taxonID	type	description	language	source
03DD11312A462A0B2AF7D297FE43FB3F.taxon	diagnosis	Diagnosis: The imago of Perisama can immediately be recognized from other Neotropical butterflies, in particular from the most closely related and similarly patterned on the upperside, Callicorini Diaethria, sharing with Perisama oblique postdiscal FW and HW submarginal greenish bands, by mostly white or yellow HW ventral ground colour, with parallel ventral median and postdiscal lines enclosing a row of blackish spots, although in some species they can be vestigial or obsolete, and contrary to Diaethria, not clustering into two groups of one or two spots (thus the common name of the latter “ eighty-eights ” or “ eighty-nines ”), and never fully developed ocelli, contrary to Callicore.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A462A0B2AF7D297FE43FB3F.taxon	description	Redescription: Adults: medium size, forewing length 16 – 23 mm (mean = 21 mm, n = 142). Head: eyes light or dark brown, setose; antennae reaching to 2 / 3 the length of costa, dorsally brown, ventrally lighter, with packs of lightly coloured scales at each flagellomere, club elongated or rounded, flattened dorso-ventrally, approximately two times thicker than shaft; labial palpi one and half the length of head, dorsally brown, ventrally lighter. Thorax: brown or black, with dorsal and ventral surface covered with long, hair-like scales, matching the DH and VH coloration, respectively; legs with light brown to pale yellow or white femur, underside covered with long hair similar to those on the underside of thorax, tibia and tarsus darker than femur, covered with light brown scales, tibial spurs present. FW triangular with outer margin slightly convex or wavy; DF ground colour black or blackish-brown with metallic blue markings; VF ground colour same as DF, sometimes with red or yellow basal patch and smaller metallic markings, the colour of apical area matches the HV ground colour; HW rounded, with more or less wavy outer margin; DH ground colour black or blackish-brown with ashen-blue and metallic blue markings; VH ground colour: various shades of white, grey, yellow or brown, with two black lines (sometimes reduced) and, sometimes, five or six dots between the lines. Abdomen: brown or black, partly covered with long hair-like scales, similar to those on thorax; male genitalia with long valvae, rounded posteriorly, uncus slightly curved ventrally, shorter than tegumen, hairy, with a sharp tip, tegumen wide and regularly domed, gnathos long and slim, almost as long as uncus, vinculum with a lobe-like, hairy process located below the valva, saccus long and thin, slightly arched ventrally, aedeagus long and thin, arched dorsally, hypandrium with a wide, semicircular proximal incision, distal end in a form of U-shaped or W-shaped fork, depending on the species, sometimes with additional spines, collar long and narrow, widening over into the triangular body; female genitalia with papillae anales slightly sclerotized interiorly, thin apophyses posteriores with club-shaped tips, ostium bursae wide and irregularly shaped; lamella antevaginalis expanded into a large pleated lobe; posterior half of ductus bursae sclerotized, followed by a wider, membranous part, widening gradually into corpus bursae; signa as two small sclerotized dots, corpus bursae oval, as long as ductus bursae. Immature stages (data available for P. oppelii, P. bomplandii and P. barnesi): larvae light green, round in cross-section, with paler sides and two parallel rows of yellow papillae on the dorsal surface with small black thorns sticking out of each papilla; head square in shape, brown, with brown, branched scoli; final instar about 2 cm long; pre-pupa pale green with orange-brown scoli, larvae spin a white silk pad on dorsal surface of a leaf; pupa emerald green, robust in shape, with small white punctuations; head with two short projections; dorsolateral keel along edge of wing pad; larval host plants Paullinia sp. (Sapindaceae) (Greeney et al. 2010) and Weinmannia sp. (Cunionaceae) (Romero 1997).	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A472A0B2AF7D185FA35F8FF.taxon	discussion	Perisama jurinei was described originally as Callicore jurinei by Guenée (1872). It was later included in the genus Perisama by Oberthür (1916). Lamas (2004) considered P. jurinei as a subspecies of P. dorbignyi (Guérin-Ménéville, [1844]), whereas Attal & Crosson du Cormier (2010) reinstated its specific status. Perisama dorbignyi is a widespread species, reported from Eastern and Western Cordillera in Colombia, eastern and southern Andean slopes in Ecuador and Peru and on eastern slopes of the Andes in Bolivia. Geographic and individual variation affects mostly the size of the red discal patch, however some differences can also be observed in the shape of the HWV lines, FWD metallic band shape and width of HWD ashen band (Fig. 3). Our phylogenetic analysis based on molecular data show that P. jurinei and P. dorbignyi form a common clade with low genetic distances, below 1.5 %. Since there are no significant differences in the morphology of male genitalia, including in the hypandrium (Fig. 4), and no evidence of the two species being sympatric, P. jurinei is here considered as a subspecies of P. dorbignyi.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A452A092AF7D4D9FC6EFCD9.taxon	discussion	Perisama ambatensis was originally described by Oberthür (1916) as a species, but was reduced to the rank of a subspecies of Perisama lebasii (Guérin-Ménéville, [1844]) by Lamas (2004), then its specific status was reinstated by Attal and Crosson-du-Cormier (2010). It can be distinguished from P. lebasii lebasii by the absence of a spearhead shaped protuberance in the extension of the green FWD cubital ray. Perisama hilara Salvin, 1869 was also described as a species. It was regarded as a subspecies of P. lebasii by Lamas (2004), and its specific status was reinstated, again, by Attal & Crosson-du-Cormier (2010). Its most prominent distinguishable character is the presence of generally small greyish-blue dots on the HWD (Fig. 5). This taxon can be easily confused with P. ambatensis. The comparison of large series of specimens allows to notice that the HWV internal, median line is more sinuous in P. hilara than in P. ambatensis, but individual variation is relatively important which sometimes makes their taxonomic identification difficult. P. hilara is relatively uncommon and reported from southern Peru and northern Bolivia. Our phylogenetic analysis shows that P. hilara, P. ambatensis and P. lebasii form a common clade, and their relative genetic distances do not exceed 1.5 %. Since there are no significant differences in male genitalia morphology (Fig. 6) and there is no evidence of their sympatry, P. hilara and P. ambatensis are formally considered here, as proposed by Lamas (2004), as subspecies of P. lebasii.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A452A092AF7D491FD76FF64.taxon	discussion	stat. rev.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A422A0E2AF7D059FC04FA37.taxon	discussion	Perisama tryphena is one of the most common species of the genus Perisama, characterised by a wide distribution, and important individual and geographic variation (Fig. 7), similar to that observed in P. bomplandii. Perisama goeringi was described as a separate species by Druce (1875) from the Venezuelan Cordillera de Mérida. Perisama affinis was described by Attal & Crosson-du-Cormier (1996) from the Department of Huánuco in Peru. Both, P. goeringi and P. affinis were regarded as subspecies of P. tryphena by Lamas (2004), and were reinstated as species by Attal and Crosson-du-Cormier (2010). Comparative analysis has shown no significant differences in male genitalia morphology (Fig. 8) of the three taxa, and molecular data also confirm they are closely related, with genetic distance varying from 0.3 - 1.2 %. Since there is also no evidence of the three taxa being sympatric, P. affinis and P. goeringi are henceforth considered as subspecies of P. tryphena.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A422A0E2AF7D011FD48FBE7.taxon	discussion	stat. rev.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A422A0E2AF7D235FDFEF847.taxon	discussion	Perisama xanthica is a very distinctive species, with a relatively low interspecific diversity. It was described as Catagramma xanthica by Hewitson (1868) and considered a subspecies of P. oppelii by Lamas (2004). Its specific status was reinstated by Attal-and-Crosson du Cormier (2010), but Zubek et al. (2015) again down-ranked it to a subspecies of P. oppelii taking into account the absence of any significant differences in the morphology of male genitalia, including the hypandrium, or wing colour patterns, except from the more vivid, yellow-orange shade of the HWV in P. xanthica. However, our molecular data clearly show that P. xanthica and P. oppelii are separate species. Their genetic distance is very high, ranging from 8.6 % – 9.4 %. Considered this, the specific status of P. xanthica is hereby formally reinstated.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A402A0C2AF7D491FC33FDCF.taxon	discussion	This taxon was described as Catagramma camelita by Hewitson (1876), placed in Perisama by Röber (1924), and cited as a separate species by most authors (e. g. Descimon 1995, Lamas 2004). It was, however, considered as a subspecies of P. alicia (Hewitson, 1868) by Attal & Crosson-du-Cormier (1996, 2010) who based their conclusions on the similarities between these two taxa in male genitalia and wing colour patterns. It would not be the case of a subspecies or individual form differing from the nominate morphotype in the ground colour of its ventral wing surface. However, in their case, both morphotypes had a red costal margin of the hind wing. On the other hand, P. camelita exhibits no costal red, whereas P. alicia does. The red costal margin is a conservative character within Perisama, and there is multiple evidence, that it is a species-specific character. Therefore, in agreement with Lamas (2004) the specific status of P. camelita is formally reinstated here.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A402A0D2AF7D1A1FB38FF2F.taxon	discussion	Attal & Crosson-du-Cormier (1996) considered P. dorbignyi aristoteles (Oberthür, 1916) as a synonym of P. dorbignyi paula (Oberthür, 1916), explaining that diagnostic features pointed out in the original description are merely individual variation. Lamas (2004) regarded P. dorbignyi aristoteles as a valid subspecies. Our comparative analysis did not show any stable morphological differences between the populations from southern Colombia which correspond to P. dorbignyi paula and northern Ecuador representing P. dorbignyi aristoteles. Therefore, we agree with Attal & Crosson-du-Cormier on the synonymy of P. dorbignyi aristoteles and P. dorbignyi paula. On the other hand, both Lamas (2004) and Attal & Crosson-du-Cormier (1996) recognize two subspecies, P. dorbignyi mola (Dognin 1899), from southern Ecuador and P. dorbignyi paula from southern Colombia and northern Ecuador. As pointed out by Attal & Crosson-du-Cormier, Oberthür has failed to mention the description of P. dorbignyi paula when describing P. mola as a new species, that is why the diagnostic characters do not allow to separate the two taxa. Attal & Crosson-du-Cormier (1996) conclude, that the most distinctive character of P. d. mola is a gap in the FWD transverse metallic band. However, the more recent comparative study of a series of specimens from Sangay National Park area, in central Ecuador, as well as from Loja and Zamora-Chinchipe provinces proved that this is not a stable character with some intermediate forms occurring in the same localities (Fig. 10). Therefore, P. dorbignyi paula is considered here as a junior subjective synonym of P. dorbignyi mola.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A402A0C2AF7D66DFEAEFADB.taxon	discussion	This taxon was described originally as Catagramma comnena (Hewitson, 1868) from Bolivia. and was included in the genus Perisama by Oberthür (1916). Its specific status was maintained by Attal & Crosson-du-Cormier who, however, at the same time report and illustrate intermediate morphotypes, between P. comnena and P. humboldti from central Peru. Lamas (2004) regarded six subspecies of P. humboldtii as subspecies of P. comnena, which were all later considered to be subspecies of P. humboldtii by Attal & Crosson du Cormier (2010). Perisama comnena is morphologically similar to P. humboldtii, and differs mostly in the shape of median and submarginal lines on the HWV, and the absence of dots in between. P. comnena has a continuous green band on the FWR, while P. humboldtii has a green inverted pipe. Comparative morphological analysis confirmed that intermediate forms occur in Peru, which, when examined separately are difficult to assign to either species. An extensive comparative investigation of male genitalia reveals that the differences between P. comnena and P. humboldtii pointed out by Attal & Crosson-du-Cormier (1996) are consistent with the spectrum of individual variation (Fig. 9). Moreover, genetic distances between the P. humboldtii and P. comnena are very low (0.5 % – 0.8 %). Therefore, P. comnena should be presently considered a subspecies of P. humboldtii. However, the large sympatry of these two taxa is somewhat puzzling, and the fact that all along the distribution of P. humboldtii, only P. comnena has a continuous green band on FWR, can suggest that they are possibly different species, close enough to hybridize regularly. Detailed molecular and morphological studies should be carried out in order to verify the taxonomical status of the subspecies of Perisama humboldtii.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A5E2A122AF7D491FB07FDEF.taxon	description	Perisama paralicia is a relatively rare species, therefore it is difficult to fully appreciate the actual its geographic individual variability. The specimens we examined differ in size of the FWV red discal patch and width of the HWD marginal ashen band. P. paralicia bonita Attal & Crosson-du-Cormier (1997) described from northern Ecuador differs from the nominate, Peruvian subspecies in the smaller size of the basal red patch and slightly narrower HWD ashen band. Simultaneously, the above authors have stated that the same characters can be observed in the nominate subspecies, as an example of extreme individual variation, which makes the validity the subspecies bonita doubtful. Moreover, our research confirmed the presence of bonita morphotype in central Ecuador and northern Peru (San Martin province). Therefore, P. paralicia bonita is considered here a synonym of P. paralicia.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A5E2A122AF7D68CFCE0FB6D.taxon	discussion	Perisama philinus was described by Doubleday ([1849]) originally in Perisama. It is one of the most distinctive species of the genus, exhibiting relatively low individual morphological variation, affecting mostly the size of the FWD metallic patch, except P. philinus nyctimene (Hewitson, 1868). Callicore saussurei Guenée, 1872 was described from Brasil but it was later established it occurs in central Peru (Rober 1924). It was considered a subspecies of P. philinus by Attal & Crosson du Cormier (1996). Putative, subspecific characters of P. philinus saussurei are slight, and expressed by somewhat more prominent discal striae and a frequently present ashen band on the HWD. Some differences between the two taxa can be also noticed in the amount and intensity of the upperside violet-blue reflection. It is weaker among the Bolivian individuals, which are somewhat duller, although the metallic colour may be spreading into the FWV cell. However, a comparative analysis of the specimens from northern Bolivia, southern and central Peru, which is where the typical morphotype is found, has shown that those characters are subject to individual variation (Fig. 11) and not correlated with geographical distribution. Therefore, P. philinus saussurei is considered here a synonym of P. philinus philinus.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A5E2A132AF7D32EFA89FE5E.taxon	discussion	Perisama tryphena is one of the most common species of Perisama with a wide geographic range and similar extent of individual variation as in P. bomplandii. Two subspecies were described so far from central and southern Ecuador, respectively: P. t. pastaza Attal & Crosson du Cormier, 1996 and P. t. pallida Attal & Crosson du Cormier, 1996. The distinctive characters of the former are: more compact and rounded FWD inverted pipe mark and thinner HWD ashen band, and in case of the latter, regular outlines of the discal part of the FWD inverted pipe mark, even thinner HWD ashen band and paler underside ground colour. Based on the given characters it is often impossible to define a particular morphotype of a single specimen. Recent data from Sangay National Park show that the extent of individual variation within particular, local populations often exceeds the alleged differences between the aforementioned subspecies (Fig. 12). Therefore, P. t. pallida is now considered a synonym of P. t. pastaza.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
03DD11312A5F2A132AF7D1DAFAEBF813.taxon	discussion	Perisama hazarma (Hewitson, 1869) is one of the most distinctive species in the genus, and one of the rarest in collections. Perisama davidi (Dognin, 1888), described as a separate species, was considered as a subspecies of P. hazarma by Attal & Crosson-du-Cormier (1996, 2010), but as a synonym of the latter by Lamas (2004). Attal & Crosson-du-Cormier (1996) pointed out some putative morphological differences, namely more golden yellow HWV ground colour, lightly alternated fringes on the HW and a more complete FWD metallic band, broken in the nominate subspecies. Recent observations in the Sangay National Park, which corresponds located to the supposed “ distributional gap ” between the nominate subspecies and P. hazarma davidi, not only prove that the actual range of the species is larger than expected, but also show that the extent of phenotypic variation of P. hazarma was underestimated (Fig. 13), and in fact the subspecies-defining characters of P. hazarma davidi simply reflect individual variation. Therefore, P. hazarma davidi is considered here a junior synonym of P. hazarma.	en	Zubek, Anna, Wahlberg, Niklas, Boyer, Pierre, Lorenc-Brudecka, Jadwiga, Mahecha-J, Oscar, Pyrcz, Tomasz (2025): Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini). Zootaxa 5636 (1): 1-34, DOI: 10.11646/zootaxa.5636.1.1, URL: https://doi.org/10.11646/zootaxa.5636.1.1
