Dr JEAN H. HUBER *
* Laboratoire d'Ichtyologie générale, Museum national d'Histoire naturelle, 43, rue Cuvier, 75231 Paris Cedex 05, France
A New species of Moema, M. quiii n.sp., is described from southern Peru, Puerto Maldonado region. It is distinguished from all other known Moema components by a unique color pattern in life for both male and female, made of small brown-bordered squares. No other Moema species is showing the diagnostic vertical bars on sides, in addition to the longitudinal pattern. The systematic status of the genus Moema is discussed: it is conservatively regarded as valid, although the author argues a personal preference to consider it as a subgenus of Trigonectes.
Une nouvelle espèce de Killi Annuel du Pérou méridional, avec un patron de coloration fascié unique, Moema quiii n.sp.
Une nouvelle espèce de Moema, M. quiii n.sp., est décrite du Pérou méridional, dans la région de Puerto Maldonado. Elle est séparée de tous les composants connus de Moema par un patron de coloration unique à la fois chez le mâle et la femelle vivants, constitué de petits carrés bordés de marron. Aucune autre espèce de Moema ne présente des barres verticales sur les flancs, en plus des marques longitudinales. Le statut systématique du genre Moema est discuté: il est considéré provisoirement comme valide, bien que l'auteur apporte des arguments personnels en faveur d'un statut de sous-genre de Trigonectes.
Peru, a small country in western South America, can be divided into 3 biogeographical regions concerning the oviparous Cyprinodont (or Killifish) fauna (Huber, 1992), with specific refugia hypothesized in the western foothills (Huber, 1998):
1- The Andes and the mountainous lakes and rivers, around 4000 meters altitude, including the large Lago Titicaca, that are dwelt by the components of the endemic genus Orestias, with 6 to 40 valid species according to authors (Huber, 2001-2002).
2- The Amazonian lowlands that correspond to the western belts of the huge Amazon river basin and that are dwelt by non-annual and annual killifishes belonging to the same groups as in the corresponding regions of Brazil, Colombia, Bolivia. The following species have been recorded (with their distribution by country between brackets with local names): Genus Aphyolebias: A. peruensis [Peru], A. rubrocaudatus [Peru], A. wischmanni [Peru], genus Fluviphylax: F. pygmaeus [Brasil (Amazonas); Colombia; Peru; Venezuela], genus Moema: M. pepotei [Bolivia; Brasil (Rondonia); Peru (?)], genus Rivulus: R. (Anablepsoides) atratus [Brasil (Amazonas); Colombia; Peru], R. (Laimosemion?) rectocaudatus [Peru], R. (Oditichthys?) christinae [Peru], R. (O.?) derhami [Peru], R. (O.?) elongatus [Peru], R. (O.?) intermittens [Peru], R. (O.?) iridescens [Peru], R. (O.?) ophiomimus [Colombia; Peru], R. (O.?) ornatus [Brasil (Amazonas); Peru], R. (O.?) peruanus [Peru], R. (O.?) rubrolineatus [Peru], R. (O.?) speciosus [Peru].
3- The foothills of the Andes, between 600 and 1200 meters altitude that are dwelt by only 3 groups of Rivulus, with counterparts in the lowlands: R. (O.?) derhami, R. (O.?) iridescens, R. (O.?) peruanus. These extend, with variations or not, to neighboring regions of the same altitude in Ecuador.
Among the Amazonian fauna, only components of Aphyolebias and Moema are annual killifishes, although recently (Huber, 1998), Rivulus rectocaudatus has been hypothesized as semi-annual. This annual or semi-annual character may seem strange in the Amazon basin where water is present everywhere, compared to the "true" landscapes of annuals (e.g. in eastern Brazil, northern Colombia and Venezuela, or in eastern Africa) that are usually dry, except during the rainy season where the full cycle of these fish takes place. However, a close analysis of their biotopes in Peru shows that they usually live in temporary biotopes, such as overflood ponds, fluvial islands, irregular creeks in flood beds where water is actually supplied for a short period in a year (and sometimes, not at all for several years).
The region of Peruvian Amazon, in terms of ichthyological collections, has been scarcely sampled except for 2 spotted areas, around Iquitos and around Pucallpa. There, a strong species diversity -speciation- has been exemplified like for Manaus, in Brazilian Mid-Amazon, with up to 7 Killifish species living sympatrically and with the species living in Iquitos usually distinct from those in Pucallpa. For example, Aphyolebias peruensis in Iquitos and A. wischmanni in Pucallpa.
The area around Puerto Maldonado was up to now even less known with 2 different samples, only, which concluded into the descriptions of 2 probably endemic species, Aphyolebias rubrocaudatus and Rivulus christinae. The new species is then the third Killifish species registered as live from the area, but a photo showing a species related to R. ornatus is available from Lance Peck and preserved material assignable to R. iridescens has been also studied (Huber, 1992) and confirmed by a photo from Lance Peck. Therefore, the speciation that is known in the Iquitos and Pucallpa areas is most probably to be met in Puerto Maldonado area, too.
Holotype: MNHN 2003-0602, male 73.4 mm SL, 99.4 mm TL. Fundo Shape, southeastern Peru, Rio Tambopata 15 Km upriver from Puerto Maldonado, 12.733°S 69.217°W, 140 m altitude. Lance R. Peck and Roberto Masias Sehue, leg. June 14, 2002, 9:00 a.m.
Paratypes: MUSM (Lima, Peru) 20295, 4 specimens, registration through Hernan Ortega, MNHNP (Asuncion, Paraguay) 33556, registration through Dario Mandelberger, 4 specimens, MNHN 2003-0603, 1 female, 1 male; BMNH 2003.6.22.1-3, 3 specimens, CAS 217427, 3 specimens; all collected with holotype. Types (9) from MNHN, BMNH and CAS have been measured and radiographed, and they have been fixed in alcohol to enable future molecular studies.
Diagnosis: A large species, with an average morphology for the genus and with extended pectoral fins in male like its congeners, but with a less pike-like morphology of the head and with a unique color pattern in both sexes, made of small, brown-bordered, squares. No other yet known Moema species is showing the diagnostic vertical bars on sides, in addition to the longitudinal pattern.
Color of Live Fish:
Male: The basic color of the flanks and the interior of the caudal, anal and dorsal fins is dark brown. Belly is somewhat lighter. On sides, 3 or 4 longitudinal continuous lines of red spots are conspicuous, each underlined and overlined by a cream yellow line and 10-12 dark brown vertical bars wipe out the longitudinal pattern where present, some being slightly oblique. The alternate longitudinal and vertical pattern produces small brown squares with 3 longitudinal lines (yellow, then orange, then yellow). A mood dependent post-opercular black blotch is available. Unpaired fins are marked by large orange and dark brown spots, more or less organized across rays. Caudal fin shows in addition an irregular dark brown line in prolongation of lower body base and a faint orange lower submargin. Both anal and dorsal fins also show a series of dark spots close to body base (3 at dorsal fin, 5 at anal). Paired fins are little darkly dotted. Head is having the 3 characteristic oblique series of yellow spots on brown background and the eye is barred with a dark oblique line, not extending beyond the orbit.
Female: The dichromism is very marked: the body is beige brown with dark longitudinal spotting that is overlined and underlined with a cream yellow edge. Three of these lines are more conspicuous, while in-between lines are thinner, more irregular, discontinuous and less dark. In addition, 7-10 vertical dark bars are present, like in male, but they are less conspicuous and restricted to between upper and lower mid sides. The unpaired fins are strongly dark spotted, rather across rays. Dark spots close to body base of anal fin are distinctive because they contrast on a light blue background, like in male. The longitudinal lighter belly zone is much broader than in male and, opposite to male, includes the lower head. The head pattern is like in male: eye barred, but the oblique pattern is less conspicuous.
Remarks regarding bars: According to Lance Peck, « they appear in sub adult males as the red pigment develops to define pattern. Adult females have the bars as faint shadows, not bold. Females appearance is of horizontal lines of spots with underlying bars. Adult males show them stronger as a result of the red pigmentation which the females lack. Juvenile sexes look alike female somewhat, but lack color as seen in adult females. Juveniles of both sexes show strong horizontal lines ».
Color In Alcohol:
Male: no bars can be seen after a while on preserved specimens exposed to dry air: bars can only be seen on very large males and females, immediately after they are pulled out from the alcoholic solution. This observation suggests that this character is exceptionally derived for this species, to the contrary to most Cyprinodonts where it is relatively primitive. Additional characters: 3 series of more prominent dark dots on sides (less so in dominant male); a post-opercular dark blotch; unpaired fins, with light, dark-ocellated, blotches; a whitish lower submargin at caudal fin; lower lip, with a dark gray underline.
Female: no bars can be seen after preservation, like in male; 3 series of more prominent dark dots on sides, like in male; a post-opercular dark blotch, like in male; unpaired fins, with dark dots and with a fin dark edge (not always in Dorsal and lower Caudal); lower lip, with a dark gray underline, like in male.
Size, proportion and formulae:
About 170mm total length, as a maximum, for the male, markedly less for the female (130 mm), measuring end of season wild collected specimens, according to Lance Peck. This is the largest Moema/Trigonectes also known so far, and also the largest Rivulinae (Huber, 2000). In South America, only members of the genera Megalebias (Aplocheilidae) and Orestias (Cyprinodontidae) include species of larger size.
Micromorphological characters: Frontal scalation is very irregular, probably of the 'D' type (rarely 'E'); the frontal neuromasts are not numerous, like for other species of Trigonectes et al. and like all Rivulus: the first anterior pair is set strongly apart, the 2 pairs of 3 supraorbital neuromasts are set in isolated holes within very shallow winding channels, the 2 pairs of 3 posterior neuromasts are fully exposed with a standard "U-shaped" organization; the pre-opercular 2 neuromasts are well exposed like in Rivulus; no neuromast or sensory buttons could be seen on the scales of sides or on fins; ctenoidy is scarce; teeth, sharp and monocuspid, are strongly recurved and very externally positioned, which together with the large mouth opening suggests a predatory behavior.
The morphological and meristic data of the 9 types in MNHN, CAS and BMNH (holotype first and in bold type; abbreviations explained in Huber,1992; * refers to "in % of SL") are, after radiophotographic confirmation, as follows:
|D||A||D/A||LL||TRAV||CIR||VER||SL (in mm)||TL*||PD*||PA*||PV*||height at anal fin*||head*||inter orbital*||eye diam.*||snout*|
Other characters: The elongated pectoral fins reach the level of tip of Ventrals in male and the level of insertion of these fins in female; the ventral fins are not elongated, unlike components of Trigonectes; number of Caudal rays varies between 34 and 36; the hypural plate is undivided, often with a narrow median (darker) ditch.
The new species cannot be placed nearer to any of the already described Moema species: none shows such a vertical pattern and all Moema species are very similar by morphology and by meristics, with similar extensions in pectoral fins of male, with identical lower orange submargin in caudal fin and with a similar post-opercular black blotch.
There are 3 distinctive color patterns in Moema males.
First, the "unequal" pattern, where some longitudinal lines on sides are more conspicuous than others (situated in-between), is seen in M. staecki (and also in the very similar M. portugali, according to Wildekamp, 2003).
Second, the "discontinuous" pattern, where series of longitudinal spots on sides are irregular and not in lines, is seen in M. piriana.
Third, the "mosaic" pattern, where posterior series of longitudinal lines become less regular to form a mosaic, is seen in M. pepotei.
Finally, a different pattern, labeled "regular", where all lines are regular and equal, up to the peduncle, like for the related Renova oscari, might be available in still poorly known populations from Ecuador and northern Peru, which may represent a distinct species.
With its "irregular" pattern, only in female, the new species may be tentatively related to the widespread species, M. staecki, from Brazil and British Guyana. However, the live pattern of the female of M. pepotei is not known either and a relationship of quiii with that species cannot be ruled out, because it is its probable vicariant (geographic neighbor).
Remarks on genus assignment to Moema:
The new species, quiii, is conservatively assigned to the genus Moema which is externally defined by a less pronounced pike-like snout than Trigonectes, the extension of pectoral fins in male, a lineated pattern in male and female, a lower orange submargin in male caudal fin, an annual behavior (Costa, 1993). The shape of the caudal fin is, unexpectedly, quite variable among Moema species: fan-like in piriana, lanceolate in some populations of staecki and spade shaped in others, without clear geographic pattern, extended in the new species, like in Aphyolebias peruensis. Thus it cannot be considered as a diagnostic character. On the other hand, in Trigonectes, snout is more pike-like and male ventral fins are extended and pectoral fins are not extended, while the lineated pattern is very similar.
However, after having studied and described the atypical species, Trigonectes aplocheiloides (Huber, 1995) and the distant population of Moema staecki from British Guyana, Moema has also been considered, with Renova, as a subgenus of Trigonectes, following a PAUP analysis of 74 characters (Huber, 1999).
With the new species, there remains only one diagnostic character for Moema, compared to Trigonectes, that is the extension of the male pectoral fins. This may be evaluated as too limited for a genus diagnosis, notably for 2 groups of allopatric phylogenetic populations replacing vicariantly each other and this is the present author's opinion that they should only be systematic subunits, notably to gather ("lump") their similarities. The name of the species would be hence Trigonectes quiii.
However, present Neotropical systematics (Costa, 1998) tend to split a lot, compared to past strategies (e.g., Parenti, 1981) and they recognize valid genera based on similar, small species groups. And to stick to the current consensus, the author has accepted with caution Moema as a valid "unstable" genus (Huber, 2000, 2001-2003).
Like other Moema, the new species shows preference for not large, stagnant temporary bodies of water (ponds, swamps, flooded areas, river islands) on a muddy bottom in the forest; dwells upper layers, near surface, in deep parts of the biotope; not very active in protected shadowed places (Huber, 2000).
According to Lance Peck, « pools form in November, March is about mid season with adult fish, pools begin to dry in May, in June most are dry or so, low water quality does not support life well and predation by birds and mammals (peccary) finishes the season. However, it varies from year to year and from pool to pool. Sometime in June there is no more water or fish. Early season collections yield about 60% females, mid season collection yield 60% males, late season yields 80% female, end of season yield 99% female; fish hunt near water edges but retreat to deep water at first sign of danger; months when the fish can be caught in the field: January to May; primary and secondary mixed forest; always in deep shade, only found in large seasonal pools with no inlet or outlets ».
Growth: According to Lance Peck, « within a pool the fish seem to all be the same age. But there is a different growth rate within individuals. In early season for instance we catch a few (2 or 3) completely adult fish, which seem to have grown at a very rapid rate, seemingly almost too fast for the time the water has been present. Within a few months all seem to be the same size ». Comment: later offspring or slower grower may also have been swallowed by predators.
Physico-chemistry and description of type locality: According to Lance Peck, « rain water with only dissolved organic compounds no sediment, acidic increasing as season progresses; temperature of air: 26-38 °C, and of water, 26 °C (on average), depth of water: 1 meter, pool bottoms taper gradually to 0 depth, type of bottom: mulm over sandy clay forest floor; many terrestrial plants (species survive and remain green throughout the inundation but many die and rot quickly) and no aquatic plants in the seasonal pools ».
Sympatric species: Early season, Aphyolebias rubrocaudatus; later season: Hoplosternum cf. pectorale, Hoplias cf. malabaricus, Erythrinus sp. Note: different, rather polychromatic, populations of A. rubrocaudatus are always found with the new species but the new species is not always found within pools of Aphyolebias.
Predators: According to Lance Peck, « Erythrinus and Hoplias feed on everything including each other. They are very aggressive predators. Erythrinus if kept together in a tank will always end up a population of one. However, do they eat juvenile Moema? Maybe not. The individuals of Hoplias and Erythrinus are not present at first. Or at least not as adult or sizable fish. We don't really see these predators until near mid season at which time the Moema are no longer juveniles».
This issue - which are probable predators of the killifishes, if any? - would be an excellent subject for someone to do a study on. These renown predators do not access the biotope directly because the biotope is not connected to any permanent water. According to Lance Peck, « perhaps eggs are held over in the soil until next season? Natives say the fish bury into the mud until the next rains. But a lack of adult fish early season casts doubt on this. But maybe a few adult fish do and reproduce with first rains? Both species, especially Erythrinus, are capable of land travel. Besides, Erythrinus are strong and resist death. When place on land they travel rapidly and efficiently in search of water. If kept in an uncovered aquarium, they exit and "walk" away towards the river. So perhaps the predators migrate from pool to pool, hatch from resistant eggs, hibernate in mud, and / or travel as fry from pool to pool in fur of mammals».
These observations should be correlated to Brosset's regarding the aggressive mimicry by the Characid Fish Erythrinus erythrinus towards Rivulus agilae in French Guyana: there, Erythrinus juveniles exhibits the characteristic pattern of the supracaudal ocellus of female Rivulus to attract the male and swallow it.
M. quiii prefers to live in tribes like Rivulus; annual development: both sexes dive, like Cynolebias et al., into substratum to lay each egg; aggressivity between males; straight, not loose posture in life, unlike Rivulus. Like Rivulus sp., M. quiii is capable of and does travel by land to change water bodies (Huber, 2000).
According to Lance Peck, « fish are good jumpers and can cover a good distance quickly when out of water; very strong and recover from dry land experiences; position in water of the new species vs. the sympatric Killifishes: active early morning and late afternoon, midday spent beneath leaf cover on bottom, near edges of pool; in aquarium shows active feeding behavior at night feeding on small fishes; on the contrary, Aphyolebias rubrocaudatus is active during daylight and goes beneath leaf cover at night; the two species which occupy the same water bodies have opposite behavior habits. The bigger Moema does feed on the smaller Aphyolebias (in an aquarium); the two fish have very similar behavior but since the bigger Moema does feed on the smaller Aphyolebias, opposite time schedules allow both to occupy the same small body of water ».
According to Lance Peck, « both Moema and Aphyolebias species live near edges but retreat to deeper water when approached from the shore. However if approached from the water side both species will take cover into leaf litter along the shore. Even to the extent of almost leaving the water. Refuge is always taken below leaf litter ».
M. quiii is only known from the type locality and from one other location along the Madre de Dios, 20 km North East of the type locality.
The genus Moema is a species flock of large to very large size, with a large range in Amazonian lowlands of Northern Brazil and its belts in Venezuela, British Guyana, Bolivia, Peru, Ecuador (at least), along the North-South (Madeira-Orinoco) and West-East (Amazon) axis. The new species lies at its western most part of range. It is northerly replaced in the Iquitos area in Peru by the "regular" yet undescribed phenotype and southerly by M. pepotei. Easterly it may be replaced by M. staecki or another unknown component. The extension of its range, apart from the Puerto Maldonado area, is unknown.
The species name, quiii, is a noun in apposition and is proposed by the discoverer Lance R. Peck, a biologist, animal photographer and exporter, based in Puerto Maldonado, as a dedication to his wife Belinda who was given the name "Qui i i" by an Ese eja shaman friend years ago.
According to him, « the name is derived from the Ese eja word "Qua' i i" which means hummingbird (picaflor). However, "Qui i i" (with a twist of sound) is the name for the "hummingbird of the water" ».
The story in more details deserves to be reported, for ethnological reasons (even if the records of epilepsy have obviously no scientific basis and Killifish are routinely eaten in Africa by locals).
The reader should remember that this vernacular language is not a language of words, it is of sounds, very subtle sounds created by humans as a language.
The Tambopata native group of Ese eja has now been reduced to less than 30 mixed families. The language is oral only. Words are invented when an unknown object is encountered. Words are created in a descriptive manor. Only one man Roberto Masias Sehue, the co-discoverer of the species, is accurately fluent in both Ese eja and Spanish and among the families, he is the only individual who cares to carry on the dialect. The older members refuse to speak the language and the younger ones don't want to learn it. Roberto Masias Sehue assisted in the compilation of a Spanish/Ese eja dictionary in 1980 (Lexico Ese-eja - Espanol, Espanol - Ese-eja, by Maria C. Chavarria Mendoza). According to him, « the author of the dictionary did not hear the delicate sounds of the language and would not/could not spell the words correctly. She omitted the sound of the letter Q from the language. Her spelling of the word for hummingbird is "Kua' i i", but the difference between Q and K is important and in fact the sound is represented by the Spanish word "Qua' i i", not "Kua' i i" ».
Now the change from "Qua' i i" to "Qui' i i": Roberto Masias Sehue explains the word is the sound of an echo. The audio call of the hummingbird is like an echo thus the trailing "i" sound. So when the hummingbird echoes, the sound transforms into the water world and changes to "Qui' i i"».
Just as hummingbirds have strong sprits according to the forest people, so do the "big red shuyu" (our killies, there), and according to natives, one should never eat killies because they possess the "madre de epilepsy" (spirit of epilepsy). If you eat them you will have seizures. This is reported to have happened one year when the people were very hungry and so went out to gather the plentiful fish and feasted on them. Everyone had seizures and many people died. The fish have never been bothered since and have no specific local name ».
Pronunciation of the name (in English): "Kwee-e-e" (in French and Spanish: "Kwi-i-i", in German: "Kui-i-i"). Common name: hummingbird killifish.
In the Aquarium:
Like other Moema sp., the species is annual, incubation in dry peat being in the order of 6 to 8 months, with sexation being reached at the age of 4 weeks and breeding at 6 weeks (Brousseau, 2002 and personal comm.); it is difficult to keep, as is the case with other Trigonectes and Moema species, because of the frequent fights between males and consecutive wounds, even if minor, and because of their short lives, rarely more than 10 months in artificial conditions (and much less in nature !). Community tanks are not advised with smaller fish because of the large size (and mouth !) of the new species and because of its predatory behavior towards the "gentle" Aphyolebias sp.
This work would not have been possible without the generous observations and collections in the field by Lance R Peck and Roberto Masias Sehue, without additional field collections by Jesus Masias Palsa, without the logistic support by Pascal Buffard (Paris), without the aquarium care at Gone Wild Peru SAC in Puerto Maldonado by Belinda Peck and Elizabeth Raine, without the advices on the manuscript by Jacques Daget from Paris MNHN and by an anonymous reviewer, without the kind help by Martine Dessoutter, Rémi Ksas and Patrice Pruvost in MNHN premises and without the kind support for types deposit by Bill Eshmeyer and David Catania (CAS), Oliver Crimmen and Tony Gill (BMNH), Hernan Ortega (MUSM) and Dario Mandelberger (MNHNP). All are warmly receiving our best thanks.
This paper is kindly dedicated to Dr Roger Brousseau, an internationally renown Californian aquarist, specialist of annual neotropical killifishes and innovative writer on killifish breeding behaviors, for his extra-ordinary welcome in San Francisco, last year with my son Nicolas, and assistance to visit the Death Valley and its Cyprinodon marvelous inhabitants.
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Huber, J.H. 2001 - 2003. Killi-Data online < http://www.killi-data.org >. A global Website dedicated to Oviparous Cyprinodontiformes or Killifish, with a Data Base on all known Taxa (files Moema, Moema pepotei, Moema staecki: 2001).
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Peck, L. 2001 - 2003. Gone Wild Peru < http://gonewild.net >. A global Website dedicated to the biology, natural history, digital cinematography, aquarist groups collection and commercial exportation of live fish from Madre de Dios department (Aquarium section).
Wildekamp, R.H. 2003. A World of Killies. Atlas of the Oviparous Cyprinodontiform Fishes of the World. Vol. 4 Amer. Killifish Assoc. Publ., in print.
male and female of the new species, Moema quiii, by Lance Peck
male and femaleof Aphyolebias rubrocaudatus by Lance Peck
male of Moema pepotei by Wolfgang Staeck
type locality biotope of the new species (also dwellt by Aphyolebias rubrocaudatus) by Lance Peck
male and female of Rivulus christinae by Lance Peck
First published and distributed between November 10 and 15. 2003, in : Huber, J.H. 2003. A New Species of annual Killifish from southeastern Peru, with a uniquely barred Color Pattern, Moema quiii n.sp. Freshwater & Marine Aquarium (FAMA), 26 (12), (November 15): 10-19, 9 figs., 1 tab.
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