La vallée du Cosnipata descend vers l’Amazonie et le Madre de Dios : confluent juste au Nord de Pilcopata
(Cliquer sur les photos pour un meilleur effet !)
Butterfly diversity in an Andene valley – Cosnipata (Kosñipata), Cuzco, S. Peru
by
Daniel Lacomme & Bruce Purser, Association des Lépidoptéristes de France
Butterfly species are related to an altitudinal gradient ranging from 500 to 3600 meters. Although the taxonomic list is not finalised, it is clear that species and sub-species of many genera change with temperature, humidity and vegetation, conditioned by the altitudinal gradient. The authors limit their summary to four genera – Morpho, Catasticta, Perisama and Pedaliodes.
The Cosnipata Valley, a tributary of the Madre de Dios, flows towards the NE and thus receives abundant humidity from the Amazon Basin. This humidity is reflected in the luxuriant vegetation which grades, almost without break, from lowland forest, through pre-montagne to high mountain « cloud forest », each biome having its particular butterfly and moth fauna. Virtually all collecting has been carried out along a gravel road which descends the valley. Study made at specific altitudes indicates important local variations in butterfly diversity which seem to be conditioned by local variations in humidity ; where the road crosses a major tributary, humidity is higher and butterflies more abundant, but where the road climbs steep slopes situated high above local streams, lower humidities coincide with markedly fewer butterflies. This tendency is confirmed by the relative abundance of butterflies along the banks of the Cosnipata, their original habitat prior to the construction of the nearby road.
The Genus Morpho (FABRICIUS, 1807) in the Cosnipata Valley
The genus Morpho (Nymphalidae) is one of the most interesting groups to relate to altitudinal distribution, ranging from the Amazonian plain to about 3000 m. Fourteen species occur in the Cosnipata Valley, including one endemic species and 5 endemic sub-species.
In addition to the close observation of the endemic M. absoloni and its exceptionally rare female (described in 1983), is the fact that there exists an altitudinal relay between species belonging to the same sub-genus. Thus, certain species of the same group are replaced by others, according to altitude :
M. telemachus – M. amphitryon (sub-genus Iphimedeia)
M. marcus – M. zephyritis – M. sulkowskyi (sub-genus Cytheritis)
M. absoloni – M. aurora (sub-genus Balakowskyna)
M. menelaus zischkai – M. godarti assarpai (via 2 or 3 successive forms), (sub-genus Grasseia)
Concerning M. godarti assarpai, a clear change in morphology affects the same sub-species ; convergent with M. menelaus on the Amazon plain, close to M. godarti didius at middle altitudes, it resembles M. godarti godarti towards 1500 m – its upper limit. This morphological variation relating to altitude, within a given species, remains to be confirmed and other equivalents should be established.
Photos of two living rare species have been made : M. cisseis and M. absoloni. Finally, the flight season at low altitude (200 – 1000 m) are grouped (May – July) while middle and high altitude species fly during the months of September to December.
The Genus Morpho (FABRICIUS, 1807) in the Cosnipata Valley (in order of classification) | |||
Species | Subgenus | Author | Range in elevation |
Morpho telemachus foucheri | Iphimedeia | LE MOULT, 1926 | Low to middle mountain |
Morpho amphitryon azurita | Iphimedeia | DUCHÊNE & BLANDIN, 2009 | Middle & high mountain |
Morpho cisseis cabrera | Iphimedeia | BLANDIN & LAMAS, 2006 | Plain & foothills |
Morpho marcus | Cytheritis | SCHALLER,1785 | Plain & foothills |
Morpho zephyritis | Cytheritis | BUTLER, 1873 | Foothills, middle mountain |
Morpho sulkowskyi lympharis |
Cytheritis | BUTLER, 1873 | High mountain |
Morpho aurora isidorssoni | Balakowskyna |
BLANDIN, 2006 | Middle & high mountain |
Morpho absoloni | Balakowskyna | MAY, 1924 | Plain & foothills |
Morpho rhetenor tello | Cypritis | BLANDIN, 2008 | Plain & foothills |
Morpho menelaus zischkai | Grasseia | FISCHER, 1962 | Plain |
Morpho godarti assarpai | Grasseia | RÖBER, 1903 | Plain, foothills, middle mountain |
Morpho deidamia grambergi | Morpho | WEBER, 1944 | Plain & foothills |
Morpho helenor theodorus | Morpho | FRUHSTORFER, 1907 | Plain & foothills |
Morpho achilles phokylides | Morpho | FRUHSTORFER, 1912 | plain & foothills |
The genus Catasticta, (Pieridae, BUTLER, 1870)
Among the many genera of pierid inhabiting the Cosnipata, we have observed 15 species of Catasticta. The authors are very grateful to Maurizio Bollino for his help in the identification of seven species. According M. Bollino, the area (from Acjanaco down along the road to Pillcopata) hosts no less than 28 species of Catasticta, including a recently described one (Catasticta pillcopata BOLLINO, 2008).
The genus Catasticta, (Pieridae, BUTLER, 1870) in the Cosnipata Valley | ||
Species | Author | Range in elevation |
Catasticta sisamnus telasco | LUCAS, 1852 | 1200 m |
Catasticta chelidonis |
HOPFFER, 1874 | 1500 m |
Catasticta prioneris | HOPFFER, 1874 | 1500 m |
Catasticta reducta boliviana | BUTLER, 1896 | 1800 m |
Catasticta corcyra | FELDER & FELDER, 1859 | 1900 m |
Catasticta pieris pieris | HOPFFER, 1874 | 2200 m |
Catasticta collina | BROWN, 1939 | 2200 m |
Catasticta manco | DOUBLEDAY, 1848 | 2250 m |
Catasticta watkinsi |
LATHY & ROSENBERG, 1912 | 2300 m |
Catasticta aureomaculata |
LATHY & ROSENBERG, 1912 | 2300 m |
Catasticta poujadei lamasi | EITSCHBERGER & RACHELI, 1998 | 2300 – 2400 m |
Catasticta toca | DOUBLEDAY, 1847 | 2500 – 2900 m |
Catasticta scurra |
RÖBER, 1924 | 2500 m |
Catasticta marcapita | RÖBER, 1909 | 2900 m |
Catasticta superba | LATHY & ROSENBERG, 1912 | 2900 m |
Ranging in altitude from 1200 to 2900 m, the genus attains maximum diversity (7 species) between 2000 and 2500 m where virtually all species are located near small creeks which cross the road (see photo) or along the banks of the Rio Cosnipata. The numerous waterfalls are frequented by Catasticta which fly in the spray and land on wet cliffs or close to running water. Their behaviour is reminiscent of certain species of the Indo-Australian genus Delias. There also exists a remarkable colour convergence between Catasticta and at least two other genera of pierid : Dismorphia niepelti and Leodonta chiriquensis, all three being parapatric between altitudes of 1500 and 2200 m.
Although there does not appear to be any significant altitudinal evolution in colour patterns, the majority of the black and white species, plus the Leodonta and Dismorphia occur below 2000 m, species above that altitude (notably C. collina & C. rileyi) tending to have dark grey dorsal patterns. The majority of species (10) were observed at precise altitudes while the remainder extended over an altitude of 200 to 400 m. The altitudinal distribution of individual species of Catasticta is remarkably limited.
The genus Perisama (Nymphalidae, DOUBLEDAY, 1847)
We have observed 10 species of Perisama in the Cosnipata Valley where maximum diversity occurs between 1500 and 1800 m i.e. slightly lower than Catasticta. Six species (alicia, philinus, comnena, hilara, jurinei & tristrigosa plus Diaethria alicia) were flying together at 1500 m, P. calamis fassli extending up to 3000 m where it was associated with Orophila cecidas, a genus which closely resembles Perisama. The following species have been observed :
The genus Perisama (Nymphalidae, DOUBLEDAY, 1847) in Cosnipata Valley | ||
Species | Author | Range in elevation |
Perisama tringa | GUÉNÉE, 1872 | 1200 – 1500 m |
Perisama alicia | HEWITSON, 1875 | 1200 – 1800 m |
Perisama hilara | SALVIN, 1869 | 1400 – 1800 m |
Perisama philinus | DOUBLEDAY, 1849 | 1500 – 1800 m |
Perisama jurinei jurinei | GUÉNÉE, 1872 | 1500 – 1800 m |
Perisama tristrigosa | BUTLER,1873 | 1600 m |
Perisama comnena | HEWITSON, 1868 | 1500 – 2500 m |
Perisama clisithera | HEWITSON, 1874 | 1800 m |
Perisama moronina | RÖBER, 1915 | 2400 m |
Perisama calamis fassli | VIETTE, 1958 | 1800 – 2900 m |
Perisama frequents gravel roads and abandoned quarries and, although more abundant near the bottoms of humid valleys, is not directly associated with streams and waterfalls. Most species have an altitudinal range of about 1000 m but this seems to vary according to the season, several species (tringa, hilara & jurenei) being considerably lower (500m) in the dry season.
There appears to be a faunal change near 1800 m, several species (tringa, philinus, comnena, hilara, jurinei & tristrigosa) occuring below 1800 m, while P. calamis fassli, P. clisithera, P. moronina and Orophila cecidas occur above. This break may be related to the ancient limit of mountain forests (currently 3500 m) which were 1500 – 2000 m lower during the last Glaciation. The three species of Perisama found above 1800 m seem to be limited to southern Peru while those below 1800 m are widely distributed (N. Peru – Ecuador). This difference suggests that the three higher altitude species have had less time to colonise laterally and thus may have evolved more recently.
The genus Pedaliodes BUTLER, 1867 (Satyrinae)
The genus Pedaliodes is endemic to the Andes. Neotropical mountain Pedaliodes may be the most divers genus of butterflies, and is among the least well-studied. LAMAS (2004) lists 246 species of which 98 were described by PYRCZ et AL has recently described 14 species.
Pedaliodes are moderately-sized butterflies whose dorsal surfaces are generally uniformly brown in colour, only the ocelli and occasional patterns on the hind wings permitting field identification.
The Satyrinae, being essentially sedentary in habit, have evolved important phylogenic differentiation conditioned essentially by the deep and complicated valley system which characterises the eastern Andes.
Fifteen species of Pedaliodes (sensu stricto) have been identified in the Cosnipata Valley between about 1000 and the Acjanaco Pass (3650 m). Collecting has enabled the establishment of two or three altitudinal zones, with local overlap and sympatry, relating to vegetal discontinuities due to orientation of the valley and to local variations in humidity due to remoteness from humid, valley floors.
Among the 19 taxa comprising the following list, 4 are recently described while 3 are currently being defined.
The genus Pedaliodes BUTLER, 1867 (Satyrinae) | ||
Species | Author | Elevation |
Pedaliodes tucca tucca | THIEME, 1905 | 1450 m |
Pedaliodes manis manis | GODMAN & SALVIN (1881) | 1600 m |
Pedaliodes phaeinomorpha | LAMAS, VILORIA & PYRCZ, 2008 | 1600 m |
Pedaliodes phrasiclea | GROSE-SMITH, 1900 | 1600 m |
Panyapedaliodes phila phila | LAMAS, 2003 | 1600 m |
Pedaliodes hewitsoni | STAUDINGER, 1897 | 1800 m |
Antopedaliodes antonia, ssp. under study | 2100 m | |
Pedaliodes pausia | (HEWITSON, 1862) | 2100 m |
Pedaliodes phrasiclea galaxias | THIEME, 1905 | 2100 m |
Pedaliodes peruviana | BUTLER, 1873 | 2400 m |
Pedaliodes antulla antulla | THIEME, 1905 | 2950 m |
Pedaliodes molesta | STAUDINGER, 1897 | 2950 m |
Pedaliodes pheres pheres | THIEME, 1905 | 3000 m |
Panayapedaliodes drymaea dryamaea | HEWITSON (1858) | 3000 m |
Pedaliodes auraria | THIEME, 1905 | 3200 m |
Pedaliodes ackeryi | PYRCZ & VILORIA, 2008 | 3300 m |
Pedaliodes phantasia phantasia | PYRCZ, 2009 | 3300 m |
Pedaliodes sp. female, under study | 3350 m | |
Punapedaliodes flavopunctata flavopunctata | (STAUDINGER, 1894) | 3350 m |
Conclusion
The four genera – Morpho, Catasticta, Perisama & Pedaliodes – change with altitude, this species evolution being typical of many peri-Andene valleys. The considerable number of species which evolve with altitude are probably one of the major factors determining the overall diversity of lepidoptera within the eastern Andes.
A detailed description of the lepidoptera of the Cosnipata Valley will appear in a thematic number of Lépidoptères, the review of the ALF (Association des Lépidoptéristes de France), early 2011.
An interesting article, but I found some misidentification of Catasticta (the genus about which I am working for many years):
photo 4 is not sinapina, but chelidonis
photo 11 is not collina, but watkinsi
photo 13 is not tamboensis, but scurra
photo 14-15 & 16 are not distincta, but aureomaculata
photo 17 is not rileya, it is watkinsi
photo 19: 1 watkinsi (top left) plus 3 toca
photo 20 & 21 are not tomyris barbara but toca
The area (from Acjanaco down along the road to Pillcopata) hosts no less than 28 species of Catasticta, including a recently described one (Catasticta pillcopata BOLLINO, 2009).
Très intéressant.
Toujours impressionnant le comportement identique et la morphologie très proche entre les Catasticta andins et les Delias papous.
Y a t-il eu des écrits sur leur séparation, leur évolution ? …
Salut,
Très bien fait, votre compte rendu sur les papillons de Kosnipata. Bravo pour les identifications des Pedaliodes. Impecable ! La derniere espèce, non décrite, est en presse actuellement, son « working name » est P. acjanacoensis LAMAS, VILORIA et PYRCZ. Je serais bien content de pouvoir vous aider à déterminer les autres espèces de la tribu Pronophilini de cette région, ou d’ailleurs. Veuillez bien consulter le site internet du muséum, ci-dessous, y compris la liste complète d’espèces de Pedaliodes d’Acjanaco (Pyrcz et al., 2009).
Adresse web du muséum : http://www.mzuj.uj.edu.pl
Jean-Claude Petit nous écrit :
« Concernant « tomyris« , il est extrêmement rare dans cette région, j’en ai attrapé un il y a deux ans, et Maurizio m’en a demandé des photos car il n’en avait jamais vu, bien qu’il ait reçu des milliers de Catasticta de là-bas ; et ce serait la ssp. subtomyris« .
Pierre Boyer :
Une première erreur concerne Pedaliodes phaeinomorpha : l’identification est bonne, mais l’altitude impossible, ou bien le papillon a été transporté là par une voiture! Ce papillon vole entre 2800 et 3300 m, dans le secteur de Wayqecha et au-dessus, mais dans la forêt des nuages, sans atteindre le pré-paramo. La dernière série en fin d’article a été décrite, c’est finalement Pedaliodes acjanaco. Le reste de l’article est bon, sauf une erreur apparaissant au niveau de la liste d’espèces, aussi bien qu’en légende de la photo. Il y a deux espèces bien différentes et éloignées phylogénétiquement, Pedaliodes phrasiclea et Pedaliodes phrasicla. Ne me demandez pas pourquoi les descripteurs ont fait une telle bêtise en utilisant des noms aussi proches, c’est comme pour les Archaeoprepona demophon et demophoon ! Quoiqu’il en soit, phrasiclea est une espèce très largement répandue en Amérique du sud à basse altitude (« Pronophilinesquement » parlant!) et monotypique, comme c’est le cas d’autres d’espèces à ces altitudes. Cette espèce là est bien représentée dans l’article et l’identification est bonne.
Pedaliodes phrasicla galaxias est une ssp de phrasicla (pas phrasiclea) propre au sud du Pérou. cette espèce aussi est largement répandue en Amérique du sud, à des altitudes à peine plus élevées que l’autre, mais présente plusieurs phénotypes différents (au moins 4 ) sur son aire de répartition. L’identification est bonne sur l’article, mais la légende fait apparaitre phrasiclea galaxias au lieu de phrasicla galaxias. C’est tout à fait pardonnable, pour quelqu’un qui n’est pas trop versé dans ce groupe, et l’ensemble de l’article est intéressant. A noter aussi que le nombre de Pedaliodes dans la vallée de Kosñipata est largement supérieur à 15, mais je ne vais pas me lancer dans le décompte total pour le moment.