards (ie: kill) any new fish introduced into their aquarium.
My own pair held their own in a tank with South American cichlids such as Acarichthys heckelii and Cichlasomataenia which were at the time of similar size. When they matured they were placed alone in a thirty gallon aquarium with an undergravel filter which had been cichlid proofed" by placing a piece of plastic window screen on top of the bottom two inches of gravel. Several rocks, pieces of driftwood and bark, and some homemade, plastic plants were present in the tank for hiding places and/ or spawning sites. The tank was lit only by the ambient room light. A rather defective heater varied the water temperature between 70° and 85° F. The water was very soft (approx. 3 DH), and the pH ranged from about 7 down to nearly 4. No attempt was made to record the temperature and pH fluctuations and correlate them with behavior.
The larger of the fish staked out a territory at one end of the tank and the smaller fish spent most of its time hiding near the other end. The fish were fed primarily Doro-Min with a supplement of frozen blood worms. The larger fish ate greedily, but the smaller only caught an occasional food stick that floated near its hiding place or ventured cautiously forth after blood
worms. Both fish grew at about the same pace, but the larger fish grew much more robust, even fat. Then one day the larger fish showed a female breeding tube.
I was a bit surprised to see that the probable male of my probable pair was actually a female. I began to worry that I either did not have a pair (as I thought that the smaller fish was unlikely to be a male) or if Idid have a pair, that they wouldn't successfully spawn (as I doubted that such a timid male would leave hiding long enough to fertilize the eggs). So, I wasn't all that excited when I finally saw eggs. Many captive female cichlids will lay eggs by themselves if a suitable mate is not available.
The eggs were laid on a nearly vertical piece of rock at the female's end of the tank. The spawning was not observed, but the behavior of the other fish on the day the eggs were found did not differ from its previous behavior. It continued to hide on the far end of the tank from the larger fish; this helped to convince me that I did not have a breeding pair of fish.
I was surprised then when the eggs, roughly 100 in number, did not fungus on the following days. After 3 days they hatched, but the fry soon disappeared, never to be seen again. Still, I now knew I had a pair and I began to condition them with more feedings of blood worms in hopes of another spawn.
On the next spawning, there were approximately 300 eggs laid in the same location as before. This time, after the eggs hatched, the number of fry began to dwindle and the male became increasingly beaten-up. I suspected that the male was eating the fry and being attacked by the female as she attempted to defend them. I siphoned a couple of dozen fry out of the tank and set them up in a ten gallon aquarium with a sponge filter in order to raise them away from the parents. I also separated the male from the female using a piece of "eggcrate" light grating placed in the middle of the tank.
The fry in the thirty gallon with the parents grew much more quickly than those in the ten gallon tank, but their numbers continued to dwindle, although at a much slower rate than before. My guess is that now and then one of the fry in the parents' tank would find its way to the male's side of the tank and be eaten by him, but I never actually witnessed this. The presence of the divider did not significantly change the general behavior of either adult fish; the male continued to spend most of his time in hiding even though he was separated from the more outgoing female.
I decided to remove the remaining fry (now down to about a dozen) from the parents' tank. They were too big to place in with the fry previously removed from the thirty gallon, so I put them in a ten gallon tank of their own. A couple of days later I observed that there was still one young fish in the parents' tank, but I decided to leave it there.
In the next couple of weeks the growth of the single juvenile remaining in the parents' tank greatly outpaced those of its siblings in the fry tanks. I decided that it could probably fend for itself with the barrier removed and I was anxious to have the adults spawn again, so I took out the egg-crate divider.
No change in behavior was noted when the divider was removed, but a couple of weeks later the male was found dead. I imagine that he was killed by the female, but this is only speculation. In any case, this ended my attempts at spawning 'Aequidens' sapayensis, for the time being. I still have the female and the one of her offspring that grew up beside her. The remaining fry were all sold or given away to friends, local aquarists, or those attending the 1994 ACA convention. I know that at least one aquarist has bred the Gold Acaras he got from me (Vinny Kutty, pers. comm.) and now has fry from them. Also, considering the information provided by Dr. Wayne Leibel (above), it might be worthwhile to look for "Green Terrors" that really aren't.
Bibliography
Conkel, D. 1990. "Fishes of the Balboan Jungle." Tropical Fish Hobbyist, 38 (8): 74ff.
Gill, T. 1858. "Synopsis of the Fresh Water Fishes of the Western Portion of the Island of Trinidad. W. r." Annals of the Lycewn of Natural History of New York. 6: 363-430.
Kner. R. and F. Steindachner. 1866. "Neue Gattungen und Arten von Fischen aus Central Amertka." Abhandlungen der Mathematisch - Phy sikalischen Classe der Koniglich Bayerischen Akademie der Wissenschaften. 10: 1-61.
Regan. C. 1903. "Descrtptions of new South Amertcan Fishes in the Collection of the British Museum." Annals and Magazine of Natural History. Ser. 7. Vol. 12: 621-630.
Stawikowski. R. and U. Werner, 1988. Die Buntbarsche der Neuen Welt: Siidamertka. Essen, West Germany.
Steindachner. F. 1878. "Zur Fisch-Fauna des MagdalenenStromes... Denkschriften der Kaiserlichen Akademie der Wissenschaften. Mathematisch Naturwissenschajtliche Classe. Wien. 39: 19-78
U.S. Office of Geography, Department of Interior. 1957 Gazetteer No. 36: Ecuador. Washington, D.C.
