Herichthys deppii (Heckel, 1840) - Rediscovering a Lost Species

Herichthys deppii breeding pair in the aquarium. Photo by Eric Hanneman.

We self-proclaimed intrepid explorers had collectively crossed the Rio Nautla four times before opportunity and motivation simultaneously appeared and propelled us upstream. The previous year, two photographs of a mystery fish (Glaser et al., 1996, page 94) had beckoned with an alluring temptation, a siren call, but the constraints of time and fatigue had prevented their appeal from fulfillment. A year later, after first traveling further south, the realization that "its now or never" awoke us from a fish collecting stupor and propelled the car again north towards the Rio Nautla.

Arriving at the river, we decided to avoid the toll by not crossing, so in front of the bridge the two rented Suburbans made a U-turn and went west on the road that followed the south bank of the Rio Nautla. The scenery was semi-tropical, the fields filled with sugar cane. After 20 minutes we came to a quaint town with neatly kept streets and buildings that appeared to be very old. It was near here (N 20d10.747m, W 96d49.579m) that we captured a Vieja sp. cf. fenestratus "Rio Nautla", an unexpected catch that extended the northern range of this genus by fifty miles and over a mountain range. The river here was wide and dirty. Going into the water to retrieve a snagged cast net, Jeff's shirt was spotted with brown material that did not easily wash out, probably debris from a sugar cane processing plant somewhere upstream. Unable to capture more fish, we headed west.

A long and dusty gravel road finally took us to a small stream. Throwing the nets yielded Mexican mollies and not much else. Some of us wanted to turn around and give up. The night before, we had driven to Veracruz to partake in the celebration of Mardis Gras. Those amongst us with the monster reputations (and hangovers) found that little sleep and the drone of the gravel under the wheels was making a case for a timid retreat home. We protested, "Just ten more kilometers." A rare vehicle coming the other way informed us that the water where we threw our nets was 'muy malo' (very bad), but that a river was just ahead. We drove on, none of us, all males, able to accurately gauge inches, much less kilometers.

We passed through a small town, Posada de la Reunion, and soon after came to a larger stream, the Rio Chapa Chapa (N 20d05.101m, W 96d53.603m). The bridge showed damage of very high water, the concrete guard rails having been bent and washed away by the water, currently some 10 feet (3 m) below. Some of us ran down to the water, momentarily seeing some fish which looked rather interesting. The stream had a moderate flow, picking up speed where it narrowed, but not enough to knock one over when crossed at its most rapid point. We caught a couple of the fish off a sandy bank, but the majority came from a deeper hole, where the debris of civilization had also settled, and came up regularly in the cast net. We threw our nets for a couple of hours, catching seven cichlids, one of which jumped from the net to the stream, resulting in a grand total of six fish. But what fish was this? Months later, after sending by email a photograph to Juan Miguel Artigas Azas, his response was, Herichthys deppii.

Searching the internet and the aquarium literature at our disposal has yielded many interesting facts about this name, but not a crystal clear answer as to whether this is that fish or not. Actually, Herichthys deppii as a species, has probably not changed or moved much since its original description as Heros deppii Heckel, 1840 by Johann J. Heckel way back in 1840 (Fishbase reference ID 2064). However, subsequent to its original discovery and description it has nearly disappeared from the collective ichthyologic consciousness. It never attained commercial demand from the aquarium trade, and perhaps for this reason has also been largely ignored until recently in current Central American cichlid literature. Aqualog 3 (Glaser et al., 1996) is the only publication in English that even has a picture of the fish, and although it is labeled as an undescribed species, Herichthys sp. "Rio Nautla", it played an important role in our "rediscovering" H. deppii. You can hardly blame the authors for the confusion. We had to go back in our literature to Seth Meek's "The Fresh-Water Fishes of México North of the Isthmus of Tehuantepec" published in 1904 to even find a reference to H. deppii. Even in this work we found no pictures to help in identification, just an abbreviated morphological description.

Young Herichthys deppii freshly collected. Photo by Jeff Cardwell.

Eventually our research led us "across the pond" to German literature where we were finally able to find additional information and photographs. Pictures were helpful in identification, but since neither of us can read German, at first we were limited in what information we could gather. However, we were fortunate to have met Mary Bailey at this year's ACA convention. Aside from being a renowned cichlid author and speaker, Mary provides translations from German to English. Her help was invaluable in our research. Mary translated Heckel's original 1840 description, the most relevant part being: "The present colour in alcohol is light rust-brown, whitish on breast and belly; on the posterior half of the body, specifically between the anus and the caudal fin, there are 6 darker vertical stripes with even intervals of the same width; there is an even darker spot on the centre of the last stripe. Each scale on the lower half of the body bears a longitudinal streak at its centre, all running in the same direction, so that there appear to be as many horizontal lines as rows of scales. The fins are the colour of the body, except that the base of the soft dorsal and anal is whitish and speckled with black between the last rays.". Heckel dedicated this beautiful species to Ferdinand Deppe of Berlin, who brought it back from his journey to México.

There are many challenges to working with preserved specimens. One of the most difficult is to account for the changes in coloration that preserved specimens undergo in comparison to live fish. Another challenge to ichthyologists of Heckel's day was that type specimens were typically deposited in the museum of the describer regardless of country of origin. Many times ichthyologists were working with the same fish but they only had drawings of preserved specimens to share with their international colleagues. Understandably, that left a lot of margin for human error. Aside from those issues many facilities of the day were so overwhelmed with new specimens that they did not practice fundamental organizational hygiene. In other words they either lost, mislabeled, or at times did not even correctly preserve many of the type specimens of newly discovered species of fish. This helps to explain why the former 'Cichlasoma' genus is still in a taxonomic state of disarray even in the year 2002.

Searching Fishbase (http://www.fishbase.org), H. deppii appears as a valid name (Kullander, 2001). Dr. Kullander wrote us, "Heros deppii is the oldest available name in Herichthys (if it is one), dating from 1840, and among the earliest names for Central American cichlids, so it cannot really be invalid. I found it a good idea to use the name in the sense of Stawikowski & Werner (Stawikowski and Werner, 1998), and it will appear as such in CLOFFSCA (Checklist of the Freshwater Fishes of South and Central America). The final roundup of this issue will have to be taken care of in due course, but I see no drawback in using the name".

When the genus Heros was assigned to South American fish, and all of the Central American cichlids formerly located within Heros were deposited into the catchall Cichlasoma genus, H. deppii was reclassified (Jordan and Everman, 1896) as Cichlasoma deppii. When Dr. Kullander (Kullander, 1983) restricted Cichlasoma to 12 South American species, with Cichlasoma bimaculatum as the type species, Heros deppii Heckel, 1840 was placed into Herichthys. H. deppii's new genus is defined by the type specimen Herichthys cyanoguttatus Baird and Girard, 1854, commonly known as the Texas Cichlid. That is how a fish species can end up being described 14 years before its genus' type species.

Ranier Stawikowski and Uwe Werner have published two books on Central American cichlids. In the first book (Stawikowski and Werner, 1985), they picture an undescribed species from near the town of Nautla, and the same fish in breeding dress below (page 116). In their second book (Stawikowski and Werner, 1998), they provide many details about H. deppii, and Mary Bailey generously translated the relevant text, paraphrased as follows:

"The type specimen is lost at the Vienna Natural History Museum, and Heckel gave the type locality simply as "México". In the Museum of Natural Science at Humboldt University in Berlin, there are eight cichlids catalogued as Heros deppii. They were probably collected in 1829, when Ferdinand Deppe, at that time palace gardener at Sanssouci (Potsdam), dispatched the specimens to Europe from Misantla on the river of the same name (Veracruz, México). He evidently donated the bulk of his collection to the Berlin Museum, but sold a few specimens to Vienna, where Johann Jakob Heckel (1790-1857) did the bulk of his work and the species was described. However, the 8 fish are apparently a mislabeled Paraneetroplus nebuliferus, 3 Vieja cf. fenestrata and four fish that based on shape are presumed to be Herichthys. The three V. cf. fenestrata may be the basis for the confusion with Heros montezuma, which may be a junior synonym for Vieja fenestrata. A reconstructed outline drawing of the lost type specimen based on Heckel's description and numerical data suggests that it may have been a member of the Rio Misantla species, and therefor we provisionally identify that species as Herichthys deppii. Heros montezuma (Heckel, 1840) might be a junior synonym, but may be linked to the Vieja types found in the same collection.

Such confusion! But even more confusing, not long after Heckel's description, the name H. deppii was synonymized with Tomocichla sieboldii (Kner and Steindachner, 1864) and this (i.e. H. deppii equals T. sieboldii) has been carried into the modern literature (Ufermann, et. al., 1987; Konings, 1989). However, T. sieboldii was described from "New Granada", the region that included Colombia, Panama and most of Venezuela. It is incredible that Kner and Steindachner could lump the fish from México, collected by Deppe on the Rio Misantla, with fish from Panama. These scientists were not working in the dark, unaware of each others efforts (see nmh-wien website). Kner, a graduated physician, worked for Heckel for three years, but then left 1839 for 10 years in Ukraine, eventually returning to Vienna in 1849 to chair the Zoology department. Heckel, not formally trained as a scientist, died in 1857. Kner threw out H. deppii as a species, synonymizing it with T. sieboldii, 7 years later. Speculation: the subordination to T. sieboldii may have been due to mislabeling of museum specimens; perhaps they were confused by the appearance of the fish after being preserved; maybe they performed just a cursory examination of the material; maybe there were other more personal reasons for their actions. We may never know the exact reason.

Heckel also described Heros montezuma from the same area of México. H. montezuma (Heckel, 1840) was also synonymized with T. sieboldii by Kner. As noted by Stawikowski and Werner, H. montezuma also may be a valid name, and could refer to any of the Vieja species we found in several rivers going north from Veracruz to Nautla. Whether these fish are truly different from Vieja fenestrata, only time and/or DNA analysis will tell.

Heckel, who was not formally trained in ichthyology, was still regarded as one of the most respected ichthyologists of his day. Much of his work is still standing the test of time nearly 145 years after his death, a true testament to the quality of his work. However, the bulk of his work was with the ichthyofauna of South rather than Central America, so his comparisons to the related but distant South American genera Hoplarchus, Hypselecara and Mesonauta are not as useful as much of his other comparative analyses. Perhaps if Heckel had had more Central American species to work with, his works may have been that much more significant to our dilemma.

Meanwhile, back in México we were packing it up. The members of the other vehicle voted to give Eric the fish. It was a grand gesture, since he was but one of the prime motivators for finding the site. Perhaps his fanaticism for swallowing stream water (sterilized afterwards by liberal doses of México's finest) convinced them that he was better mollified than confronted. We slacked our thirst in the village Posada de la Reunion. They had an outdoor sports bar, with two pool tables, a soccer game on TV, a taco stand, and a cooler with ice-cold beer (see Morfitt, 2001). We spent many pesos and an hour or two there before driving east and then south, back to the hotel at Boca de Oveja.

A week later the fish were safely at home. The H. deppii were placed in a 55 gallon tank with a bunch of highland swordtails (Xiphophorus alvarezi). The smaller swordtails didn't last very long, so the H. deppii were moved into another tank. There, after 8 weeks, they spawned for the first time at about 3" TL. Remarkably, there was initially little color change in the brood caring fish, compared to Herichthys carpintis "Laguna de la Puerta", found farther to the north, or especially compared to the geographically closer Herichthys sp. "turquoise", which exhibit dramatic color changes, turning from the normal turquoise, to nearly completely white and black markings. As the H. deppii matured, their spawning colors changed too. The H. deppii female eventually gained the dark lower mask characteristic of H. cyanoguttatus, but the male got only a dusky chin. During this time one of the H. deppii was killed. The main pair was removed and promptly spawned. The fry were divided into 4 tanks. The fry are fairly easy to raise. They accept baby brine shrimp and other fine foods. Like other Central American cichlid fry, they need clean water. When overcrowded, they can be prone to developing bloat, which in fry is basically incurable, so do not try to raise too many in too small of a tank.

All five remaining adults were then placed into a 100 gallon tank with four Astatheros rostratus, a large male Thorichthys maculipinnis, and a pair of Cryptoheros sp. "Honduran red points". The dominant pair of H. deppii spawned again but lost the fry the day after they were free swimming. The large T. maculipinnis was removed, and things really changed. The pair of H. deppii spawned again. A pair of A. rostratus began to come into their spawning coloration with a blackening of the area under their operculum and began to guard a territory close by. The red points spawned at the far end of the tank. Then a second pair of H. deppii took over the flower pot formerly occupied by the T. maculipinnis and laid eggs. By now the female H. deppii had fairly normal Herichthys breeding color, though she was not solid black caudally, and in the tail retained some stripes. The male only turned black in the chin, his body only showing the striped pattern, though lighter than the female. Instead of the fry being eaten right away, the dominant pair of H. deppii guarded the babies for a week. The colors of the adults continued to change, the unpaired fins having taken on a rosy hue. With the pair currently guarding a spawn of several hundred fry, they appear very dusky, almost gray, totally unlike their youthful appearance, and totally unlike any other Herichthys we have ever seen.

So is this fish Herichthys deppii? It is basically the same fish as pictured in Stawikowski and Werner (1985) page 116, Stawikowski and Werner (1998) page 333, Aqualog 3 (Glaser et al., 1996) page 94, and probably the undescribed "Cichlasoma sp." on page 189 in Konings (1989). Stawikowski and Werner (1998) say that the reconstructed line drawing of H. deppii, based on Heckel's 1840 measurements, matches the Herichthys types found in the collection of Deppe in the Berlin Museum. But most telling is the original description by Heckel, "Each scale on the lower half of the body bears a longitudinal streak at its centre, all running in the same direction, so that there appear to be as many horizontal lines as rows of scales." This time, the pictures speak for themselves.

Our research to "rediscover" this new and yet very old species first led us across the back country of the East coast of Central México. Then, on a literary trail from the United States to England, Sweden, Germany, and Switzerland in our efforts to verify that what we had actually discovered was what we had originally set out to find. The mixture of Herichthys, Vieja, and Paraneetroplus in the collections of Deppe suggest that more surprises await future explorers. We found a Vieja species with Herichthys deppii in the Rio Nautla, and further south in the Rio Antigua, we found Vieja fenestrata and Thorichthys maculipinnis, but not Paraneetroplus, which is, however, in Deppe's collection. It is what we did not yet find that will propel these self-proclaimed intrepid explorers south again.

We would like to extend our thanks to Mary Bailey, Sven Kullander, Michi Tobler and Juan Miguel Artigas Azas for their gracious assistance in our research. Also thanks to Rusty Wessel, Dan Woodland, Joe Middleton, James Maney, Steve Lundblad, Craig Morfitt, Ian Tapp, Jeff Cardwell, Jason Barrett, Randy Parnham, and Viral Surati for being our companions.