European Journal of Taxonomy 71: 1-66

http://dx.doi.org/10.5852/ejt.2014.71

www. europeanj ournaloftaxonomy. eu

2014 • Patrick Blandin et at.

ISSN 2118-9773

This work is licensed under a Creative Commons Attribution 3.0 License.

Monograph

urn:lsid:zoobank.org:pub:F667B43B-86A3-4D7D-9A22-021E52CDA30F

Revisiting the Andean butterfly Eryphanis zolvizora group

(Lepidoptera, Nymphalidae): one or several species?

Patrick BLANDIN 1 , Roger BRISTOW 2 , Andrew NEILD 3 , Juan Carlos DE SOUSA 4 , Yuvinka

GARECA 5 & Blanca HUERTAS 3 6

Museum National d’Histoire Naturelle, Entomologie, 57 rue Cuvier, 75005 Paris, France

E-mail: [email protected] (corresponding author)

2 2 Sid Bank, Sid Fane, Sidmouth, Devon EX10 9AW, UK

E-mail: devonbristows@btinternet. com

3 The Natural History Museum, Entomology Department, Cromwell Road, SW7 5BD, London, UK

E-mails: [email protected] . [email protected]

4 Museo del Instituto de Zoologia Agricola, Facultad de Agronomia, Universidad Central de Venezuela,

Maracay, Apdo. 4579, Aragua, Venezuela

E-mail: [email protected]

5 Museo de Historia Natural Alcide d’Orbigny, Entomologia, CP4324, Av. Potosi N°1458 (esq. Av.

America), Cochabamba, Bolivia

E-mail: [email protected]

1 urn:lsid:zoobank.org:author:F79EECF9-4E10-4826-BA99-01DElEB0DD0E

2 um:lsid:zoobank.org:author:E9AD6AAl-2C47-4358-B606-0A84042D872D

3 urn:lsid:zoobank.org:author:268E6AD0-5E7A-4073-B530-6CE39D963474

4 urn:lsid:zoobank.org:author:68F7180E-1695-4716-B794-1900B02881D8

5 um:lsid:zoobank.org:author:17D6BF6C-A94B-4057-8057-454C3DF0CEA2

6 urn:lsid:zoobank.org:author:F96FCD0A-4A22-4782-B077-4588E193134A

Abstract. Eryphanis zolvizora (Hewitson, 1877) is a rare Andean endemic butterfly, described from

Bolivia, which has been historically classified either as a unique species, or as part of a group of three

allopatric species from Bolivia, Ecuador and Colombia. In this paper, the group is revised using more than

200 specimens housed in 35 European and North and South American public and private collections. For

the first time, the presence of the group in Western Ecuador and Venezuela is confirmed, and important

data on Peruvian populations are provided. In some populations, individual variations of genitalia are

observed. Nevertheless, male genitalia allow the distinction of four geographical groups. Considering

also habitus characters, eight taxa are distinguished and considered to be subspecies, of which five are

new: Eryphanis zolvizora inca ssp. nov., Eryphanis zolvizora chachapoya ssp. nov., Eryphanis zolvizora

casagrande ssp. nov., Eryphanis zolvizora reyi ssp. nov., and Eryphanis zolvizora isabelae ssp. nov. In

the present state of knowledge, these taxa are allopatric, except for a possible geographic overlap in

central Peru, where data are insufficient to prove sympatry. The “several subspecies vs. several species”

dilemma is discussed, considering its impact for conservation action and policies.

Keywords. Morphinae, Brassolini, Eryphanis zolvizora group, subspecies, taxonomic inflation.

European Journal of Taxonomy 71: 1-66 (2014)

Blandin P., Bristow R., Neild A., Carlos de Sousa J., Gareca Y. & Huertas B. 2014. Revisiting the Andean butterfly

Eryphanis zolvizora group (Lepidoptera, Nymphalidae): one or several species? European Journal of Taxonomy

71: 1-66. http://dx.doi.org/10.5852/eit.2014.71

Introduction

Within the butterfly family Nymphalidae, the sub-family Morphinae includes two tribes, the Morph ini

and Brassolini, both endemic to the Neotropical region. The former includes three genera as follows:

Antirrhea Hiibner, 1822, Caerois Hiibner, 1819 and Morpho Fabricius, 1807, the last the most familiar,

as it includes big and bright blue species that are emblematic of the Amazons. Following the most recent

checklist (Casagrande 2004), the tribe Brassolini comprises 18 genera including the genus Caligo Hiibner,

1819 (the well known large Owl butterflies). Since Fruhstorfer’s review (1912), SticheFs catalogue

(1932), and before Casagrande’s checklist, five genera have been revised. More recently, Penz (2008)

produced a phylogenetic review of the genus Eryphanis Boisduval, 1870. This genus was created for

three species which have been included previously in the genus Pavonia Godart, 1824 (an invalid name

according to Hemming 1967), or in the genus Caligo Hiibner, 1819, or even in the genus Opsiphanes

Doubleday, 1849. All species and subspecies currently recognized by Casagrande (2004) were known

as early as 1912 (Fruhstorfer 1912). Between SticheFs catalogue and Casagrande’s checklist, the only

changes concern the name of a species, E. polyxena (Meerburgh, 1780) being a junior synonym of E.

automedon (Cramer, 1775), and one subspecies included in the synonymy of another subspecies within

E. polyxena. Moreover, Casagrande (2004) indicated no new subspecies or species to be described.

Thus, the systematics of Eryphanis could be considered as well established at that time.

This consensus was challenged by Penz (2008), who studied male and female genitalia for the first

time in this genus. She separated E. lycomedon (C. Felder & R. Felder, 1862) from E. automedon , E.

bubocula (Butler, 1872) from??, aesacus (Herrich-Schaffer, 1850), E. opimus Stau dinger, 1887 from A 1 .

zolvizora (Hewitson, 1977), and together with DeVries she described E. greeneyi Penz & DeVries, 2008,

a new species close to E. zolvizora and E. opimus , these three species forming the Eryphanis zolvizora

group.

Penz’s work (2008) illustrates the taxonomic issues that arise with uncommon taxa distributed

throughout immense regions such as the Neotropics. Should slightly different specimens collected in

localities separated by hundreds or thousands of kilometres be considered as subspecies of a unique,

polytypic species, or as distinct species? In such little-known groups, the delimitation of species is

variable, depending on the ‘lumper’ or ‘splitter’ taxonomist’s culture. The Eryphanis zolvizora group

provides an interesting model to address this issue, which is a matter of intense debate (amongst many

others, see for example Cracraft 2000 and James 2010). Beyond theoretical considerations on the nature

of species, its evolutionary significance and its ecological role, practical questions arise: by raising

subspecies to species level, splitters contribute to ‘taxonomic inflation’, with potential consequences on

conservation policies (Isaac et al. 2004).

Eryphanis zolvizora (Hewitson, 1877) was described from two Bolivian specimens lacking precise

locality data. Staudinger (1887) described Eryphanis opimus as a different species, based on three

specimens, two from Manizales, central Cordillera, Colombia, and one from Chanchamayo, central Peru.

Stichel (1904; 1909) placed the two taxa in one species, and added specimens from Peru and Ecuador

to the nominate sub-species. Fruhstorfer (1912) followed Stichel’s classification. He indicated that E. z.

zolvizora occurs in Peru, but without more precise locality details. Considering E. z. opimus , he wrote

that in Ecuador and Peru a rather similar variety exists, which was not sufficiently studied, even though

Staudinger outlined some weak differences. Stichel (1932) maintained that the nominate subspecies

included the Peruvian and Ecuadorian populations, a position followed by D’Abrera (1987). Casagrande

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

(2004), in her checklist, also recognised only one species, divided into the nominate subspecies and

opimus as a Colombian subspecies. Penz (2008) showed that there are clear morphological differences

in male genitalia between Bolivian and Colombian specimens, and she observed differences in female

genitalia between Bolivian and Ecuadorian specimens. She also took into consideration wing shape and

ornamentation. Despite having studied a very limited number of specimens (3 33 , 1 ? from Bolivia,

7 33 , 4 $? from Ecuador, and only 1 3 from Colombia), she divided the group into three species. It

is true that genitalia differences are used to support separation at the specific level, but the possibility

that geographical variations of genitalia exist within a single species cannot be excluded. Therefore, in

the case of the Eiyphanis zolvizora group, we think that plausible taxonomic decisions cannot be taken

without knowing the characteristics of populations distributed throughout its range from Bolivia to

Venezuela, and especially Peru.

Fruhstorfer (1912) considered Eryphanis zolvizora as a very rare species, reflected in the small numbers

of specimens housed in most public and private collections. Here, we aim to provide a comparative study

using a large set of 249 specimens (197 33 , 52 $ $), from European and North and South American

public and private collections. Firstly, we present a geographical survey of the Eryphanis zolvizora

group. It exists throughout the Central and Northern Andes, from Bolivia to Venezuela, a country where

the existence of zolvizora- like populations had not been reported before. We also confirm the existence

of populations in western Ecuador. Data on altitudinal distribution are discussed. We dissected many

specimens to compare genitalia, and we specify differences between populations from different areas.

Moreover, we discovered individual variations of some genitalia characters. Several new taxa are

recognised. We discuss two competing taxonomic arrangements - several species vs. one polytypic

species - which are possible in the present state of knowledge.

Material and methods

A historical survey of the discovery of the Eryphanis zolvizora group is provided in Appendix 1 and a

detailed list of the specimens studied here is provided in Appendix 2. For the purpose of this study, we

divided Peru into a southern part, including Puno, Madre de Dios and Cusco Departments, a central

part, including Junin and Pasco Departments, and a northern part, including Huanuco, La Libertad,

Cajamarca, San Martin and Amazonas Departments. Private and public collections visited are listed and

their respective acronyms used throughout the text are indicated below:

AMNH

ANSP

BMNH

CAS

CBF

FLMNH

HAGAM

IAvH

ICNUN

IRSN

JCS

JFLC

LACM

MCC

American Museum of Natural History, New York, USA

Andrew Neild collection, London, UK

Academy of Natural Sciences, Philadelphia, USA

Natural History Museum, London, UK

California Academy of Sciences, San Francisco, USA

Coleccion Boliviana de Fauna, La Paz, Bolivia

McGuire Center for Lepidoptera and Biodiversity, Florida Museum of

Natural History, University of Florida, Gainesville, USA

Francisco Pinas collection, Quito, Ecuador

Herbert Art Gallery & Museum, Coventry, UK

Instituto Alexander von Humboldt, Bogota, Colombia

Instituto de Ciencias Naturales, Bogota, Colombia

Institut Royal des Sciences Naturelles, Brussels, Belgium

Juan Carlos De Sousa Coelho collection, Funchal, Madeira

Jean-Fran 9 ois Le Crom collection, Bogota, Colombia

Julian Salazar collection, Manizales, Colombia

Natural History Museum of Los Angeles County, USA

Mauro and Clara Costa collection, Caracas, Venezuela

European Journal of Taxonomy 71: 1-66 (2014)

MECN

MHNC

MHNNKM

MIZA

MJP

MNHN

MPM

MZUJ

NNM

RAMM

UFPC

UMO

USNM

ZMHB

ZMUA

Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador

Museo de Historia Natural Alcide d’Orbigny, Entomologia, Cochabamba,

Bolivia

Museo de Historia Natural Noel Kempff Mercado, Santa Cruz, Bolivia

Museo del Instituto de Zoologia Agricola, Maracay, Venezuela

Museo de Historia Natural ‘Javier Prado’, Universidad Nacional Mayor de

San Marcos, Lima, Peru

Museum National d’Histoire Naturelle, Paris, France (CG: collection generale;

PBB: collection Patrick Blandin (Brassolini); PBGL: specimens collected

during Patrick Blandin and Gilbert Lachaume studies in northern Peru)

Milwaukee Public Museum, Wisconsin, USA (pictures provided by C. Penz)

Zoological Museum of the Jagiellonian University, Krakow, Poland

Nationaal Natuurhistorisch Museum, Leiden, The Netherlands

Peter Wilson collection, Cornwall, UK

Romero Family collection, Maracay, Venezuela

Royal Albert Memorial Museum, Exeter, UK

Stephane Attal collection, Paris, France

Universidade Federal do Parana, Curitiba, Parana, Brazil

Oxford University Museum of Natural History, Oxford, UK

National Museum of Natural History, Smithsonian Institution, Washington DC,

USA

Zoologisches Museum der Humboldt-Universitat, Berlin, Germany

Zoologisch Museum, Amsterdam, The Netherlands

Other abbreviations used throughout the text:

DFW

DHW

FWL

PLT

VFW

VHW

dorsal, upperside

dorsal, upperside forewing

dorsal, upperside hindwing

forewing

fore wing length (measured from the base to the tip of the wing)

holotype

hindwing

length

lectotype

paralectotype

paratype

syntype

ventral, underside

ventral, underside forewing

ventral, underside hindwing

When necessary, authors of this paper are indicated by corresponding initials: PB (Patrick Blandin); RB

(Roger Bristow); AN (Andrew Neild); JCSC (Juan Carlos De Sousa Coelho); YG (Yuvinka Gareca);

BH (Blanca Huertas).

In addition to the characters which appeared important to Penz (2008), we have studied several

additional characters, such as the colour of the androconial patch in the anal area of the hindwing, which

varies between Eryphanis species and sometimes between subspecies (unpublished observations). We

recognize the following characters which may have a diagnostic role (Figs 1,2):

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

a) importance of projection of Cut on HW, evaluated by the value in millimetres (mm) of the index P

= LCul-((LM3+LCu2)/2), where LM3, LCul and LCu2 are the length between the tips of the veins

M3, Cul and Cu2 and the point of origin of Cu2 at the cubital cell vein (Fig. la);

b) presence (vs. absence or incompleteness) on DFW of an orange mark between veins R5-M1,

extending the submarginal band towards the anterior wing margin, thus forming Penz’s “proximal

branch” of the submarginal band (Fig. lb);

c) size and shape of the orange marks forming the submarginal band on DFW (Fig. lc);

d) development of violet-blue iridescence on the dorsal surface (Fig. Id);

e) length and colour of the male androconial patch in the anal area of DHW (Fig. le);

f) ground colour of the ventral surface, and colour of the median area, between the proximal and distal

white stripes (Fig. 2f) (we compared recently collected specimens to old specimens collected more

Fig. 1 . Characters studied on the habitus (recto) and their major variations, a. HW projection, estimated

by P = LCul-((LM3+LCu2)/2) in milimetres). b. Orange mark between veins R5-M1. c. Size and shape

of the orange marks, d. Violet-blue iridescence, e. Male androconial patch.

European Journal of Taxonomy 71: 1-66 (2014)

than a century ago, and we did not observe significant differences between ventral surfaces, but we

observed the opposite on dorsal surfaces, which are paler in old specimens);

g) shape and size of the isolated white spot situated approximately at the middle of the VFW cell,

between Cul and Cu2 (Fig. 2g);

h) presence and size of a white spot at the angle formed by the cubital vein of the VFW cell and Cu2

(Fig. 2h);

i) presence (vs. absence or incompleteness) on the VFW of a connection (‘bridge’) between the basal

and median white stripes in cell Cu2-2A (Fig. 2i);

j) length on the VHW of posterior extension of the proximal and distal white stripes (Fig. 2j);

Fig. 2. Characters studied on the habitus (verso), and their major variations, f. Ground colour of the

ventral surface and of the median area. g. Isolated white spot on FW cell. h. White spot at the angle

formed by the cubital vein of the FW cell and Cu2. i. ‘Bridge’ between white stripes in FW cell Cu2-2A.

j. Length on HW of posterior extension of the white stripes, k. Black designs in the HW cell, and black

curved line anterior to the Cul-Cu2 ocellus. 1. Ring around the HW costal ocellus, m. Widest diameter

® of the HW Cul-Cu2 ocellus.

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

k) on the VHW, prominence (vs. absence or incompleteness) of the black designs in the cell, and of a

black, curved line between veins Cul and Cu2, anterior to the Cul-Cu2 ocellus (Fig. 2k);

l) presence (vs. absence or incompleteness) of a white ring around the VHW costal ocellus;

completeness or incompleteness of the inner black ring (Fig. 21);

m) size of the VHW Cul-Cu2 ocellus, evaluated by its widest diameter ® (parallel to Cul), measuring

from the outer margin of the prominent dark brown ring (Fig. 2m).

Most of the studied specimens were photographed. Photographs were taken with different cameras and

in different light conditions. Despite our best efforts to correct them, there are still disparities in colour

dorsal arch

B 1

B 2

Fig. 3. Characters studied on genitalia. Al. Male from Bolivia (Eryphanis zolvizora zolvizora (Hewitson,

1877); MNHN, PBB 2286). A2. Male from Central Peru (La Suiza, Pasco; UFPC; PT of Eryphanis

zolvizora chachapoya ssp. nov.; picture by M. M. Casagrande). Bl. Female from Bolivia (Eryphanis

zolvizora zolvizora (Hewitson, 1877); MNHN, CG). B2. Female from Venezuela (Rancho Grande,

Aragua; PT of Eryphanis zolvizora isabelae ssp. nov.; 118-JCSC; picture by J.-C. de Sousa).

Table 1. Geographical origins and numbers of the studied specimens.

European Journal of Taxonomy 71: 1-66 (2014)

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

Table 2. Altitudinal range of the Eryphanis zolvizora group.

Country

Lower reliable

altitudes

Higher reliable

altitudes

More or less doubtful low altitudes

Bolivia

1000 m

2600 m

Buena Vista; Chapare <400 m

Peru

1300-1400 m

2800 m

Chirimayo, “1000 ft”

Ecuador

1500 m

2163 m

R. Arajuno, 700 m; Napo, 465 m

Santo Domingo (Pichincha), 200 m

Colombia

900 m

2500 m

Buenaventura-Queremal, 200 m

Puerto Narino , 150 m

Venezuela

1150 m

2100 m

balance, contrast, and saturation, and relatively subtle colour differences are not always obvious on the

photographs of variable quality that we have reproduced on our composite figures.

Measurements of FWL, length of the androconial patch, LM3, LCul, LCu2 and ® were taken directly

from specimens, or from scaled pictures, the precision being ± 0.5 mm. Where necessary, we indicate

the number (n) of measurements, and minimum (min) and maximum (max) values. On some damaged

specimens, LCul could not be measured and the P index could not be calculated. The specimens on

which measurements were made are indicated in Appendix 2.

Genitalia were prepared using a 10% solution of KOH and stored in a glycerol/70% ethanol solution

(3:1). Structures were observed using an optical stereo microscope with light and white field. Descriptive

terms (Fig. 3) are those used by Penz (2008). The specimens which were dissected are indicated in

Appendix 2.

Results

Geographic distribution and altitudinal range

In a synthesis of brassoline biogeography, Blandin (1977) included E. zolvizora in the group of species

associated with Andean montane forest and provided a map showing its distribution from Bolivia to

Colombia, indicating a few areas along the Andes. Even today, tens if not hundreds of kilometres separate

“neighbouring” localities. However, thanks to our survey of more than 150 specimens with correct

data (Table 1), a regional pattern becomes apparent. The zolvizora group is distributed in the whole

Tropical Andes Hotspot, as defined in Mittermeier et al. (2004). In the Central Andes, it exists along the

eastern Andean slopes from Bolivia (Santa Cruz Department) to the Peruvian north-eastern cordilleras

(San Martin and Amazonas Departments). In the Northern Andes, it extends along the Ecuadorian and

Colombian eastern cordilleras, and then along the Cordillera de Merida and the Cordillera de la Costa in

Venezuela. In western Ecuador, its existence is confirmed, and it extends northwards along the Pacific

side of the Colombian western cordillera. The E. zolvizora group is also represented in Colombia on the

oriental slopes of the western cordillera, in the central cordillera (at least on its western slopes), and on

both sides of the eastern cordillera.

Most reliable altitudinal data range from 900 to 2800 m (Table 2; see Appendix 2 for detailed data).

Fassl (1915a) indicated E. opimus on both sides of the western Colombian cordillera at heights between

2000 and 2500 m. In Venezuela, most specimens were taken between about 1150 and 1500 m elevation,

although this is most likely a reflection of the lack of access to higher elevations at each site rather than

an indication of the true upper limit of the species, as a few specimens have been collected between 1900

European Journal of Taxonomy 71: 1-66 (2014)

to 2100 m. These altitudes correspond to cloud forests in the broadest sense. These ecosystems exist, in

large mountain ranges as the Andes, between 1500 and 3500 m (Brown & Kappelle 2001). According to

some authors, cloud forests can be observed as low as 1300 m, for example on eastern Peruvian slopes

(Brack Egg & Mendiola Vargas 2004). These cloud forests are characterized by bamboos belonging

to the genus Chusquea. Harold Greeney reared E. greeneyi on Chusquea scandens at the Yanayacu

Biological Station, c. 2100 m, in eastern Ecuador (Penz 2008). It is likely that the distribution of the

E. zolvizora group depends on the altitudinal range and geographical extent of cloud forest Chusquea

species.

Given this information, lower elevations are most likely the result of mislabelling. From Bolivia, two

specimens in the FLMNH collected in 1925 and 1994 are labelled ‘Buenavista’, a lowland locality

(under 400 m) in Santa Cruz Department. We have confirmed that zolvizora populations do exist in

the Andean slopes of Santa Cruz Department, but not in the vicinity of Buena Vista. Perhaps there was

a confusion with Bellavista, where E. zolvizora has been collected recently. There is also one male

in MNHN (PBB 260), probably mislabelled, from “Chapare 300 m”. A Peruvian specimen (BMNH)

is labelled “Chirimayo, 1000 ft, Ockenden, July 1901”; it is likely that there is a typographic error,

as Ockenden’s data are generally reliable. From eastern Ecuador, some specimens with altitudes as

low as 700 m (a specimen labelled “Rio Arajuno”, Napo Province, Ecuador, in MNHN) and 465 m

(a specimen labelled [Puerto] Napo, Napo Province, Ecuador; PW) almost certainly are mislabelled.

It should be noted that in Ecuador (and other countries), data from commercial collectors are often

unreliable. The female labelled from Santo Domingo, 200 m, in Pichincha Province, is supposed to

have been collected at a higher altitude along the road to Quito. A Colombian specimen (IAvH), labelled

“Buenaventura-Queremal, 200 m, Valle de Cauca”, was collected by Schmidt-Mumm, who often made

errors, indicating with apparent precision (altitude, longitude, latitude) incorrect localities (J.-F. Le

Crom, pers. comm.). We consider that this specimen was probably collected in the mountains, along the

road from Buenaventura to Cali. There is also a male labelled from Puerto Narino, in the Amazonian

lowlands, which is certainly mislabelled.

However, one cannot exclude the possibility that cloud forests may exist at altitudes lower than 1000 m,

in orographic situations favouring cold and hyper-humid local conditions. For example, in Venezuela,

along the Rio Frio in the eastern El Tama range, one of the authors (AN) regularly but infrequently

encountered and received reports of lower montane forest species as low as 650 m. This seems to be

related to local topography, with extremely steep slopes and very humid and luxuriant valleys. The

observations above allow us to stress the fact that the Eryphanis zolvizora group exhibits significant

variation in altitudinal range.

Morphological characters

Male habitus (Figs 4A-F, 5A-F, 6A-D)

FWL generally varies between 55 and 67 mm in all populations. Only one dwarf specimen was

encountered, from Barinitas (Venezuela), with a FWL of 52 mm; it has not been included in the

measurement statistics.

The hindwing projection is reduced in Bolivian specimens (average P = 3.3, cf. Table 3) and medium

(average P = 4.4) in the southern and central Peruvian specimens. The northern Peruvian specimens and

4 specimens from central Peru (San Francisco, “Rio Perene”, La Suiza, Huancabamba) have a strong

projection, as well as northern Peruvian and eastern Ecuadorian specimens (average P = 5.6 and 5.5,

respectively). Colombian specimens from the valley of the Rio Cauca and Venezuelan specimens have

reduced or medium projections, the average values of P ranging from 3.6 to 4.3 mm. In other regions

of the Northern Andes (western Ecuador and Narino Department on one hand, north of the Colombian

Cordillera Oriental on the other), individual values of P range from 4.0 to 5.5 mm

BLANDIN P. et a!., Revision of the Andean Eryphanis zolvizora group

Fig. 4. Habitus of males. Red spot: holotype (HT) or lectotype (LT). Yellow spot: syntype (ST) orparatype

(PT). A. Eryphanis zolvizora zolvizora (Hewitson, 1877), LT (Bolivia; BMNH). B. The southernmost

known specimen of E. z. zolvizora (Manchones, Santa Cruz, Bolivia; MHNC). C. Eryphanis zolvizora

inca ssp. nov., HT (Aguas Calientes, Cuzco, Peru; MJP). D. Eryphanis zolvizora chachapoya ssp. nov.,

HT (Huamanpata, Amazonas, Peru; MJP). E. Eryphanis z. chachapoya ssp. nov., ST of Eryphanis

opimus Staudinger, 1887, from Chanchamayo, Junin, Peru (BMNH). F. PT of E. z. chachapoya ssp.

nov. with small spots (Alto Nieva, Amazonas, Peru; MNHN, PBGL 706).

European Journal of Taxonomy 71: 1-66 (2014)

Fig. 5. Habitus of males. Red spot: holotype (HT) or lectotype (LT). A. Eryphanis zolvizora greeneyi

Penz & DeVries, 2008, stat. rev., HT (Yanayacu, Napo, Ecuador; BMNH). B. E. z. greeneyi , specimen

with increased forewing spots (Baeza, Napo, Ecuador, Pinas). C. Eryphanis zolvizora casagrande

ssp. nov., HT (Altaquer, Narino, Colombia; ICNUN). D. E. z. casagrande from Western Ecuador (Las

Gralarias, Pichincha; FLMNH). E. Eryphanis zolvizora opimus (Staudinger, 1887), LT (Manizales,

Colombia; ZMHB). F. E. z. opimus , specimen with reduced spots (Cauca, Colombia; MNHN, CG).

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

The ground colour of the male dorsal surface is a dark brown (faded into a paler brown in old specimens).

There is a violet-blue iridescence on the median part of the wings. This iridescence is very faint in

Bolivian specimens, notably on the HW, and more pronounced in Venezuelan specimens, notably from

the Cordillera de la Costa. Specimens from southern Peru have a more brilliant violet than Bolivian

males, on both the FW and HW; however, there are individual variations. Specimens from northern Peru

and eastern and western Ecuador are intermediate. In most Colombian specimens, the iridescence is

notably pronounced, but generally less than in Venezuelan specimens.

On the DFW, orange marks form a submarginal band. Bolivian males have a supplementary orange

mark in R5-M1 (sometimes not very pronounced) forming a proximal branch to this submarginal

band. This mark appears, more or less faded, in a majority of specimens from southern Peru. It is more

often absent in other populations, or very faint in some Colombian specimens. The other orange marks

Fig. 6. Habitus of males. Red spot: holotype (HT). Yellow spot: paratype (PT). A. Eryphanis zolvizora

reyi ssp. nov., HT (La Chimenea, Barinas, Venezuela; MIZA). B. E. z. reyi ssp. nov., PT (Charala,

Colombia; MNHN, PBB 2321). C. Eryphanis zolvizora isabelae ssp. nov., HT (Choroni, Aragua,

Venezuela; R, to be donated to MIZA). D. E. z. isabelae ssp. nov., PT; specimen with reduced forewing

spots (Choroni, Aragua, Venezuela; R).

European Journal of Taxonomy 71: 1-66 (2014)

Table 3. Variation of the hindwing projection index P in males (n = number of individuals on which

measurements have been made, directly or on scaled pictures).

Country

P (mm)

Average value

Minimum value

Maximum value

Bolivia

3.3

2.0

4.5

Southern Peru and 2 specimens from

central Peru

4.4

3.7

5.8

Northern Peru and 4 specimens from

central Pern

5.6

4.8

6.5

Eastern Ecuador

5.5

4.6

6.5

Pichincha, Western Ecuador

4.4

Narino, Western Colombia

5.5

Cauca Valley, Colombia

3.9

3.0

4.5

North of eastern cordillera, Colombia

4.9

4.8

5.0

El Tama and Merida ranges,

Venezuela

3.6

2.5

4.5

Cordillera de la Costa, Venezuela

4.3

3.5

5.5

forming the submarginal band vary widely in size. In Bolivia and southern Peru, males have generally

large marks; on veins M3, Cul and Cu2 they are clearly separated (often more widely in southern

Peru). In northern Peru the marks can be very large, as in several specimens from Amazonas, but also

small, as in several specimens from San Martin and Huanuco. The same variability exists in Ecuador,

Colombia and Venezuela. We hypothesize that this variation is a matter of intra-population variation,

as clearly illustrated by specimens from the Santo Domingo Valley (Barinas, Venezuela). In Colombian

and Venezuelan specimens, where the marks are large, they are often crescent shaped, and they are

barely separated by the veins, which remain brown-black.

The length of the androconial patch in the HW anal fold varies from 5.0 to 9.0 mm (Table 4). Considering

the variation between specimens of the same geographical group, we cannot conclude that there is any

geographical variation other than in the Venezuelan Cordillera de la Costa, where the androconial length

is clearly longer. Moreover, in this population, the androconial patch is pale greyish-brown, while it is

a bright creamy, pale citrus yellow in all other populations, except a few specimens that also exhibit a

pale greyish-brown.

The ventral background colour is ochre-brown, with little geographical variation. In Bolivian and

southern Peruvian specimens, the ochre-brown often shows a yellowish tinge on the distal part of the

wings, notably on the FW margins. The median area, between the white stripes, presents a contrasting

darker ochre-brown colour. In northern Peru a majority of specimens exhibits a darker pattern: the FW

margins are less yellow, and the median area is generally a dark chocolate-brown. Eastern Ecuadorian

specimens are often more yellowish than northern Peruvian ones. Colombian and Venezuelan specimens

have a lighter, yellowish aspect; the median area is lighter than in specimens from other countries.

However, these differences do not have a definite value: for example, in northern Peru and Ecuador

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Table 4. Variations of the length of the male HW androconial patch (n = number of individuals on which

measurements have been made, directly or on scaled pictures).

Country

Length of the androconial patch (mm)

average

minimum

maximum

Bolivia

7.1

6.0

8.0

Southern Pern and 2 specimens from

central Peru

7.4

6.4

8.5

Northern Pern and 4 specimens from

central Peru

7.3

5.0

9.0

Eastern Ecuador

6.8

5.8

8.0

Pichincha, Western Ecuador

8.3

Narino, Western Colombia

7.5

Cauca Valley, Colombia

8.1

9.0

North of eastern cordillera, Colombia

7.6

6.6

8.5

El Tama and Merida ranges,

Venezuela

7.0

5.8

7.5

Cordillera de la Costa, Venezuela

9.0

8.3

9.5

Table 5. Variations of the larger diameter (O) of the Cul-Cu2 ocellus on male ventral hindwing, (n =

number of individuals on which measurements have been made, directly or on scaled pictures).

Country

O (mm)

Average value

Minimum value

Maximum value

Bolivia

12.0

10.5

14.7

Southern Peru and 2 specimens from

central Peru

12.1

10.1

14.7

Northern Peru and 4 specimens from

central Peru

10.0

8.5

11.5

Eastern Ecuador

9.5

8.0

10.8

Pichincha, Western Ecuador

10.5

Narino, Western Colombia

10.1

Cauca valley, Colombia

10.7

9.5

11.9

North of eastern cordillera, Colombia

10.0

9.9

10.0

El Tama and Merida ranges,

Venezuela

9.5

7.0

11.0

Cordillera de la Costa, Venezuela

8.7

7.0

10.5

European Journal of Taxonomy 71: 1-66 (2014)

as well, there are some specimens where the median area is not a dark chocolate-brown, but a lighter

tobacco ochre-brown, as in southern Peru on the one hand, and as in Colombia on the other.

Both the VFW and VHW are crossed by two conspicuous white stripes. On the VFW, they are clearly

separated in the cell Cu2-2A in Bolivian and southern Peruvian specimens, the inner margin of the distal

stripe just forming a small tooth. In northern Peruvian and eastern Ecuadorian specimens, this tooth

is often more developed, forming a dark projection which can connect with the outside black margin

of the proximal stripe; exceptionally, this black connection is more or less filled with white scales. In

specimens from western Ecuador and Narino in south-western Colombia, the connection is complete,

forming a white bridge with black margins. On the contrary, Colombian specimens from the Cauca

Valley have just a short tooth, as in Bolivian males. In males from the north of the eastern Colombian

cordillera and from Venezuela, the tooth is elongated and more often reaches the proximal stripe. On

the VHW, the white stripes generally extend far posterior to the Cul-Cu2 ocellus, especially the distal

stripe. However, in the Narino males the two stripes are shorter, reaching no more than the anterior edge

of the ocelli.

Within the VFW cell, there is an isolated white patch next to the base of Cul-Cu2. In specimens from

the Cordillera de la Costa (northern Venezuela), this patch is generally much larger than in any other

population, and roughly triangular or in the form of an inverted heart. At the angle formed by the cubital

vein of the cell and Cu2, there is in the population of the Cordillera de la Costa an elongated white spot,

which is always much smaller or even absent in all other populations.

In Bolivia and southern Peru, the VHW costal ocellus is circled by a brown, incomplete ring, a little

paler than the median area. In northern Peru, this ring is lighter, and more or less covered by white

scales, thus contrasting more strongly with the dark-brown median area. This is also the case in eastern

Ecuadorian specimens, where the ring is often almost completely covered with bright white scales. In the

only western Ecuadorian male that we have studied, the costal ocellus is circled by an almost complete

white ring. It is the same in males from south-western Colombia (Narino). In Colombian specimens

from Cauca Valley, the ring is generally a poorly contrasting pale brown, with sometimes a few whitish

scales. On the contrary, specimens from the eastern cordillera and from Venezuela (Cordillera de Merida

and Cordillera de la Costa) have whitish scales more developed. In all populations, there is an inner

black circle, which is generally complete, notably in specimens from the Cauca Valley, but which is

interrupted along the costa in specimens from the Cordillera de Merida.

The size of the VHW Cul-Cu2 ocellus is variable in all populations. However, it is clearly larger in

Bolivian and southern Peruvian specimens. It is notably smaller in northern Peru and the Northern

Andes, having the smallest values in northern Venezuela (Table 5).

The dark brown lines in the VHW cell, and the dark brown curved line above the Cul-Cu2 ocellus,

are well developed in Bolivian specimens. This is also the case for southern Peruvian specimens, but

the dark brown curved line is sometimes faded, or even absent. In northern Peruvian specimens, all

these lines are strongly faded, if not completely absent. They are not very well developed in eastern

Ecuadorian specimens, being sometimes as faded as in northern Peru. On the contrary, they are well

developed in western Ecuadorian, Colombian and Venezuelan specimens.

Female habitus (Figs 7A-F, 8A-B)

FWL generally varies between 61 and 68 mm in all populations. The general colour on both dorsal and

ventral surfaces is lighter than in males, and there is usually an intense, median, violet-blue sheen to

both FW and HW. Bolivian specimens have a fainter iridescence than all other females. On the contrary,

in the northern Venezuelan populations, the dorsal colour is lighter, and more bluish, than in other

populations. Other characters vary between populations as in males. In the females from Narino (south-

BLANDIN P. et a!., Revision of the Andean Eryphanis zolvizora group

Fig. 7. Habitus of females. Yellow spot: paratype (PT). A. Eryphanis zolvizora zolvizora (Hewitson,

1877) (San Jacinto, Bolivia; BMNH). B. Eryphanis zolvizora chachapoya ssp. nov., PT (San Jose de

Molinopampa, Amazonas, Peru; MNHN, PBGL 198). C. Eryphanis zolvizora greeneyi Penz & DeVries,

2008, stat. rev. (Ecuador; JFLC). D. Eryphanis zolvizora casagrande ssp. nov., PT (LaPlanada, Narino,

Colombia; IAvH). E. E. z. casagrande ssp. nov. (Huigra, Chimborazo, Ecuador; ANSP). F. Eryphanis

zolvizora opimus (Staudinger, 1887) (Rio Agua (presumably Rio Aguacatal), Valle del Cauca, Colombia;

BMNH).

European Journal of Taxonomy 71: 1-66 (2014)

Fig. 8. Habitus of females. Yellow spot: paratype (PT). A. Eryphanis zolvizora reyi ssp. nov., PT (La

Chimenea, Barinas, Colombia; MIZA). B. Eryphanis zolvizora isabelae ssp. nov., PT (Colonia Tovar,

Aragua, Venezuela; MCC).

Fig. 9. Inter- and intra-individual variations of the extremities of valvae in a population of E. zolvizora

chachapoya ssp. nov. from San Jose de Molinopampa, Amazonas, Peru (MNHN).

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

Fig. 10. Male genitalia. A. Eryphanis zolvizora zolvizora (Caranavi, Bolivia; MNHN, PBB 2286). B.

Eryphanis zolvizora inca ssp. nov., PT (Llactohuaman, Cusco, Peru; MJP).

Fig. 11. Gnathos and valva extremity in various specimens of Eryphanis zolvizora inca ssp. nov. A. PT

from Santo Domingo, Puno, Peru (BMNH 8223). B. PT from San Torenzo, Cusco, Peru (MNHN, PPB

2115). C. HT from Aguas Calientes, Cusco, Peru (MJP). D. PT from Alfamayo, Cusco, Peru (MNHN,

PBB 2184). E. PT from Calabaza, Junin, Peru (MNHN, PBB 2308). F. Specimen from Oxapampa,

Pasco, Peru (MJP).

European Journal of Taxonomy 71: 1-66 (2014)

western Colombia) and Huigra (western Ecuador), on the VFW, a complete connection between the

white stripes forms a white “bridge”; the VHW white stripes are longer than in males, but they remain

very slightly shorter than in most other females.

Male genitalia (Figs 3A1-A2, 9, 10A-B, 11A-E, 12A-J, 13A-I, 14A-F)

In all specimens, the uncus is humped; the gnathi are more or less quadrangular or trapezoid, with long,

narrow and sharp prongs; the valvae are long and globally narrow; dorsally, they show a longitudinal,

clearly delimited ridge bearing a row of spines. The phallus has a more or less perpendicular prong

anterior to its distal opening, and the distal end of the phallus forms a prong beyond the opening.

Diagnostic characters are provided by the gnathi (size and shape) and by the valvae (shape; number

and arrangement of spines). It should be emphasized that we have observed individual variations of

valvae characters within some populations. Moreover, there are commonly intra-individual differences

between the right and left valvae, with variations in spine number (Fig. 9).

Fig. 12. Male genitalia of various specimens of Eryphanis zolvizora chachapoya ssp. nov. A. PT from

San Jose de Molinopampa, Amazonas, Peru (MNHN, PBGF 190). B. PT from Alto Nieva, Amazonas,

Peru (MNHN, PBB 2157). C. Specimen from San Augustin, San Martin, Peru (MNHN, PBGF 521).

D. PT from Cumpang, Fa Fibertad, Peru (MJP). E. HT from Huamanpata, Amazonas, Peru (MJP).

F. Specimen from Carpish, Huanuco, Peru (MNHN, PBB 2311). G. Specimen from San Francisco,

Chanchamayo, Junin, Peru (MNHN, PBB 1407). H. PT from Fa Suiza, Pasco, Peru (UFPC). I. PT from

Huancabamba, Pasco, Peru (BMNH 8224). J. PT from Cushi, Pasco, Peru (BMNH 8225).

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

In Bolivian specimens, the valva is characterized by a subterminal swelling, where the ridge has 2-3

strong teeth; beyond the swelling, there is a smooth, rounded gap, followed toward the extremity by a

row of small, regular spines. The gnathos is relatively small, the ventral region being reduced, with the

prong quite close to the ventral margin (Fig. 3A1).

In specimens from southern and central Peru, the valva is regular, without swelling; the ridge bears

a row of more than 6 spines, the size of which varies more or less regularly towards the extremity;

the gnathos, as in Bolivian specimens, is smaller than in specimens from northern Peru (Fig. 11A-E).

The northernmost specimens presenting these features are labelled “Rio Perene” (Junin Department;

genitalia MJPZOTV 005PB) and “Oxapampa” (Pasco Department; genitalia MJPZOTV 007PB).

In specimens from northern Peru (Huanuco, Ta Libertad, Amazonas and San Martin Departments)

the gnathos is large, and its ventral region is strongly developed. The valva axis has no swelling, or

only a weak one, but the ridge more often has a pronounced swelling, bearing 2-3 very strong spines;

distally, there is more often a clear gap, followed by a short distal row of short spines, the size of which

reduces towards the extremity (Fig. 12A-J). In central Peru, a specimen labelled “Rio Perene” (genitalia

MJPZOTV 06PB) and three other specimens (San Francisco, Chanchamayo, Junin, MNHN PBB 1407;

Fig. 13. A-C. Male genitalia of Eryphanis zolvizora greeneyi Penz & DeVries, 2008, stat. rev. A.

Specimen from Rio Blanco, Ecuador (MNHN, PBB 729). B. Specimen from Rio Blanco, Ecuador

(MNHN, PBB 537). C. Specimen from Rio Blanco, Ecuador (MNHN, PBB 257). - D-F. Male genitalia

of Eryphanis zolvizora casagrande ssp. nov. D. HT from Altaquer, Narino, Colombia (ICNUN). E. PT

from Ricaurte, Narino, Colombia (IAvH; drawing by Jean-Fran 90 is Le Crom, JFL 336). F. Specimen

from Las Gralarias, Pichincha, Ecuador (FLMNH). - G-I. Male genitalia of Eryphanis zolvizora opimus

(Staudinger, 1887). G. Specimen from Manizales, Caldas, Colombia (BMNH 8228). FI. Specimen from

Armenia, Quindio, Colombia (MNHN, CG). I. Specimen from Pereira, Risaralda, Colombia (BMNH

8227).

European Journal of Taxonomy 71: 1-66 (2014)

Huancabamba, Pasco, BMNH VIAL 8224; and Oxapampa, La Suiza, Pasco, UFPC) present genitalia

with similar characteristics (Figs 3A2, 12H).

In eastern Ecuadorian specimens the valva is regular, without swelling; the ridge bears a row of more

than 6 spines, the size of which varies more or less regularly towards the extremity. The gnathos is

smaller than in specimens from northern Peru, more or less of the same size as those in Bolivia and

southern Peru (Fig. 13A-C). The only male we studied from western Ecuador (Fig. 13F), the two males

from Narino (Fig. 13D-E), other Colombian males (Fig. 12G-I) and Venezuelan males (Fig. 14A-F)

have similar characteristics, but the gnathos can be larger. Taking into account probable individual

variability of valva features, it is not possible to separate either Colombian specimens or Venezuelan

specimens from Ecuadorian specimens.

Female genitalia (Figs 3B1-B2, 15A-0)

The sterigma forms a dorsal arch, with two symmetric projections directed inward. These projections

are generally divided in one dorsal branch and one shorter ventral branch, of which size and shape are

Fig. 14. A-C. Male genitalia of Eryphanis zolvizora reyi ssp. nov. A. PT from La Mina, Barinas,

Venezuela (R, 115-JCSC). B. PT from San Isidro, Barinas, Venezuela (MNHN, PBB 2324). C. PT

from Charala, Santander, Colombia (MNHN, PBB 2321). D-F. Male genitalia of Eryphanis zolvizora

isabelae ssp. nov. D. PT from Choroni, Aragua, Venezuela (R, 112-JCSC). E. PT from Choroni, Aragua,

Venezuela (MIZA, 117-JCSC). F. PT from Choroni, Aragua, Venezuela (R, 111-JCSC).

BLANDIN P. et a!., Revision of the Andean Eryphanis zolvizora group

Fig. 15. Female sterigma. A. Eryphanis zolvizora zolvizora (Hewitson, 1877) (Bolivia; MNHN, CG).

B-H. Eryphanis zolvizora chachapoya ssp. nov. B. PT from Cushi, Pasco, Peru (BMNH 8226). C. PT

from Carpish, Huanuco, Peru (MNHN, PBB 2285). D. Specimen from Mallqui, Huanuco, Peru (MNHN,

PBB 2327). E. PT from San Jose de Molinopampa, Amazonas, Peru (MNHN, PBGL 198). F. PT from

Nieva, Amazonas, Peru (MNHN, PBGL 596). G. PT from Alto Nieva, Amazonas, Peru (MNHN, PBGL

519). H. Specimen from Alto Nieva, Amazonas, Peru (MNHN, PBGL 520). I. Eryphanis zolvizora

greeneyi Penz & DeVries, 2008, stat. rev., specimen from Valladolid, Zamora-Chinchipe, Ecuador

(MNHN, PBB 2288). J. Eryphanis zolvizora casagrande ssp. nov., PT from La Planada, Narino,

Colombia (IAvH; drawing by Jean-Frant^ois Le Crom, JFL 337). K. E. zolvizora opimus (Staudinger,

1887), specimen from ‘Colombie’ (MNHN, CG). L. Eryphanis zolvizora reyi ssp. nov., PT from La

Mina, Barinas, Venezuela (R, 116-JCSC). M-O. E. zolvizora isabelae ssp. nov. M. PT from Choroni,

Aragua, Venezuela (R, 114-JCSC). N. PT from Rancho Grande, Aragua, Venezuela (MIZA, 118-JCSC).

O. PT from La Llanada, Distrito Federal, Venezuela (MNHN, PBB 2322).

European Journal of Taxonomy 71: 1-66 (2014)

variable, probably at the intra-populational level; moreover, specimens sometimes present differences

between the two ventral branches. The shape, positioning and orientation of the dorsal branches seem to

provide diagnostic characters, but, as there are individual variations, more specimens should be studied

for definite conclusions.

Bolivian females (2 specimens dissected) have projections with long, narrow and sharp dorsal branches,

directed dorsally towards the internal side of the arch; their extremities can cross (Fig. 15A). The

Bolivian specimen studied by Penz (2008, fig. 2G) shows similar characteristics. Unfortunately, no

southern Peruvian female was available for dissection. In northern Peruvian females (7 specimens

dissected; Fig. 15B-H), the pointed dorsal branches are generally shorter than in Bolivian females, and

their extremities are more distant from the internal side of the arch. Nevertheless, there are individual

variations, as illustrated by three specimens from the Nieva Valley (Amazonas Department): the

sterigma of one specimen is rather similar to those of Bolivian females (Fig. 15H). In the southern

Ecuadorian female from Valladolid (Fig. 151), the dorsal branches are close to the internal side of the

arch; moreover, they have notably irregular outlines; these features are rather similar to those of the E.

greeneyi female figured by Penz (2008, fig. 2H). In the female from Narino, the dorsal branches are

rather similar to those of E. greeneyi , with irregular outlines, but they are more distant from the internal

side of the arch (Fig. 15J). In a Colombian female probably coming from the Cauca Valley (Fig. 15K),

the dorsal branches have a regular outline similar to northern Peruvian females, but their extremities

are notably less distant from the internal side of the arch; nevertheless, the differences with one of the

Peruvian females (cf. Fig. 15H) are weak. In one female from the Santo Domingo Valley in western

Venezuela, the inward projections are thicker; consequently, the dorsal branches are shorter, but their

extremities cross close to the internal side of the arch (Fig. 15L); however, in another female, also from

the Cordillera de Merida, the sterigma is fairly similar to that of the female from the Cauca Valley. On

the contrary, in females from the Cordillera de la Costa, the projections are narrower, and the bases of the

dorsal branches are widely separated from the internal side of the arch; nevertheless, their extremities

are close to it, and can cross (Fig. 15M-0).

One or several species: taxonomic decision

Considering male genitalia, four groups are clear-cut: a Bolivian group, a southern Peruvian group, a

northern Peruvian group, and a Northern Andean group, including populations from Ecuador, Colombia

and Venezuela. Female genitalia possibly allow the separation of a Bolivian group, a northern Peruvian

group, an eastern Ecuador population, and a Colombian-Venezuelan population (and, possibly, the

population from the Cordillera de la Costa). However, as individual variations do exist, as observed in

females from northern Peru, more specimens are needed to check the diagnostic role of female genitalia.

Consequently, we have established the diagnoses without indicating the characters of female genitalia

(but they are described under description sections).

When wing characters are taken into account, the Bolivian, southern and northern Peruvian groups

are confirmed, but the Northern Andean group must be divided into an eastern Ecuadorian group, a

western Ecuadorian and south-western Colombian group, a group in the Cauca Valley, a group in the

northern Colombian eastern cordillera and the Venezuelan Cordillera de Merida, and a group in northern

Venezuela (Cordillera de la Costa). We consider these eight groups as different taxa.

In the present state of knowledge, these geographical groups seem allopatric. Contact or transition zones

are not documented. In central Peru, a few specimens suggest a possible geographical overlap between the

southern and the northern Peruvian groups. In MJP (coll. Paul Martin), there are two specimens labelled

“Rio Perene”, one belonging to the southern group, the other to the northern group; moreover, in the same

collection, there is a specimen of the southern group labelled “Oxapampa”, and we know a specimen

from the northern group labelled “La Suiza, Oxapampa” (UFPC). Unfortunately, the localities of the

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

three specimens in MJP are unreliable (Gerardo Lamas, pers. comm.), making any definite conclusion

impossible. However, from the area of Chanchamayo, a classic locality in the Junin Department, there is

a specimen of the southern group labelled “Chanchamayo” in the AMNH, and a specimen of the northern

group labelled “San Francisco, Chanchamayo” in the MNHN. Moreover, Staudinger’s opimns syntype

from Chanchamayo (Fig. 4E) probably belongs to the northern group. Therefore, we can conclude that

the two groups are both represented in a limited area, and that a geographical overlap is not impossible.

If we consider male genitalia, currently used to separate species in butterfly systematics, it is obvious

that Bolivian specimens show clear differences with specimens from southern Peru and other areas.

Southern Peruvian specimens have some similarities with specimens from the Northern Andes.

Specimens from northern Peru have some unique features in the valvae. Therefore, there are arguments

to consider at least four species. However, habitus characters show affinities between Bolivian and

southern Peruvian specimens on the one hand, and between Bolivian and Northern Andean specimens

on the other. There are also strong similarities between northern Peruvian and eastern Ecuadorian

specimens. As a consequence, separations suggested by male genitalia do not fit exactly with separations

suggested by the habitus. Therefore, as no sympatry is documented between any pair of neighbouring

geographical groups, we chose to classify the E. zolvizora group as one species, divided into eight

subspecies: Eryphanis zolvizora zolvizora (Hewitson, 1877) limited to Bolivia; a new subspecies for

southern Peruvian populations; another for northern Peruvian populations; E. zolvizora greeneyi (Penz

& DeVries, 2008), stat. rev., for eastern Ecuador; a new subspecies for western Ecuador and south¬

western Colombia (Narino Department); E. zolvizora opimus Staudinger, 1887, from the Cauca Valley;

a new subspecies for the north of Colombian eastern cordillera and western Venezuela (Cordillera de

Merida); and a new subspecies for the Venezuelan Cordillera de la Costa.

Systematics

Class Hexapoda Blainville, 1816

Order Lepidoptera Linnaeus, 1758

Superfamily Papilionoidea Latreille, 1802

Family Nymphalidae Rafinesque, 1815

Subfamily Morphinae Newman, 1834

Tribe Brassolini Boisduval, 1836

Genus Eryphanis Boisduval, 1870

The arrangements of the taxonomy of the Eryphanis zolvizora group made in this revision are listed

below, modifying the arrangements published by Casagrande (2004) and Penz (2008):

Eryphanis zolvizora (Hewitson, 1877)

zolvizora zolvizora (Hewitson, 1877)

zolvizora opimus Staudinger, 1887

zolvizora greeneyi Penz & DeVries, 2008, stat. rev.

zolvizora inca Blandin ssp. nov.

zolvizora chachapoya Blandin ssp. nov.

zolvizora casagrande Bristow ssp. nov.

zolvizora reyi Bristow, Neild, De Sousa & Huertas ssp. nov.

zolvizora isabelae Neild & De Sousa ssp. nov.

Taxa are presented below in a South-North arrangement because the species was described from

Bolivia. Diagnoses are detailed for males, as they have been studied in larger numbers than females.

Important diagnostic characters are shared by male and female habitus, except the characters of the male

androconial patches. Male genitalia also provide important diagnostic characters. Because of the low

European Journal of Taxonomy 71 : 1-66 ( 2014 )

number of dissected females, and the existence of some important individual variations, female genitalia

do not provide obvious diagnostic characters.

Eryphanis zolvizora zolvizora (Hewitson, 1877)

Figs 3A1, 4A-B, 7A, 10A, 15A

Pavonia zolvizora Hewitson, 1877. Illustrations of new species of exotic butterflies selected chiefly

from the collections ofW. Wilson Saunders and William C. Hewitson. Vol. IV, London, John Van Voorst,

(Morphinae: Pavonia II, plate [6]).

Caligo zolvizora - Kirby 1877: 847. — Weymer & Maasen 1890: 62.

Pavonia zolvizora -Kirby 1879: 110.

Eryphanis zolvizora zolvizora - Stichel 1904: 33; 1909: 157; 1932: 157. —Fruhstorfer 1912: 312. —

Casagrande 2004: 203.

Eryphanis zolvizora AD’Abrera 1987: 406. —Penz 2008: 14, figs 1G, 2G, 4G, 5G, 6G (re-description).

Diagnosis

Poorly developed HW projection (Figs la, 4A-B); moderately developed oblique orange spot on DFW

between the R5 and Ml veins (Figslb, 4A-B); VHW costal ocellus circled by a brown, incomplete ring

without white scales (Figs 21, 4A-B); large Cul-Cu2 ocellus (average widest diameter: 12mm) (Figs

2m, 4A-B). The valva is characterized by a subterminal swelling, where the ridge has 2-3 strong teeth;

beyond, a smooth rounded gap followed by a row of small regular spines. Gnathos is relatively small, its

ventral region being reduced (Figs 3A1, 10A).

Type material

Lectotype, here designated

S, two white labels, printed and written in black: //Bolivia (Buckley) [hand written] Hewitson Coll.

79-69. Pavonia [printed] zolvizora Hew. 2 [hand written]// B.M. Type No. Rh [printed] 6110 Pavonia

zolvizora S Hew. [hand written]// (BMNH).

Paralectotype, here designated

S, one label, hand written in black: //Hewitson 1// (BMNH).

Type locality

Bolivia.

Description

Male

The FWL varies from 57 to 67 mm. The hindwing projection is poorly developed (average P = 3.3; n

= 14; cf. Table 3). Between the R5 and Ml veins there is an oblique orange spot forming the proximal

branch of the submarginal band. The DFW five submarginal orange spots are large, but clearly separated

by the veins (Figs lc, 4A-B). Their form and size are variable (however, the M3-Cul spot is generally

triangular, rarely crescent-like), but they are never strongly reduced. The wings on the dorsal surface are

dark brown, with a weak violet iridescence on FW and absent on HW (Fig. 4A-B). The androconial patch

in the anal area of the DHW is 6.0-8.0 mm long (cf. Table 4); its colour is a bright creamy, pale citrus

yellow, sometimes weakly greyish, thus less bright (Fig. 4A-B). The underside is strongly patterned

(Fig. 4A-B); the ochre-brown background often shows a yellowish tinge on the distal part of the wings,

notably on FW margins; the colour of the median area is a contrasting ochre-brown. In the VFW cell,

the isolated white spot is more often small or medium; however, it is sometimes elongated and in contact

with a chain of whitish spots connected with the white distal limit of the cell (Figs 2g, 4A-B). There is no

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

white spot in the angle formed by the cubital vein of the VFW cell and Cu2, or it is extremely small (Figs

2h, 4A-B). On the VFW, the two white stripes are clearly separated in the cell Cu2-2A; the inner margin

of the distal stripe, which generally forms a small, blunt tooth, is underlined by a thick black line (Figs

2i, 4A-B). On the VHW, the white stripes extend far beyond the Cul-Cu2 ocellus, notably the distal

one, which reaches vein 2A (Figs 2j, 4A-B). Dark brown lines are well developed in the HW cell, as

well as a dark brown curved line above the Cul-Cu2 ocellus (Figs 2k, 4A-B). The VHW costal ocellus

is circled by a brown, incomplete ring, a little paler than the median area; there are no white scales on

this ring; the black ring is generally complete (Fig. 4A-B). The Cul-Cu2 ocellus is large (Fig. 4B); its

widest diameter (O) generally varies between 10.5 and 14.5 mm (cf. Table 5; NB: in the lectotype, the

ocellus is relatively small, see Fig. 4A, an uncommon feature). The genitalia do not present significant

individual variations.

Female

The FWT varies from 62 to 68 mm. The hindwing projection is as reduced as in males. The basal 2/3 rd

of the dorsal surface is a lighter brown than in males, with a distal violet tinge on the DFW and DHW,

brighter than in the male (Fig.7A). On the DFW, the orange spots exhibit the same characteristics as in

males. The ventral surface can be lighter than in males, notably on the distal parts (Fig. 7A), but some

specimens are hardly different from males. The sterigma has inward projections with short, pointed

ventral branches (sometimes with 2 points) and long, narrow dorsal branches which are dorsally oriented

and can cross in the middle (Fig. 15A).

Flight periods

Eryphanis z. zolvizora has been collected from January to May and from September to November.

Distribution

Bolivia. For a long time, the distribution of E. z. zolvizora was poorly known, as most specimens had

been essentially collected near the roads from Cochabamba to Villa Tunari (Cochabamba Department),

and in some localities in the “Coroico - Caranavi” area (La Paz Department). Recent collecting by

Bolivian entomologists has provided new localities, notably in Santa Cruz Department, where the area

of Manchones represents the southernmost locality (ca. 18°45’S) known. Reliable altitudes range from

1000 m to 2600 m. We suppose that E. z. zolvizora exists locally in all cloud forests along the northern

part of the Bolivian Andes. Referring to Fruhstorfer (1912), Penz (2008) suggests that perhaps E.

zolvizora (E. z. zolvizora according to our conception) reaches southern Peru. However, all specimens

we know from southern Peru (Puno Department) belong to E. z. inca ssp. nov. Nevertheless, it should be

noted that there is no information about populations existing near the Bolivian-Peruvian border.

Remarks

The species name zolvizora (originally described in the genus Pavonia ) was authored by Hewitson in

his Illustrations of new species of exotic butterflies selected chiefly from the collections of W. Wilson

Saunders and William C. Hewitson , issued in five volumes between 1852 and 1877 (Lamas et al. 1995).

The date of publication of zolvizora is variously given as 1876 (e.g. in Stichel 1909) or 1877 (e.g. in

Lamas et al. 1995), and the volume of the publication is given as IV (published 1867-1871 according

to the date on the title page) (Stichel 1932) or V (published 1872-1876 according to the date on the title

page) (Stichel 1909; Lamas et al. 1995). The contents of the 5 volumes of ‘Illustrations ’ were issued,

unpaginated (1286 pp - including blank pages and 300 plates), in a series of unnumbered parts. The

text for zolvizora is on the reverse of the page with Pavonia seleucida which has ‘Published January 1 st

1877’ printed at the bottom. Two copies of'Illustrations ’inspected by the authors (at the USNM Library

available online ( www.biodiversitvlibrarv. org ) and in the UMO Library, Oxford), have the text preceded

by two plates. However, two copies inspected at the BMNH, including Lord Rothschild’s bequest copy,

European Journal of Taxonomy 71: 1-66 (2014)

have the text before the plates. The first plate is of P. seleucida (with the page note ‘W.C. Hewitson

del et lith Oct. 1876’), the second of P. zolvizora (with the page note ‘W.C. Hewitson del et lith Nov.

1876’). The original copy acquired by the BMNH has the date of publication of the text of P. zolvizora

pencilled in as ‘l.i.1877’ (recorded also in Griffin 1932). The USNM copy has the dates of publication

of the plates pencilled in as Jan. 1 1877 (presumably after Griffin 1932). This date for plates and text

would suggest that they belong to Vol. V, but in the Systematic Index for all five volumes Pavonia I ([pi.

5], P. seleucida ) and II ([pi. 6], P. zolvizora) are clearly part of Vol. IV, supposedly published 1867-71.

However, in Vol. 5, under ‘CORRECTIONS’ it states that ‘ An index is given by which the binder will

be able to arrange the plates'. It is possible that some copies of The Illustrations were bound in the

order that they were issued or acquired by the different libraries, which could account for Pavonia being

included in Vol. V, rather than in Vol. IV where the Index indicates it was intended to be published. In

summary, as the plates and parts in the various volumes of the 4 sets examined were not bound in order

as they were issued, but according to the order laid down by Hewitson, it seems to us to be correct to

regard P. zolvizora as part of Vol. IV (as in all 4 copies examined), and we regard the name P. zolvizora

to have been published on the 1 Jan. 1877 (but see Tamas et al. 1995).

The specimen identified as the type of E. zolvizora at the BMNH, a male collected by C. Buckley

(Fig. 4A), has no precise locality, being labelled as follows: “Bolivia (Buckley) Hewitson Coll. 79-69.

Pavonia zolvizora Hew. 2”. Following Kirby’s catalogue of Hewitson’s Butterflies at the BMNH (Kirby

1879: 110), it is known that there is another specimen labelled ‘Hewitson 1’, part of the type series at

the BMNH; it was found during recent curation conducted by BH. Therefore, these two specimens are

syntypes. The specimen we have chosen to be the lectotype was formerly considered to be the holotype.

Consequently, the second specimen is a paralectotype.

Eryphanis zolvizora inca Blandin ssp. nov.

Figs 4C, 10B, 11

Diagnosis

On the DFW, the R5-M1 orange mark is more or less faded, sometimes missing (Fig. 4C); HW projection

slightly pronounced (Fig. 4C); VHW costal ocellus is circled by a brown incomplete ring without white

scales (Fig. 4C); on VHW, the widest diamaeter of the Cul-Cu2 ocellus averages 12mm (Fig 4C). The

valva is regular, without swelling; the dorsal ridge bears a row of more than 6 spines, the size of which

varies more or less regularly; gnathos is small (Figs 10B, 11).

Etymology

The holotype was collected by Dr. Gerardo Lamas close to Machu Picchu, the emblematic site of the

famous Inca civilization, in the Cusco area.

Type material (14 specimens)

Holotype

PERU: c?, white label printed in black: // PERU - CU [Cusco]- Aguas Calientes 2050 m 1309-7231

[13°09’S 72°31’W] 21.X.2001 [21 Oct. 2001] G. Lamas// (MJP).

Paratypes

PERU: S, II S. Domingo, Carabaya [Peru] 6000 ft. [13°50’S 69°40’W], Janv.01 [Jan. 1901] Wet season

(Ockenden) // BMNH (E) #806244 // B.M.(N.H) Rhopalocera VIAL No. 8223 // (BMNH); $, II S.

Domingo, Carabaya [Peru], 4500 ft. [13°50’S 69°40’W], 01 [1901], Dry season, [Ockenden] //BMNH

(E) #808079 // B.M.(N.H) Rhopalocera VIAL No. 8576 // (BMNH); S, II S. Domingo, Carabaya [Peru],

4500 ft., [13°50’S 69°40’W], 01 [1901], Dry season. [Ockenden] // BMNH (E) # 808078 // (BMNH);

S, II Inca Minas, Puno [Peru], [13°50’S 69°40’W], 1 Jul. 1900 // (AMNH); c?, // Chirimayo, [Puno] S.

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

E. Peru. VI.01 [Jun. 1901] Dry [dry season] (Ockenden) 1000 ft [13°27’S 70°18’W] // BMNH (E) #

808081 // (BMNH); // Inambari, [Puno] Peru // BMNH (E) # 525986 // (BMNH); II San Lorenzo,

Rio Marcapata, Cusco, Perou [Peru] // PBB 2115 // (MNHN, PBB); II Llacatahuaman, Quebrada

Bagre [Cusco, Peru], 1700 m, [12°52’S 73°30’W], 27/07/1998 [27 Jul. 1998], G. Valencia leg. (Cusco)

// (MJP); (?, // San Pedro, Cusco [Peru], 1400 m, [13°03’S 71°33 , W], 4-8/xi 2007 [4-8 Nov. 2007], F. &

A. West leg. // (MJP); S, I I Alfamayo, route [road] Ollantaytambo-Quillabamba, Cusco, Perou [Peru],

M. Cabrera leg. // PBB 2184 // (MNHN, PBB); S, I I Perou Marchu Pijchu [sic] [Peru, Machu Picchu]

alt. 2000m 10/7/1934 [10 Jul. 1934] A. Larichesy // IRSNIG 10 375 // IRSSNB; II PE. [Peru] 10 12

87 [10 Dec. 1987] Macchu Picchu [sic] [Machu Picchu] Coll. J. Dubois // (MNHN, CG); II Calabaza,

PampaHermosa, Prov. Satipo, Junin, Perou [Peru] 2200 m, [11°29’2.02”S 74°47 , 39.25 ,, W], Avril 2008

[Apr. 2008] // PBB 2308 // (MNHN, PBB).

Other material examined

Several specimens are not included as paratypes (data in Appendix 2), notably two males in MJP,

labelled respectively “Rio Perene (Junin)” (genitalia MJPZOLV 005PB, MJP) and “Oxapampa (Pasco)”

(genitalia MJPZOLV 007PB) as their localities are not reliable (Gerardo Lamas, pers. comm.).

Description

The FWL varies from 60 to 65 mm The holotype FWL is 62 mm The hindwing projection (average P =

4.4; n = 13; cf. Table 3) is more pronounced than in E. z. zolvizora , and less pronounced than in specimens

from northern Peru (E. z. chachapoya ssp. nov.). On the DFW the R5-M1 orange mark is more or less

faded, sometimes missing. The other orange marks are generally large; the violet iridescence is more

pronounced on FW and HW than in E. z. zolvizora and in E. z. chachapoya ssp. nov.; the androconial

patch is a creamy, pale citrus yellow (Fig. 4C). The background colour of the ventral surface, as in E.

z. zolvizora , is less dark than in E. z. chachapoya ssp. nov.; on the VFW, the isolated white spot in the

cell is of medium size, more or less oval, or irregular; there is no white spot in the angle formed by the

cubital vein of the cell and Cu2, or it is very small; the inner margin of the distal white stripe on VFW,

between Cu2 and 2A, generally forms a small, blunt tooth (Fig. 4C). On the VHW, the white stripes

extend far beyond the Cul-Cu2 ocellus, notably the distal one, which reaches vein 2A; in the cell the

dark brown lines are generally developed as in E. z. zolvizora , but the dark brown curved line above the

Cul-Cu2 ocellus is sometimes absent; the costal ocellus is circled by a brown incomplete ring, a little

paler than the median area and without white scales; the black ring is generally complete (Fig. 4C). On

average, the VHW Cul-Cu2 ocellus is as large as in E. z. zolvizora (O average value: 12.1 mm; n = 13;

cf. Table 5) and larger than in all other subspecies. The genitalia do not present significant individual

variations.

Flight periods

Specimens have been collected in January, April, June, July and October.

Distribution

Peru. E. z. inca ssp. nov. extends in southern and central Peru from Puno Department to Junin Department,

and possibly to the south of Pasco Department.

European Journal of Taxonomy 71: 1-66 (2014)

Eryphanis zolvizora chcichapoya Blandin ssp. nov.

Figs 3A2, 4D-F, 7B, 9, 12, 15B-H

Diagnosis

HW projection is strongly pronounced (Figs la, 4D, F); on the DFW the R5-M1 orange patch is often

missing or very reduced (Fig. 4D-F); on the VFW the two white stripes are generally separated in the

cell Cu2-2A, but the inner margin of the distal stripe forms a projection towards the outer black margin

of the proximal stripe (Fig. 4D-F); the dark brown lines on the VHW cell and the curved line above

the Cul-Cu2 ocellus are faint or absent (Fig. 4D-F); the VHW costal ocellus is circled by a clear ring

contrasting with the median area; in many specimens it is widely covered with white scales (Fig. 4D-F);

large VHW Cul-Cu2 ocellus (average widest diameter: 10 mm). The valva axis has no swelling, or only

a weak one; the ridge usually with a pronounced swelling, bearing 2-3 very strong spines; towards the

extremity there is more often a gap, followed by a short distal row of short spines diminishing towards

the extremity; gnathos is large and its ventral region is strongly developed (Fig. 12).

Etymology

Named after the pre-Inca Chachapoya civilization which existed in northern Peru, mainly in the present

Amazonas and San Martin Departments; the Chachapoyas were also called “Warriors of the Clouds”.

Type material (23 specimens)

Holotype

PERU: S, a white label printed in black ink: // PERU, AM [Amazonas], Valle de Huamanpata, Lejia

0620/7727 [6 o 20’S 77°27’W] 2150m 23.X.2005 [23 Oct. 2005] L. Campos // (MJP).

Paratypes

PERU: II San Francisco, Chanchamayo, Junin, Perou, Jan. 1964, Mme Harris leg. // PBB 1407 //

(MNHN, PBB); S, H Oxapampa, La Suiza 2180m Estado Pasco, Peru [10°36’S 75°29’W] 5-13.Junio

2002 [5-13 Jun. 2002] Bernhard Wenczel // (UFPC); II Huancabamba, 6800 ft, [10°23 , S 75°33’W],

Pasco [Peru] // BMNH VIAL 8224 // BMNH (E)# 806245 // (BMNH); II Cushi [Pasco, Peru], 1820

m, (W. Hoffmans) // BMNH (E)# 806246; VIAL 8225 // (BMNH); II Cushi [Pasco, Peru], 1900 m,

[9°58’S 75°42’W], (W. Hoffmans) // (BMNH); $ 9 II Panao Huanuco-Peru 3.000 m, [9 0 54’S 75°58’W],

1-1982 [Jan. 1982] Schunlce Leg // (UFPC); II Carpish, 2300 m, Huanuco, Perou, Jan. 2006 // PBB

2284 // (MNHN, PBB); $ 9 II Cumpang, entre Tayabamba y Ongon, 2400-2700 m, [8°16’S 77°00’W],

21/10/79 [21 Oct. 1979], T. Parker, leg. (La Libertad) [Peru] // genitalia MJPZOLV 004PB // (MJP);

//Environs de Mendoza, [06°23’S 77°27W], Amazonas, Perou, Jun. 1994, B. Calderon leg.//PBB 1900

// (MNHN, PBB); <$, II San Jose de Molinopampa, 2200-2400 m, [06° 12’ 77°34’W], 2005, Amazonas,

Perou, B. Calderon leg. // PBGL 166 // (MNHN, CG); c?, // San Jose de Molinopampa, 2200-2400 m,

[06°12’ 77°34’W], 2008, Amazonas, Perou // PBB 2309 // (MNHN, PBB); II Le long de Falto rio

Nieva, 2250 m, [05°42’40”S 77°47’15”W], 4-9 Mar. 2009, Amazonas, Perou, Jose Ananias Tafur leg.

// PBGL 471 // (MNHN, CG); //Alto Nieva, 1900 m, [05°39’S 77°47’W], sans date [without date],

Amazonas, Perou // PBB 2157 // (MNHN, PBB); $ 9 //Alto Nieva, 1900 m, [05 o 39’S 77°47’W], Dec.

2012, Amazonas, Perou, Joel Clever Pintado leg. //PBGL 706 // (MNHN, CG); //Villa Hermosa, 1500

m, [05°36’S 77°47’W], Dec. 2010, Amazonas, Perou, Joel Clever Pintado leg. // PBGL 594 // (MNHN,

CG); ?, // Cushi, 1900 m, [9°58’S 75°42’W], Prov. Pasco Peru (W. Hoffmans) // BMNH (E)# 806247

VIAL 8226// (BMNH); $, //Carpish, 2300 m, Huanuco, Perou, Jul. 2005 //PBB 2285 //(MNHN, PBB);

5 , // San Jose de Molinopampa, 2200-2400 m, [6°12’S 77°34’W], 5 Jan. 2007, Amazonas, Perou, B.

Calderon leg. // PBGL 198 // (MNHN, CG); ?, //Alto Nieva, 1900-2000 m, [5°39 , 00”S 77 o 46’30”W],

Mar. 2010, Amazonas, Perou, Joel Clever Pintado leg. // PBGL 519 // (MNHN, CG); $, // Nieva, 1900

m, [5°40’S 77°36’W], Sep. 2011, Amazonas, Perou, Joel Clever Pintado leg. // PBGL 596 // (MNHN,

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

CG); §, rio Jenesis [sic], env. 1400 m, [5°32-33’S 77°48-49’W], 11/2012 [Nov. 2012], Amazonas, Peru,

Joel Clever Pintado leg. // (MJP); §, // San Augustin, 1400-1500 m, [5°47’S 77°30’W], Setiembre 2012

[Sep. 2012], San Martin, Peru, Joel Clever Pintado leg. // (MJP).

Other material examined

Data on the 21 specimens not included in the paratypes are given in Appendix 2. Among them, a male

in MJP, labelled “Rio Perene (Junin)” (genitalia MJPZOLV 006PB) has an unreliable locality (Gerardo

Lamas, pers. comm.). One of the syntypes of E. z. opimus from Chanchamayo (in ZMHB) has the HW

projection as well developed as in 77 z. chachapoya ssp. nov. (estimated value ofP index: 5.8). The inner

projection of the distal white stripe on VFW is well developed. The ring of the VHW costal ocellus is

whitish and contrasts strongly with the brown median area, and the diameter of the Cul-Cu2 VHW

ocellus is 10.5 mm. These features are comparable with E. z. chachapoya ssp. nov., but not with E. z.

inca ssp. nov.

Description

Male

The FWL varies from 55 to 65 in in The holotype FWL is 61 mm Average P = 5.5 (n = 27; cf. Table 3).

On the DFW the R5-M1 orange patch is often missing or very reduced. The other orange marks are well

separated; their size is very variable; the violet iridescence is less pronounced than in E. z. inca ssp.

nov.; the androconial patch is a bright creamy, pale citrus yellow (Fig. 4D-F). The background colour

of the ventral surface is generally darker than in E. z. inca ssp. nov.; the isolated white spot in the VFW

cell is of medium size, more or less oval, or irregular; there is no white spot in the angle formed by the

cubital vein of the VFW cell and Cu2, or it is extremely small (Fig. 4D-F). On the VFW the two white

stripes are generally separated in the cell Cu2-2A; nevertheless, the inside margin of the distal stripe

forms a marked inner projection, which often connects with the outside black margin of the proximal

stripe (Fig. 4E); exceptionally (3 of 35 specimens), the projection is filled with white scales and forms a

complete bridge between the two white stripes, as in E. z. casagrande ssp. nov. (Fig. 2i). On the VHW,

the white stripes extend far beyond the Cul-Cu2 ocellus, notably the distal one, which reaches vein 2A.

The dark brown lines on the VHW cell and curved line above the Cul-Cu2 ocellus are very faint or

even absent (Fig. 4D-F). The VHW costal ocellus is circled by a clear ring contrasting with the median

area; moreover, in many specimens it is widely covered with white scales (Figs 21, 4D-F). On average,

the VHW Cul-Cu2 ocellus (O average value: 10.0 mm; n = 31; cf. Table 5) is clearly smaller than in E.

z. inca ssp. nov. (Fig. 4D-F). The valva axis has no swelling, or only a weak one; the ridge more often

has a pronounced swelling, bearing 2-3 very strong spines; towards the extremity, there is more often a

clear gap, followed by a short distal row of short spines, the size of which reduces towards the extremity;

gnathos is large, and its ventral region is strongly developed (Fig. 12). Individual variations of the valvae

have been observed within a local population, but diagnostic characters are not blurred (cf. Fig. 9).

Female

The FWL varies from 61 to 67 mm. Diagnostic characters correspond to those of the males (Fig. 7B).

The violet iridescence is limited but bright. The underside is lighter than in males. The dorsal branches

of the sterigma inner projections are generally shorter than in E. z. zolvizora and E. z. greeneyi , and they

are more separated from the internal side of the arch; nevertheless, there are individual variations, and

one specimen has the sterigma similar to that of E. z. zolvizora specimens (Fig. 15B-H).

Flight periods

Specimens have been collected in January, March, June, July, September, October, November and

December.

European Journal of Taxonomy 71: 1-66 (2014)

Distribution

Peru. E. z. chachapoya ssp. nov. is distributed from Junin Department (central Peru) to Amazonas and

San Martin Departments (northern Peru).

Remarks

This taxon is created for populations from central to northern Peru, which are clearly distinct from E. z.

zolvizora and E. z. inca ssp. nov. considering wing characters as well as genitalia. Wing characters are

closer to the Ecuadorian E. z. greeneyi , but the male genitalia are clearly distinct.

Eryphanis zolvizora greeneyi Penz & DeVries, 2008 stat. rev.

Figs 5A-B, 7C, 13A-C, 151

Eryphanis greeneyi Penz & DeVries, 2008; Penz 2008: 15-17, figs 1H, 2H, 4H, 5H, 6H.

Diagnosis

HW projection strongly pronounced (Fig. 5A); on the DFW the R5-M1 orange patch is usually missing

or very reduced; the isolated white spot in the VFW cell is medium to large (Fig. 5A-B); on the VFW,

in space Cu2-2A, the inner tooth of the distal stripe forms a black point which may just connect with the

outer black margin of the proximal stripe (Fig. 5B); in the VHW cell and above the Cul-Cu2 ocellus,

faint dark brown lines are generally visible; the VHW costal ocellus has a narrow, bright white outer

ring (Fig. 5A-B). The valva axis has no swelling; the dorsal ridge has a row of more than 6 spines,

diminishing towards the extremity (Fig. 13A).

Type material

Holotype

ECUADOR: S, three labels: // Ecuador, Napo, Yanayacu Biological Station, 5km W of Cosanga,

May 2007// reared on Chusquea scandens, H. F. Greeney// Eryphanis grteeneyi Penz and DeVries

HOFOTYPE// (BMNH).

Paratypes

Detailed information in Penz (2008). ECUADOR: 5 SS, Napo Prov.,Yanayacu Biological Station

(BMNH); Provincia Napo (MECN); Provincia Napo (AMNH); Rio Blanco, near Banos, (DeVries

Collection, USA); Balzapampa, Ecuador (FFMNH). 4 § §, Napo, Yanayacu Biological Station (BMNH);

Napo Prov., Yanayacu Biological Station (MECN); Napo, Biol. Yanayacu (AMNH); Provincia Napo,

San Isidro (DeVries Collection, USA).

Type locality

Ecuador, Napo Province, Cosanga.

Description

Male

The FWL varies from 55 to 66 mm (our data, and according to Penz 2008). Average P = 5.5 (n = 11; cf.

Table 3), as in E. z. chachapoya ssp. nov., differing in this way from other subspecies from the Northern

Andes. On the DFW the R5-M1 orange patch is more often missing or very reduced; the other orange

marks are well separated, at least by the veins; their size is very variable; the violet iridescence is fainter

than in E. z. chachapoya ssp. nov.; the androconial patch is a bright creamy, pale citrus yellow (Fig.

5A-B). The underside is close to that of E. z. opimus as it is generally lighter, with a more yellowish tint,

than in E. z. chachapoya ssp. nov. (Fig. 5 A); however, there are intermediate specimens, which resemble

more closely specimens of E. z. chachapoya ssp. nov. (Fig. 5B). On the VFW, the isolated white spot in

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

cell is medium to large, with an irregular, more or less quadrangular, triangular or oval shape; there is no

white spot in the angle formed by the cubital vein of the cell and Cu2, or it is very small (Fig. 5 A); on

the VFW, in space Cu2-2A, the inner tooth of the distal stripe is well developed, forming a black point

which in some specimens forms a narrow black bridge with the outside black margin of the proximal

stripe (Fig. 5A-B). On the VHW, the white stripes extend far beyond the Cul-Cu2 ocellus, notably the

distal one, which reaches vein 2A; in the cell and above the Cul-Cu2 ocellus, the dark brown lines

are generally visible, but very faint in some specimens; the VHW costal ocellus has a narrow, bright

white outer ring (sometimes the white scales do not cover the ring entirely); the black ring is generally

complete (Fig. 5A-B); the widest diameter of the Cul-Cu2 ocellus has an average value of 9.5 mm (n

= 11; cf. Table 5). Genitalia exhibit weak individual variations (Fig. 13A-C). The valva axis has no

swelling, and the dorsal ridge bears a row of more than 6 spines, decreasing in size more or less regularly

towards the extremity; in that way, E. z. greeneyi strongly differs from E. z. chachapoya ssp. nov., while

it is similar to Colombian and Venezuelan subspecies. The gnathos is smaller than in E. z. chachapoya

ssp. nov., but its size probably falls within the range of individual variations in E. z. opimus.

Note. The five orange spots that comprise the FW band vary in size. In the holotype from Yanayacu, the

three lower spots are crescentic and just touch; in others, the spots are smaller, less obviously crescentic

and do not touch. Both types of FW band are seen in Rio Blanco specimens and it is assumed that they

represent individual variation. A more extreme example is the male figured by Pinas (2004, figs 653 and

654) from Baeza, where the FW spots are large and contiguous (Fig. 5B). As Baeza lies fairly close to

Yanayacu, it is assumed that this phenotype falls within the variation of E. z. greeneyi.

Female

The FWT varies from 64 to 68 mm (our data and after Penz 2008). Considering the habitus (Fig. 7C),

the diagnostic characters correspond to those of the males, except the size of the Cul-Cu2 ocellus, which

can be large (O = 11.5 mm in a female from Valladolid). The violet iridescence is limited but bright on

both the FW and HW. The underside is lighter than in males. The dorsal branches of the sterigma are

close to the internal side of the arch. Moreover, they have marked irregular outlines; in that way, E. z.

greeneyi differs from other subspecies.

Flight periods

E. z. greeneyi appears to fly throughout the year, with May being the commonest month of capture.

Distribution

Ecuador. Eryphanis z. greeneyi extends throughout eastern Ecuador (possibly extending into southern

Colombia) at altitudes generally between 1500 and 2220 m, but there are some records (if genuine)

which suggest that it may occur as low as 600 m. It is well known from the much-collected area of

the upper Rio Pastaza between the Rio Blanco and Rio Verde and extends to the extreme south of

Ecuador (Podocarpus, Valladolid and Cordillera Llagunillas). A specimen in MZUJ is labelled from El

Oro Province, near Balsas; however, it is likely that this specimen had been mislabelled, as it belongs to

material donated by K. Jasinski (Keith Willmott, pers. comm.).

Remarks

The taxon Eryphanis greeneyi , described as a distinct species, was based on 7 males and 4 females, of

which several were reared by H.F. Greeney at the Yanayacu Biological Station, near Cosanga, Ecuador

(Napo Province). The holotype (a male) and a female paratype are deposited in the BMNH. The name

was given in honour of the naturalist Harold Francis Greeney III, who first reared this taxon.

Two males labelled ‘Caqueta’ in the Le Crom collection were probably collected on the road from

Altamira to Florencia at a height of between 1500 and 2000 m (J.F. Le Crom, pers. comm). No other E.

European Journal of Taxonomy 71: 1-66 (2014)

zolvizora specimen is known from eastern slopes of the Colombian Cordillera Oriental. These two males

are somewhat darker than most other Colombian specimens. Geographically, they are closest to E. z.

greeneyi from Ecuador, to which we putatively attach them; however, the HW projection at Cul is only

poorly developed and there is no narrow white line surrounding the costal HW ocellus.

Eryphanis zolvizora casagrande Bristow ssp. nov.

Figs 5C-D, 7D-E, 13D-F, 15J

Diagnosis

HW projection only poorly developed; no R5-M1 orange mark on DFW (Fig. 5C); there is a wide bridge

in space Cu2-2A on the VFW formed by a complete white connection between the white stripes (Figs 2i,

5C); the isolated white spot in the VFW cell is quite large and semicircular; the narrow outer white stripe

on the VHW only just extends down to the smaller of the lower ocelli; the broader inner white stripe only

extends as far as the larger of the lower ocelli (Fig. 5C); the dark brown lines in the VHW cell and curved

line above the Cul-Cu2 ocellus are well developed, the distal lines form two conjoined elongate circles

(Fig. 5C). The valva axis has no swelling; the dorsal ridge has a row of more than 6 spines, diminishing

towards the extremity (Fig. 13D).

Etymology

Named after Prof. Dr. Mima Casagrande (Universidade Federal do Parana, Curitiba, Parana, Brazil) in

recognition of all her work on the brassolines.

Type material (3 specimens)

Holotype

COFOMBIA: S, four white labels printed in black: // Narino, Barbacoas, Altaquer, Reserva Rio Nambi.

[1°15’N 78°07’W] Julio 24 de 1995 [24 Jul. 1995] G. Andrade-C. Feg. Altitud: 1425 m. // GAC: 7615//

ICN-MHN-F 14791 // ICN 011101 // (ICNO).

Paratypes

COFOMBIA: $ II Narino Ricaurte Fa Planada 01°05 , 06 ,, N 77°53 , 06 ,, W, 1800 m 8.ix.l994 [8 Sep.

1994], Julian Salazar, Feg. SM-4385 // IAvH 8238 // genit JFF 336 // (IAvH); $ // Narino Fa Planada

ViaHondon 1°15’N 78°15’W 1930 m Malaise 16 x. 2.xi. 2000 [16 Oct.-2 Nov. 2000], G. Oliva Feg. M.

139 // IAvH-E 67776 // genit JFF 337 // (IAvH).

Other material examined

Because so few specimens have been seen from the west of Colombia (2 c?c?, 1 $) and Ecuador, (1 (?, 2

5 5) it is possible that, when more specimens are examined, the Ecuadorian population may prove to be

a separate taxon. Therefore, the following specimens are not included in the type series:

ECUADOR: //Pichincha, Reserva Fas Galarias [sic], Nunbird fr. 0°0’34”S, 78°43 , 50”W, 27.iv.2011

[27 Apr. 2011]/T Kell/#008/FFMNH-MGCF-15-ll-26/DNA Voucher lep-04376 (FFMNH); $, //

Huigra, 1220 m, [2°18’S 78°59’W] Nov. 1926, Coxey// (ANSP); §, //Pichincha Prov., Santo Domingo

de los Colorados, alt. 200 m, Apr. 1982 // (FACM).

A male labelled “EKWADOR [sic], Prov. Zamora-Chinchipe, Valladolid, 17 05 1998 [17 May 1998],

leg. K. Jasinski” (MZUJ), exhibits the white bridge on VFW, and the broader inner white stripe on VHW

only extends as far as the larger of the lower ocelli, as in E. z. casagrande ssp. nov. Other features are

consistent with the above diagnosis. Keith Willmott (pers. comm.) commented that the locality may be

not reliable, and we suspect an inversion of labels with the E. z. greeneyi- like specimen, in the same

collection, which bears a label from the western El Oro Province.

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

In addition, we have seen a web image of a male at Tandayapa Lodge (1600 m, Nov. 28 th 2007, W.

Cook), and there is a picture of a male (presumably) in Silva (2011), taken at the Rio Guajalito Reserve

(1900 m, 12 Nov. 2009).

Description

Male

The FW length in the holotype is 58 mm, in the paratype 62 mm The values of the P index are estimated

at 5.5 mm in the holotype and 4.4 mm in the Ecuadorian specimen from Las Gralarias. These values

are close to those observed in E. z. greeneyi', possibly, the HW projection should be more pronounced

than in E. z. opimus. There is no R5-M1 orange mark on DFW; the orange DFW band is either moderate

(HT; Fig. 5C) or well developed and composed of five spots of variable size which are just contiguous,

being separated only by the veins; in that way, E. z. casagrande ssp. nov. may differ from E. z. greeneyi ,

where the spots are often separated. The iridescence is extremely faint. The androconial patch of the

holotype is less lemon-coloured than in E. z. opimus (Fig. 5C); however, the paratype male and the

single Ecuadorian male examined have a very pale cream androconial patch (Fig. 5D). The ground

colour of the verso is lighter, more yellowish than in E. z. chachapoya ssp. nov. The isolated white

spot in the VFW cell is quite large and semicircular. On the VFW, there is no white spot in the angle

formed by the cubital vein and Cu2; the complete white connection between the white stripes in space

Cu2-2A delimits half-a-circle (up against Cu2) with a chestnut-brown inner core and a dark brown outer

ring (Fig. 5C). The narrow outer white stripe on the VHW only just extends down to the smaller of the

lower ocelli; the broader inner white stripe only extends as far as the larger of the lower ocelli. The dark

brown lines in the VHW cell and curved line above the Cul-Cu2 ocellus are well developed; the distal

lines form two conjoined elongate circles (Fig. 5C). The costal ocellus on the VHW has a narrow, off-

white outer ring; the black ring is almost complete. In the holotype, on the VHW, the largest diameter

of Cul-Cu2 ocellus measures 10.1 mm (cf. Table 5); it is weakly larger in the paratype; thus, there is no

indication of a difference with either E. z. greeneyi or E. z. opimus. Genitalia are similar to those of E. z.

greeneyi and other subspecies from the Northern Andes, while they strongly differ from the genitalia of

E. z. chachapoya ssp. nov. (Fig. 13D-F).

Note. One striking feature of the male from Las Gralarias, Ecuador, is the circular orange spot with a

large dark brown core in space M3-Cul of the dorsal forewing band (Fig. 5D). This is unique amongst

all the E. zolvizora specimens examined. However, a smaller dark brown point is present in one male

from the Rio Aguacatal (eastern slope of the Colombian western cordillera), collected by Fassl (USNM),

a male from “Cali” in MNHN (PBB 1406), and two males of E. z. chachapoya ssp. nov. from the upper

Rio Nieva (Amazonas, Peru) in MNHN (PBGL 471 and 518).

Female

The paratype female FWL is 62 mm ; Ecuadorian females range from 60 to 67 mm The ventral surface

shows the same diagnostic features as the male. However, on VHW, the inner white stripe goes slightly

beyond the ocelli, but remains shorter than in all other subspecies (Fig. 7D). The markings in the VHW

cell of the Huigra female are rather faint, but this specimen is fairly old and worn (Fig. 7E). The dorsal

branches of the inward projections of the sterigma are similar to those of E. z. greeneyi , as they have

irregular outlines, but they are more distant from the internal side of the arch, as in E. z. chachapoya ssp.

nov. and E. z. opimus (Fig. 15J).

Flight period

Specimens have been caught or seen in April, July, September, October and November.

European Journal of Taxonomy 71: 1-66 (2014)

Distribution

Colombia. Narino Department of south-west Colombia between 1380 and 1930 m.

Ecuador. E. z. casagrande ssp. nov. presumably extends in suitable habitat throughout the western

slopes from the Colombian border to the extreme south of the country at heights of 1200 m to 1800 m;

the Santo Domingo specimen at 200 m is an anomaly and may be from a higher altitude along the road

to Quito.

Remarks

Specimens of E. zolvizora from south-western Colombia and western Ecuador are characterized by

a wide ‘bridge’ in space Cu2-2A on the VFW formed by a complete white connection between the

white stripes, delimiting half-a-circle (up against Cu2) with a chestnut-brown inner core and a dark

brown outer ring; in that way, E. z. casagrande ssp. nov. differs from its near neighbours E. z. opimus

in the Cauca Valley, Colombia, and E. z. greeneyi in the Oriente of Ecuador, as well as from all other

subspecies, except E. z. chachapoya ssp. nov., where we know 3 (of 35) specimens having a white

bridge (MNHN: PBB 2310, PBGL 521 and PBGL 705). We consider that the presence of a wide, white

bridge is the principal diagnostic character of E. z. casagrande ssp. nov., but suggesting a closer affinity

with E. z. chachapoya ssp. nov. than E. z. greeneyi. However, the male genitalia are similar to those of

E. z. greeneyi.

There are specimens of E. zolvizora collected in the 19 th and early 20 th century labelled from the western

Ecuadorian village of ‘Balzapamba’. Like many specimens supposedly from this locality, they are from

eastern Ecuador and are in fact E. z. greeneyi.

Eryphanis zolvizora opimus Staudinger, 1887

Figs 5E-F, 7F, 13G-I, 15K

Eryphanis opimus Staudinger, 1887: 217.

Eryphanis zolvizora opimus - Stichel 1904: 33;1909: 157, fig. 36; 1932: 75. —Fruhstorfer 1912: 312.

— D’Abrera 1987: 406-407. — Casagrande 2004: 203.

Eryphanis opimus (Staudinger, 1887) [sic] - Penz 2008: 15-15, figs II, 41, 51 (status revised and re¬

description).

Diagnosis

HW projection only poorly developed; a faint orange spot between the R5-M1 veins is present in some

specimens; the DFW band generally with five well developed, almost contiguous, crescent-shaped spots

(Fig. 5E) (however, there are some specimens with small, sometimes blurred, disjunct spots; cf. Fig.

5F); on the VFW, in space Cu2-2A, the inner margin of the distal white stripe forms a projection that

does not reach the black border of the proximal white stripe; in the VHW cell the dark brown lines and

curved line above the Cul-Cu2 ocellus are well developed (Fig. 5E-F). The valva axis has no swelling;

the dorsal ridge has a row of more than 6 spines, dimimishing towards the extremity (Fig. 13G).

Type material

Lectotype, here designated

COLOMBIA: S, dirty white, rectangular label: //Manizales, Cauca [handwritten in dark brown ink,

followed by an indecipherable abbreviation: S?hl.]// (ZMHB).

Paralectotype, here designated

COLOMBIA: S, dirty white, rectangular label: //Manizales, Cauca [handwritten in dark brown ink,

followed by an indecipherable abbreviation: S?hl.]// (ZMHB).

Type locality

Manizales (Colombia).

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

Description

Male

The FWL varies from 58 to 66 mm. Average P = 3.9 (n = 6; cf. Table 3); it is possible that there is no

significant difference with other populations from the Northern Andes, except E. z. greeneyi. In some

specimens, there is a faint orange spot between the R5-M1 veins, making a very weak projection of the

proximal branch of the submarginal band (Fig. 5E); this feature has not been observed in other specimens

from the Northern Andes. The DFW band varies from well developed, with five barely contiguous

spots, to five small disjunct spots in some specimens, that we consider to be individual variations, like

similar specimens from northern Peru, Ecuador and Venezuela. The violet iridescence is a little more

pronounced than that of E. z. greeneyi , but less than in E. z. reyi ssp. nov. and E. z. isabelae ssp. nov.

(cf. Fig. 6). The androconial patch is a bright creamy, pale citrus yellow. The background colour of

the ventral surface has a lighter yellowish aspect than in E. z. chachapoya ssp. nov. (Fig. 5E). On the

VFW, the isolated white spot in the cell is medium to large, with a variable, roughly oval, triangular or

crescent-like shape; there is sometimes a small white spot in the angle formed by the cubital vein and

Cu2 (Fig. 5E-F); in space Cu2-2A, the inner margin of the distal white stripe forms a projection that

does not reach the black border of the proximal white stripe, and is often less pronounced than in E. z.

greeneyi (Fig. 5E-F). On the VHW, the white stripes extend far beyond the Cul-Cu2 ocellus, notably the

distal one, which reaches vein 2A; in the cell, the dark brown lines and curved line above the Cul-Cu2

ocellus are well developed; the costal VHW ocellus has an incomplete pale circle, sometimes partially

covered with white scales; the black ring is complete and sometimes thicker than in any other subspecies

(Fig. 5E-F). The VHW Cul-Cu2 ocellus (® average value: 10.7 mm; n = 6; cf. Table 5) is larger, on the

average, than that of E. z. greeneyi and E. z. reyi ssp. nov., and much larger than in E. z. isabelae ssp.

nov. Genitalia exhibit weak individual variations (Fig. 13G-I). The valva is regular, without a swelling;

the ridge bears a row of more than 6 spines, the size of which varies more or less regularly; the gnathos

is larger than those in E. z. zolvizora , and sometimes smaller than in E. z. chachapoya ssp. nov.

Female

Only 6 females are known (labelled from ‘Calima Valley, Cauca River’; Calima, 45km W of Buga;

Rio Agua; ‘Rio Aguatal’; Colombia Ost; and ‘Colombie’). The FWL varies from 66 to 70 mm. The

diagnostic characters correspond to those of the males (Fig. 7F). On the dorsal surface, the FW band is

composed of five narrow crescentic orange spots that just touch each other (the band is slightly narrower

than that of most E. z. greeneyi females). There is the faintest hint of a sixth orange spot between the

R5-M1 nerves (Fig. 7F). There is a strong purplish sheen in the middle of both the FW and HW. As in

the male, on the VFW, in space Cu2-2A, the inner margin of the distal white stripe forms a projection

that does not reach the black border of the proximal white stripe (an exception is the female from Rio

Aguacatal), unlike in E. z. reyi ssp. nov. and E. z. isabelae ssp. nov. The dorsal branches of the inward

projections of the sterigma have a regular outline, in this way being different from E. z. greeneyi and

more similar to E. z. chachapoya ssp. nov., but their extremities are notably less distant from the internal

side of the arch than in females from northern Peru (Fig. 15K); nevertheless, the differences with one

of the E. z. chachapoya ssp. nov. females (cf. Fig. 15H) are weak. Female genitalia of E. z. opimus do

differ markedly from those of other Northern Andean females, except one female of E. z. reyi ssp. nov.

which has thicker inward projections and shorter dorsal branches (cf. Fig. 15L).

Flight periods

The only ten dates that we have are from May to November.

European Journal of Taxonomy 71: 1-66 (2014)

Distribution

Colombia. The only specimens with reliable data are from the Cauca Valley, and from Rio Aguacatal

(this name has often been misspelt on data labels as Aquatil, Aquatal, Aguatal, Aguaca, and similar), on

eastern slopes of the western cordillera. It is unfortunate that, of the many specimens of E. z. opimus that

we have seen, only five reliable localities can be confirmed in the Cauca Valley: Calima Valley, on the

west side of the valley (i.e. east side of the western cordillera); and Armenia, Pereira, Manizales, and

Mesopotamia, on the east side of the valley (i.e. west side of central cordillera). Fassl (1915a: 10) shows

opimus on both sides of the western cordillera at heights between 2000 and 2500 m, but we have not

seen any E. zolvizora specimen from these altitudes on the west side of the western cordillera, although

specimens labelled ‘Cali, 1000 nT in the MNHN and MPM were probably collected along the road from

Cali to Buenaventura on the Pacific slope, as well as the ‘Buenaventura-Queremal’ male in the IAvH.

Reliable altitudes range from 900 to 2500 m.

Remarks

The taxon E. opimus was described by Staudinger (1887), on the basis of three syntypes (males), two

from Manizales (Colombia) and one from Chanchamayo (Peru). After Stichel (1904), it was considered

as a subspecies of E. zolvizora , until Penz (2008) revised its status. The name opimus has always

been used for Colombian specimens (Stichel 1904, 1909, 1932; Fruhstorfer 1912; Fassl 1914, 1915a,

1915b; D’Abrera 1987). Staudinger noted differences between the Peruvian syntypic specimen and the

Colombian ones. It should be noted that none of the syntypes match the figured specimen (fig. 36) in

Stichel (1909). To stabilize the situation, we have designated as lectotype the undamaged Manizales

syntype. The other Manizales specimen is therefore a paralectotype. As indicated previously, it is likely

that the syntype from Chanchamayo belongs to E. z. chachapoya ssp. nov.

Eryphanis zolvizora reyi Bristow, Neild, De Sousa & Huertas ssp. nov.

Figs 6A-B, 8A, 14A-C, 15L

Diagnosis

HW projection poorly developed; no orange mark between veins R5-M1 (Fig. 6A-B). The white spot

near the middle of the VFW cell is relatively equidimensional, isolated, medium to large (Fig. 6A-B).

On the VFW, the white vertical stripes in cell Cu2-2A are generally connected by a thin black horizontal

line, thus distinguishing E. z. reyi ssp. nov. from E. z. opimus ; the black ring of the anterior VHW ocellus

is often broken at the costal margin (Fig. 6A). The valva is regular, without swelling; the ridge bears a

row of more than 6 spines, the size of which varies more or less regularly (Fig. 14A).

Etymology

We name this subspecies to honour the memory of the Venezuelan Tepidopterist Rafael Fernando Rey

Cardenas (1958-2010). He was an adept field collector, with a special interest in butterflies of the family

Pieridae. He placed great emphasis on collecting in the State of Tachira, around San Cristobal, as well

as in El Tama National Park, where he discovered a remarkable new montane species of Catasticta (C.

revancha - see Rey & Pyrcz 1996). Fernando Rey also generously contributed to AN’s research for the

book series The Butterflies of Venezuela (Neild 1996; 2008). An obituary by one of the authors (AN) and

Tomasz Pyrcz (MZUJ) is currently in preparation.

Type material (30 specimens)

Holotype

VENEZUELA: <$, two white labels printed in black: // VENEZUELA, Barinas, La Chimenea, 1500m,

[8°49N 70°31 ’W] 1-6-VI-1973 [1-6 Jun. 1973] //J. Salcedo col.// (MIZA).

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Paratypes

VENEZUELA: 3, H Anzoategui, Qda Guazo, Venezuela, Lara, 1440 m, [9°36’N 69°53’W], 13-16 VI

72, [13-16 Jun. 1972] J. Salcedo & F. Zambrano// (MIZA); §, // Anzoategui, Qda Guazo, Venezuela,

Lara, 1440 m, [9°36’N 69°53’W], 13-16 VI 72, [13-16 Jun. 1972] J. Salcedo & F. Zambrano// (MIZA);

3, // Barinas, La Mina above San Isidro, 1450 m, [8°50’N 70°34’W], III-1988 [Mar. 1988], leg Romero

// (AN); // Barinas, San Isidro, Las Minas, 1475 m, [8°50’N 70°34’W], 07-SEP-2002 [7 Sep. 2002],

Col. Juan C. de Sousa C. // (JCSC); 3, II Barinas, San Isidro, Las Minas, 1500 m, [8°50’N 70°34’W], 10

Oct.2010. Col. Mauro Costa // (JCSC); $, // Barinas, Qda. La Soledad, Via Barinitas - Sto. Domingo,

[8°49’N 70°3LW], 5 Apr. 1985. Col. Mauro Costa//(JCSC); c?, //Barinas, La Soledad, [8°49 , N70°31 , W],

5-IV-1985 [5 Apr. 1985] // (MCC); 3, II Barinas, San Isidro, Mina, 1500 m. [8°50’N 70°34’W], 04-X-

2009 [4 Oct. 2009] // (MCC); 3, II Barinas, San Isidro, Mina, 1500 m. [8°50’N 70°34’W], 04-X-2009

[4 Oct. 2009] // (MCC); 3, II Barinas, San Isidro, Mina, 1500 m. [8°50’N 70°34 , W], 09-XI-2009 [9

Nov. 2009] // (MCC); II Barinas, San Isidro, Mina, 1500 m. [8°50’N 70°34’W], 12-XII-2009 [12

Dec. 2009] // (MCC); ?, // Barinas, San Isidro, Mina, 1500 m. [8 o 50’N 70°34’W], 05-1-2011 [5 Jan.

2011] (MCC); ?, //VENEZUELA, Barinas, La Chimenea, 1500m, [8°49N 70°3UW]l-6-VI-1973 [1-6

Jun. 1973] // J. Salcedo col.// (MIZA); 3,1 I Barinas, San Isidro, Vallee du Rio Santo Domingo, 1500 m,

[8°50’N 70°34’W], 9-XI-2010 [9 Nov. 2010], M. Costa & S. Attal // (MNHN, PBB 2324); 3, //Barinas,

San Isidro, Vallee du Rio Santo Domingo, 1500 m, [8°50’N 70°34’W], 9-XI-2010 [9 Nov. 2010], M.

Costa & S. Attal // (MNHN, PBB 2325); 3, H Barinas, Barinitas, [La Mina de San Isidro, 1450 m,

[8°50’N 70°34’W], X-87 [Oct. 1987], C.f.R // 115-JCSC // (R); $, // Barinas, Barinitas, [La Mina de

San Isidro, 1450 m, [8°50’N 70°34 , W], X-87 [Oct. 1987], C.f.R // (R); 3, II Barinas, Barinitas, [La

Minade San Isidro, 1450 m, [8°50’N 70°34’W], X-87 [Oct. 1987], C.f.R// (R); §, //Barinas, Barinitas,

[La Mina de San Isidro, 1450 m, [8°50’N 70°34’W], X-87 [Oct. 1987], C.f.R // (R); ?, // Barinas,

Barinitas, [La Mina de San Isidro, 1450 m, [8 o 50’N 70°34’W], X-88 [Oct. 1988], C.f.R // 116-JCSC

// (R); 5, //Barinas, Barinitas, [LaMina de San Isidro, 1450 m, [8°50’N 70°34’W], X-88 [Oct. 1988],

C.f.R // (R); 3,1 I Barinas, Barinitas, [La Mina de San Isidro, 1400 m, [8°50’N 70°34’W], XI-95 [Nov.

1995], C.f.R // (R); $, // Barinas, Barinitas, [La Mina de San Isidro, 1400 m, [8°50’N 70°34’W], XI-95

[Nov. 1995], C.f.R // (R); $, // Merida, Paramo El Molino, SE of Sta Cruz de Mora, 1600 m, [8°16’N

71°34’W] 03.1.2001 [3 Jan. 2001], Leg. M. Costa // (AN); II Tachira, Sierra de El Tama, Via Delicia,

1900 m, [7°36’N 72°26’W], III-87 [Mar. 1987] C.f.R// (R); 3, //Tachira, Sierra de El Tama, Via Delicia

(Pabellon), 1700 m, [7°36’N 72°26’W], IV-1990 [Apr. 1990] C.f.R //(R).

COLOMBIA: 3, Santander, El Cerro-La Luchata, 29/06/2006, 06°37 , 75 ,, N 73°18 , 88 ,, W, 1950-2050

m, posada on vegetacion bosque primero, B. Huertas, C. Rios y J. Arias leg., Proyecto YARE (IAvH); 3,

as preceding specimen, but 30/06/2006; 3, Santander, Charala, 2000 m, juillet 2008 (MNHN).

Description

Male

The FWL varies from 58 to 62 mm, excluding one male from Colombia with an unusually small FWL

of 52 mm (which has been excluded from our statistical measurements). The holotype FWL is 59.5 mm.

The hindwing projection (average P = 3.6; n = 15; cf. Table 3) is rather similar to that of E. z. zolvizora

and E. z. opimus. There is no orange mark between veins R5-M1 (Fig. 6A-B). The holotype and some

of the paratypes have five submarginal orange spots which are contiguous at the veins (Fig. 6A-B).

However, the form and size of these spots are variable and in some specimens they are indistinct and

disjunct. The violet iridescence on the dorsal surface is stronger than in E. z. opimus , but it is usually a

little weaker than in E. z. isabelae ssp. nov., especially on the DFW (Fig. 6A-B). The bright creamy, pale

citrus androconial patch is shorter, on average, than in E. z. opimus and E. z. isabelae ssp. nov. (cf Table

4); it should be noted that, in one specimen from the Sierra de El Tama, the colour of the patch is a pale

greyish-brown, as in E. z. isabelae ssp. nov. Ground colour of the ventral surface as in other subspecies

of the Northern Andes, except some darker specimens of E. z. greeneyi (Fig. 6A-B). The shape of the

European Journal of Taxonomy 71: 1-66 (2014)

isolated white spot near the middle of the VFW cell is irregular but relatively equidimensional, and

its size varies from medium to large; in this way, E. z. reyi ssp. nov. (Fig. 6A-B) is clearly different

from E. z. isabelae ssp. nov. (Fig. 6C-D) and other subspecies. On the VFW, the white spot at the basal

angle of cell Cul-Cu2 is generally very small (Fig. 6A), or absent (Fig. 6B). On the VFW, the white

vertical stripes in cell Cu2-2A are generally connected by a thin black horizontal line (Fig. 6A-B),

which immediately distinguishes E. z. reyi ssp. nov. from E. z. opiums, in which this bridge is absent.

On the VHW, the white stripes extend far posterior to the Cul-Cu2 ocellus. On the VHW, the black

markings in the cell and the curved line anterior to the Cul-Cu2 ocellus are fainter than in E. z. opimus

(Fig. 6A-B). In most specimens, the anterior HWV ocellus is encircled by an outer pale/off white circle,

contrasting with the background more than in E. z. opimus ; in addition, the black ring is often broken at

the costal margin (Fig. 6A), a difference with E. z. opimus and E. z. isabelae ssp. nov. On average, the

VHW Cul-Cu2 ocellus (O average value: 9.5 mm; n = 15; cf. Table 5) is probably a little smaller than

in E. z. opimus. The valva is similar to those of other subspecies of the Northern Andes (Fig. 14A-C);

the gnathos is clearly smaller than in E. z. chachapoya ssp. nov. and generally slightly smaller than in

E. z. opimus.

Female

Female FWT length varies from 63 to 66.5 mm (mean = 64.8 mm ; n= 8). The principal diagnostic

characters are the same as in males (Fig. 8A). On the VFW, the black line at the base of VFW cell M3-

Cul is well-defined, while it is absent or nearly so in females of E. z. isabelae ssp. nov. The forewing

band is composed of five well developed, crescentic, submarginal orange spots which are contiguous

at the veins. There is a violet-blue dorsal iridescence on both the FW and HW, almost similar to that of

E. z. opimus , but usually not so extensive or as blue as that of E. z. isabelae ssp. nov. The genitalia of

the two females we have dissected exhibit some differences. In one specimen (El Molino, Merida), the

sterigma is close to that of the females of E. z. opimus and E. z. casagrande ssp. nov., as well those of E.

z. isabelae ssp. nov. females: the inward projections are short and not very wide, and the dorsal branches

are large. In the other specimen (Barinitas, Barinas; Fig. 15L), the inward projections are wider and

larger; consequently the dorsal branches are shorter; most likely this is a matter of individual variation.

Flight periods

Dated specimens are for March to June, and September to November.

Distribution

Colombia. Santander Province of northern Colombia at almost 2000 m.

Venezuela. Cordillera de Merida in north-western Venezuela, from approximately 1400 to 1900 m on

either side of the Cordillera.

Remarks

Specimens of this taxon were discovered during an expedition by the MIZA in 1972 to the Cordillera de

Merida, in north-western Venezuela, where a series of males and females were collected. More recently,

E. z. reyi ssp. nov. was observed at sunny intervals resting on herbaceous vegetation, by one of the

authors (BH) and colleagues in the pristine forest (surrounded by regenerated forest) at Ta Tuchata, in

the Serrania de los Yariguies, Santander, during field expeditions of the YARE Project, and 2 specimens

were collected (see details in Huertas & Donegan 2006). La Luchata is the type locality of another

recently described butterfly (Huertas et al. 2009).

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

Eryphcinis zolvizora isabelae Neild & De Sousa ssp. nov.

Figs 6C-D, 8B, 14D-F, 15M-0

Diagnosis

HW projection moderately developed. No orange mark between veins R5-M1. The androconial patch

is on average longer (mean: 9 mm) than in all other subspecies and is pale greyish-brown (Fig. 6C-D).

The violet iridescence on the FW is generally slightly stronger and distally more extensive than in E.

z. reyi ssp. nov. The white patch in the VFW cell next to the base of Cul-Cu2 is larger by 50%-100%

than other subspecies (Fig. 6C-D). On the VFW, the white Cu2-Cul spot in the basalmost angle formed

by the cubital vein and Cu2 is relatively large and always present (Fig. 6C-D). On the VFW, the black

line at the base of cell M3-Cul is usually diffuse or poorly-defined in males (Fig. 6C-D), and diffuse or

nearly absent in females (Fig. 8B), a difference with all other subspecies. The valva is regular, without

swelling; the ridge bears a row of more than 6 spines, the size of which varies more or less regularly.

Etymology

We name this subspecies in honour of the memory of Isabel Montesino de Romero (1931-2008). The

Romero family collected the vast majority of the type series of both the Venezuelan subspecies described

herein. The late Isabel, and her children Francisco (“Paco”) and Maria de la Concepcion (“Conchita”),

very kindly helped us in many ways during our protracted research for this paper, and we use this

opportunity to thank them profusely for their great kindness and generosity.

Type material (12 specimens)

Holotype

VENEZUELA: S, white label handwritten in black ink: // Choroni, 1450. [Via Choroni 1450 m]

[10°2UN 67°34’W] 11-78 [Feb. 1978]. C. f. R. s. [collected by familia Romero] // (R, to be donated to

MIZA).

Paratypes

VENEZUELA: II Choroni (arbol). [Via Choroni, Aragua] [10°2UN 67°34’W], 6-70. [Jun. 1970] C.

f. R. // 111-JCSC // (R); & // Choroni. 1450. [Via Choroni, Aragua, 1450 m] [10°2UN 67°34’W], 9-71.

[Sep. 1971] C. flia. Romero // 117-JCSC // (MIZA); & // Choroni 15. [Via Choroni, Aragua] [10 o 22’N

67°35’W], 10-72. [Oct. 1972] C. f. R. // 112-JCSC // (R); & // Choroni, 1250. [Via Choroni, Aragua,

1250m] [10°22’N67°35’W], 9-76. [Sep. 1976] C. f. R. //(R); $, //R. Grande. [Rancho Grande, Aragua,

1150 m] [10°20’N 67°40’W], 8-65. [Aug. 1965] C. Eduardo // 118-JCSC // (MIZA); $, // [Choroni 21.

[Via Choroni, Aragua] [10°2UN 67°34’W], 8-70. [Aug. 1970] C. f. R. 1550 m. // 113-JCSC // (R); ?, //

Choroni. 1.400 s. [Via Choroni, Aragua, 1400 m] [10°22’N 67°35’W], V-79.[May 1979] C. f. R. // (R);

?, // Choroni 28. [Via Choroni, Aragua] [10°22’N 67°35’W], 28-9-76. [28 Sep. 1976] C. f. R. 1150 m.

Cara Norte // 114-JCSC // (R); §, //La Llanada. 1.650 mts. [Parque Nacional el Avila, Distrito Federal]

[10°33’N 66°56’W], 29 Oct. 1989 // (MNHN, PBB 2322); $, // Colonia Tovar. 2.100 mts. [10°24’N

67°19’W], IV-93. [Apr. 1993] C. F. R. Edo. Aragua // (R); $, // Col. Tovar, 2.100 m, via Capachal.

[Aragua] [10°24’N 67°19’W], 02-04-2010 [2 Apr. 2010] //(MCC).

Other material examined

Included in the measurements but excluded from the type series: c?, // Mexico // Druce Coll, ex Kaden

Coll. // Eryphanis zolvizora Hew. Godman-Salvin Coll. 1919.-6. // BMNH (E) #808093 // (BMNH).

European Journal of Taxonomy 71: 1-66 (2014)

Description

Male

The FWL varies from 57 to 66 mm (mean: 63.3 mm ; n = 6). The holotype FWL is 60 mm The hindwing

projection is developed to an average degree (P = 4.1; n = 5; cf. Table 3), similar to specimens from

Colombia. There is no orange mark between veins R5-M1 (Fig. 6C-D). The holotype (Fig. 6C) and some

of the paratypes have five submarginal orange spots which meet at the veins (but these are separated

from each other by the vein itself, which is dark brown); however, although the form and size of these

spots are variable, they are well-developed in all but one of the known specimens; therefore, it is likely

that this character falls into the variation observed in populations from the Northern Andes and northern

Peru. The violet iridescence on the DFW is generally slightly stronger and distally more extensive

than in E. z. reyi ssp. nov. and E. z. opimus. The length of the DHW androconial patch is on average

longer than in all other subspecies, with a mean of 9.0 mm (n = 6; cf. Table 4), and the patch is pale

greyish-brown (Fig. 6C-D), while it is a bright creamy white in all other subspecies, except in some rare

individuals. Ground colour of the ventral surface as in other subspecies of the Northern Andes (except

some darker specimens in E. z. greeneyi). The white patch on the FWV cell next to the base of Cul-Cu2

is roughly subtriangular or in the form of an inverted heart-shape; it is distinctly larger (50%-100%)

than in any other subspecies (Fig. 6C-D). On the VFW, the white Cu2-Cul spot in the basalmost angle

formed by the cubital vein and Cu2 is relatively large and always present (Fig. 6C-D). On the VFW, the

black line at the base of cell M3-Cul is usually diffuse or poorly defined in males (it is narrow, but well

defined, in all other subspecies). On the VFW, the white vertical stripes in cell Cu2-2A are generally

connected by a thin black horizontal line (Fig. 6C-D). On the VHW, the black designs in the cell and

the curved line anterior to the Cul-Cu2 ocellus are fainter than in E. z. opimus, but almost similar to

those of E. z. reyi ssp. nov. In most specimens, the anterior VHW ocellus is encircled by an outer pale/

off-white circle, contrasting with the background more than in E. z. opimus', the black ring is complete

(Fig. 6C-D), being dissimilar to E. z. reyi ssp. nov. in which it is usually broken at the costal margin. On

average, the VHW Cul-Cu2 ocellus (O average value: 8.7 mm; n = 5; cf. Table 5) is probably similar

to that of E. z. reyi ssp. nov. and perhaps of E. z. greeneyi , but smaller than in all other subspecies. The

valva is regular, without swelling; the ridge bears a row of more than 6 spines, the size of which varies

more or less regularly (Fig. 14D-F); the gnathos is generally slightly smaller than in E. z. opimus ; there

is no noticeable difference with E. z. reyi ssp. nov.

Female

Female FWL length varies from 63 mm to 68.5 mm (n = 6). The dorsal and ventral surfaces show the

same diagnostic features as in the male. The violet-blue dorsal iridescence on both the FW and HW is

usually a more extensive, lighter and bluer tint than the one observed on E. z. reyi ssp. nov. (Fig. 8B).

The inward projections of the sterigma are narrower than in E. z. opimus and E. z. reyi ssp. nov. and

the bases of the dorsal branches are widely separated from the internal side of the arch (Fig. 15M-0);

nevertheless, the extremities of these branches are close to it and clearly cross in 3 of the 4 dissected

specimens; in this way, E. z. isabelae ssp. nov. possibly differs from other subspecies.

Flight periods

Specimens have been collected in February, April, May, and June, but the majority were taken in August,

September and October. With the exception of February, which is usually (but not always!) dry, these

months fall within the rainy season.

Distribution

Venezuela. This taxon appears to be endemic to the Cordillera de la Costa in northern Venezuela, where

it has been recorded from approximately 1150 m to at least 2100 m.

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

General Discussion

Together with ornithologists, lepidopterists contributed to the ‘Trinomial Revolution’ in Zoology,

‘subspecies’ being acknowledged as an official nomenclatural rank (Mallet 2007). However, even in

birds, which are far better known than any other major taxon, species concepts and limits are the subject of

continual debate (Cracraft 2000; Alstrom et al. 2008; James 2010). There is no consensus in ornithology

on how to treat subspecies, with some authorities preferring phylogenetic concepts and the elimination of

many subspecies altogether (e.g. Zink 2004) and others noting the importance of this status (e.g. Patten

& Unfit 2002; Remsen 2010). Subspecies have been defined as “a breeding population that occupies

a distinct segment of the geographic range of its species and that is measurably distinct in phenotype,

genotype, or both” (James 2010). The most frequent modem usage for subspecies in ornithology is for

diagnosable allopatric populations with small non-clinal differences from other allopatric populations,

but where the differences are not considered equivalent to those between sympatric species (Remsen

2010). This concept generally corresponds to the practice of delimiting subspecies among lepidopterists.

Despite this, many entities have been described as subspecies for geographical forms recognizable only

to their author (Descimon & Mallet 2009), thus cluttering checklists and contributing to taxonomic

inflation. Descimon & Mallet (2009), arguing for a revival of the subspecies rank in a modern way,

emphasized ‘cohabitation’ as the ‘touchstone’ of all criteria for species separation: when overlapping

distinct populations produce unimodal frequency distributions of morphological traits and genotypes,

subspecies should be designated, while separate species must be recognized if frequency distributions

are bimodal. However, the cohabitation criterion is efficient only if overlapping populations can be

accurately sampled.

The Eryphanis zolvizora group offers a case study of these issues. Our study represents the most

comprehensive survey of this group ever accomplished. Specimens have been collected over a period

of more than 150 years, in a relatively small number of localities scattered along the Andes. Even in

northern Peru, where a special effort has been made recently, only 20 males and 6 females were obtained

from 2005 to 2012: E. zolvizora butterflies are obviously unco mm on in the field. Despite this limited

information, we have decided to recognize eight taxa distributed from Bolivia to Venezuela (Fig. 16).

Having studied a large number of male genitalia, we observed individual variation between right and left

valvae, and individual variation of valvae and gnathi. However, these limited variations do not blur the

existence of distinctive genitalia structures corresponding to geographical groups of populations. Within

the genus Eryphanis , male genitalia present strong differences between several previously described

species, as illustrated by Penz (2008). Therefore, considering the differences in male genitalia between

E. zolvizora zolzivora and E. z. inca ssp. nov.; E. z. inca ssp. nov. and E. z. chachapoya ssp. nov.; and

E. z. chachapoya ssp. nov. and E. z. greeneyi , it would have been logical to acknowledge that these taxa

correspond to different species. However, we considered here the eight taxa of the Eryphanis zolvizora

group as subspecies, for two reasons. First, limits between taxa indicated by male genitalia characters

and by wing characters do not always coincide. Second, in the present state of knowledge, there are

no known contact zones between any E. zolvizora taxa; therefore, there is no information relevant to

assessing the cohabitation criterion.

Thus, we have preferred to adopt a lumper’s attitude, assuming that clear differences in male genitalia

do not necessarily imply separate species. It should be emphasized that such differences exist within

taxa from the Central Andes (Bolivia and Peru), while there is no significant difference between taxa

from the Northern Andes (Ecuador, Colombia, Venezuela). Therefore, if male genitalia differences

support species separation. Central Andean taxa are better candidates for species rank than Northern

Andean taxa. For example, data suggest that E. z. inca ssp. nov. and E. z. chachapoya ssp. nov. perhaps

overlap in central Peru. If their cohabitation were demonstrated, it is possible that the distribution of

European Journal of Taxonomy 71: 1-66 (2014)

Fig. 16. Geographical distribution of Eryphanis zolvizora subspecies. Unreliable localities are excluded.

Male genitalia of specimens illustrating the different subspecies are represented.

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

their morphological characters would be bimodal, supporting the existence of two species (Descimon &

Mallet 2009). Similarly, geographical information would be necessary to check the spatial relationships

between E. z. zolvizora and E. z. inca ssp. nov.: if cohabitation with bimodal distributions of characters

were revealed, two species should be recognized. Moreover, the precise geographical relationships

between E. z. chachapoya ssp. nov. and the Ecuadorian subspecies E. z. greeneyi and E. z. casagrande

ssp. nov. must be studied from northern Peru to the western and eastern Andean slopes of southern

Ecuador. As male genitalia of E. z. chachapoya ssp. nov. differ from those of the Ecuadorian subspecies,

the cohabitation of populations with a bimodal distribution of characters is a plausible hypothesis.

Between subspecies from the Northern Andes, differences concern several subtle wing characters that

often have only a ‘statistical value’. However, we decided to create three new taxa to emphasize plausible

evolutionary divergences between populations associated with different geographical areas. In so doing,

we implicitly apply the phylogenetic concept. Should someone decide to raise these taxa to species rank,

this decision will contribute to taxonomic inflation, as defined by Isaac et al. (2004) “for cases in which

many existing subspecies are raised to species level”.

Taxonomic inflation has important influences on conservation (Agapow et al. 2004; Isaac et al. 2004;

Mallet 2007). Species are the ‘currency’ for conservation policies, which generally consider species

richness, number of endemic species and number of endangered species in order to choose and manage

conservation areas. When a polytypic species is split into several species, these species evidently have

more restricted geographic ranges and decreased abundance. Thus, the number of endemic and fragile

species increases. However, the primary source of taxonomic inflation is the creation of new taxa

belonging to the ‘species-group’ in the sense of the International Code of Zoological Nomenclature

(art. 45.1), whatever may be the level initially chosen - either species or subspecies. As emphasized

by Mallet (2007), conservationists are increasingly aware of the conservation of biodiversity at all

hierarchical levels, “including well-marked subspecies”. Thus, from a pragmatic viewpoint, any taxon

described as a ‘species-group entity’ should have the same potential interest. As a consequence, the

taxonomist’s first responsibility is to decide whether specimens from a particular area can be named:

in so doing, he creates a ‘potential conservation entity’. As underlined by Agapow et al. (2004), to use

measures other than species counts, for example subspecies counts, “forces us to ask what it is that we

are trying to preserve”.

The Eryphanis zolvizora group is endemic to the Tropical Andes, the world’s richest Biodiversity

Hotspot (Mittermeier et al. 2004). This hotspot encompasses a very fragmented geographical system

of cordilleras and valleys, resulting from a complex geodynamic history, which certainly influenced

the diversification of butterflies (Blandin & Purser 2013). Cloud forests form an ecological belt which

could develop only when mountain ranges attained sufficient altitudes. For example, such altitudes

were probably attained far longer than 10 million years ago in Bolivia (Barnes & Ehlers 2009), while

the Colombian eastern cordillera only attained 1500-2000 m a.s.l. by between 5 and 2 million years

ago (Gregory-Wodzicki 2000). Therefore, it is possible that cloud forest butterfly lineages, from one

range to another, have different ages. In this context, it seems logical that the level of morphological

divergence between E. zolvizora populations may differ, depending on whether they inhabit old or recent

ranges. Thus, by recognizing within the Eryphanis zolvizora group subspecies with different levels of

morphological divergence, we underline the evolutionary heterogeneity of the Tropical Andes Hotspot,

and the interest to preserve cloud forests not only at the core of each entity’s area, but also in transition

areas where evolutionary hypotheses could be checked.

Acknowledgements

Part of this work was realized in the framework of the project “Geographie de la Biodiversite des

Lepidopteres dans le nord du Perou”, which received financial support from the Museum national

European Journal of Taxonomy 71: 1-66 (2014)

d’Histoire naturelle (Paris) for several years from 2005 (Programme Pluri-Formations “Etat et structure

phylogenetique de la biodiversite actuelle et fossile”). Field work was performed with permission of

Peruvian authorities. We warmly thank Dr. G. Lamas (MJP, Lima), for access to collections, loan of

specimens, information and comments, and G. Lachaume (Paris) for collaboration in the field and for

many stimulating discussions.

_ r

The YARE expeditions in Colombia were supported by: the Royal Geographical Society, BP Conservation

Programme (BirdLife International, Conservation International, Flora & Fauna International, Wildlife

Conservation Society), Fondo para Accion Ambiental, Fundacion Omacha, Conservation International

Colombia (Becas Iniciativa de Especies Amenazadas - Jorge Ignacio ‘El Mono’ Hernandez-Camacho),

Rio Tinto pic, Duke of Edinburgh, the Percy Sladen Memorial Fund (Linnean Society), Fundacion

ProAves, Game Conservancy Trust, World Pheasant Association, Tropical Andean Butterflies Diversity

Project, Carter Ecological, Universidad Industrial de Santander, Universidad de Caldas, Universidad de

Tolima and Gobernacion de Santander. Special thanks to J.J Arias and C. Rios.

Ackowledgements are due to the Museum national d’Histoire naturelle (Paris) and the Natural History

Museum (London) for letting us facilities there such as collections, labs and librairies, and to the

Biodiversity Library for facilitating access to key research resources.

The authors acknowledge the invaluable help from each of the following collection owners and curators,

and colleagues, for providing access to the collections and specimens, loan of specimens, authorization

for taking photographs, and for assisting with information, photographs or drawings, or help in the field:

G. Andrade (Colombia, Bogota, ICNUN). - S. Attal (France, Paris). - H. Azurduy, L. Gonzales and T.

Vidaurre (Bolivia, Natura Fundation, Santa Cruz). - M.M. Casagrande (Brasil, UFPR, Curitiba). - J.

Clavijo and J. DeMarmels (Venezuela, Maracay, MIZA). -M. and C. Costa (Venezuela, Caracas). -R. de

Vos (The Netherlands, Amsterdam, ZMA). - P. Grootaert (Belgium, Brussels, IRSN). - J. Hogan (United

Kingdom, Oxford, UMO). - J.-F. Le Crom (Bogota). - J. Ledezma and A. Valdivia (Bolivia, Santa Cruz

de la Sierra, MHNNKM). - C. Medina and A. Parrales (Colombia, Bogota, IAvH). - W. Mey (Germany,

Berlin, ZMHB). - H. Morgenroth (United Kingdom, Exeter, RAMM). - C. Penz (USA, University of

New Orleans). - Padre F. Pinas (Ecuador, Quito). - T. Pyrcz (Poland, Krakow, MZUJ). - R.G. Robbins

and B. Harris (USA, Washington D.C., USNM). -F. and C. Romero (Venezuela, Maracay). - J. Salazar

(Colombia, Manizales). - Jaime Sarmiento (Bolivia, La Paz, CBF). - X. Silva (Ecuador, Quito). - P.

Thompson (United Kingdom, Coventry, HA GAM). -B. van Bekkum-Ansari (The Netherlands, Leiden,

Naturalis). - J. Weintraub (USA, Philadelphia, ANSP). - Keith Willmott (USA, Gainesville, FLMNH).

- P. Wilson (United Kingdom, Cornwall). - W. Xie (USA, Los Angeles, LACM).

The authors also thank two anonymous referees for their useful comments and suggestions.

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Manuscript received: 6 August 2013

Manuscript accepted: 28 October 2013

Published on: 3 January 2014

Topic editor: Koen Martens

Desk editor: Natacha Beau

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium;

Natural history Museum of Denmark, Copenhagen, De nm ark

European Journal of Taxonomy 71: 1-66 (2014)

Appendix 1

The discovery of the E. zolvizora group: a historical and geographical survey

In Bolivia, the first specimens of E. zolvizora apparently were collected by Clarence Buckley, who

travelled in the country in 1873-75 (Sclater & Salvin 1879, Vane-Wright 1991, Paynter 1992). The

specimen currently considered as the holotype (Fig. 4A), along with one other with the same label data,

were bequested to the BMNH in 1879 by W.C. Hewitson (Kirby 1879; BMNH registry records). Later,

other specimens were collected at the end of the 19 th century by the Garlepp brothers, who worked for

the German entomologist and trader Otto Staudinger. Gustav Garlepp collected in various localities,

mostly in the departments of Santa Cruz (1890) and Cochabamba (1890-1892). In the BMNH, there is

a female he collected at San Jacinto in Cochabamba department (Fig. 7A). Otto Garlepp joined him in

1893, and they collected from 1893 to 1897, and in 1904, in La Paz department (Berlepsch 1901, Paynter

1992, www.zobodat.at/D/runD/D/cacheD/personen_details.php?nr=7091 accessed 30 Jul. 2013). Other

historical specimens were collected by Anton Heinrich Fassl, who worked for two years, from the end

of 1912, in the upper Beni basin (La Paz department) (Fassl 1920). More recent specimens from this

area, labelled “Caranavi”, were provided by insect dealers. Specimens from Cochabamba department

labelled “El Palmar”, collected in the 1950s, were perhaps provided by Francisco Steinbach, or by

Rudolf Zischlca, from the locality “El Palmar”, along the ancient road from Cochabamba to Villa Tunari.

At the end of the 1970s, Nicolas Kuscevic sent specimens to one of the authors (PB) from localities along

the new road joining Cochabamba to Villa Tunari, but without precise data. Nevertheless, he regularly

collected near Incachaca at 2000 m (Nicolas Kuscevic, pers. comm.), and his E. zolvizora specimens are

with little doubt from this locality. More recently, a few specimens from Santa Cruz, Cochabamba and

La Paz departments have been collected by Bolivian entomologists; the southernmost known specimen

(Fig. 4B), from Manchones, Santa Cruz department, by one of the authors (YG).

For the purpose of this study, we divided Peru into a southern part, including Puno, Madre de Dios

and Cusco departments, a central part, including Junin and Pasco departments, and a northern part,

including Huanuco, La Libertad, Cajamarca, San Martin and Amazonas departments. Southern and

central specimens are few, and only males are known; nevertheless, they provide reliable information.

The oldest specimen we know - a syntype offs, opimus Staudinger, 1887 (Fig. 4E) - was collected in the

Chanchamayo valley (Junin department), in 1885, by Franz Thamm, a German collector who worked for

Otto Staudinger (Lamas 1981). In the very south, specimens were collected by the Englishman George

Richard Ockenden, who made important butterfly collections in Puno from 1900 to 1906 (Lamas 1981).

There are a few other old specimens from Chanchamayo/Perene (Junin) and Oxapampa (Pasco) in the

Paul Ferdinand Martin collection (MJP). Two males and one female from Cushi near Pozuzo (northern

Pasco department, close to Huanuco), were collected by the German Wilhelm Hoffmanns, who worked

in central Peru from 1901 to 1904, for Lord Rothschild (Lamas 1981). Some specimens from Cusco and

Junin departments have been collected more recently by Peruvian entomologists (Fig. 4C), as well as

by trade collectors. Northern males and females are known from different localities. From time to time,

specimens from Huanuco department are provided by traders; they were generally collected in the area

of Carpish, south of Tingo Maria. A few specimens have been collected by Peruvian entomologists in La

Libertad and Amazonas departments (Fig. 4D). The best infonnation has been gathered for Amazonas

populations (Mendoza, San Jose de Molinopampa and Alto Nieva areas), during Patrick Blandin and

Gilbert Lachaume surveys (2005-2012) (Figs. 4F, 7B). Unfortunately, no material is known from the

department of Cajamarca nor from the Cordillera del Condor, which forms the border with Ecuador.

Perhaps the oldest Ecuadorian specimens were collected by Marc de Mathan, a French collector who

did a tremendous amount of work in northern Peru and Ecuador, between 1879 and 1909 (Lamas 1981).

Another French collector, Paul Dognin, who lived in Loja, indicated the presence of Eryphanis opimus

(according to his determination) in the surroundings (Dognin 1894). Campos (1927) referred to this

BLANDIN P. et al. , Revision of the Andean Eryphanis zolvizora group

information in his list of Ecuadorian brassolines. Nevertheless, populations from south-eastern Ecuador

(Zamora-Chinchipe province) are poorly known: one female from Parque Podocarpus, Loja (Pinas

2004, figs. 655-656), one male and one female collected by Stephane Attal (SA, MNHN); two males

collected at the end of the 1990s by Artur Jasinslci in Zamora-Chinchipe province (MZUJ), and five

males collected by Keith Willmott in 1997 and 2006 (FLMNH). Blandin & Descimon (1975) quoted

two males from eastern Ecuador (Tungurahua and Napo provinces), and provided drawings of dorsal

and ventral surfaces. In fact, a large number of specimens are known from these provinces, where many

people have collected, notably the Velastegui family, since the 1960s. However, many specimens lack

precise locality data. Fortunately, Penz (2008) provided detailed information on the population studied

by Harold Greeney at the Yanayacu Biological Station (Napo) ( Eryphanis greeneyi Penz & DeVries,

2008; Fig. 5A).

Blandin & Descimon (1977), in a survey of Brassolini from western Ecuador, were unaware of the

existence of “ zolvizora” populations. Old specimens of zolvizora labelled from the western village of

‘Balzapamba’, like many supposedly from this locality, are from eastern Ecuador. The first specimen

with good locality data, a female (Fig. 7E), was caught in 1926 by C. Judson Coxey at Huigra (in

ANSP), but this specimen, until recently, has been overlooked. There is a female collected in 1982

from ‘Santo Domingo’ (probably Tinalandia) in LACM. In MZUJ, there is a male collected by Artur

Jasinslci in 1997, labelled from Balsas, El Oro province; however, this specimen is probably mislabelled

(Keith Willmott, pers. comm.); it is likely that it was collected in the Zamora-Chinchipe province. Keith

Willmott infonned us (pers. comm.) of one male from the Tandayapa Bird Lodge (Pichincha province),

on the 28 Nov. 2007, observed by Will Cook; web images of this specimen are available on two sites

www.carolinanature.com/pix/ecuador/leps3.html and www.tandava.com [accessed 30 Jul. 2013], A

male (presumably), attracted to a drinking trough for hummingbirds, was observed and photographed

on the 12 Nov. 2009, in the Rio Guajalito Reserve, at 1900 m on the old road from Quito to Santo

Domingo (Pichincha) (Xavier Silva, pers. comm.; Silva, 2011). Finally, Tim Kell caught a male (Fig.

5D) atReservaLas Gralarias (Pichincha) in Apr. 2011.

Colombia has been historically poorly sampled in comparison with other countries rich in butterflies.

Label data found on specimens of Eryphanis are often somewhat vague. Therefore, we decided to refer

to the following areas: the western slopes of the western cordillera; the Cauca valley and the Magdalena

valley, indicating, where possible, their westesrn or eastern slopes; the eastern slopes of the eastern

cordillera. The western slopes of the Cauca valley correspond to the eastern side of the western cordillera

- generally the Cali area -, and its eastern slopes to the western side of the central cordillera.

Specimens labelled “Bogota” have no reliable location. Eryphanis opimus Staudinger, 1887 was

described from Manizales (Fig. 5E). Anton Heinrich Fassl, who travelled in the country from 1908-1912,

quoted Eryphanes [sic] opimus Stgr. in association with Opsiphanes camena and Caligo oberthuri [sic]

in the western cordillera, without indicating on which side (Fassl 1914), but later he noted opimus on

both sides (Fassl 1915a). In the USNM there is one male he collected, labelled ‘Rio Aguacatal, 2000 m’,

located on eastern slopes north-west of Cali (Fassl 1914, 1915a). Presumably from the same area, but

labelled ‘Rio Aguatal’ and ‘Rio Agua’, there are two known females (in UFPC and BMNH) (Fig. 7F).

Fassl (1911) did not mention opimus in the central cordillera, despite the fact that a few specimens had

been collected in the 1880s. Ultimately, Fassl wrote that E. opimus was very rare on the eastern slopes

of the eastern cordillera (Fassl 1915b). To our knowledge, the oldest Colombian specimen, labelled

‘Bogota’ (UMO), was collected by the British Vice Consul Edward W. Mark between 1848 and 1857.

Today, Colombian populations are known through a few specimens from a limited number of localities.

The majority came from the Cauca valley (western slopes and eastern slopes). Two males, in the MNHN

and MPM, are labelled “Cali, 1000 m”. These were apparently provided by a trader who stamped all

European Journal of Taxonomy 71: 1-66 (2014)

specimens with this “locality”. Nonetheless they might have been collected on the road from Cali to

Buenaventura. There is a female in the MNHN labelled “Colombie, 1901, H. Donckier, Coll. Boullet”,

and two males from the same collection labelled “Cauca, 1887, H. Ribbe” and “Cauca, 1903, H. Ribbe”.

It is presumed that these specimens simply labelled “Cauca” are from the Cauca valley. A male and

female (HAGAM) are labelled from Calima Valley, ‘Cauca River’ [sic], and are therefore probably from

the eastern side of the western cordillera. There is one male (JS) from the western side of the western

cordillera, km 55. Three specimens (lc?, IAvH; l<f, 1$ ICNUN; Figs. 5C, 7D), have been collected

in the department of Narino. Information concerning the Magdalena valley and the eastern side of the

eastern cordillera remains limited. Two males labelled “Caqueta” (JFTC), almost certainly came from

the eastern cordillera on the road between Altamira and Florencia between 1500 and 2000 m (J.-F. Te

Crom, pers. comm.). Recently in 2005, one of the authors (BH) and colleagues discovered a population

in the Serrania de los Yariguies, Santander department, Municipio Galan (2f, IAvH), in the Magdalena

valley, on the western side of the eastern cordillera, and there is one specimen from Charala (Santander)

in MNHN (Fig. 6B).

The first Venezuelan specimen known to us was collected in 1965 by Eduardo Rodriguez and the Romero

family in the Cordillera de la Costa (MIZA). Eater, in the 1970s, other specimens were collected in this

cordillera, the majority by the Romero family in Aragua state on the road from Maracay to Choroni, in

the west-central Cordillera de la Costa (Figs. 6C, 8B). In 1972, during an expedition by the MIZA in

the Cordillera de Merida, a pair was collected by Juvenal Salcedo and Felix Zambrano near Anzoategui

in Lara state. Two further populations were subsequently discovered by the Romero family in the mid

1980s in the Santo Domingo valley in Barinas state, on central eastern slopes of the Cordillera de

Merida (Figs. 6A, 8A), and in Tachira state in the northern Sierra de El Tama, in the area known as

Matamula and Pabellon. More recently a female was collected in Vargas state above Maiquetia at 1650

m (incorrectly labelled “La Llanada”; unidentified collector) on the Camino viejo de Los Espanoles to

Caracas (MNHN), and a population was located in the area around Colonia Tovar in the east-central

Cordillera de la Costa (Aragua state) by the Romero family and Mauro Costa. In 2001 a further population

was located by Mauro Costa, this time in Merida state in the central valley of the Cordillera de Merida,

approximately 55 km SW of the city of Merida. While the number of known specimens - especially

females - from Venezuela is relatively high compared with other countries, the data indicate that, with

rare exceptions, specimens are caught singly and often years apart, indicating great rarity.

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Appendix 2

List of the studied specimens of Eryphanis zolvizora

Departments or Provinces have been included (between brackets). Unless they have been provided

by collectors, approximate coordinates have been added where possible. Unless otherwise indicated,

altitude is given in meters.

■ = Specimens on which measurements have been taken directly. ♦ = Specimens on which measurements

have been taken on scaled pictures. * = Specimens of which genitalia have been dissected.

Eryphanis zolvizora zolvizora (Hewitson, 1877)

BOLIVIA: Santa Cruz and Cochabamba departments

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Camino a Manchones 17 Apr. 2011,

Y. Gareca leg. (Santa Cruz)

18°45’8.76”S

64°0 , 30.72”W

1695

MHNC

Bellavista, 11-15 Oct. 2007, J. Lede,

L. Gomez and A. Alcoba (Santa Cruz)

18°12’00”S

63°45’00”W

1382

MHNNKM#21376

MHNNKM

Bellavista, 11-15 Oct. 2007, J. Lede,

L. Gomez and A. Alcoba (Santa Cruz)

18°12’00”S

63°45’00”W

1382

MHNNKM#21377

MHNNKM

Sivingal, Prov. Florida, 8 Mar. 1991,

W. Rojas leg. (Santa Cruz)

6140

MHNNKM

Buenavista, 21 Feb. 1994 (Santa Cruz)

FLMNH

Buenavista, 06 Apr. 1925 (Santa Cruz)

FLMNH

Rio Juntas, 1890, G. Garlepp leg.

(Cochabamba)

17°07’S

65°16’W

1000

ZMHB

El Palmar, Apr. 1950 (Cochabamba)

17°06’S

65°17’W

1600

PBB 1408

MNHN

El Palmar, Nov. 1958 (Cochabamba)

17°06’S

65°17’W

1600

FLMNH

Em*

Probably Incachaca, N. Kuscevic leg.

(Cochabamba)

17°14’S

65°49’W

2000

PBB 971

MNHN

Chapare, Oct. 1949 (Cochabamba)

300

PBB 260

MNHN

Villa Tunari - Cochabamba, Sillar

Bajo, 17 Feb. 2009, T. Pircz leg.

(Cochabamba)

1450

MZUJ

Yendo a Chocolatal 12-17 Feb. 2009,

Y. Gareca, T. Pircz and J. Wojtusiak

leg. (Cochabamba)

17°08’19.6”S

65°42’43.9 , ’W

1518

MHNC

Carmen Pampa 25-28 Sep. 2001, C.

Flamel and A. Pascall (Cochabamba)

16°37’06”S

66°28’50” W

1820

MHNC

Locotal, 1891, G. Garlepp leg.

(Cochabamba)

17°1LS

65°39’W

2600

ZMHB

San Jacinto, G. Garlepp leg.

(Cochabamba)

17°10’S

65°44’W

6-8000ft

BMNH(E)#808088

BMNH

El Palmar, Apr. 1947 (Cochabamba)

17°06’S

65°17’W

1600

FLMNH

cp*

Probably Incachaca, Kuscevic leg.

(Cochabamba)

17°14’S

65°49’W

2000

PBB 1061

MNHN

European Journal of Taxonomy 71 : 1-66 ( 2014 )

Eryphanis zolvizora zolvizora (Hewitson, 1877)

BOLIVIA: La Paz department

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

R. Tanampaya, 1894, Garlepp leg.

(La Paz)

16°21’46.38”S

67°35’34.73”W

1146

BMNH(E)#808077

BMNH

Chaco, 1894, Garlepp leg. (La Paz)

16°19’59.76”S

67°47’59.67”W

2000-3000

ZMHB

Tunquini-El Chairo, 2003, F. Guerra

leg. (La Paz)

16°12’28.42” S

67°52 , 59.92” W

1600

MHNNKM6037

MHNNKM

Nor Yungas PN-ANMI Cotapata Est.

Biologica Tunquini, Sep. 1999, J.

Corro leg. (La Paz)

16°12’28.42” S

67°52 , 59.92” W

1600

CBF-1125

CBF

<?♦

San Antonio, Mar. 1896, Garlepp leg.

(La Paz)

17°14’S

65°19’W

1200

BMNH(E)#808071

BMNH

San Antonio, 1896, Garlepp leg.

(La Paz)

17°14’S

65°19’W

1800

ZMHB

Coroico, 1914, A H. Fassl leg. (La Paz)

16°10’S

67°44’W

1200

BMNH(E)#808076

BMNH

<?♦

Coroico, A H. Fassl leg. (La Paz)

16°10’S

67°44’W

1200

Coleqao Julius Arp

N°18/537

UFPC

<?♦

Coroico, A H. Fassl leg. (La Paz)

16°10’S

67°44’W

1200

USNM

Coroico (La Paz)

16°10’S

67°44’W

1200

BMNH(E)#808073

BMNH

Coroico, E.H. Page leg. (La Paz)

16°10’S

67°44’W

1200

BMNH

Coroico (La Paz)

16°10’S

67°44’W

1200

RAMM

Zongo, Rio 16 Mar. 1979, Schmitt leg.

(La Paz)

15°47’S

67°44’W

AMNH

Farinas (La Paz)

15°55’S

67°47’W

RAMM

<?■

“Caranavi” (La Paz)

15°56 , 02”S

67°27’42”W

PBB2153

MNHN

(?■*

“Caranavi” (La Paz)

15 o 56’02”S

67°27’42”W

PBB 2286

MNHN

(?■*

“Caranavi” (La Paz)

15°56’02”S

67°27’42”W

PBB 2287

MNHN

Nor Yungas, Villa Incahuara, Caranavi,

C. Tello Egg leg. (La Paz)

15°23 , 37.81”S

67°40’25.30”W

CBF-4529

CBF

<?■

Nord Yungas 26-27 Jan. 1984, T.

Porion leg. (La Paz)

1600

PBB 1410

MNHN

Las Yungas, Nov. 1990 (La Paz?)

FLMNH

Bolivie, Garlepp leg. (Staudinger

1895) (La Paz)

BMNH(E)#808074

BMNH

(j)*

Coroico, A H. Fassl leg. (La Paz)

16°10’S

67°44’W

1200

BMNH(E)#808089

VIAL 8577

BMNH

Coroico, A H. Fassl leg. (La Paz)

16°10’S

67°44’W

1200

BMNH(E)#808090

BMNH

Coroico, A H. Fassl leg. (La Paz)

16°10’S

67°44’W

1200

Coleqao Julius Arp

N° 18/538

UFPC

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Eryphanis zolvizora zolvizora (Hewitson, 1877)

BOLIVIA: La Paz department (continued)

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Zongo, Pacollo, 8 Feb. 1991 (noche),

R. Coreticona leg. (La Paz)

16°02’29”S

67°58 , 26”W

CBF-4530

CBF

cp*

Rio Songo, 1910, A H. Fassl leg. (La

Paz)

15°43’S

67°41’W

MNHN

Eryphanis zolvizora zolvizora (Hewitson, 1877)

BOLIVIA: specimens without precise or reliable locality

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Bolivia (Buckley) Hewitson

Coll. 79-69.

Pavonia zolzivora Hew, 2

BMNH(E)#705086

BM TYPE 6110

SYNTYPE

E. zolvizora

BMNH

[Bolivia] Hewitson 1

SYNTYPE E.

zolvizora

BMNH

Bolivia

BMNH(E)#808091

BMNH

Bolivia

BMNH(E)#808092

BMNH

Bolivia

BMNH(E)#808072

BMNH

Bolivia (Levick)

BMNH(E)#808075

BMNH

Bolivia

Colegao Julius Arp

N°18/536

UFPC

Bolivie

PBB 1409

MNHN

Em*

Bolivie (0. Staudinger, 1902)

MNHN

S+

Bolivia

USNM

Bolivia

RAMM

Negnejahuza, May 1926

AMNH

European Journal of Taxonomy 71: 1-66 (2014)

Eryphanis zolvizora inca Blandin, ssp. nov.

PERU: Puno : Cuzco, Junin and Pasco departments

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

(?♦*

Santo Domingo, Carabaya, Jan. 1901

Wet season, G.R. Ockenden leg.

(Puno)

13°50’S

69°40’W

6000 ft

BMNH (E) #

806244

VIAL 8223

BMNH

<?♦*

Santo Domingo, Carabaya, 1901 Dry

season, G.R. Ockenden leg. (Puno)

13°50’S

69°40’W

4500 ft

BMNH (E) #

808079

VIAL 8576

BMNH

(?♦

Santo Domingo, Carabaya, 1901 Dry

season G.R. Ockenden leg. (Puno)

13°50’S

69°40’W

4500 ft

BMNH (E) #

808078

BMNH

Inca Minas, 1 Jul. 1900 (Puno)

13°50’S

69°40’W

AMNH

Carabaya (Puno)

14°05’S

70°27’W

AMNH

Inambari (Puno)

BMNH (E)#

525986

BMNH

<?♦

Chirimayo, Jun. 1901 Dry season, G.R.

Ockenden leg. (Puno)

13°27’S

70°18’W

1000 ft

BMNH (E) #

808081

BMNH

San Lorenzo, Rio Marcapata (Cusco)

PBB2115

MNHN

<?♦

San Pedro, 4-8 Nov. 2007, F. & A.

West leg. (Cusco)

13°03’S

71°33’W

1400

MJP

<?♦*

Aguas Calientes, 21 Oct. 2001, G.

Lamas leg. (Cusco)

13°09’S

72°31’W

2050

Genitalia

MJPZOLV 003PB

MJP

<?♦

Marchu Pijchu [sic] A. Larichesy leg.

(Cusco)

2000

IG 10 375

IRSN

Macchu Picchu [sic], 10 Dec. 1987, J.

Dubois leg. (Cusco)

MNHN,

<?♦*

Llacatahuaman, Quebrada Bagre, 27

Jul. 1998, G. Valencia leg. (Cusco)

12°52’S

73°30’W

1700

Genitalia

MJPZOLV 001PB

MJP

c7-*

Alfamayo, route Ollantaytambo-

Quillabamba, M. Cabrera leg. (Cusco)

PBB2184

MNHN

<?■*

Calabaza, Pampa Hermosa, prov.

Satipo, Apr. 2008 (Junin)

11°29’2.02 S

74°47’39.25”W

2200

PBB 2308

MNHN

Chanchamayo, (Junin)

11°04’S

75°19’W

1500

AMNH

<?♦*

Rio Perene (Junin)

Genitalia

MJPZOLV 005PB

MJP

<?♦*

Oxapampa (Pasco)

Genitalia

MJPZOLV 007PB

MJP

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Eryphanis zolvizora chachapoya Blandin, ssp. nov.

PERU: Junin, Pasco, Huanuco and La Libertad departments

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Chanchamayo, 1885, F. Thamm leg.

(Junln)

11°04’S

75°19’W

SYNTYPE

E. opimus

BMNH

San Francisco, Chanchamayo, Jan.

1964, Mme Harris leg. (Junin)

PBB 1407

MNHN

<?♦*

Rio Perene (Junin)

Genitalia

MJPZOLV 006PB

MJP

<?♦*

Oxapampa, La Suiza, 5-13 Jun. 2002,

B. Wenezel leg. (Pasco)

2180

UFPC

<?♦*

Huancabamba (Pasco)

10°23’S

75°33’W

6800 ft

BMNH (E)#

806245 VIAL 8224

BMNH

<?♦*

Cushi, W. Hoffmans leg. (Pasco)

09°58’S

75°42’W

1820

BMNH (E)#

806246 VIAL 8225

BMNH

Cushi, W. Hoffmans leg. (Pasco)

09°58’S

75°42’W

1900

BMNH

Cushi, W. Hoffmans leg. (Pasco)

09°58’S

75°42’W

1900

BMNH (E)#

806247 VIAL 8226

BMNH

<?♦

Panao, Jan. 1982, Schunke leg.,

(Huanuco)

09°54’S

75°58’W

3000

UFPC

Carpish, (Huanuco)

09°43’S

76°06’W

PBB 259

MNHN

(?■*

Carpish, Jan. 2006 (Huanuco)

09°43’S

76°06’W

2300

PBB 2284

MNHN

(?■*

Carpish, Apr. 2008 (Huanuco)

09°43’S

76°06’W

2800

PBB 2311

MNHN

(?■*

Probably Carpish (Huanuco)

1500-2000

PBB 2326

MNHN

(j)*

Carpish, Jul. 2005 (Huanuco)

09°43’S

76°06’W

2300

PBB 2285

MNHN

Cp H*

Mallqui, (Huanuco)

1500-2000

PBB 2327

MNHN

<?♦*

Cumpang, entre Tayabamba y Ongon,

21 Oct. 1979, T. Parker, leg. (La

Libertad)

08°16’S

77°00’W

2400-2700

Genitalia

MJPZOLV 004PB

MJP

Eryphanis zolvizora chachapoya Blandin, ssp. nov.

PERU : Amazonas, San Martin

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Mendoza area, Jun. 1994, B. Calderon

leg. (Amazonas)

06°23’S

77°27W

PBB 1900

MNHN

< Sn*

Mendoza area, Nov. 2006, B. Calderon

leg. (Amazonas)

06°23’S

77°27W

PBB 2289

MNHN

<?♦*

Valle de Huamanpata, Lejia, 23 Oct.

2005, L. Campos leg. (Amazonas)

06°20’S

77°27’W

2150

Genitalia

MJPZOLV 002PB

MJP

San Jose de Molinopampa, 20 May

2005

B. Calderon leg. (Amazonas)

06°12’S

77°34’W

2200-2400

PBGL 153

MNHN

(?■*

San Jose de Molinopampa, 2005

B. Calderon leg. (Amazonas)

06°12’S

77°34’W

2200-2400

PBGL 166

MNHN

European Journal of Taxonomy 71: 1-66 (2014)

Eryphanis zolvizora chachapoya Blandin, ssp. nov.

PERU : Amazonas, San Martin (continued)

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

(?■*

San Jose de Molinopampa, 20 May

2005 B. Calderon leg. (Amazonas)

06°12’S

77°34’W

2200-2400

PBGL 190

MNHN

(?■*

San Jose de Molinopampa, Mar. 2005

B. Calderon leg. (Amazonas)

06°12’S

77°34’W

2200-2400

PBGL 296

MNHN

(?■*

San Jose de Molinopampa, 2008,

B. Calderon leg. (Amazonas)

06°12’S

77°34’W

2200-2400

PBB 2309

MNHN

Oso Perdido, Dec. 2011, J.C. Pintado

leg. (Amazonas)

05°43’S

77°50-51’W

2200-2400

PBB 2324

MNHN

(?■*

Le long de l’alto rio Nieva, 4-9 Mar.

2009 (Amazonas)

05°42’40”S

77°47’15”W

2250

PBGL 471

MNHN

Le long de l’alto rio Nieva, 9 Mar.

2009 (Amazonas)

05°42’40”S

77°47 , 15”W

2250

PBGL 518

MNHN

(?■*

Alto Nieva (Amazonas)

05°39’S

77°47’W

1900

PBB 2156

MNHN

(?■*

Alto N ieva (Amazonas)

05°40’S

77°46’W

1900

PBB 2157

MNHN

Alto Nieva, Dec. 2012, J. C. Pintado

leg. (Amazonas)

05°40’S

77°46’W

1900

PBB 2325

MNHN

(?■*

Nieva, Mar. 2010, J.C. Pintado leg.

(Amazonas)

05°40’S

77°46’W

1900

PBGL 592

MNHN

(?■*

Villa Hermosa, vallee du rio

Nieva, Dec. 2010, J.C. Pintado leg.

(Amazonas)

05°36’S

77°47’W

1500

PBGL 594

MNHN

<?■

Nieva, Sep. 2011, J.C. Pintado leg.

(Amazonas)

05°40’S

77°46’W

1900

PBGL 595

MNHN

San Jose de Molinopampa, 15 Jan.

2007, B. Calderon leg. (Amazonas)

06°12’S

77°34’W

2200-2400

PBGL 198

MNHN

<^)*

Alto Nieva, Mar. 2010, J.C. Pintado

leg. (Amazonas)

05°40’S

77°46’W

1900-2000

PBGL 519

MNHN

(j)*

Alto Nieva, camino a Villa Hermosa,

Feb. 2010, J.C. Pintado leg.

(Amazonas)

05°40’S

77°46’W

1900-2000

PBGL 520

MNHN

Rio Jenesis, Nov. 2012, J. C. Pintado

leg. (Amazonas)

05°32-33’S

77°48-49’W

1400-1600

MJP

Nieva, Sep. 2011, J.C. Pintado leg.

(Amazonas)

05°40’S

77°46’W

1900

PBGL 596

MNHN

El Choferrito, Alto Mayo, Oct. 2010,

J.C. Pintado leg. (San Martin)

05°40’S

77°45’W

1800

PBGL 593

MNHN

Sm*

Jorge Chavez, Alto Mayo, 9 Nov. 2006

(San Martin)

5°40-41’S

77°43-44’W

1300-1400

PBGL 470

MNHN

(?■*

Jorge Chavez, Alto Mayo, (San Martin)

5°40-4ES

77°43-44’W

1300-1400

PBB 2283

MNHN

(7-*

Jorge Chavez, Alto Mayo, Sep. 2007

(San Martin)

5°40-41’S

77°43-44’W

1300-1400

PBB 2310

MNHN

<?■*

San Augustin, Naranjos area, 4 Nov.

2006 (San Martin)

5°47’S

77°30’W

1400

PBGL 521

MNHN

San Augustin, Naranjos area, Sep.

2012 (San Martin)

5°47’S

77°30’W

1400

MJP

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Eryphanis zolvizora greeneyi Penz & DeVries, 2008

EASTERN ECUADOR: Zamora-Chinchipe, Morona-Santiago and Pastaza provinces

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Cordillera Llagunillas, Rd. San

Andreas - Calderon, 20 May 1998

(Zamora-Chinchipe)

2000-2600

MUZJ

Romerillos, ridge to west, 1 Nov. 1997,

K. R. Willmott (Zamora-Chinchipe)

4°11.0’S

78°56.50W

1600

FLMNH

San Francisco, casa de Arcoiris, km

24 Loja-Zamora rd., 14 Sep. 2006,

K.R. Willmott, R. Aldaz leg. (Zamora-

Chinchipe)

3°59.30’S

79°5.58’W

2000-2100

FLMNH# 111778

FLMNH

San Francisco, casa de Arcoiris, km

24 Loja-Zamora rd., 27 Oct. 2006,

K.R. Willmott, R. Aldaz leg. (Zamora-

Chinchipe)

3°59.30’S

79°5.58’W

2000-2100

FLMNH# 111779

FLMNH

San Francisco, casa de Arcoiris, km

24 Loja-Zamora rd., 27 Oct. 2006,

K.R. Willmott, R. Aldaz leg. (Zamora-

Chinchipe)

3°59.30’S

79°5.58’W

2000-2100

FLMNH# 111776

FLMNH

San Francisco, casa de Arcoiris, km 24

Loja-Zamora rd., 2 Nov. 2006, K.R.

Willmott leg. (Zamora-Chinchipe)

3°59.30’S

79°5.58’W

2000-2100

FLMNH# 111777

FLMNH

Rio San Francisco (Rio Zamora), 12

Oct. 2006, S. Attal leg. (Zamora-

Chinchipe)

03°58’S

79°05’W

2000

cp *

Valladolid, 03 Jan. 2002, S. Attal leg.

(Zamora-Chinchipe)

04°34’S

79°08’W

PBB 2288

MNHN

Parque Podocarpus, 1 May 1999, M.

Kling (Loja)

04°06’S

78°57’W

2000

0656

Macas-Guamote, Sept. 1999, Estevez

leg. (Morona-Santiago)

02°16’S

78°10’W

1500

MECN

Puyo area, Feb. 1996 (Pastaza)

PBB 2328

MNHN

Eryphanis zolvizora greeneyi Penz & DeVries, 2008

EASTERN ECUADOR: Tungurahua, Napo and Sucumbios provinces

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

8m*

Machay, Jan. 1994 (Tungurahua)

01°24’S

78°18’W

1800-2200

PBB 1899

MNHN

8m*

Machay, 14 Feb. 1995 (Tungurahua)

01°24’S

78°18’W

1800-2200

PBB 2166

MNHN

Machay, 1 Aug. 1997 (Tungurahua)

01°24’S

78°18’W

1800-2200

MUZJ

8m*

Rio Blanco, 3 May 1972 (Tungurahua)

01°23’S

78°20’W

1700

PBB 729

MNHN

8m*

Rio Blanco, San Jose, 10 Jun. 1967

(Tungurahua)

01°23’S

78°20’W

1700

PBB 257

MNHN

8m*

Rio Blanco, 25 Apr. 1966

(Tungurahua)

01°23’S

78°20’W

1700

PBB 537

MNHN

8 ♦

Rio Blanco, 25 Aug. 1937

(Tungurahua)

01°23’S

78°20’W

1800

USNM

European Journal of Taxonomy 71: 1-66 (2014)

Eryphanis zolvizora greeneyi Penz & DeVries, 2008

EASTERN ECUADOR: Tungurahua, Napo and Sucumbios provinces (continued)

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Rio Blanco (Tungurahua)

01°23’S

78°20’W

1800

<?♦

Rio Blanco, near Banos, 17 Apr. 1956

(Tungurahua)

01°23’S

78°20’W

1650

scaled picture,

C. Penz

MPM

Rio Blanco, 28 Dec. 1938

(Tungurahua)

01°23’S

78°20’W

BMNH(E)#808100

BMNH

Rio Verde, 6 May 2001 (Tungurahua)

01°24’S

78°17’W

1500

Rio Verde, E. Ecuador, M.G. Palmer

(Tungurahua)

01°24’S

78°17’W

5000ft

BMNH(E)#808082

BMNH

Ambato, Anda Vazconez (Tungurahua)

BMNH(E)#808084

BMNH

Ambato, Anda Vazconez (Tungurahua)

BMNH(E)#808085

BMNH

Env. d’Ambato. R. P. Blanc

(Tungurahua)

BMNH(E)#808086

BMNH

Equateur Or. de Banos a Canelos. M.

de Mathan 9 Oct. 1894. (Tungurahua/

Pastaza)

BMNH(E)#808083

BMNH

Rio Verde, E. Ecuador, M.G. Palmer

(Tungurahua)

01°24’S

78°17’W

5000ft

BMNH(E)#808097

BMNH

Yunguilla, Juil. 1937 (Tungurahua)

01°12’S

78°20’W

1900-2000

AMNH

Biscaya (= Viscaya), 7 May 1996

(Tungurahua)

01°20’34”S

78°24’07”W

MUZJ

(?■*

Rio Arajuno, 27 Mar. 1968 (Napo)

01°14’S

77°42’W

700

PBB 258

MNHN

El Arrayan, Baeza, 28 Dec. 1997,1.

Manzano (Napo)

2000

0654

<?♦

Yanayacu Biological Station, 5km W

of Cosanga, May 2007, reared by H.F.

Greeney, (Napo)

00°35,9’S

77°53,4’W

2163

HOLOTYPE

E. greeneyi

BMNH(E)#808102

BMNH

(?♦

Yanayacu Biological Station, reared,

2006, H. Greeney, (Napo)

00°35,9’S

77°53,4’W

2163

BMNH

Napo

465

Yanayacu Biological Station, May

2007, reared by H.F. Greeney, (Napo)

00°35’S

77°53’W

2163

PARATYPE

E. greeneyi

BMNH(E)#808103

BMNH

Rio Palmar, km 16.5 La Bonita-

Rosa Florida rd., 23 Nov. 1996, K.R.

Willmott leg. (Sucumbios)

0°25.10’N

77°32.20’W

1200

FLMNH

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Eryphanis zolvizora greeneyi Penz & DeVries, 2008

EASTERN ECUADOR: specimens without precise or reliable locality

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Balsas, 14 Apr. 1997, K. Jasinsly

leg. (mislabelled? K. Willmott pers.

comm.)

800-1100

MUZJ

Balzapamba, Ecuador occ.

(mislabelled)

FLMNH

Balzapamba, Ecuador occ.

(mislabelled)

USNM

Equateur, ‘Balzapamba’. Prov. de

Bolivar. M. de Mathan Oct. 1893 -

Feb. 1894 (mislabelled)

BMNH(E)#808098

BMNH

Balzapamba (mislabelled)

BMNH

Ecuador

AMNH

Ecuador

Ecuador (Levick collection)

BMNH(E)#808087

BMNH

Ecuador

BMNH(E)#808101

BMNH

Ecuador

BMNH(E)#808099

BMNH

Ecuador

NNM

Ecuador

ZMUA

Oriente

AMNH

Ecuador, Velastegui leg.

JFLC

Eryphanis zolvizora uncertain ssp., cf. greeneyi Penz & DeVries, 2008

COLOMBIA: southern Cordillera Oriental

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

2<?<?

[Altamira-Florencia road], Caqueta,

Mar. 1987, Urbina leg.

1500-2000

JFLC

Eryphanis zolvizora greeneyi Penz & DeVries, 2008

COLOMBIA: specimen without reliable locality

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Puerto Narino, 20-30 Jul. 1980

(Amazonas)

3°45’N

70°20’W

150

LACM

European Journal of Taxonomy 71: 1-66 (2014)

Eryphanis zolvizora casagrande Bristow, ssp. nov.

WESTERN ECUADOR: Pichincha and Chimborazo provinces

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

<?♦*

Reserves Las Gralarias, 27 Apr. 2011,

T.Kell (Pichincha)

0°0’39”S

78°43’50”W

1800-2050

151126

FLMNH

Santo Domingo [Tinalandia ?], Apr.

1980 (Pichincha)

200

LACM

Tandayapa Lodge, 28 Nov. 2007, Will

Cook (Pichincha)

0°10’S

78°46’W

1600

Web image

c7?

Rio Guajalito Reserve, 12 Nov.

2009,old road from Quito to Santo

Domingo (Pichincha)

1900

Picture

p. 101 in

Silva 2011

Huigra, Nov. 1926 (Chimborazo)

02°18’S

78°59’W

1220

ANSP

Eryphanis zolvizora casagrande Bristow, ssp. nov.

ECUADOR: specimen without reliable locality

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Valladolid, 17 May 1998, K.

Jasinski leg. (Zamora-Chinchipe)

(mislabelled?)

MUZJ

Eryphanis zolvizora casagrande Bristow, ssp. nov.

COLOMBIA: Narino department

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

<?♦*

Altaquer, Rio Nambi, Jul. 1995

(Narino)

01°15’N

78°07’W

1380

GAC 7615 ICN-

MNH-L 14791

ICNUN

Ricaurte, La Planada, 8 Sep. 1994,

Julian Salazar leg. (Narino)

01°05’06”N

77°53’06”W

1800

IAvH 8238 S.M.

4385 Gen. JFL 336

IAvH

cp*

Reserva La Planada, via Hondon

Trampa : Malaise H2, 16 Oct.-2 Nov.

2000 G.. Oliva leg. (Narino)

01°15’N

78°15’W

1930

IAvH-E 67776

Gen. JFL 337

IAvH

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Eryphanis zolvizora opimus Staudinger, 1887

COLOMBIA: Caldas, Risaralda, Antioquia and Valle del Cauca departments

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Manizales (Caldas)

05°05’N

75°32’W

2500

SYNTYPE E.

opimus

ZMHB

Manizales (Caldas)

05°05’N

75°32’W

2500

SYNTYPE E.

opimus

ZMHB

Manizales. A.M. Patino. (Caldas)

05°05’N

75°32’W

BMNH(E)#806249

VIAL 8228

BMNH

Col. (Cauca) Distrito de Pereira.

Roman Valencia. 1886 (Risaralda)

BMNH(E)#806248

VIAL 8227

BMNH

Col. (Cauca) Distrito de Pereira.

Roman Valencia. 1886 (Risaralda)

BMNH(E)#808096

BMNH

Mesopotamia (Antioquia)

05°5LN

75°16’W

900

AMNH

Em*

Vallee du Quindio, environs Armenia

Mme Farre Duchartre leg. 1930

(Quindio)

04°25’N

75°23’W

1800

MNHN

Cauca Valley

BMNH(E)#806250

VIAL 8229

BMNH

Em*

Cauca, H. Ribbe, 1887

MNHN

Em*

Cauca C. Ribbe, 1903

MNHN

<?■*

“Cali”, 11 Aug. 1974 (Valle del Cauca)

1000

PBB 1406

MNHN

<?♦

Cali, 23 Sep. 1964

J.R. Neidhoefer Collection

1000

07-32 dissected by

C. M. Penz

MPM

Calima Valley, Cauca River, Nov. 1988

(Greenwood) (Valle del Cauca)

03°53’N

76°36’W

1300

HAGAM

E ♦

Rio Aguacatal, Colomb. West Cord.

Coll. Fassl (Valle del Cauca)

03°34’N

76°40’W

2000

USNM

Calima Valley, Cauca River, 19 May

1988 (Greenwood) (Valle del Cauca)

03°53’N

76°36’W

1300

HAGAM

R. Aguatal (presumably rio Aguacatal,

Valle del Cauca)

1800

Colegao Julius

Arp N° 18/540

UFPC

R. Agua (presumably rio Aguacatal,

Valle del Cauca)

BMNH

Valle, km 55, west side, 3 Oct. 1985,

J. Salazar

European Journal of Taxonomy 71: 1-66 (2014)

Eryphanis zolvizora opimus Staudinger, 1887

COLOMBIA: specimens without precise or reliable locality

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Buenaventura-Queremal 4 May 1985,

Schmidt-Mumm leg. (mislabelled?)

03°37’N

76°53’W

200

IAvH

Bogota, Colombia

BMNH(E)#808094

BMNH

Bogota

BMNH(E)#808095

BMNH

Bogota, between 1848 and 1857,

Edward. W. Mark, British Vice Consul

UMO

Bogota

USNM

cp*

Colombie, H. Donckier, 1901

MNHN

Eryphanis zolvizora reyi Bristow, Neild, De Sousa & Huertas ssp. nov.

COLOMBIA: northern Cordillera Oriental

Sex

Data

Coord.

Alt.

a.s.l.

Reference

Collection

8**

El Cerro-La Luchata, Municipio

Galan, 29 Jun. 2006, B. Huertas, J.C.

Rios & J.J. Arias leg., Proyecto YARE

(Santander)

06° 37’75” N

73° 18’ 88” W

1950-2050

IAvH

El Cerro-Ea Luchata, Municipio

Galan, 30 Jun. 2006, B. Huertas, J.C.

Rios & J.J. Arias leg., Proyecto YARE

(Santander)

06° 37’75” N

73° 18’ 88” W

1950-2050

IAvH

8m*

Charala Jul. 2008 (Santander)

06°15’N

73°05’W

2000

PBB 2321

MNHN

Eryphanis zolvizora reyi Bristow, Neild, De Sousa & Huertas ssp. nov.

VENEZUELA: Tachira, Merida, Barinas and Lara states

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

Sierra de El Tama, Via Delicia, Mar.

1987 (Estado Tachira)

7°36’N

72°26’ W

1900

(7-

Sierra de El Tama, Via Delicia

(Pabellon), Apr. 1990 (Edo. Tachira)

7°36’N

72°26’W

1700

cj)*

Paramo El Molino, SE of Sta. Cruz de

Mora, NW Venezuela. 3 Jan. 2001, M.

Costa leg. (Merida)

8°16’N

71°34’W

1600

Ea Mina, above San Isidro, Mar. 1988,

Legit Romero (Barinas)

8°50’N

70°34’W

1450

(7-

Barmitas [La Mina de San Isidro],

Oct. 1987. Estado Barinas.

8°50’N

70°34’W

1450

Barmitas. [La Mina de San Isidro],

Oct. 1987. (Barinas)

8°50’N

70°34’W

1450

115- JCSC

Barinitas [La Mina de San Isidro],

Nov. 1995. (Barinas)

8°50’N

70°34’W

1400

8m*

Vallee du Rio Santo Domingo, 16

Nov. 2009. M. Costa & S. Attal leg.

(Barinas)

8°50’N

70°34’W

1500

PBB 2323

MNHN

BLANDIN P. et al ., Revision of the Andean Eryphanis zolvizora group

Eryphanis zolvizora reyi Bristow, Neild, De Sousa & Huertas ssp. nov.

VENEZUELA: Tachira, Merida, Barinas and Lara states (continued)

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

2<?<?

San Isidro, Vallee du Rio Santo

Domingo, 9 Nov. 2010. M. Costa & S.

Attal leg. (Barinas)

8°50’N

70°34’W

1500

PBB 2324

PBB 2325

MNHN

San Isidro, Las Minas, 7 Sep. 2002,

Col. Juan C. De Sousa C. (Barinas)

8°50’N

70°34’W

1475

JCS

San Isidro, Las Minas, 10 Oct. 2010,

Col. Mauro Costa. (Barinas)

8°50’N

70°34’W

1500

JCS

San Isidro (Mina), 4 Oct. 2009,

(Barinas)

8°50’N

70°34’W

1500

MCC

San Isidro (Mina), 4 Oct. 2009,

(Barinas)

8°50’N

70°34’W

1500

MCC

San Isidro (Mina), 9 Nov. 2009,

(Barinas)

8°50’N

70°34’W

1500

MCC

San Isidro (Mina), 12 Dec. 2009

(Barinas)

8°50’N

70°34’W

1500

MCC

La Soledad, 5 Apr. 1985 (Barinas)

8°49’N

70°31’W

1000

MCC

La Chimenea. 1-6 Jun. 1973, J.

Salcedo col. (Barinas)

8°49’N

70°31’W

1500

MIZA

Barinitas [La Mina de San Isidro], Oct.

1987 (Barinas)

8°50’N

70°34’W

1450

<^)*

Barinitas [La Mina de San Isidro], Oct.

1988 (Barinas)

8°50’N

70°34’W

1450

116-JCSC

Barinitas [La Mina de San Isidro], Oct.

1988 (Barinas)

8°50’N

70°34’W

1450

Barinitas [La Mina de San Isidro],

Nov. 1995 (Barinas)

8°50’N

70°34’W

1400

San Isidro [Mina], 5 Jan. 2011

(Barinas)

8°50’N

70°34’W

1500

MCC

Qda. La Soledad, Via Barinitas - Sto.

Domingo. 5 Apr. 1985, Col. Mauro

Costa (Barinas)

8°49’N

70°31’W

1000

JCS

La Chimenea. 1-6 Jun. 1973, J.Salcedo

col. (Barinas)

8°49’N

70°3LW

1500

MIZA

Anzoategui, Qda. Guazo, 13-16

Jun. 1972, J. Salcedo. L. Zambrano

(Lara)

9°36’N

69°53’W

1440

MIZA

Anzoategui, Qda. Guazo, 13-16 Jun.

1972, J. Salcedo. L. Zambrano (Lara)

9°36’N

69°53’W

1440

MIZA

European Journal of Taxonomy 71: 1-66 (2014)

VENEZUELA: Eryphanis zolvizora isabe/ae Neild & De Sousa ssp. nov.

Sex

Data

Coord.

Alt. a.s.l.

Reference

Collection

(?■*

Via Choroni, Jun. 1970 (Aragua)

10°2EN

67°34’W

1450

111- JCSC

(?■*

Via Choroni, Oct. 1972 (Aragua)

10°22’N

67°35’W

1250

112- JCSC

( 7 -

Via Choroni, Sep. 1976 (Aragua)

10°22’N

67°35’W

1250

c7-

Via Choroni, Feb. 1978 (Aragua)

10°2EN

67°34’W

1450

(?■*

Choroni, Sep. 1971. C. flia. Romero

(Aragua)

10°2EN

67°34’W

1450

117- JCSC

MIZA

<^>*

Via Choroni, Aug. 1970 (Aragua)

10°2EN

67°34’W

1550

113-JCSC

Via Choroni, May 1979 (Aragua)

10°22’N

67°35’W

1400

(j)*

Via Choroni, cara norte, 28 Sep. 1976

(Aragua)

10°22’N

67°35’W

1150

114- JCSC

cp*

Rancho Grande, Aug. 1965, C.

Eduardo (Aragua)

10°20’N

67°40’W

1150

118- JCSC

MIZA

Colonia Tovar, Apr. 1993 (Aragua)

10°24’N

67°17’W

2100

Coloma Tovar, via Capachal, 2 Apr.

2010 (Aragua)

10°24’N

67°19’W

2100

MCC

cp*

La Llanada [Parque Nacional El

Avila], 29 Oct. 1989 (Distrito Federal)

10°33’N

66°56’W

1650

PBB 2322

MNHN

Eryphanis zolvizora isabelae Neild & De Sousa ssp. nov.

VENEZUELA: specimen without reliable locality

Sex

Data

Coord.

Alt.

a.s.l.

Reference

Collection

“Mexico”

BMNH[E]#808093

BMNH