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NOTA LEPIDOPTEROLOGICA

Volume 38 No. 1 • Sofia, 15.06.2015 • ISSN 0342-7536

Sylvain Cuvelier, Morten Molgaard. Pseudochazara amymone

(Lepidoptera, Nymphalidae) in Albania: Variability analysis,

androconial scales and new distributional data 1-22

Oleg E. Berlov, Ivan N. Bolotov. Record of Borearctia menetriesü

(Eversmann, 1846) (Lepidoptera, Erebidae, Arctiinae) larva on

Aconitum rubicundum Fischer (Ranunculaceae)

in Eastern Siberia 23-27

Martina Sasic, Milos Popovic, Sylvain Cuvelier, Milan Duric,

Filip Franeta, Martin Gascoigne-Pees, Toni Koren, Dirk Maes,

Branko Micevski, Nikola Micevski, Morten S. Molgaard,

Chris van Swaay, Irma Wynhoff, Rudi Verovnik. Contribution to

the knowledge of the butterfly fauna of Albania 29^15

Kari Nupponen, Matti Ahola, Marko Nieminen, Urmas Jürivete.

Biology and distribution of the declining moth Ethmia pyrausta

(Pallas, 1771), with description of the larva

(Gelechioidea, Depressariidae, Ethmiinae) 47-58

Enrique Garcia-Barros. Multivariate indices as estimates of dry body

weight for comparative study of body size in Lepidoptera 59-74

Matti Ahola, Martti Kuisma, Reima Leinonen, Hannu Saarenmaa,

Kimmo Silvonen. Description of immature stages of

Xestia brunneopicta (Matsumura, 1925), with a key to the mature

larvae of the European species of Xestia (Pachnobia)

(Lepidoptera, Noctuidae) 75-88

Thomas J. Simonsen, Ole Karsholt, Malcolm J. Scoble.

In Memoriam: Niels Peder Kristensen (1943-2014) 89-102

Raymond Guenin. Book Review: Die Widderchen des Iran

[The bumet moths of Iran]

-NlAfî

JUL 2 8 2015

103-105

/ /DDADltô

Nota Lepi. 38(1) 2015: 1-22 | DOI 10.3897/nl.38.9230

Pseudochazara amymone (Lepidoptera, Nymphalidae) in Albania:

Variability analysis, androconial scales and new distributional data

Sylvain Cuvelier1, Morten Molgaard2

1 Vlaamse Vereniging voor Entomologie, Werkgroep Dagvlinders, Diamantstraat 4, B-8900 leper, Belgium; Sylvain.

cuvelier@pandora. be

2 Gertrud Rasks Vej 86, DK-92 10 Aalborg S0, Denmark; msm2@stofanet.dk

http://zoobank.org/mElDBS-F263-4CDB-9CB5-C14C6CD2Fm

Received 26 January 2014; accepted 9 January 2015; published: 29 January 2015

Subject Editor: Zdenek Fric.

Abstract. For the first time a comparison of variable external characters of a series of males and females

of Pseudochazara amymone (Brown, 1976) from southern Albania is conducted. Pseudochazara amymone,

flying together with P. mniszechii tisiphone (Brown, 1980), was local and quite common in steep valleys on

ophiolite substrate on two separate mountains, one of which is a recently discovered locality by Eckweiler

(2012), while the other one is a new locality. An analysis of external characters of all specimens from the two

localities suggests no statistically significant differences. In the field, patrolling P. amymone males are easily

distinguished from P. mniszechii tisiphone males but this is not the case for females, and therefore we pro-

vide determination keys for males and females of these two species. These are based on a statistical analysis

of a specimen series from one Albanian P. mniszechii tisiphone population compared with all P. amymone

in this study. Photographs of androconia, copula and some extreme forms of P. amymone are presented. To

encourage further research in this poorly explored country a map is included, showing all historical records of

Papilionoidea from literature, including our own observations.

Samenvatting. Voor het eerst wordt een vergelijking gepubliceerd van de variabele uiterlijke kenmerken van

een reeks mannetjes en wijfjes Pseudochazara amymone (Brown, 1976) uit Zuid Albanie. Pseudochazara

amymone was lokaal en vrij algemeen in steile valleien op ophioliet gesteente in twee gescheiden gebergten

en vloog samen met P. mniszechii tisiphone (Brown, 1980). Aan de auteurs werd bevestigd dat het eerste

gebied de recent, door Eckweiler (2012) gevonden plaats is. Het tweede gebied is nieuw. Een analyse van de

uitwendige kenmerken van alle exemplaren uit de twee gebieden suggereert geen significante verschillen. In

het veld kunnen patrouillerende P. amymone mannetjes gemakkelijk onderscheiden worden van P. mniszechii

tisiphone mannetjes maar dit is niet het geval bij de wijfjes. Daarom zijn determinatiesleutels voor beide taxa

opgenomen (zowel voor mannetjes als wijfjes). Deze zijn gebaseerd op een statistische analyse van een Alba-

nese P. mniszechii tisiphone populatie met alle P. amymone in deze Studie. Foto’s van de androconia, copula

en sommige extreme vormen van P. amymone worden getoond. Om verder onderzoek aan te moedigen in dit

zwak onderzocht land is een landkaart opgenomen die alle historische Papilionoidea gegevens inclusief onze

eigen observaties weergeeft.

Resumé. For forste gang gennemfores en sammenligning af de varierende eksteme kendetegn pâ en sérié

hanner og hunner af Pseudochazara amymone (Brown, 1976) fra det sydlige Albanien. Pseudochazara amy-

mone forekom sammen med P. mniszechii tisiphone (Brown, 1980) lokalt, men ret almindeligt, i stejle dale

med ofiolitiske mineraler i to adskilte bjergomrâder, hvoraf det ene er en nyligt opdaget lokalitet af Eckweiler

2

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

(2012), og det andet er en ny lokalitet. En analyse af aile eksemplarers eksteme kendetegn fra de to omrâder

viser ingen statistisk signifikante forskelle. I feiten kan patruljerende P. amymone hanner let adskilles fra P.

mniszechii tisiphone hanner, men dette er ikke tilfældet med hunneme, og derfor gives bestemmelsesnogler

til hanner og hunner af disse to arter. Disse er baseret pâ en statistisk analyse af en sérié eksemplarer fra en

albansk P. mniszechii tisiphone population og alle P. amymone. Fotos af duftskæl, pairing og visse ekstreme

former af P. amymone præsenteres. For at tilskynde andre til at foretage videre undersogelser i dette sâ dârligt

udforskede land inkluderes et udbredelseskort over alle hidtidige fund af Papilionoidea nævnt i litteraturen,

inklusive vore egne fimd.

Introduction

Brown’s Grayling, Pseudochazara amymone (Brown, 1976) was discovered by John Brown in

early July 1975 in NW Greece (type locality: “mountains just north of Ioannina”) based on four

males (Brown 1976) and years later a single female (oral communication). Since then, despite

many efforts, almost all searches for this butterfly in Greece have turned out negative and no other

voucher specimens are available. A lot of strange rumours, describing a rocket speed flight and

strange nuptial behaviour, often followed these negative searches (Cuvelier 2010). Since its dis-

covery, P. amymone has been the subject of speculation like hardly any other butterfly species in

Europe and a myth has been created around it. Its taxonomic status is still uncertain. Because of the

close resemblance in the genitalia of Albanian amymone with Turkish Pseudochazara mamurra

(Herrich-Schaffer, [1846]), Eckweiler (2012) treated amymone as a subspecies of P. mamurra.

DNA analysis might shed additional insights on the taxonomic position of the taxon amymone and

is underway by an independent group (oral comm. Verovnik). In this article we chose to follow

Fauna Europaea (de Jong 2013), which gives this taxon species status.

Albania is a country that, due to its political (50 years of communist regime and civil war) and

infrastructural situation, only recently became open for travelling and lepidopterological investiga-

tion. Large parts of the country remain unexplored. It is not surprising that P. amymone remained

undiscovered in Albania until recently. Here the faunistical elements of central Europe meet with

those from the Balkans, the Mediterranean and Asia Minor.

Eckweiler had the idea to start searching for P. amymone in Albania in 2010. In July 2010 he

discovered a first single P. amymone male and, in July 2011, five further males and one female in

southern Albania. Before the publication of his observations, this discovery was again surrounded

by mysterious communications. But at least this time strong proof of its existence, supported by the

photographs of voucher specimens, was soon given (Eckweiler 2012). A short message on Facebook

from van Swaay during the summer of 20 1 2 also mentioning a few P. amymone from Albania was

the only other evidence known to us at that time. This message included a photograph of a male

Pseudochazara , sitting with closed wings that looked quite different from Pseudochazara mniszechii

tisiphone (Brown, 1980). But even with both sources, the locality remained obscure and the given

information again supported the extreme rarity of the butterfly. The article (Eckweiler 2012) made

some suggestions concerning a potentially wider distribution area in Albania than the single undis-

closed locality where the P. amymone had been found and the need for further surveys. Based on

all of this, a joint trip to Albania was planned by the authors with the objective to search for further

evidence and to study the biology of P. amymone in the country.

Nota Lepi. 38(1): 1-22

3

As this taxon is often associated with Turkish Pseudochazara mamurra (Herrich-Schäffer,

1852) (Gross 1978; Tolman and Lewington 1997; Eckweiler 2004; Tshikolovets 201 1; Eckweiler

2012), we used Google Earth to search for potential localities of P. amymone similar to Turkish

habitat photographs (Hesselbarth et al. 1995). With the available good resolution of satellite

photographs it is possible to recognize the colour of the geological substrates together with a lot

of topographical details. Such places seemed quite common in Albania and were far too numerous

for a dedicated search. Pamperis (1997) also mentioned that in one Greek locality P. amymone

is sympatric with P. mniszechii tisiphone , but flying at the end of the flight time of P. mniszechii

tisiphone. This suggests a geology of dark red soil as in typical Pseudochazara biotopes. The

habitat photograph from the Eckweiler paper (2012) also was suggestive of such soils and the

pink flowers were an interesting clue for future field research. A geological map from Monjoie et

al. (2008) helped us focus our research strictly on the south-eastern Albanian province of Korçë.

On Google Earth these areas, mentioned as ophiolite nappes in the maps, looked a lot like some

Turkish biotopes with steep, dark red, dry slopes in river valleys. Combined with the information

about altitude, this enabled us to be very selective concerning target areas.

Our second objective was to explore areas in Albania that had never been explored before for

butterflies. Before this field trip, we searched for all historical data from the sparse literature about

Albanian butterflies. Maps of species and a global distribution map clearly showed how poor is

the coverage for this country. We also intended to survey other areas in the provinces of Korçë,

Kolonjë, Përmet, Tepelenë and Skrapar to increase the knowledge of the butterfly fauna from

Albania in general. The detailed results of all our own observations will be published in a future

faunistic publication.

Abbreviations

AL: androconium length; AB: androconium breadth; A: ratio AL/AB; FW: forewing; HW: hindwing; MM: Morten Schnei-

der Molgaard; N°: number; N/A: not applicable; oc.: ocelli; SC: Sylvain Cuvelier; SD: standard deviation; subm.: submar-

ginal; UNS: underside; UPS: upperside; Var: variable.

Material and methods

Sample collecting and database construction

In two localities in the Albanian province Korçë (Boboshtiçë and Gjergjeviçë) males and females

of P. amymone and P. mniszechii tisiphone were netted by both authors. A search for all potential

references on the butterflies of Albania was made during the preparation for the trip. The rele-

vant publications were gathered from different sources in order to build a database including as

much historical data as possible (Abadjiev and Beshkov 1996a, b; Alberti 1965; Beshkov 1994;

Beshkov 1995; Beshkov and Misja 1995; Gaskin 1990; Misja and Kurrizi 1984; Moucha 1963a,

b; Murraj 1972; Plöciennik et al. 2009; Popescu-Gorj 1971; Rebel 1913; Rebel 1918; Rebel and

Zerny 1931; Verovnik and Popovic 2013a, b). Only the data from all species with rather precise

indications of the locality were included in an Excel spreadsheet with coordinates in decimal

degrees that were defined with Google Earth and an online coordinate conversion tool (Montana

State University 2014). All data from our personal observations were included in this database.

During our surveys, coordinates were obtained in the field with a GPS (Garmin Etrex Legend). A

4

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

map of Albania was adapted for use with DMAP distribution mapping software to produce dis-

tribution maps per species and one global coverage map of all Albanian butterfly species. During

history the borders of Albania have changed. Some historical data now in fact concern localities

that are situated in Montenegro, Kosovo and Macedonia. To be as complete as possible, we have

maintained these observations in the coverage map.

Study of external characters

Since the discovery of P. amymone by Brown (1976), as far as is known to the present authors, there

have been only 10 male and two female voucher specimens collected and included in publications.

Pseudochazara species are very variable and difficult to identify. No comparative studies on the

external characters of a good number of P. amymone specimens have been published so far and we

had no precise idea about the variability of the external characters of both sexes.

The male holotype, figured in black and white, was for a long time the only documented picture

of this species (Brown 1976). In his article, Brown described the external characters based on a

very small series of four males: “ Upperside similar to graeca but wings more rounded and with no-

tably broad clear orange postdiscal bands more or less broken by grey-brown ground colour along

v4 of forewing and enclosing blind black oc. in S 2, 5 and minute black oc. in S 3, 4 on forewing.

Sex brand inconspicuous. Hindwing sometimes with small black ocellus in S 2 and dark grey sub-

marginal line broken by orange along veins. Marginal grey band thin (1-2 mm wide). Underside

ground colour pale yellow-grey but variable. Hindwing irrorate with darker scales and indistinct

striae. Forewing length 26-27 mm. Female. Unknown.”

Luckily, the two original photographs from this publication were available and they allow a

better comparison. For this purpose, Jos Dils (Belgium) kindly provided the two photographs of the

male holotype (Brown 1976) (Fig. 1). On the underside of the photograph it is written: “WATSONI

? Clench & CHS SCH” looking like a link to a Pseudochazara species from Afghanistan (Fig. 1).

However, as we never received a reply from Brown, it is not possible to fully understand this detail.

Recently, two prepared male specimens were figured by Eckweiler (2012) that look different

from the holotype but with such a small sample size it is difficult to estimate if this is within the

normal range of variability. Concerning females, for a long time there was only one figure (Tol-

man and Lewington 1 997) but recently a first photograph of the upper- and underside of a single

female from Albania became available (Eckweiler 2012). Even fewer photographs of the butterfly

in nature have been published and for all these documents there remained a degree of uncertainty

concerning the final determination (Cuvelier 2010, Eckweiler 2012).

In the field, fresh males of P. amymone look quite different when flying than P. mniszechii

tisiphone and identification is possible in a fair number of cases. As Pseudochazara species almost

never sit with open wings, reliable identification in the field is often based on the underside of the

wings and for both sexes it is difficult and depends on the freshness of the butterflies. Therefore we

also sampled P. mniszechii tisiphone (Figs 5a-h) at Boboshtiçë, where this butterfly was extremely

common, in order to compare the two taxa and to obtain determination keys for males and females

of both species.

Potential variables of the external characters were selected and included in an Excel workbook

containing separate worksheets per species and gender. After this first selection, the colour of

the fringes was discarded as a variable due to the difficulty of formulating measurable criteria.

Nota Lepi. 38(1): 1-22

5

iiimmmrimmmmutlllilllllll HlnnTïlïl HrmrmmtrriTmilTiUTnnT^^

Figure 1. Upper and underside of the P. amymone holotype photographs, holotype collected in the mountains just

N. of Ioânnina, Epiros, Greece, 650m, 1 0.vii. 1975 (photograph: SC).

Drawings of measurement on UPS and UNS are included in Appendix 1. The following variables

were analyzed: UPS FW: Var 1: length of FW from apex to point of attachment to thorax, fringes

included (mm); Var 2: visual assessment of the oc. in S2 and S5 (blind= 0, white pupil= 1); Var 3:

visual assessment of the number of spots in S3 and S4 (0, 1 or 2); Var 4: width of the submarginal

band across the centre of the ocellus in S2 (mm); Var 5: Var 4/Var 1 (%); Var 6: visual assessment

of the conspicuous sex brand in cell (absent= 0, present= 1). UPS HW: Var 7: visual assessment

of the number of oc. in the submarginal area (0, 1,2 or 3); Var 8: width of the submarginal band

along vein 3 (mm); Var 9: Var 8/Var 1 (%). UNS FW: Var 10: visual assessment of the oc. in S2

and S5 (blind= 0, white pupil =1); Var 1 1 : visual assessment of the pale area from ocellus in S5 to

the cell (uniform= 0, contrasted = 1); Var 12: visual assessment of the number of spots in S3 and

S4 (0, 1 or 2); Var 13: visual assessment of the marginal line (diffuse= 0, sharp= 1); Var 14: visual

assessment of the fine black line-shape markings in the basal area of the cell (absent= 0, present^

1); Var 15: length of the ocellus in S5 (mm); Var 16: shortest distance from the white centre of the

ocellus in S5 to the margin (mm); Var 17: Var 15/Var 1 (%); Var 18: Var 15/Var 16 (%); Var 19:

Var 16/Var 1 (%). UNS HW: Var 20: visual assessment of the number of oc. in the submarginal

area (0, 1,2 or 3); Var 21 : visual assessment of the median band (absent^ 0, present^ 1);

6

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

We photographed the upper- and underside of all male and female specimens of P. amymone and P.

mniszechii , each of us in our personal reference collections. A scale bar was included with each spec-

imen (Appendix lb). Each digital image was imported into Paint Shop Pro v. 6.02. A straight vector

line was drawn on the butterfly in its own layer to measure the exact length of the desired parameter.

Afterwards, the vector line was rotated into horizontal position and then moved onto the scale bar

under the butterfly, making it possible to measure length in millimetres, at an accuracy level of 0.25

mm. The whole dataset was used for two analyses: a) a variability study between the two P. amymone

populations and b) a comparison between all P. amymone and P. mniszechii tisiphone. Statistical anal-

ysis was performed with StatSoft STATISTICA 12. The Mann- Whitney U test was used to test for

differences at 0.05 significance level in a two-tailed test.

Study of androconial scales

Androconial scales were removed from the upperside of the forewings of one P. amymone and one

P. mniszechii tisiphone and photographed with a calibrated 5 megapixel Dino-Lite AD-70 13MZT

digital microscope with adjustable magnifications. The length and breadth of the androconia were

measured according to the description by Wakeham-Dawson (2000) but at maximum magnifica-

tion (x500).

Cartography

DMAP, distribution mapping software: http://www.dmap.co.uk/. — Dr. Alan Morton, Blackthorn

Cottage, Chawridge Lane, Winkfield, Windsor, Berkshire, SL4 4QR, UK.

Results

Field notes

On 15.vii.20 13, late in the afternoon, we started our search for P. amymone in a narrow valley with

very steep slopes near Boboshtiçë (province of Korçë). On the dark red-grey slopes our attention

was attracted by cushions of pink flowers that we identified as Acantholimon echinus (L.) Boiss.

(Plumbaginaceae) and the whole area looked very similar to the published habitat photograph

(Eckweiler 2012). The rest of the day we explored this area and found some P. mniszechii tisi-

phone, but we had not had a glimpse of P. amymone. We were, however, quite certain that we had

to be near its habitat.

The next morning, 16.vii.2013, we entered deeper into the river valley and searched at 1 1 00—

1200 m altitude on SSW exposed steep rocky slopes with parts of loose gravel. On this ophiolite

substrate (Fig. 2a) with characteristic red-grey colour, scattered tall grasses were growing, but

otherwise the area was almost devoid of vegetation (Fig. 2b). Here P. amymone had just emerged

and males (Fig. 2d) were already flying quite commonly. We observed the males showing a typical

territorial behaviour: patrolling and chasing away other males. The females were searching for

nectar sources and egg-laying places in the scattered tall grasses that were present in the biotope.

The rumours of a rocket speed flight and strange nuptial behaviour appeared not to be true. The

species is not shyer than other species in the genus Pseudochazara.

Nota Lepi. 38(1): 1-22

7

Figure 2. a-b. Habitat of P. amymone , Boboshtiçë, Albania, 16.vii.2013. c. Habitat of P. amymone, Gjerg-

jeviçë, Albania, 18.vii.2013. d. S P ■ amymone, Boboshtiçë, Albania, 16.vii.2013 (photographs: MM), e. S

P. amymone caught by a crab spider, Boboshtiçë, Albania, 18.vii.20 13. f. T. onustus holding a $ P. amy-

mone, Boboshtiçë, Albania, 18.vii.2013 (photographs: SC), g. Copula of P. amymone, Boboshtiçë, Albania,

16.vii.2013 (coll. & photograph: SC), h. Copula of P. mniszechii tisiphone, Gjergjeviçë, Albania, 18.vii.2013

(photograph: MM).

8

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

On 18.vii.20 13, early morning, we again visited this locality and observed that some males were al-

ready getting worn. One male P. amymone (Fig. 2e), seen from afar sitting with open wings on flowers

of A. echinus , had been caught by a crab spider (Fig. 2f) that was identified by Rop Bosmans (Belgium)

as Thomisus onustus (Walckenaer, 1805), a common species in the Balkans. This butterfly also shows

typical injuries on the hind wings caused by lizards that were common in the habitat. Females were

already more numerous than two days earlier. Here, P. amymone flies sympatrically with P. mniszechii

tisiphone. The males of these two species are easily distinguished in the field, but this is not the case

for females. Flying males of P. amymone look smaller and also show a much more pronounced orange

and black contrast. Although the females of P. amymone are a little smaller, this cannot be observed in

the field. They also do not exhibit the contrasting orange and black colours and this makes it difficult in

the field to distinguish females of these two species.

In Boboshtiçë, we observed the first ever known copula (Fig. 2g) and a pale form (Figs 3g-h) of

a male P. amymone. After our trip, Eckweiler confirmed (oral comm., August 2013) that this was

the locality where he originally discovered P. amymone in Albania.

During these explorations, on 18.vii.2013, we were able to extend the known distribution of P.

amymone by approximately 25 km to the west, as the crow flies, as we discovered it in another

remote mountain range in the westernmost part of the province of Korçë. This mountain range is

physically separated from Boboshtiçë by a 10 km broad river valley, at 850 m altitude.

Entering a remote valley, we observed steep rocky slopes orientated to the SSW on ophiolite

substrate, as we had seen in Boboshtiçë. On climbing these slopes we immediately observed a

large population of P. amymone. The new biotope is situated near Gjergjeviçë (Fig. 2c) and in

the upper part of the known altitudinal distribution (Pamperis 2009) of P. amymone , being at an

altitude of 1200-1400 m. It has the same characteristics as the biotope at Boboshtiçë, except for

one major difference: the presence of scattered bushy vegetation whereas the habitat at Boboshtiçë

is completely open. We observed that P. amymone near Gjergjeviçë is also sympatric with and

even outnumbering P. mniszechii tisiphone and here a copula of this species was photographed

(Fig. 2h). The flight period of P. amymone was apparently the same as in Boboshtiçë, due to

the general freshness of most of the specimens and the presence of good numbers of females as

observed in the morning of the same day at Boboshtiçë.

Variability of P. amymone from two Albanian populations

To our big surprise, we noticed P. amymone was not rare at all in the two biotopes (Boboshtiçë and

Gjergjeviçë) and that the habitat in both localities was very large but difficult, if not impossible,

to explore. Nevertheless we sampled enough voucher specimens to get a better idea of the range

of variation in external characters and to make a comparative study on the habitus of these two

separate populations of P. amymone.

There are two limitations of our dataset which require further attention in the future when new

populations are discovered. The sample size of 38 males and 19 females for such variable butter-

flies remains suboptimal and the sample size of the two localities is not equal. The range and mean

of all variables for the two populations is given in Table 1. Males and females are figured (Figs

3a-h, 4a-f). All the variables of the two populations clearly overlap and there was not a single

variable showing clear differences between the two populations. For these reasons the measure-

Nota Lepi. 38(1): 1-22

9

Figure 3. Variability in P. amymone. a-b. S typical upper- and underside, Boboshtiçë, Albania, 16.vii.2013

(coll. & photograph: SC), c-d. $ typical upper- and underside, Gjergjeviçë, Albania, 18.vii.2013 (coll. & pho-

tograph: MM), e-f. S aberration, upper- and underside, Boboshtiçë, Albania, 16.vii.2013 (coll. & photograph:

10

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

Figure 4. Variability in P. amymone. a-b. $ typical P. amymone upper- and underside, Gjergjeviçë, Albania,

18.vii.2013. c-d. $ typical P. amymone upper- and underside, Boboshtiçë, Albania, 16.vii.2013. e-f. $ dark

form upper- and underside, Gjergjeviçë, Albania, 18.vii.2013. (Coll. & photographs: SC).

ments from the two populations can be pooled together for the comparison between Albanian P.

amymone and P. mniszechii tisiphone.

Two P. amymone specimens exhibited asymmetry between right and left side. This was the case

for one male with one spot in S3-S4 on the left FW UPS and no marking on the right FW. One

female had one black ocellus on the left side of the HW UNS and two on the right side.

There seem to be a few marked differences between P. amymone from Albania and the ori-

ginal description by Brown (1976). It is clear that the holotype (Fig. 1) is not fresh and probably

the paratypes were even more worn. A few butterflies in our study that were less fresh also have

a clearer appearance and tend to become more orange in the postdiscal bands of the FW UPS. We

Nota Lepi. 38(1): 1-22

11

Figure 5. Variability in P. mniszechii tisiphone. a-b. Typical S upper- and underside, Boboshtiçë, Albania,

16.vii.20 13. c-d. $ typical upper- and underside, Boboshtiçë, Albania, 16.vii.2013. e-f. $ typical upper- and

underside, Boboshtiçë, Albania, 16.vii.2013. g-h. $ dark form upper- and underside, Boboshtiçë, Albania,

18.vii.2013. (Coll. & photographs: SC).

12

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

Table 1. Measurements of P. amymone from Boboshtiçë versus P. amymone from Gjergjeviçë.

Nota Lepi. 38(1): 1-22

13

have clearly shown that many characteristics are much more variable within a single population

than described by Brown (1976). We presume, until more material becomes available, that Greek

P. amymone falls within the given range of variability.

Determination keys between P. amymone and P. mniszechii tisiphone

The maximal range and mean of all variables for the two species are shown in Table 2. An over-

view of the statistical significance status of all variables is shown in Table 3. Some variables of P.

amymone and P. mniszechii tisiphone did not overlap at all. Some differences are present in both

sexes, others are only present in males. Below we list the variables that can be used in distinguish-

ing P. amymone and P. mniszechii tisiphone.

Var 13: UNS FW submarginal line in both sexes (Figs 6a-b). This line is always sharp in P. amy-

mone and diffuse in P. mniszechii tisiphone.

Var 14: UNS FW basal area of the cell in both sexes (Figs 6c-d). There are always black linear

markings inside this area in P. amymone which are absent in P. mniszechii tisiphone.

Var 6: UPS FW sex brand position in males (Figs 6e-f). P. amymone has a black sex brand over the

whole cell and the androconial field is extending to the inner margin of the FW. P. mniszechii

tisiphone does not have a sex brand in the cell and the androconial field is covering only half

of the cell towards the inner margin of the FW.

Var 7: UPS HW number of oc. in males (Figs 6g-h). There is 1 ocellus in P. amymone and 2 oc. in

P. mniszechii tisiphone. Females of the two species have both a range of 1 to 2 oc.

The wingspan of the females was in many cases useful in the field for the identification but one

P. mniszechii tisiphone female falls in the upper range of P. amymone.

14

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

Table 2. Measurements of pooled data of P. amymone versus P. mniszechii tisiphone.

Nota Lepi. 38(1): 1-22

15

Table 3. Overview of the statistical significance of Mann- Whitney U tests of all variables of P amymone

versus P. mniszechii tisiphone. Males DF = 51, females DF = 37. P values below 0.05 are shown in bold.

Androconial scales of P. amymone and P. mniszechii tisiphone

The dense sex band in the FW cell makes it difficult to isolate the androconial scales (Fig. 7a) of P.

amymone. Scales with quite different shapes, sometimes bright silver-grey (Fig. 7b), were found in

this area and created confusion. The type of these scales is unclear to the authors.

16

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

Figure 6. Visualization of the significant difference between P. amymone (a, c, e, g) and P. mniszechii

tisiphone (b, d, f, h). Var 13 (a-b), Var 14 (c-d), Var 6 (e-f), Var 7 (g-h).

According to the criteria of Gross, the androconial scales of P. amymone (Fig. 7c) are in general

of type 7 (Gross 1978) but few were found where the diameter of the lamina decreased immediately

from the basal stalk tasowards the apex. These are more closely resembling a type 6 androconial

scale. AL= 0.42 mm, AB= 0.05 mm and A= 8.4. These data are near the values for P. mamurra in

Wakeham-Dawson and Kudma (2000). Also the shape of the scales from P. amymone falls within

the ranges of P. mamurra (Wakeham-Dawson and Kudma 2000; Wakeham-Dawson and Kudma

2005). The androconia of P. amymone are much larger and of a different shape than P. graeca (Wake-

ham-Dawson, 2000). The androconial scales oTP. mniszechii tisiphone (Fig. 7d) are different in shape,

transitional type 5-6 with dimensions: AL= 0.33 mm, AB= 0.027 mm and A= 12.19.

Coverage of historical data and personal observations

Additional field work in other localities of the province of Korçë and in the provinces Kolonjë,

Përmet, Tepelenë and Skrapar fill an important gap in the documented distribution of the butterflies

from south-eastern Albania. The coverage map (Fig. 8) shows historical data (red dots), the two

areas where P. amymone was found (blue letters B and G) and all new places that were surveyed

by the authors (green dots).

Discussion and conclusion

The discovery of P. amymone on a new isolated mountain and the fact that large parts of Albania

with similar geological origins still remain unexplored suggest that P. amymone might be more

widely distributed than previously thought. With further field research on slopes in steep river

valleys, not necessarily with ophiolites, the species will undoubtedly be discovered at new sites.

The butterfly probably has a more restricted and fragmented distribution in Greece due to the

less frequently present favoured type of geological substrate. This idea is already supported by the

almost complete lack of evidence despite the many efforts of numerous lepidopterists since the

discovery of the butterfly by Brown (1976). Using geological data, we suggest that the research in

Nota Lepi. 38(1): 1-22

17

Figure 7. a. UPS FW sex brand of <$ P. amymone (x 250), Boboshtiçë, Albania, 16.vii.2013. b. UPS FW

scales in sex brand of S P ■ amymone (x 500), Albania, 16.vii.2013. c. Androconial scale of S P ■ amymone

(x 500), Boboshtiçë, Albania, 16.vii.2013. d. Androconial scale of S P ■ mniszechii tisiphone (x 500), Bobo-

shtiçë, Albania, 16.vii.2013. (Coll. & photographs: SC).

Greece should be extended because areas with ophiolite substrate are present over a wider part of

continental Greece, whereas they are hardly present near Ioannina. Understanding more fully the

habitat requirements of P. amymone , it now looks possible to elucidate the mystery of its wherea-

bouts in Greece in order to undertake any conservation measures if needed.

During the examination of the Albanian material it became clear that P. amymone is a very

variable species and that the original description (Brown 1976) did not cover the whole range of

variability, partly because of the very restricted number of studied butterfly vouchers. Pamperis

(1997: 348-349, fig. 3.7.10; 2009: 499, fig. 3.5.1 1) shows figures with characteristic marks on the

underside of P. amymone in comparison with other Greek species of Pseudochazara. The black

and white figure of P. amymone focuses on a black base of the FIW UNS and a pale dentate line in

the postdiscal area with more contrast than in other Pseudochazara species. This black base is not

a striking feature of the Albanian specimens. Darker grey scales in that area are sometimes visible

in both P. amymone and P. mniszechii tisiphone. The dentate line was visually assessed as Var.

2 1 showing overlap and for both species this variable was sometimes scored as absent. In external

18

Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

Figure 8. Coverage of historical data and personal observations: map of Albania indicating Papilionoidea

observations from literature (•) and from observations by the authors (®). B: Boboshtiçë; G: Gjergjeviçë:

localities of P. amymone.

Nota Lepi. 38(1): 1-22

19

features, Pamperis (1997: 351; 2009: 500) in analogy with Brown (1976), focuses on the orange

brown band in the postdiscal area.

The only feature for Greek P. amymone that seems different is the presence of the broad and

clear orange postdiscal bands. Only for older butterflies from Boboshtiçë and Gjergjeviçë is there a

tendency to paler orange postdiscal bands. This potential difference should be documented by stud-

ying material from new localities including additional material in this dataset to increase the sample

size. Adding material from Greece to get an idea about the full range of the external characters of

this taxon seems mandatory. A few other criteria seem specific to P. amymone in comparison with

P. mniszechii tis iphone. It became clear that P. mniszechii tis iphone is a very variable species too.

Despite clear differences in the androconial scales, it would be interesting to make an analysis

of external characters with the very similar P. graeca, a species that has never been found in the

same locality with P. amymone.

The androconial scale of P. amymone falls within the range of the different subspecies of P. ma-

murra and this result supports the treatment by Eckweiler of P. amymone as a potential subspecies

of P. mamurra. Even though androconia have been used as a taxonomic character for distinguish-

ing species of the genus Pseudochazara, no comment is given here as independent DNA analysis

is ongoing and will be published soon.

We encourage entomologists to visit Albania during different periods of the year to do research

not only for P. amymone but also to survey large parts of the country that are poorly explored for

butterflies. It will certainly help to significantly improve knowledge about the distribution of many

taxa in the south-western Balkans.

Acknowledgements

We are grateful to Rudy Swennen for his valuable help with geological data of Albania. We also thank Jos

Dils for providing the photographs of the P. amymone holotype, Rop Bosnians for the determination of the

crab spider and Wolfgang Eckweiler for his confirmation of the locality where he discovered P. amymone.

We are indebted to Willy De Prins, Roger Vila and Enrique Garcia-Barros for suggestions about the study of

androconial scales and to Luc Merveillie for his help on the cartography. We express our gratitude to Vlad

Dinca and Leonardo Dapporto for fruitful discussions about the final draft before submission. We thank the

reviewers for their comments that helped to fine-tune the submitted draft.

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Sylvain Cuvelier & Morten Molgaard: Pseudochazara amymone...

Appendix 1

Variables of P. amymone and P. mniszechii tisiphone. a. UPS variables of S P ■ amymone , Boboshtiçë, Al-

bania, 16.vii.20 13. b. UPS variables of <$ P. mniszechii tisiphone , Boboshtiçë, Albania, 16.vii.2013. c. UNS

variables of S P • mniszechii tisiphone , Boboshtiçë, Albania, 16.vii.2013. d. UPS variables of $ P. mniszechii

tisiphone, Boboshtiçë, Albania, 16.vii.2013. e. Sex brand of S P ■ amymone (x30), Boboshtiçë, Albania,

16.vii.2013. f. UNS pale area from ocellus in S5 towards cell, P. amymone (x32), Boboshtiçë, Albania,

16.vii.20 13. (Coll. & photographs: SC).

Nota Lepi. 38(1) 2015: 23-27 | DPI 10.3897/nl.38.8664

Record of Borearctia menetriesii (Eversmann, 1846) (Lepidoptera,

Erebidae, Arctiinae) larva on Aconitum rubicundum Fischer

(Ranunculaceae) in Eastern Siberia

Oleg E. Berlov1, Ivan N. Bolotov2

1 The State Nature Reserve «Baikalo-Lensky», Ministry of Natural Resources and Environment of the Russian Federation,

Baikalskaya 291 -B, 664050 Irkutsk, Russian Federation; olegberlov@narod.ru

2 Institute of Ecological Problems of the North, the Ural Branch of the Russian Academy of Sciences, Severnoy Dviny Emb.,

23, 163000 Arkhangelsk, Russian Federation; inepras@yandex.ru

http://zoobank.org/865C86A0-7354-42C4-9AFD-26BAA37A3E6A

Received 28 September 2014; accepted 28 November 2014; published: 30 January 2015

Subject Editors: Jadranka Rota.

Abstract. In this note we report the first record of Borearctia menetriesii (Eversmann, 1846) (Erebidae: Arc-

tiinae) larva on a native host plant, Aconitum rubicundum Fischer (Ranunculaceae). This aconite species is a

close relative of A. lycoctonum, which is widespread across Eurasia, but has a scattered distribution in Fen-

noscandia. The majority of B. menetriesii localities are situated within the distribution range of A. lycoctonum

and other aconite taxa, which are diverse and widespread in the Eastern Palaearctic. However, only two of the

six westernmost B. menetriesii localities in Finland are in accordance with sporadic records of A. lycoctonum.

Our record confirms that B. menetriesii is a polyphagous species like most other boreal Arctiinae. We have

expanded the list of a few Lepidoptera species which can use Aconitum spp. as suitable host plants despite the

fact that they are poisonous for insects because of high alkaloid content.

Introduction

The Menetries’s tiger moth Borearctia menetriesii (Eversmann, 1846) (Erebidae: Arctiinae) is the

most enigmatic representative among the Palaearctic arctiine moths. The biology of this large and

colorful species is poorly known because of its extremely low abundance throughout its distribu-

tion range (Lappi et al. 2004; Dubatolov 2010; Bolotov et al. 2013). Only single specimens were

found in the majority of known localities, and sometimes the records are separated from each other

by many decades (Bolotov et al. 2013).

Krogerus (1944) experimentally identified three available host plants in Finland, including

Taraxacum spp. (Asteraceae), Plantago ssp. (Plantaginaceae) and Polygonum ssp. (Polygonaceae).

In a preliminary report on the food preference of the larvae, Saarenmaa (2014) lists 15 plant

species which B. menetriesii larvae preferred or accepted during experiments, including Larix spp.

(Pinaceae), Rubus chamaemorus L., R. idaeus L., R. saxatilis L. and Potentilla palustris (L.) Scop.

(Rosaceae), Menyanthes trifoliata L. (Menyanthaceae), Rumex crispus L., Polygonum persicaria

L. and P. lapathifolium L. (Polygonaceae), Plantago major L. (Plantaginaceae), Rib es rubrum L.

(Grossulariaceae), Salix phylicifolia L. (Salicaceae), Taraxacum officinale Weber (Asteraceae),

Vaccinium uliginosum L. (Ericaceae) and Viola riviniana Rchb. (Violaceae). He noted that the larch

species might be a significant food plant over the majority of the B. menetriesii range. However, all

those data are based exclusively on these laboratory experiments. There is the unique observation

24

Berlov & Bolotov: Record of Borearctia menetriesii (Eversmann, 1846) ...

Figure 1. The last instar larva of B. menetriesii on Aconitum rubicundum Fischer, Baikalo-Lensky Nature

Reserve, 9.viii.2013 (photo: O. E. Berlov).

in natural habitat in Finland in June 1920 of a larva having climbed a spruce trunk (Krogerus 1944).

Here we report the first record of a feeding larva on a native host plant in the Baikalo-Lensky State

Nature Reserve, Eastern Siberia.

Observations

Locality: Eastern Siberia, the Baikal Lake Area, the Bolshoy Anay River terrace, 53056’19”N,

107°24’35”E, ca 770 m alt., mixed coniferous taiga forest with \\Qxb-Equisetum-moss plant cover

(locality description and photo: Suppl, material 1: Table SI, Fig. SI). A last instar larva of B. me-

netriesii was collected alive on Aconitum rubicundum Fischer (Ranunculaceae) 9.viii.2013 (Figs

1-3) and was placed in a cage that was taken to the Irkutsk city. In captivity, the larva had a daytime

feeding activity and consumed only fresh A. rubicundum leaves which we had collected from the

same locality as the larva. The leaves were completely eaten by 15.viii.2013. Unfortunately, we

could not find any aconite species in the city surroundings. The larva did not accept Taraxacum spp.

and Plantago spp. leaves which we placed in the cage and it was found dead on 22.viii.20 13. An

additional larva was captured dead in a pitfall trap at the same locality on 10.viii.2013. The collected

larvae were 32-35 mm long.

Discussion

The observed host plant, A. rubicundum , is distributed in Central and Eastern Siberia, and is closely

related to the widespread Eurasian A. lycoctonum (Malyschev and Pechkova 1993) and might even

represent its eastern subspecies (Ivanova 1978). These two species (or subspecies) were separated

Nota Lepi. 38(1)2015: 23-27

25

Figures 2-3. Aconitum rubicundum Fischer, the host plant of B. menetriesii , Baikalo-Lensky Nature Reserve. 2.

An inflorescence, upstream of the Pravaya Kirenga River, 14.vii.2006 (photo: N.V. Stepantsova). 3. A leaf at the

B. menetriesii locality, 9.viii.2013 (photo: O. E. Berlov).

on minor diagnostic features, particularly the location and density of hairs on the stem and leaf

blade; both have identical chromosome number (2n = 16) (Malyschev and Pechkova 1993). All

Russian B. menetriesii localities are situated within the distribution range of A. lycoctonum and

other aconite taxa, which are especially diverse and widespread in the Eastern Palaearctic, including

26 species in Siberia and 37 species in the Russian Far East (Jalas and Suominen 1989; Malyschev

and Pechkova 1993; Kharkevich 1995; Bolotov et al. 2013). In boreal Russia, various aconite spe-

cies are abundant in the plant cover of river valleys and humid alpine meadows (Peshkova 1985;

Malyschev and Pechkova 1993; Kharkevich 1995) where B. menetriesii most frequently occurs

(Bolotov et al. 2013). For example, A. lycoctonum is one of the dominant plant species in the B.

menetriesii habitat in the Sotka River Valley, Arkhangelsk Region (Bolotov et al. 2013). However,

A. lycoctonum has a scattered distribution in Finland (Jalas and Suominen 1989; Lampinen et al.

2014), and only two of the six Finnish B. menetriesii localities are near sites where this plant species

was recorded (Suppl, material 1: Fig. 2S).

Aconite species have a strong insecticidal activity (Yuan et al. 2012) because of their high

alkaloid content (Azimova and Yunusov 2013). Eighteen alkaloids were isolated from A. lycoc-

tonum (Azimova and Yunusov 2013). A. rubicundum contains at least nine diterpenoid alkaloids

(Nishanov et al. 1991).

The HOSTS database (Robinson et al. 2010) listed only 16 Lepidoptera species feeding on

Aconitum spp. The majority of these species are polyphagous (12 of them), including Euproctis

similis (Fuessly, 1775), a unique Erebidae representative. According to other sources (Vorbrodt

and Müller-Rutz 1914; Freina and Witt 1987; Bellmann 2003), there are two Arctiinae species

recorded on Aconitum spp., Arctia flavia (Fuessly, 1779) on A. lycoctonum ssp. vulparia (Rchb.)

Nyman and Diaphora sordida (Hübner, 1803) on A. napellus Linnaeus.

26

Berlov & Bolotov: Record of Borearctia menetriesii (Eversmann, 1846) ...

Our record confirms that B. menetriesii is a polyphagous species like most other boreal Arctii-

nae (Dubatolov 1 990), but additional experiments are needed for an appropriate evaluation of the

role of Aconitum spp. as a host plant for European populations of B. menetriesii.

Acknowledgements

The authors are grateful to Dr. N. V. Stepantsova, a botanist of the Baikalo-Lensky Nature Reserve,

for help in identification of A. rubicundum , and to Dr. A. Zilli, Dr. J. Rota and an anonymous re-

viewer for valuable comments on the manuscript.

References

Azimova SS, Yunusov MS (2013) Natural Compounds: Alkaloids. Springer, New York, 80 pp. doi:

10.1007/978-1-4614-0560-3

Bellmann H (2003) Der neue Kosmos Schmetterlingsfuhrer - Schmetterlinge, Raupen und Futterpflanzen.

Kosmos, Auflage, 448 pp.

Bolotov IN, Gofarov MY, Kolosova YS, Frolov AA (2013) Occurrence of Borearctia menetriesii (Evers-

mann, 1 846) (Erebidae: Arctiinae) in Northern European Russia: a new locality in a disjunct species range.

Nota lepidopterologica 36(1): 65-75.

Dubatolov VV (1990) Tiger moths (Lepidoptera, Arctiidae: Arctiinae) of South Siberian mountains (report

2). In Zolotarenko GS (Ed.) Arthropods and helminths, Fauna of Siberia Series. Nauka Publisher, Novo-

sibirsk, 139-169. [In Russian]

Dubatolov VV (2010) Tiger-moths of Eurasia (Lepidoptera, Arctiidae) (Nyctemerini by R. de Vos & V. V.

Dubatolov). Neue Entomologische Nachrichten 65: 1-106.

Freina dJJ, Witt TJ (1987) Die Bombyces und Sphinges der Westpalearktis (Bd. 1-2). Forschung & Wissen-

schaft Verlag GmbH, München, 708 pp.

Ivanova MM (1978) Flora of the Upper Angara Valley. In: Malyshev LI, Peshkova GA (Eds) Flora of the

Transbaikalia. Nauka, Novosibirsk, 174-242. [In Russian]

Jalas J, Suominen J (1989) Atlas Florae Europaeae - Distribution of vascular plants in Europe (Vol. 8.) Nym-

phaeaceae to Ranunculaceae. The Committee for Mapping the Flora of Europe and Societas Biologica

Fennica Vanamo, Helsinki, 261 pp.

Kharkevich SS (1995) Plantae Vascularis Orientis Extremi Sovietici (Vol. 7). Nauka, Saint Petersburg, 395

pp. [In Russian]

Krogerus H (1944) Das Vorkommen von Callimorpha menetriesi Ev. in Fennoskandien, nebst Beschreibun-

gen der verschiedenen Entwicklungsstadien. Notulae Entomologicae 24 (3^t): 79-86.

Lampinen R, Lahti T, Heikkinen M (2014) Kasviatlas 2013 [Atlas of the distribution of vascular plants in

Finland 2013]. Helsingin Yliopisto, Luonnontieteellinen keskusmuseo, Helsinki, http://www.luomus.fi/

kasviatlas [accessed 21. xi.20 14]

Lappi E, Mikkola K, Ryynänen J (2004) Idänsiilikäs Borearctia menetriesii , tervetuloa takaisin! [Welcome

back Borearctia menetriesii ]. Baptria 29 (1): 28-29.

Malyschev LI, Pechkova GA (1993) Flora Sibiriae (Vol. 6) - Portulacaceae - Ranunculaceae. Nauka, Novo-

sibirsk, 3 10 pp. [In Russian]

Nishanov AA, Sultankhodzhaev MN, Yunusov MS, Kondraf ev VG (1991) Alkaloids of Aconitum rubricun-

dum. Chemistry of Natural Compounds 27(3): 349-352. doi: 10.1007/BF00630324

Pakkanen P, Wettenhovi J (2014) Borearctia menetriesii in Finland, http://perhoset.nettitieto.fi/historia/arcti-

inae/bor-menetriesi.htm [accessed 21. xi.20 14]

Peshkova GA (1985) The plant cover of Siberia (the Baikal area and Transbaikalia). Nauka, Novosibirsk, 145

pp. [In Russian]

Nota Lepi. 38(1) 2015: 23-27

27

Robinson GS, Ackery PR, Kitching IJ, Beccaloni GW, Hernandez LM (2010) HOSTS - A Database of the

World’s Lepidopteran Hostplants. Natural History Museum, London, http://www.nhm.ac.uk/hosts [Ac-

cessed 20.xi.2014]

Saarenmaa H (2014) Conservation ecology of Borearctia menetriesii. http://bormene.myspecies.info [Ac-

cessed 22.xi.20 14]

Vorbrodt vK, Müller-Rutz J (1914) Die Schmetterlinge der Schweiz (Bd. 2). Druck and Verlag von K.J.

Wyss, Bern, 726 pp.

Yuan CL, Wang XL, Yang DS (2012) Insecticidal bioactivity of diterpenoid alkaloids from Aconitum sino-

montanum Nakai. Modem Agrochemicals 3: 40^43.

Supplementary material 1

The collection locality of Borearctia menetriesii larvae in Eastern Siberia and records of Aconitum ly-

coctonum and Borearctia menetriesii in Finland.

Authors: Oleg E. Berlov, Ivan N. Bolotov

Explanation note: Table SI, Figs S1-S2.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

Nota Lepi. 38(1) 2015: 29^15 | DOI 1 0.3897/nl.38.88 1 4

Contribution to the knowledge of the butterfly fauna of Albania

Martina Sasic16, Milos Popovic2, Sylvain Cuvelier3, Milan Buric2,

Filip Franeta2, Martin Gascoigne-Pees4, Toni Koren5, Dirk Maes6,

Branko Micevski7’8, Nikola Micevski8, Morten S. Molgaard9,

Chris van Swaay6, Irma Wynhoff6, Rudi Verovnik610

1 Croatian Natural History Museum, Demetrova 1, HR- 10000 Zagreb, Croatia

2 HabiProt, Bulevar Osloboâenja 106/34, 11040 Belgrade, Serbia

3 Vlaamse Vereniging voor Entomologie, Werkgroep Dagvlinders, Diamantstraat 4, B-8900 leper, Belgium

4 2 Barretts Close, Stonesfield, Oxfordshire 0X29 8PW, United Kingdom

5 University of Primorska, Science and Research Centre, Institute for Biodiversity Studies, Giordana Bruna 6, SI-6310

Izola, Slovenia

6 Butterfly Conservation Europe (BCE), PO. Box 506, NL-6700 AM Wageningen, The Netherlands

7 Ss. Cyril and Methodius University, Faculty of Natural Sciences and Mathematics, Department of Animal Taxonomy

and Ecology, 1000 Skopje, Macedonia

8 Macedonian Entomological Society (ENTOMAK), Blvd. ASNOM 58, 2-4, 1000 Skopje, Macedonia

9 Nordjysk Lepidopterologklub, Gertrud RasksVej 86, DK-92 1 0 Aalborg S0, Denmark

10 University of Ljubljana, Biotechnical Faculty, Department of Biology, Vecna pot 111, SI- 1000 Ljubljana, Slovenia

http://zoobank.org/7FACABAA-FABE-484C-8C4A-E6573BCB55El

Received 24 October 2014; accepted 18 February 2015; published: 4 March 2015

Subject Editor: Zdenek Frie.

Abstract. Albanian insect fauna is one of the least studied in Europe. In 2012 and 2013 surveys were under-

taken with the aim of improving the knowledge of the distribution of butterflies, particularly in the southern

part of the country. This research has resulted in the publication of three new species records for Albania.

Here we add two new species to the list of native butterflies of Albania, Melitaea ornata Christoph, 1 893 and

Cupido alcetas (Hoffmannsegg, 1804). We recorded a total of 143 species including several confirmations of

historical published records.

The total number of species has consequently increased to 198, which is comparable with butterfly diversity

in neighbouring countries. Unlike its neighbours, Albania has preserved many of its traditional agricultural prac-

tices and consequently its rich fauna has been well protected during the last decades. However, with the opening

up of the country to outside influences this will undoubtedly change as the process of intensification has already

started in more populated coastal areas. It is therefore imperative to identify important butterfly areas in need of

conservation and to take decisive measures to preserve traditional agricultural practices.

Introduction

Albania is a European country in the south-eastern Mediterranean region. Its total area is 28,748 km2,

with 28.5% of its surface area exceeding 1000 m in altitude making it one of the most mountainous

countries in Europe. It has diverse landscapes, ranging from high mountains in the north and east

to an extensive coastline in the west. The climate benefits from both Mediterranean and Central

European influences, with mean January temperatures ranging between -3° to 10°C and mean July

temperatures varying between 17° to 25°C. Rainfall ranges from 600 mm to over 3000 mm in high

30

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

mountain areas (Weatheronline 2014). Albania is in the contact zone between Central European

and Mediterranean fauna and is a part of the Mediterranean biodiversity ‘hotspot’ (Cuttelod et

al. 2008) with exceptionally rich fauna and flora (MMPAU 2007; Radford et al. 2011). A recent

revision of the butterfly fauna of Albania resulted in an updated checklist of 196 species (Verovnik

and Popovic 2013a), and with possible additional species to be discovered it is one of the richest

butterfly countries in Europe.

Lack of interest in butterflies by the local community, inadequate funding and political isolation dur-

ing Communist times has left the butterfly fauna of Albania amongst the least studied in the Balkans.

Southern Albania in particular has never been extensively studied and only the accounts of a few scien-

tific surveys have been published (Gaskin 1990; Abadjiev and Beshkov 1996a; Abadjiev and Beshkov

1996b; Misja 2005; Verovnik and Popovic 2013b; Cuvelier and Molgaard 2015). This paper provides

additional information on the distribution of butterflies in Albania, listing and discussing the species

that have been recorded during the last two years of field surveys. It is a continuation of a recent initi-

ative to increase the knowledge of butterfly diversity and distribution in Albania providing a platform

for further butterfly research in this country (Verovnik and Popovic 2013a, 2013b). A comparison of

Albanian fauna with its neighboring countries is presented, and the threats, as a result of the transition

from traditional to modem agricultural practices, are discussed.

Methods

The surveys of butterfly fauna, carried out by several groups of researchers, started in July 20 1 2

and continued in 2013. Butterflies were observed, photographed and identified in the field, with

only a few specimens collected for further study and identification. Butterfly identification was

based on Tolman and Lewington (2008) and Lafranchis (2004). Additionally, Pieris balcana

Lorkovic, 1970 was identified consulting the website of Ziegler (2013), and Melitaea ornata

Christoph, 1893 was confirmed using DNA barcoding gene COI (Verovnik, unpublished data).

Male genitalia measurements were taken only from collected specimens of Leptidea sinapis (Lin-

naeus, 1758) (Hubrechts 2013; Maes, unpublished data). Taxonomy and nomenclature follow van

Swaay et al. (2010) and/or Fauna Europaea.

We compared the number of species observed in Albania to the number of species observed in

neighbouring countries. The total number of species in Albania was compiled from all available

data, excluding species that are not native to the region (sensu IUCN 2012). The number of species

observed in neighbouring countries is in accordance with the Red List of European Butterflies (van

Swaay et al. 2010).

The study took place in five southern Albanian counties (Korçë, Elbasan, Gjirokastër, Fier and

Berat) concentrating mainly on the mountain regions of Mali i Moravës, Gramoz (Mali i Gramozit),

Ostrovicë, Devoll River Gorge, Mt. Tomorri and on Mt. Nemërçkë near Gjirokastër. In total 68 local-

ities were visited, but these were subsequently grouped into 30 larger locations (Fig. 1):

1. Ohrid lake, close to the Village of Urahë (41°03’45”N; 20°37’28”E; 811 m). Road verges, rocky

slopes with shrubs.

2. Korçë, Drenovë, gorge NE of the village (40°35’19”N; 20°48’25”E; 1075 m). Dry rocky slopes

with limited vegetation cover.

Nota Lepi. 38(1): 29^45

31

0

Key

" State border

• Locations

10 20 30 km

Figure 1. Map of Albania with the position of study locations.

3. Korçë, Drenovë, gorge SE of the village (40o34’29”N; 20°47’59”E; 1075 m). Dry rocky slopes

with limited vegetation cover.

4. Korçë, Drenovë, Parku Kombëtar Bredhi i Drenovës, SE of the village (40°34’0 1 ”N; 20°49’00,,E;

1 170 m). Forests and forest clearings close to a stream and open, rocky habitats in the lower

parts of the valley.

5. Korçë, Boboshtiçë, valleys and gorges E of the village (40o32’59”N; 20°46,45”E; 1040 m). Dry

rocky slopes with limited vegetation cover.

6. Korçë, Boboshtiçë, on the road to Dardhë (40°3ri5”N; 20o47’57”E; 1565 m). Forests, forest

clearings and meadows close to the main road.

7. Korçë, Lavdar, in the valley E of the village (40°36’05”N; 20°40’08”E; 992 m). Open, rocky

habitats with limited shrubs and trees, meadows.

8. Voskopojë, Gjergjevicë, small gorge on the road E of the village (40o35’07”N; 20°34’53”E;

1269 m). Dry rocky slopes with shrubs and grasses.

9. Voskopojë, Lekas Village (40°36’0r’N; 20°30’52”E; 991 m). Dry rocky slopes with shrubs

and trees, meadows.

10. Voskopojë, along the road NW of the Village of Tudis (40°37,18”N; 20°29’2r’E; 1204 m).

Dry rocky slopes with shrubs and trees, meadows.

11. Voskopojë, along the road SW of the Village of Marjan (40°33’57”N; 20o28’45”E; 1225 m).

Flowery meadows with shrubs and trees.

32

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

12. Voskopojë, Mali i Ostroviçës (40°33,30”N; 20°26’59”E; 1231 m). Flowery meadows on slopes.

13. Devoll Gorge, W of the small town of Maliq, before the gorge (40°43’39”N; 20°39’45”E; 825

m). Dry rocky habitats with limited vegetation cover.

14. Devoll Gorge, along the road E of the Village of Strelcë (40°43’24”N; 20°32’40”E; 689 m). Dry

rocky slopes with limited vegetation cover.

15. Devoll Gorge, on the road Gjinkas-Moglicë (40°42’22”N; 20°25’20”E; 508 m). Dry rocky

slopes with limited vegetation cover on calcareous terrain.

16. Devoll Gorge, on the road Moglicë-Bratilë (40°44’03”N; 20°19’59”E; 385 m). Dry rocky slopes

with limited vegetation cover.

17. Gramsh, Grabove e Posthme, in the gorge below the village (40°46’33”N; 20°21’47”E; 880 m).

Dry rocky slopes with limited vegetation cover, overgrown slopes.

18. Gramsh, Lenie, in the village and along the stream below (40°45’57”N; 20°23’40,,E; 992 m).

Orchards, overgrown gravel stream beds.

19. Gramsh, Maja e Valamarës, on the ridge S of the summit (40°45’43”N; 20°27’07”E; 2088 m).

High mountain grasslands (some parts intensively grazed), forest fragments and rocky terrain.

20. Berat, Mali i Tomorrit foothills, E of the Village of Poliçan (40°36,01,,N; 20°08’13”E; 662 m).

Dry rocky slopes with shrubs and trees.

21 . Berat, Mali i Tomorrit, south facing slopes below the mountain ridge (40°38’06”N; 20°09,46”E;

2339 m). Alpine scree slopes with limited grass cover.

22. Berat, Drobonik, along the road S of the village (40°40’ 16”N; 19°57’38”E; 416 m). Open wood-

lands.

23. Berat, Gllavë (40°29’05”N; 19°58,34”E; 909 m). Dry rocky slopes with shrubs and trees.

24. Permet, Bejkollare (40°2r26”N; 20°18’02”E; 926 m). Dry rocky slopes with shrubs and trees,

meadows.

25. Tepelene, at the entrance of the gorge, close to the Village of Kelcyre (40°18’04,,N; 20°07’47”E;

261 m). Dry, calcareous terrains, ruderal areas.

26. Gjirokaster, Cajupi (40°1 r31”N; 20°10’25,,E; 1387 m). Dry, calcareous terrains partially cov-

ered with low shrubs, pastures.

27. Gjirokaster, Sheper ridge (40° 1 1 ’27”N; 20°20’30”E; 1698 m). Dry, mountain grasslands, rocky

slopes.

28. Ersekë, along the road from Leskovik to Ersekë (40°12’36”N; 20°37’57”E; 1098 m). Dry flow-

ery meadows.

29. Ersekë, Rehove, lower slopes of Gramoz Mts. above the village (40°20’00”N; 20°43’43”E; 1547

m). Grasslands, rocky terrains and pastures.

30. Ersekë, Rehove, at the ridge of the Gramoz Mts. (40°19’59”N; 20°45’7”E; 2147 m). High moun-

tain grasslands, pastures and rocky terrain.

Results

During our field surveys in Albania, we recorded a total of 143 butterfly species, from 66 genera and

5 families. Overall it is a total of 1415 records from 68 locations. Cupido alcetas (Hoffmannsegg,

1 804) and Melitaea ornata were recorded for the first time in Albania. A single male specimen of C.

alcetas was observed in the vicinity of Lavdar Village, in dense grassland close to the forest edge. A

single worn female of M. ornata was collected above the gorge SE of Drenovë Village.

Nota Lepi. 38( 1 ): 29-A5

33

The list of recorded species from southern Albania with localities depicted as numerals from the

methods section and observation dates in brackets following each locality:

Family Hesperiidae

1. Pyrgus armoricanus (Oberthür, 1910) Observations: 16 (22.vii.20 13)

2. Pyrgus serratulae (Rambur, 1839) Observations: 27 (25.vii.20 13), 29 (12.vii.2012)

3. Pyrgus cinarae (Rambur, 1839) Observations: 6 (11 .vii.2012), 26 (24.vii.2013), 27 (25.vii.2013)

4. Spialia orbifer (Hübner, 1823) Observations: 2 (2 1 .vii.201 3), 3 (21 .vii.2013), 4 (2 1 .vii.201 3),

5 (ll.vii.2012), 6 (1 1. vii.2012), 7 (21. vii.2013), 8 (17.vii.2013), 10 (17.vii.2013), 11 (19.

vii.2013), 12 (19.vii.2013), 15 (22.vii.2013), 16 (23.vii.2013), 22 (22.vii.2013), 27 (25.vii.2013)

5. Spialia phlomidis (Herrich-Schäffer, 1845) Observations: 2 (21.vii.2013), 3 (21. vii.2013), 15 (22.

vii.2013)

6. Muschampia proto (Ochsenheimer, 1808) Observations: 25 (24.vii.2013)

7. Carcharodus alceae (Esper, 1780) Observations: 3 (21.vii.2013), 4 (2 1 .vii.201 3), 5 (1 1. vii.201 2),

15 (22.vii.2013), 16 (22.vii.201 3), 25 (24.vii.2013), 26 (24.vii.20 13), 27 (25.vii.2013)

8. Carcharodus lavatherae (Esper, 1783) Observations: 2 (21.vii.2013), 3 (2 1 .vii.201 3), 5 (11.

vii.2012)

9. Carcharodus floccifera (Zeller, 1847) Observations: 4 (21.vii.2013), 12 (19.vii.2013), 27 (25.

vii.2013), 30 (12.vii.2012)

10. Carcharodus orientalis Reverdin, 1913. Observations: 5 (1 1. vii.2012), 20 (21.vii.2013), 21 (21.

vii.2013)

11. Erynnis tages (Linnaeus, 1758) Observations: 3 (21. vii.201 3), 4 (21 .vii.201 3), 5 (11 .vii.2012,

18.vii.2013), 7 (2 1 .vii.201 3), 10 (17.vii.2013), 12 (19.vii.2013), 15 (22.vii.2013), 16 (22.vii.2013,

23.vii.2013), 17 (23.vii.2013), 24 (20.vii.2013), 26 (24.vii.2013), 27 (25.vii.2013)

12. Erynnis marloyi (Boisduval, 1834) Observations: 23 (22.vii.201 3), 26 (24.vii.2013), 27 (25.

vii.2013)

13. Thymelicus acteon (Rottemburg, 1775) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 4 (21.

vii.2013), 5 (ll.vii.2012), 6 (1 1. vii.201 2), 14 ( 1 0.vii.20 1 2), 22 (22.vii.2013), 23 (22.vii.20 13),

24 (20.vii.2013)

14. Thymelicus lineola (Ochsenheimer, 1808) Observations: 2 (21. vii.201 3), 3 (21. vii.201 3), 4 (21.

vii.2013), 5 (15.vii.2013, 16.vii.2013), 6 (1 l.vii.2012), 7 (2 1 .vii.20 1 3), 11 ( 1 9.VÜ.20 1 3), 15

(10.vii.2012), 27 (25.vii.2013), 29 (12.vii.2012)

15. Thymelicus sylvestris (Poda, 1761) Observations: 2 (21. vii.20 13), 5 (11 .vii.2012), 6 (11.

vii.2012), 8 (17.vii.2013), 9 (17.vii.2013), 10 ( 1 7.VÜ.20 1 3), 14 (10.vii.2012), 16 (23.vii.2013),

21 (21.vii.2013)

16. Hesperia comma (Linnaeus, 1758) Observations: 27 (25.vii.2013)

17. Ochlodes sylvanus (Esper, 1777) Observations: 2 (21. vii.20 13), 4 (21. vii.20 13), 5 (1 1. vii.20 12),

6 (11 .vii.20 1 2), 11 (19.vii.2013), 13 (10. vii.201 2)

Family Papilionidae

18. Parnassius apollo (Linnaeus, 1758) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 5 (11.

vii.2012, 15.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 1. vii.2012), 8 (17.vii.2013, 1 8.VÜ.20 1 3),

12 (19.vii.2013), 16 (22.vii.2013), 19 (23.vii.2013), 30 (12.vii.2012)

34

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

19. Parnassius mnemosyne (Linnaeus, 1758) Observations: 12 (19.vii.2013), 21 (21.vii.2013,

25.vii.2013), 30 (12.vii.2012)

20. Papilio machaon Linnaeus, 1758. Observations: 1 (15.vii.2013), 3 (21 .vii.2013), 5 (11.

vii.2012, 16.vii.2013), 7 (21.vii.2013), 8 (17.vii.2013), 14 (10.vii.2012, 22.vii.2013), 15 (22.

vii.2013), 16 (22.vii.2013, 23.vii.2013), 17 (23.vii.2013), 20 (2 1 .vii.201 3), 21 (21.vii.2013,

25.vii.2013), 22 (22.vii.2013), 24 (20.vii.2013), 25 (24.vii.2013), 26 (24.vii.20 13), 27 (25.

vii.2013), 28 (14.vii.2013), 29 (12.vii.2012)

21. Iphiclides podalirius (Linnaeus, 1758) Observations: 1 (15. vii.201 3), 2 (21. vii.201 3), 3 (21.

vii.2013), 5 (ll.vii.2012, 18.vii.2013), 6 (1 l.vii.2012), 7 (21. vii.201 3), 9 (17.vii.2013),

10 (17.vii.2013), 11 (19.vii.2013), 13 (10.vii.2012), 14 (10.vii.2012, 22.vii.2013), 15 (10.

vii.2012, 22.vii.2013), 16 (22.vii.2013, 23.vii.2013), 17 (23.vii.2013), 20 (21. vii.2013), 22

(22.vii.2013), 23 (22.vii.20 13), 24 (20.vii.2013), 25 (24.vii.201 3), 26 (24.vii.2013), 27 (25.

vii.2013), 28 (20.vii.2013), 29 (12.vii.2012)

Family Pieridae

22. Aporia crataegi (Linnaeus, 1758) Observations: 3 (2 1 . vii.20 1 3), 4 (2 1 .vii.20 1 3), 6 ( 1 1 .vii.20 1 2),

8 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 14 (10.vii.2012), 29 (12.vii.2012)

23. Pieris rapae (Linnaeus, 1758) Observations: 2 (21. vii.20 13), 4 (21. vii.20 13), 5 (11 .vii.20 1 2,

15.vii.2013), 6 (1 l.vii.2012), 7 (2 1 .vii.20 1 3), 1 0 (17.vii.2013), 12 (19.vii.2013), 13 (10.

vii.2012), 15 (10.vii.2012), 16 (22.vii.2013, 23.vii.2013), 18 (23. vii.20 13), 19 (23.vii.2013),

22 (22.vii.2013), 23 (22.vii.20 13), 24 (20.vii.2013), 25 (24.vii.2013), 26 (24.vii.20 13), 27

(25.vii.2013), 30 (12.vii.2012)

24. Pieris mannii (Mayer, 1851) Observations: 2 (2 1 .vii.20 1 3), 3 (2 1 .vii.20 1 3), 6 (1 l.vii.2012),

10 (17.vii.2013), 12 (1 9.VÜ.20 1 3), 14 (22.vii.20 13), 15 (10.vii.2012), 16 (22.vii.2013,

23.vii.2013), 17 (23.vii.2013), 18 (23.vii.2013), 19 (23.vii.2013), 21 (25.vii.2013), 22 (22.

vii.2013), 25 (24.vii.2013)

25. Pieris ergane (Geyer, 1828) Observations: 2 (21. vii.20 13), 8 (17.vii.2013), 14 (22.vii.20 13),

16 (23.vii.2013), 17 (23.vii.2013), 21 (2 1 .vii.20 1 3), 27 (25.vii.2013)

26. Pieris balcana Lorkovic, 1970. Observations: 6 (1 l.vii.2012)

27. Pieris napi (Linnaeus, 1758) Observations: 8 (18.vii.2013), 10 (17.vii.2013), 1 1 (19.vii.2013),

14 (22.vii.20 13), 16 (23.vii.201 3), 18 (23.vii.2013), 19 (23.vii.2013), 21 (21. vii.20 13), 27

(25.vii.2013), 28 (20.vii.2013)

28 . Pontia edusa (Fabricius, 1777) Observations: 3 (21. vii.20 13), 4 (21. vii.20 13), 5 (1 l.vii.2012),

6 (1 l.vii.2012), 9 (17.vii.2013), 10 (17.vii.2013), 13 (10.vii.2012), 14 (10.vii.2012), 15 (10.

vii.2012, 22.vii.2013), 16 (22.vii.2013, 23.vii.2013), 22 (22.vii.2013), 25 (24.vii.201 3), 26

(24.vii.2013), 27 (25.vii.2013), 29 (12.vii.2012), 30 (12.vii.2012)

29. Euchloe ausonia (Hübner, 1804) Observations: 3 (21. vii.20 13)

30. Colias aurorina (Herrich-Schäffer, 1850) Observations: 5 (1 l.vii.2012), 6 (1 l.vii.2012), 26

(24.vii.2013), 27 (25.vii.2013), 29 (12.vii.2012), 30 (12.vii.2012)

31. Colias alfacariensis Ribbe, 1905. Observations: 2 (21. vii.20 13), 3 (21. vii.20 13), 4 (21.

vii.2013), 5 (1 l.vii.2012, 16.vii.2013, 18.vii.2013), 6 (1 l.vii.2012), 7 (2 1 .vii.201 3), 8 (17.

vii.2013, 18.vii.20 13), 9 ( 1 7.vii.20 13), 10 (17.vii.2013), 1 1 (19.vii.2013), 13 ( 1 0.vii.20 1 2), 14

(10.vii.2012, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 17 (23.vii.2013), 20 (2 1 .vii.20 1 3),

24 (20.vii.2013), 27 (25.vii.2013), 28 (20.vii.2013), 29 ( 1 2.VÜ.20 1 2), 30 ( 1 2.VÜ.20 1 2)

Nota Lepi. 38(1): 29-45

35

32. Colias croceus (Fourcroy, 1785) Observations: 2 (21. vii.20 13), 3 (21.vii.2013), 4 (21.vii.2013),

5 (ll.vii.2012, 15.vii.2013, 16.vii.2013,18.vii.2013), 6 (1 1 .vii.2012), 7 (21. vii.20 13), 8 (17.

vii.20 13, 18.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 13 (10.vii.2012),

14 (10.vii.2012, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 16 (22.vii.2013, 23.vii.2013),

17 (23.vii.2013), 18 (23.vii.2013), 19 (23.vii.2013), 20 (21. vii.20 13), 21 (25.vii.2013), 22

(22.vii.2013), 23 (22.vii.20 13), 24 (20.vii.2013), 25 (24.vii.20 13), 26 (24.vii.2013), 27 (25.

vii.20 13), 28 (20.vii.20 13), 29 (12.vii.2012), 30 (12.vii.2012)

33. Gonepteryx rhamni (Linnaeus, 1758) Observations: 5 (1 1. vii.2012), 6 (1 1. vii.20 12), 7 (21.

vii.20 13), 12 (19.vii.2013), 14 (22.vii.2013), 19 (23.vii.2013), 21 (25.vii.2013), 27 (25.

vii.20 13), 30 (12.vii.2012)

34. Gonepteryx cleopatra (Linnaeus, 1767) Observations: 27 (25.vii.20 13)

35. Gonepteryx farinosa (Zeller, 1847) Observations: 26 (24.vii.20 13)

36. Leptidea sinapis (Linnaeus, 1758) Observations: 2 (2 1 .vii.20 1 3), 3 (2 1 .vii.20 13), 4 (21.

vii.2013), 5 (ll.vii.2012, 15.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 1. vii.2012), 7 (21.

vii.20 13), 8 (17.vii.2013, 18.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 13

(10.vii.2012), 14 (10.vii.2012, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 16 (22.vii.2013,

23.vii.2013), 17 (23.vii.2013), 18 (23.vii.20 13), 20 (21.vii.2013), 22 (22.vii.20 13), 23 (22.

vii.2013), 24 (20.vii.2013), 25 (24.vii.20 13), 26 (24.vii.2013), 27 (25.vii.2013), 28 (20.

vii.2013), 29 (12.vii.2012)

37. Leptidea duponcheli (Staudinger, 1871) Observations: 3 (21. vii.20 13), 4 (21. vii.20 13), 5 (1 1.

vii.2012), 6 (1 1. vii.2012), 7 (21. vii.20 13), 14 (10.vii.2012), 15 (10.vii.2012, 22.vii.2013)

Family Lycaenidae

38. Thecla betulae (Linnaeus, 1758) Observations: 16 (22.vii.2013, 23.vii.2013)

39. Favonius quercus (Linnaeus, 1758) Observations: 29 (12.vii.2012)

40. Satyrium acaciae (Fabricius, 1787) Observations: 4 (2 1 .vii.2013), 5 (1 1. vii.20 12), 6 (11.

vii.2012), 11 (19.vii.2013), 15 (10.vii.2012), 29 (12.vii.2012)

41. Satyrium ilicis (Esper, 1779) Observations: 6 (1 1. vii.20 12), 13 (10.vii.2012), 15 (10.vii.2012)

42. Satyrium spini (Denis & Schiffermüller, 1775) Observations: 2 (21. vii.20 13), 3 (21. vii.20 13), 6(11.

vii.2012), 9 (17.vii.20 13), 11 (19.vii.2013), 15 (10.vii.20 12), 21 (25.vii.2013), 27 (25.vii.2013)

43. Satyrium w-album (Knoch, 1782) Observations: 6 (1 1. vii.2012), 9 (17.vii.2013)

44. Callophrys rubi (Linnaeus, 1758) Observations: 6 (1 1. vii.2012)

45. Lycaena phlaeas (Linnaeus, 1761) Observations: 2 (21. vii.20 13), 3 (21. vii.20 13), 4 (21.

vii.2013), 5(1 l.vii.2012), 6(1 l.vii.2012), 13 (10.vii.2012), 15 (10.vii.2012, 22.vii.2013), 16

(22.vii.2013), 21 (21. vii.20 13), 25 (24.vii.2013), 26 (24.vii.2013), 27 (25.vii.2013)

46. Lycaena dispar (Haworth, 1802) Observations: 5 (1 l.vii.2012)

47. Lycaena virgaureae (Linnaeus, 1758) Observations: 6 (1 l.vii.2012), 12 (19.vii.2013), 21 (25.

vii.2013), 29 (12.vii.2012)

48. Lycaena tityrus (Poda, 1761) Observations: 2 (21. vii.20 13), 5 (1 l.vii.2012), 12 (19.vii.2013),

16 (22.vii.20 13), 29 (12.vii.2012)

49. Lycaena alciphron (Rottemburg, 1775) Observations: 2 (21. vii.20 13), 3 (21. vii.20 13), 4 (21.

vii.2013), 5 (ll.vii.2012), 6 (1 l.vii.2012), 12 (19.vii.2013), 14 (10.vii.2012)

50. Lycaena thersamon (Esper, 1784) Observations: 6 (1 l.vii.2012), 14 (10.vii.2012), 15 (10.

vii.2012)

36

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

51. Lycaena candens (Herrich-Schäffer, 1844) Observations: 30 (12.vii.2012)

52. Lampides boeticus (Linnaeus, 1767) Observations: 14 (10.vii.2012), 27 (25.vii.2013)

53. Leptotes pirithous (Linnaeus, 1767) Observations: 5 (11 .vii.2012), 7 (21.vii.2013), 15 (10.

vii.2012), 24 (20.vii.2013)

54. Tarucus balkanica (Freyer, 1844) Observations: 25 (24.vii.20 13)

55. Cupido minimus (Fuessly, 1775) Observations: 2 (21.vii.2013), 4 (21.vii.2013), 5 (15.vii.2013,

16.vii.2013), 6 (1 1. vii.2012), 11 (19.vii.2013), 12 (19.vii.2013), 19 (23.vii.20 13), 27 (25.

vii.2013)

56. Cupido osiris (Meigen, 1829) Observations: 7 (21.vii.2013), 13 (10.vii.2012), 14 (10.vii.2012)

57. Cupido alcetas (Hoffmannsegg, 1804) Observations: 7 (21. vii.2013)

58. Celastrina argiolus (Linnaeus, 1758) Observations: 3 (21. vii.2013), 4 (21. vii.2013), 5 (15.

vii.2013), 6 (1 1. vii.2012), 7 (21. vii.2013), 10 (17.vii.2013), 14 (10.vii.2012), 15 (10.vii.2012),

17 (23.vii.2013), 18 (23.vii.2013), 25 (24.vii.20 13), 26 (24.vii.2013)

59. Phengaris alcon (Denis & Schiffermüller, 1775) Observations: 6 (1 1. vii.2012), 27 (25.vii.2013)

60. Phengaris avion (Linnaeus, 1758) Observations: 4 (21. vii.2013), 6 (1 1. vii.2012)

61. Iolana iolas (Ochsenheimer, 1816) Observations: 6 (1 l.vii.2012), 7 (21. vii.2013), 14 (10.

vii.2012), 27 (8-19.vii.2013)

62. Scolitantides orion (Pallas, 1771) Observations: 16 (23.vii.20 13)

63. Pseudophilotes vicrama (Moore, 1865) Observations: 3 (21.vii.2013), 5 (1 1. vii.2012), 6 (11.

vii.2012), 8 (18.vii.2013), 10 (17.vii.2013), 15 (10.vii.2012), 27 (25.vii.2013)

64. Plebejus sephirus (Frivaldzky, 1835) Observations: 27 (25.vii.2013)

65. Plebejus argus (Linnaeus, 1758) Observations: 5 (1 1. vii.2012, 16.vii.2013), 10 (17.vii.2013),

11 (19.vii.2013), 13 (10.vii.2012), 14 (10.vii.2012), 15 (10.vii.2012), 23 (22.vii.201 3), 24

(20.vii.2013), 28 (20.vii.2013)

66. Plebejus idas (Linnaeus, 1761) Observations: 2 (2 1 .vii.2013), 4 (21. vii.2013), 6 (11 .vii.2012),

7 (21.vii.2013), 8 (17.vii.2013), 16 (22.vii.2013), 18 (23.vii.2013), 23 (22.vii.2013), 27 (25.

vii.2013), 29 (12.vii.2012)

67. Aricia eumedon (Esper, 1780) Observations: 12 (19.vii.2013)

68. Aricia agestis (Denis & Schiffermüller, 1775) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 4

(21.vii.2013), 5 (1 l.vii.2012, 18.vii.2013), 6 (1 1 .vii.2012), 7 (21. vii.2013), 12 (19.vii.2013),

13 (10.vii.2012), 14 (10. vii.2012), 15 (10.vii.2012, 22.vii.2013), 16 (22.vii.2013, 23.vii.2013),

18 (23.vii.2013), 22 (22.vii.2013), 23 (22.vii.2013), 24 (20.vii.2013), 25 (24.vii.20 13), 26

(24.vii.2013), 27 (25.vii.2013), 29 (12.vii.2012)

69. Aricia artaxerxes (Fabricius, 1793) Observations: 5 (16.vii.2013), 6 (1 l.vii.2012), 21 (25.

vii.2013)

70. Aricia anteros (Freyer, 1838) Observations: 12 (19.vii.2013)

71. Cyaniris semiargus (Rottemburg, 1775) Observations: 4 (21. vii.2013), 5 (1 l.vii.2012), 6 (11.

vii.2012), 11 (19.vii.2013), 12 (19.vii.2013), 16 (23.vii.2013), 30 (12.vii.2012)

72. Polyommatus damon (Denis & Schiffermüller, 1775) Observations: 8 (17.vii.2013), 12 (19.

vii.2013)

73. Polyommatus ripartii (Freyer, 1830) Observations: 3 (21 .vii.2013), 4 (21. vii.2013), 5 (11.

vii.2012, 16.vii.2013), 6 (1 l.vii.2012), 7 (21.vii.2013), 8 (17.vii.2013), 10 (17.vii.2013), 11

(19.vii.2013), 15 (22.vii.2013), 27 (25.vii.2013)

Nota Lepi. 38(1): 29^15

37

74. Polyommatus admetus (Esper, 1783) Observations: 2 (21.vii.2013), 3 (21.vii.2013),4 (21.

vii.2013), 5 (ll.vii.2012, 18.vii.2013), 7 (21.vii.2013), 10 (17.vii.2013), 14 (10.vii.2012,

22.vii.2013), 15 (22.vii.2013), 18 (23.vii.2013), 23 (22.vii.2013), 26 (24.vii.2013), 27 (25.

vii.2013), 29 (12.vii.2012)

75. Polyommatus escheri (Hübner, 1823) Observations: 9 (17. vii.2013), 10 (17.vii.2013), 14 (10.

vii.2012, 22.vii.2013), 15 (22.vii.2013), 16 (22.vii.2013), 28 (20.vii.2013)

76. Polyommatus amandus (Schneider, 1792) Observations: 5 (1 l.vii.2012), 6 (1 1 .vii.2012), 12

(19.vii.2013), 14 (10.vii.2012)

77. Polyommatus thersites (Cantener, 1835) Observations: 3 (21. vii.2013), 4 (21. vii.2013), 9 (17.

vii.2013), 15 (10.vii.2012), 23 (22.vii.201 3), 27 (25.vii.2013), 29 (12.vii.2012)

78. Polyommatus dorylas (Denis & Schiffermüller, 1775) Observations: 3 (21. vii.2013), 4 (21.

vii.2013), 6 (11. vii.2012), 16 (23.vii.2013), 26 (24.vii.20 13), 27 (25.vii.2013)

79. Polyommatus daphnis (Denis & Schiffermüller, 1775) Observations: 2 (21. vii.2013), 3 (21.

vii.2013), 4 (21. vii.2013), 5 (1 1 .vii.2012), 6 (1 1 .vii.2012), 7 (21. vii.2013), 9 (17.vii.2013),

11 (19.vii.2013), 14 (10.vii.2012), 15 (10.vii.2012, 22.vii.2013), 16 (22.vii.2013), 20 (21.

vii.2013), 24 (20.vii.2013), 27 (25.vii.2013), 28 (20.vii.2013)

80. Polyommatus coridon (Poda, 1761) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 4 (21.

vii.2013), 5 (1 1. vii.2012), 7 (21. vii.2013), 9 (17.vii.2013), 16 (23.vii.2013), 26 (24.vii.2013)

81. Polyommatus bellargus (Rottemburg, 1775) Observations: 4 (21. vii.2013), 5 (1 1. vii.2012),

12 (19.vii.2013), 14 (10.vii.2012), 15 (10.vii.2012, 22.vii.2013), 16 (23.vii.2013), 26 (24.

vii.2013), 27 (25.vii.2013), 28 (20.vii.2013)

82. Polyommatus icarus (Rottemburg, 1775) Observations: 2 (21. vii.2013), 3 (21. vii.2013),

4 (21.vii.2013), 5 (1 1 .vii.2012, 15.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 1. vii.2012), 7

(21.vii.2013), 8 (17.vii.2013), 9 ( 1 7.VÜ.20 1 3), 10 ( 1 7. vii.20 1 3), 11 (19.vii.2013), 12 (19.

vii.2013), 13 (10.vii.2012), 14 ( 1 0.vii.20 1 2, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013),

16 (22.vii.2013, 23.vii.2013), 17 (23.vii.2013), 18 (23.vii.2013), 19 (23.vii.2013), 20 (21.

vii.2013), 21 (21.vii.2013, 25.vii.2013), 22 (22.vii.2013), 23 (22.vii.2013), 24 (20.vii.2013),

25 (24.vii.2013), 26 (24.vii.2013), 27 (25.vii.2013), 28 (20.vii.2013), 29 (12.vii.2012), 30

(12.vii.2012)

83. Polyommatus eros (Ochsenheimer, 1808) Observations: 30 (12.vii.2012)

Family Nymphalidae

84. Libythea celtis (Laicharting, 1782) Observations: 21 (25. vii.20 13), 26 (24.vii.20 13)

85. Apatura iris (Linnaeus, 1758) Observations: 6 (1 1. vii.2012), 8 (18.vii.2013)

86. Apatura ilia (Denis & Schiffermüller, 1775) Observations: 13 (10.vii.2012)

87. Limenitis reducta (Staudinger, 1901) Observations: 4 (21. vii.20 13), 6 (1 1. vii.20 12), 8 (17.vii.2013),

1 1 (19.vii.2013), 14 (22.vii.201 3), 24 (20.vii.2013),26 (24.vii.20 13), 27 (25.vii.2013)

88. Nymphalis antiopa (Linnaeus, 1758) Observations: 1 1 (19.vii.2013), 21 (25.vii.20 13)

89. Nymphalis polychloros (Linnaeus, 1758) Observations: 11 (19.vii.2013)

90 . Aglais io (Linnaeus, 1758) Observations: 4 (21. vii.20 13), 6 (1 l.vii.2012), 12 (19.vii.2013), 14

(10.vii.2012), 19 (23 .vii.20 1 3), 21 (2 1 .vii.20 1 3, 25.vii.2013)

91. Aglais urticae (Linnaeus, 1758) Observations: 6 (1 l.vii.2012), 12 (19.vii.20 13), 19 (23.

vii.2013), 21 (21.vii.2013, 25.vii.2013), 30 (12.vii.2012)

38

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

92. Vanessa atalanta (Linnaeus, 1758) Observations: 11 (19.vii.2013), 14 (10.vii.2012), 18 (23.

vii.2013), 21 (25.vii.2013), 26 (24.vii.20 13), 27 (25.vii.2013), 30 (12.vii.2012)

93. Vanessa car dui (Linnaeus, 1758) Observations: 2 (21.vii.2013), 4 (21. vii.2013), 5 (11 .vii.2012,

15.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 1 .vii.2012), 7 (2 1 .vii.201 3), 8 ( 1 7.VÜ.20 1 3,

18.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 12 (19.vii.2013), 13 (10.

vii.2012), 14 (10.vii.2012), 18 (23.vii.2013), 19 (23. vii.201 3), 20 (21.vii.2013), 21 (21.

vii.2013, 25.vii.2013), 22 (22.vii.20 13), 23 (22.vii.2013), 24 (20.vii.2013), 26 (24.vii.2013),

27 (25.vii.2013), 28 (20.vii.201 3), 29 (12.vii.2012)

94. Issoria lathonia (Linnaeus, 1758) Observations: 3 (2 1 .vii.20 13), 4 (2 1 .vii.20 1 3), 5 (1 1 .vii.20 12),

6 (1 1 .vii.2012), 10 (1 7.VÜ.20 1 3), 11 (19.vii.2013), 14 (10.vii.2012), 18 (23.vii.2013), 19(23.

vii.2013), 21 (21.vii.2013, 25.vii.2013), 26 (24.vii.201 3), 27 (25.vii.2013), 29 (12.vii.2012),

30 (12.vii.2012)

95. Polygonia c-album (Linnaeus, 1758) Observations: 4 (2 1 .vii.20 1 3), 6 (ll.vii.2012), 7 (21.

vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 15 (22.vii.20 13), 17 (23.vii.2013), 18 (23.

vii.2013), 26 (24.vii.2013), 27 (25.vii.2013)

96. Argynnis pandora (Denis & Schiffermüller, 1775) Observations: 2 (21.vii.2013), 6 (11.

vii.2012), 14 (10.vii.2012), 21 (25.vii.2013), 29 (12.vii.2012)

97. Argynnis paphia (Linnaeus, 1758) Observations: 2 (21. vii.20 13), 3 (21. vii.20 13), 4 (21.

vii.2013), 5 (18.vii.2013), 6 (1 1. vii.20 12), 7 (21. vii.20 13), 8 (17.vii.2013), 14 (10.vii.2012), 15

(10.vii.2012), 16 (23.vii.2013), 24 (20.vii.2013), 27 (25.vii.2013), 28 (20.vii.2013)

98. Argynnis aglaja (Linnaeus, 1758) Observations: 4 (2 1 .vii.201 3), 6 (11 .vii.2012), 9 (17.

vii.2013), 10 (17.vii.2013), 12 (19.vii.20 13), 26 (24.vii.2013), 27 (25.vii.2013)

99 .Argynnis adippe( Denis & Schiffermüller, 1775) Observations: 4 (21. vii.20 13), 6 (1 1. vii.20 12),

9 (17.vii.2013), 10 (17.vii.2013), 15 (10.vii.2012)

100. Argynnis niobe (Linnaeus, 1758) Observations: 4 (21. vii.20 13), 6 (1 1. vii.2012), 12 (19.

vii.2013), 19 (23.vii.2013), 24 (20.vii.2013), 27 (25.vii.2013)

101. Brenthis hecate (Denis & Schiffermüller, 1775) Observations: 5 (16.vii.2013)

102. Brenthis daphne (Bergsträsser, 1780) Observations: 4 (21. vii.20 13), 5 (1 1. vii.20 12), 6 (11.

vii.2012), 14 (10.vii.2012)

103. Boloria graeca (Staudinger, 1870) Observations: 30 (12.vii.2012)

104. Boloria euphrosyne (Linnaeus, 1758) Observations: 19 (23.vii.2013)

105. Boloria dia (Linnaeus, 1767) Observations: 2 (21. vii.20 13), 5 (11 .vii.2012)

106. Melitaea phoebe (Denis & Schiffermüller, 1775) Observations: 2 (21. vii.20 13), 4 (21.

vii.2013), 5(1 l.vii.2012), 6(1 l.vii.2012), 8 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013),

14 (10.vii.2012), 19 (23.vii.2013), 22 (22.vii.20 13), 25 (24.vii.20 13), 29 (12.vii.2012)

107. Melitaea didyma (Esper, 1779) Observations: 3 (21. vii.20 13), 4 (21. vii.20 13), 6 (1 l.vii.2012),

8 (17.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 15 (10.vii.2012), 21 (21.

vii.2013), 25 (24.vii.2013), 26 (24.vii.20 13), 29 (12.vii.2012)

108. Melitaea trivia (Denis & Schiffermüller, 1775) Observations: 15 (10.vii.2012), 16 (23.

vii.2013), 21 (25.vii.2013), 23 (22.vii.20 13), 27 (25.vii.2013), 30 (12.vii.2012)

109. Melitaea athalia (Rottemburg, 1775) Observations: 4 (21.vii.2013), 6 (1 l.vii.2012), 7 (21.

vii.2013), 11 (19.vii.2013), 17 (23.vii.2013), 29 (12.vii.2012)

1 10. Melitaea ornata Christoph, 1893. Observations: 3 (21. vii.20 13)

111. Euphydryas aurinia (Rottemburg, 1 775) Observations: 15 (1 0.vii.2012)

Nota Lepi. 38(1): 29^5

39

112. Melanargia galathea (Linnaeus, 1758) Observations: 3 (21.vii.2013), 4 (21.vii.2013), 5

(ll.vii.2012, 15.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 l.vii.2012), 7 (21.vii.2013), 8 (17.

vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 13 (10.vii.2012), 14 (10.

vii.2012, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 16 (22.vii.2013, 23.vii.2013), 17 (23.

vii.2013), 19 (23.vii.2013), 21 (21.vii.2013, 25.vii.2013), 22 (22.vii.2013), 23 (22.vii.2013),

24 (20.vii.2013), 28 (20.vii.2013), 29 (12.vii.2012), 30 (12.vii.2012)

1 13. Melanargia russiae (Esper, 1783) Observations: 8 (17.vii.2013, 18.vii.2013), 12 (19.vii.2013),

21 (21.vii.2013, 25.vii.2013), 26 (24.vii.20 13), 27 (25.vii.2013)

114. Melanargia larissa (Geyer, 1828) Observations: 2 (21.vii.2013), 3 (21.vii.2013), 4 (21.

vii.2013), 5 (11. vii.2012, 15.vii.2013, 16.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 l.vii.2012),

7 (21. vii.2013), 8 (17.vii.2013, 18.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 13 (10.

vii.2012), 14 (10.vii.2012, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 16 (22.vii.2013,

23.vii.2013), 17 (23.vii.2013), 20 (21. vii.2013), 21 (21.vii.2013, 25.vii.2013), 23 (22.

vii.2013), 24 (20.vii.2013), 26 (24.vii.2013), 27 (25.vii.2013), 28 (20.vii.2013), 29 (12.

vii.2012), 30 (12.vii.2012)

115. Hipparchia syriaca (Staudinger, 1871) Observations: 1 (15. vii.2013), 9 (17.vii.2013), 24 (24.

vii.2013)

116. Hipparchia fagi (Scopoli, 1763) Observations: 2 (21. vii.2013), 5 (16.vii.2013, 18.vii.2013),

8 (17.vii.2013, 18.vii.2013), 15 (22.vii.2013), 16 (23.vii.2013), 26 (24.vii.20 13), 27 (25.

vii.2013), 27 (20.vii.2013), 29 (12.vii.2012)

117. Hipparchia senthes (Fruhstorfer, 1908) Observations: 15 ( 1 0.vii.20 1 2), 17 (23.vii.2013), 29

(12.vii.2012)

1 18. Hipparchia statilinus (Hufnagel, 1766) Observations: 16 (22.vii.2013, 23.vii.2013)

119. Chazara briseis (Linnaeus, 1764) Observations: 2 (21.vii.2013), 3 (21.vii.2013), 5 (11.

vii.2012, 16.vii.2013), 7 (21.vii.2013), 8 (18.vii.2013), 14 (10.vii.2012, 22.vii.2013), 15 (22.

vii.2013), 26 (24.vii.2013), 27 (25.vii.2013)

120. Pseudochazara anthelea (Hübner, 1824) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 5 (16.

vii.2013, 18.vii.2013), 8 (18.vii.2013), 14 (10.vii.2012), 15 (22.vii.20 13), 16 (22.vii.2013), 28

(14.vii.2013)

121. Pseudochazara mniszechii (Herrich-Schäffer, 1851) Observations: 2 (21. vii.2013), 3 (21.

vii.2013), 5 (1 l.vii.2012, 15.vii.2013, 16.vii.2013, 18.vii.2013), 8 (17.vii.2013, 18.vii.2013),

28(14. vii.2013)

122. Pseudochazara amymone Brown, 1976. Observations: 2 (21. vii.2013), 3 (21. vii.2013), 5(11.

vii.2012, 16.vii.2013, 18.vii.2013), 8 (18.vii.2013), 14 (10.vii.2012, 22.vii.2013), 16 (22.

vii.2013, 23.vii.2013)

123. Satyrus ferula (Fabricius, 1793) Observations: 2 (21. vii.2013), 5 (1 l.vii.2012, 16.vii.2013), 8(17.

vii.2013), 16 (23 .vii.2013), 21 (21.vii.2013, 25.vii.2013), 26 (24.vii.20 13), 27 (25.vii.2013)

124. Brintesia circe (Fabricius, 1775) Observations: 1 (15.vii.20 13), 3 (21. vii.2013), 4 (21.

vii.2013), 5 (1 l.vii.2012, 16.vii.2013, 18.vii.2013), 6 (1 l.vii.2012), 7 (21.vii.2013), 8 (17.

vii.2013, 18.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 14 (10.vii.2012,

22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 17 (23.vii.2013), 19 (23.vii.2013), 21 (21.

vii.2013), 22 (22.vii.2013), 23 (22.vii.2013), 24 (20.vii.2013), 25 (24.vii.2013), 26 (24.

vii.2013), 27 (25.vii.2013), 28 (20.vii.2013), 29 (12.vii.2012)

125. Arethusana arethusa (Denis & Schiffermüller, 1775) Observations: 3 (21. vii.2013)

40

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

126. Erebia medusa (Denis & Schiffermüller, 1775) Observations: 12 (19.vii.2013), 19 (23.

vii.2013), 21 (21.vii.2013, 25.vii.2013), 30 (12.vii.2012)

127. Erebia gorge (Hübner, 1804) Observations: 21 (21. vii.2013, 25.vii.2013)

128. Erebia rhodopensis Nicholl, 1900. Observations: 30 (12.vii.2012)

129. Erebia ottomana Herrich-Schäffer, 1847. Observations: 12 (19.vii.2013), 19 (23.vii.20 13),

30 (12.vii.2012)

130. Erebia mêlas (Herbst, 1796) Observations: 19 (23.vii.2013), 21 (21.vii.2013, 25.vii.2013),

30 (12.vii.2012)

131. Maniola jurtina (Linnaeus, 1758) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 4 (21.

vii.2013), 5 (ll.vii.2012, 15.vii.2013, 16.vii.2013, 18.vii.2013), 6 (1 1 .vii.2012), 7 (21.

vii.2013), 8 (17.vii.2013, 18.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013),

13 (10.vii.2012), 14 (10.vii.2012), 15 (10.vii.2012), 16 (22.vii.20 13), 17 (23.vii.2013), 18

(23. vii.2013), 21 (21. vii.2013), 23 (22.vii.2013), 24 (20.vii.2013), 25 (24.vii.20 13), 26 (24.

vii.2013), 27 (25.vii.2013), 28 (20.vii.2013), 29 (12.vii.2012)

132. Hyponephele lycaon (Rottemburg, 1775) Observations: 2 (21. vii.2013), 5 (1 1. vii.2012,

16.vii.2013, 18.vii.2013), 8 (17.vii.2013, 18.vii.2013), 9 (17.vii.2013), 10 (17.vii.2013), 11

(19.vii.2013), 26 (24.vii.20 13), 27 (25.vii.2013), 29 (12.vii.2012)

133. Hyponephele lupina (Costa, 1836) Observations: 8 (17.vii.2013)

134. Aphantopus hyperantus (Linnaeus, 1758) Observations: 4 (21. vii.2013)

135. Pyronia tithonus (Linnaeus, 1767) Observations: 7 (21. vii.2013), 13 (10.vii.2012), 14 (10.

vii.2012), 15 (22.vii.2013), 17 (23 .vii.2013), 18 (23. vii.2013), 20 (21. vii.2013), 24 (20.vii.2013)

136. Coenonympha rhodopensis Elwes, 1900. Observations: 12 (19.vii.2013), 19 (23.vii.2013), 30

(12.vii.2012)

137. Coenonympha pamphilus (Linnaeus, 1758) Observations: 2 (21. vii.2013), 3 (21. vii.2013), 5 (11.

vii.2012), 7 (21.vii.2013), 10 (17.vii.2013), 11 (19.vii.2013), 13 (10.vii.2012), 14 (10.vii.2012,

22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 22 (22.vii.2013), 23 (22.vii.20 13), 24 (20.vii.2013),

25 (24.vii.2013), 26 (24.vii.20 13), 27 (25.vii.201 3), 28 (20.vii.2013)

138. Coenonympha arcania { Linnaeus, 1761) Observations: 2 (21.vii.2013), 3 (21. vii.2013), 4 (21.

vii.2013), 5 (ll.vii.2012), 6 (1 1 .vii.20 12), 8 (1 7.VÜ.20 1 3), 10 (1 7.VÜ.20 1 3), 1 1 ( 1 9.VÜ.20 1 3),

12 (19.vii.2013), 16 (22.vii.20 13), 17 (23.vii.2013)

139. Coenonympha orientalis Rebel, 1910. Observations: 6 (1 1. vii.20 12), 12 (19.vii.2013), 19 (23.

vii.2013), 30 (12.vii.2012)

140. Pararge aegeria (Linnaeus, 1758) Observations: 2 (21. vii.20 13), 4 (21. vii.20 13), 7 (21.

vii.2013), 18 (23.vii.2013), 27 (25.vii.2013)

141. Lasiommata megera (Linnaeus, 1767) Observations: 2 (2 1 .vii.20 1 3), 3 (21. vii.20 13), 5 (11.

vii.2012, 16.vii.2013, 18.vii.2013), 7 (21. vii.20 13), 8 (17.vii.2013, 1 8.VÜ.20 1 3), 10 (17.

vii.2013), 11 (19.vii.2013), 14 (10.vii.2012, 22.vii.2013), 15 (10.vii.2012, 22.vii.2013), 16

(22.vii.2013, 23.vii.2013), 17 (23.vii.2013), 19 (23.vii.2013), 21 (21. vii.2013, 25.vii.2013),

22 (22.vii.2013), 23 (22.vii.2013), 24 (20.vii.2013), 25 (24.vii.2013), 26 (24.vii.2013), 27 (25.

vii.2013), 28 (20.vii.2013), 30 (12.vii.2012)

142. Lasiommata maera (Linnaeus, 1758) Observations: 2 (21. vii.2013), 6 (1 1. vii.2012), 15 (10.

vii.2012, 22.vii.2013), 16 (22.vii.2013, 23.vii.2013), 17 (23.vii.2013), 21 (2 1 .vii.20 1 3), 23

(22.vii.2013), 25 (24.vii.2013), 26 (24.vii.2013), 27 (25.vii.2013)

143. Kirinia roxelana (Cramer, 1777) Observations: 27 (25.vii.20 13)

Nota Lepi. 38(1): 29-45

41

2*

c

3

O

O

Serbia

Romania

Montenegro

Greece

Republic of Macedonia

Croatia

Bulgaria

Bosnia and Herzegovina

Albania

0 50 100 150 200 250

Number of species observed

Figure 2. The total number of butterfly species recorded in Albania compared with its neighbouring countries.

If we include all the published records, the total number of butterfly species recorded in Albania

has risen to 198, which equates to 41% of the total European butterfly fauna. Compared with its

neighbouring countries (Fig. 2) only Greece (229), and Bulgaria (215) have more recorded species,

while other countries have similar diversity.

Discussion

During the two years of field studies in southern Albania, five new species were recorded for the

country. Colias aurorina , Apatura iris and Pieris balcana were observed during the first survey in

2012 (Verovnik and Popovic 2013b). Cupido alcetas and Melitaea ornata were added in 2013. The

presence of both species in Albania is not unexpected, although C. alcetas is rare in the neighbour-

ing Republic of Macedonia (Schaider and Jaksic 1989) and Greece (Pamperis 2009). This butterfly

could be easily overlooked due to its similarity with C. decoloratus and C. argiades , both of which

have previously been recorded from Albania (Rebel and Zemy 1931). As only historical publica-

tions are available for reference, misidentifications cannot be excluded. C. alcetas is possibly more

widespread in Albania as it frequents a variety of habitats (Pamperis 2009) preferring more humid,

sheltered or overgrown biotopes along streams or rivers. M. ornata , on the other hand, is possibly

more widespread in the southern Balkans with several records from the Republic of Macedonia

(Verovnik et al. 2010; Verovnik 2012), Serbia (Jaksic 2011) and Croatia (Koren and Stih 2013).

However, it is very similar to the more widely distributed M. phoebe, and therefore easily over-

looked (Toth et al. 2013). Identification from studying the overwintering larvae of both species is

usually required to confirm its presence (Russell et al. 2007; Töth and Varga 2010).

Additionally, we confirm the presence of Erebia rhodopensis in Albania. Our record, from the

Gramoz Mts., is the first authenticated record for the country. Its presence in the Gramoz Mts. was

not unexpected, as it is common on the Greek side of the same mountain range (Pamperis 2009).

42

Sasic et al. : Contribution to the knowledge of the butterfly fauna of Albania

The species had previously been reported in Albania from Mt. Kobilica in the Shar Mts. (Rebel

and Zemy 1931). However this mountain currently lies on the border between Kosovo and the

Republic of Macedonia. There is a possibility that it is also present on the Albanian part of the Shar

Mts., further west of Mt. Kobilica.

Among other species that have been recorded one of the most notable is the Balkan endemic

Pseudochazara amymone which has only recently been discovered in Albania (Eckweiler 2012).

Its distribution in Greece still remains unknown, although it has been reported from several sites

(Pamperis 2009). Based on our surveys, more detailed information is now available on its distri-

bution, threats (Verovnik et al. 2014), habitat selection, life cycle, morphology and variability

(Gascoigne-Pees et al. 2014; Cuvelier and Molgaard 2015).

In addition to those discovered by Verovnik and Popovic (2013b), two other colonies of Colias

aurorina were discovered in 2013 on calcareous ridges east of Gjirokaster on Mt. Nemëçkë (Loc.

26) and Mt. Lunxhërisë (Loc. 27), extending the known range of this species in Albania by 50

kilometres to the west.

Albania has a similar number of species in comparison to its neighbours (Fig. 2), highlighting

the importance of this region for butterfly conservation. The additional number of butterfly species

recorded in Greece and Bulgaria can be explained by the fact that more faunistic surveys have been

carried out in these countries and they both have a much larger surface area. Greece, in particular,

supports many local species found only on its offshore islands close to mainland Turkey, and these

species are absent from the rest of Europe (Pamperis 2009). More detailed and well organized

surveys in Albania should certainly result in a more complete list of butterflies for this country.

In particular, the mountains in the northern part of the country which experience a more conti-

nental climate may harbour some additional species such as Leptidea juvernica (Williams, 1946),

Neptis sappho (Pallas, 1771), Limenitis populi (Linnaeus, 1758), Limenitis Camilla (Linnaeus, 1764),

Melitaea diamina (Lang, 1789), Melitaea arduinna (Esper, 1783), Nymphalis vaualbum (Denis &

Schiffermüller, 1775), and Minois dryas (Scopoli, 1763), whilst higher up in the mountains Plebejus

optilete (Knoch, 1781), Erebia alberganus (De Prunner, 1798) and Pyrgus andromedae (Wallengren,

1853) could also be discovered. Additionally, early spring surveys of the gorges in the eastern part of

the country could provide new records, potentially of Anthocharis damone (Boisduval, 1 836), Euchloe

pennia (Freyer, 1852) and Pseudophilotes bavius (Eversmann, 1832).

As traditional low intensity farming is economically non-profitable, many parts of the Balkan

Peninsula have suffered from rural depopulation resulting in an aging population. Abandonment

of rural communities has resulted in the breakdown of traditional agricultural practices (Karoglan

Todorovic 2013), especially low intensity cattle farming. Historically, traditional grazing and

mowing have created semi natural habitats supporting a diversity of species including butterflies.

Abandonment of agriculture and the decline in the number of livestock has resulted in the shrink-

ing of species-rich grasslands and, consequently, biodiversity loss (van Swaay et al. 2012). How-

ever, the situation in Albania is complex. Statistically, Albanian rural communities are character-

ized by large number of small farms and the smallest average farm and plot size of all the Balkan

countries (Kazakova and Stefanova 2010). Modernisation of agricultural practices has not been

implemented, especially in the mountainous parts of the country, where traditional cattle grazing

is still carried out. However, Albania is now open to the agricultural practices adopted by other

European countries and it is only a matter of time before changes will take place, resulting in the

Nota Lepi. 38(1): 29—45

43

loss of the preserved mosaic of habitats. Urgent measures regarding nature conservation in Albania

are therefore needed as neglecting the situation would almost certainly lead to a dramatic reduction

of its native fauna and flora.

It is of paramount importance to complete the faunal list and to initiate nature conservation

guidelines, especially when adopting new agricultural policies. With respect to butterflies, results

from faunistic surveys would help pave the way for new initiatives regarding butterfly conserva-

tion with the prospect of implementing a network of Prime Butterfly Areas (PB As; see van Swaay

and Warren 2003) . Mali i Moravës, Gramoz (Mali i Gramozit), Devoll River Gorge, Mt. Tomorri

and areas on Mt. Nemercke are among top candidates for PBAs, but there are many more areas to

be identified. We hope that this contribution will stimulate more people to study the rich flora and

fauna of Albania.

Acknowledgements

We wish to express our thanks to Heribert Hanisch for his additional records of I. iolas and P. mniszechii. Fund-

ing for the initial survey in 2012 was in part provided by the Mohamed bin Zayed Species Conservation Fund.

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Nota Lepi. 38(1) 2015: 47-58 | DPI 10.3897/nl.38.9034

Biology and distribution of the declining moth Ethmia pyrausta

(Pallas, 1771), with description of the larva (Gelechioidea,

Depressariidae, Ethmiinae)

Kari Nupponen1, Matti Ahola2, Marko Nieminen3, Urmas Jürivete4

1 Merenneidontie 19 D, FIN-02320 Espoo, Finland; email: Kari.Nupponen@kolumbus.fi

2 Metsdnreunantie 27 G, FIN-85900 Reisjdrvi, Finland; email: MJ.Ahola@kotinet.com

3 Department of Biosciences, University of Helsinki, PO Box 65 (Viikinkaari 1), FI— 00014 University of Helsinki, Finland

4 Moora Umb 8, EE-11625 Tallinn, Estonia; email: urmas@kolmabi.ee

http://zoobank.org/90EF92E0-D58D-439C-957A-90735EC15611

Received 27 November 2014; accepted 8 January 2015; published: 17 March 2015

Subject Editor: Erik van Nieukerken.

Abstract. Records of Ethmia pyrausta (Pallas, 1771) from the Baltic countries, the British Isles and Fennos-

candia are listed. All known aspects of habitat requirements, larval biology and adult behaviour, mostly based

on our own observations in the field, are described. Instructions for conservation and habitat management are

presented. The larva is described and illustrated in detail.

Introduction, material and methods

Ethmia pyrausta (Pallas, 1771) (Figs 1, 2) is one of the rarities in the European fauna of the sub-

family Ethmiinae. The species occurs sporadically in the hemiboreal zone in western and central

parts of the Palaearctic region. Most of the records are old and the species is considered to be

declining at least in its European distribution range. Although the larval host plant(s) and the flight

period of the adult are known, there are very scarce data available on the behaviour of larvae and

adults, as well as on the preferred habitats of the species.

During 2002-2006, the present occurrence and status of the populations of all protected Lep-

idoptera species was evaluated on the Aland Islands in the SW Finnish archipelago by Faunatica

Oy (Nupponen et al. 2007). One of the fifteen focal species was E. pyrausta , which was known

to have occurred on the islands, but no confirmed records of the species existed since the 1950s.

One abundant population was discovered in 2005 by Kari Nupponen in the central part of the main

island, where there were two studies of the larval behaviour (in July 2005 and 2006; Figs 3, 4). The

species was also recorded close to the southern coast of Aland, where a single male was observed

in early June, 2006, by Erkki and Leena Laasonen. Ene and Urmas Jürivete discovered another

abundant population of E. pyrausta from SE Estonia in 2007, and adult behaviour was studied

there in May 2008.

The description of the larva (below, Figs 5-10) is based on two caterpillars from Finström,

Aland Island (25.vii.2006, see Table 1). The larvae were preserved in ethanol in the field, and later

studied by Matti Ahola. The hypopharyngeal complex, mandibles and labrum of the larval head

48

Nupponen et al: Biology and distribution of the declining moth Ethmia pyrausta...

Figure 1. Adults of Ethmia pyrausta (Pallas, 1771) (Finland, Al: Finström, e.L, larvae found 25.vii.2006 on

Thalictrum flavum).

were dissected and mounted on slide. The chaetotaxy was studied from larvae in alcohol, and the

living larva (Fig. 4) was photographed to show the habitus. Naming of the setae follows Flinton

(1946) as interpreted by Ahola and Silvonen (2005).

Distribution

Ethmia pyrausta was described from the Samara region, the eastern part of European Russia, in the

1 8th century (Sattler 1967). There are two recent records of E. pyrausta from the steppes of the Ural

Mountains (Nupponen in press) and another one from Uljanovsk district (W. Mey, pers. comm.),

but apparently the species is very rare in the Volgo-Ural region. In Russia, the species is known

to occur widely but sporadically in the hemiboreal zone, from Karelia in the west to the Baikal

region in the east (Sinev 2008). It is also known from Mongolia (Ulan-Bator) and China (Kuldzha,

Xinjiang) (Dubatolov et al. 1997, Dubatolov 2014). In western Europe, E. pyrausta occurs only in

Scotland, Sweden, Finland and the Baltic countries.

In Scotland, E. pyrausta is restricted to the Highlands. It was known by a single specimen dis-

covered in May 1 853 on the banks of the River Shin, until two specimens were unexpectedly found

in 1996 in the Cairngorms (about 1000 m a.s.l.) (Anonymous 2014; Kimber 2014). Subsequently

several further specimens were found, one at Loch Vrotachan on the NNW end of Caimwell, Ab-

erdeenshire (810 m a.s.l., 28.V.2001), one at the River Averon close to Loch Morie, East Ross-shire

(8.V.2008), one on the slopes of Ben Griam Mor in 2012, and 15 specimens in Croick Estate (24.

iv.-31.v.2014) (Anonymous 2014).

In Sweden, E. pyrausta has declined severely. It has been recorded in eight provinces in the

central part of the country (Gustafsson 2012). However, only from two provinces, viz. Uppland

and Dalarna, are there rather recent records. Here the moth occurs along the River Dalälven, but no

records are known from the most recent years (Nils Ryrholm, pers. comm.).

In Finland, E. pyrausta occurs with certainty only on the Aland Islands, where it is apparently

declining due to habitat loss. Most records are from the 1940s and 1950s, and many populations

have vanished since (Nupponen et al. 2007). Since the 1950s, there are only confirmed records

from two localities. Additionally, there is a single record of the species from the southern coast

of the Finnish mainland (Helsinki, 1 larva, 1946; Hyönteistietokanta 2014), thus its occurrence in

southern Finland cannot be excluded.

Nota Lepi. 38(1): 47-58

49

Figure 2. Female of Ethmia pyrausta in resting posture (Finland, Al: Finström, e.l., larva found 25.vii.2006).

Figure 3. Moist meadow in Finström, central Âland Islands. Habitat of Ethmia pyrausta (photo: K. Nup-

ponen).

50 Nupponen et al. : Biology and distribution of the declining moth Ethmia pyrausta...

Figure 4. Larva of Ethmia pyrausta on Thalictrum flavum (Finland, Al: Finström, 19.vii.2005) (photo: K.

Nupponen).

In the Baltic countries, the species occurs sporadically in Latvia (Sulcs and Sulcs 1978, Savenkov

and Sulcs 2010) and Estonia (Nolcken 1871, Petersen 1924, Jürivete and Öunap 2008). There is also

one locality for the species in northern Lithuania close to the Latvian border (Povilas Ivinskis, pers.

comm.). The Finnish and Baltic records of E. pyrausta known to us are listed in Table 1 .

Description of the larva

Larvae of the genus Ethmia have a chaetotaxy generally typical of Lepidoptera, with one excep-

tion: D2 setae of abdominal segment 9 are laterad of D1 unlike other Gelechioidea, but similar

Nota Lepi. 38(1): 47-58

51

Table 1. Records of Ethmia pyrausta (Pallas, 1771) from Finland and the Baltic countries.

ESTONIA

LATVIA

52

Nupponen et al. : Biology and distribution of the declining moth Ethmia pyrausta...

LITHUANIA

to Cryptolechiinae (Kaila 2004, Heikkilä et al. 2014). Setae Al-3 and LI on head form a nearly

straight line, and secondary setae are present on abdominal SV groups, including both the prolegs

and the 9th abdominal segment. E. pyrausta differs from other ethmiine species by having second-

aries only on 9th abdominal segment.

Head morphology: Head semiprognathous, rather rounded, surface smooth but not shining,

frontoclypeus slightly longer than epicranial suture, adfrontal suture joined to epicranial suture

before vertical notch. Six stemmata present on each side, nearly equal in size but stemma 2 slightly

smaller, stemmata 5 and 6 in line with caudal margin of antennal socket. Spinneret tubular, taper-

ing distad and proximad, about three times as long as wide. Labial palpi slender, segment Lpsl two

times longer than wide, seta Lpl about twice as long as segment Lps2, seta Lp2 as long as Lpsl

(Fig. 5). Stipular setae shorter than Lp2 of labial palpi, position on chitinised part of prementum.

Hypopharynx largely bare, median and lateral parts of posterior region covered with tiny spines

(Fig. 6). Laciniogalea of maxillae with stout sensilla, Ss2 thicker than Ssl on galeal lobe and Stl

thinner than St2-3. Maxillary palpi with stout third segment, longer than second one (Fig. 7). Cut-

ting margin of mandible with very tiny ventral tooth and with straight and smooth edge of second

dorsal tooth. Other teeth unspecialized. Three inner ridges present on inner surface of mandible.

Labrum with low and rather large notch, seta LR1 situated on level with LR2, setae LR5 and LR6

separately on line with seta LR4 (Fig. 8).

Chaetotaxy: Position of PI setae on level with AF2 on head, distance Pl-Pl shorter than

P2-P2, setae Al, A2 and A3 situated straight on line. Setae D1 and D2 of prothoracic shield close

to each other, seta SD2 also on shield but SD1 not. Three L setae and two SV setae present on

prothorax; LI distinctly ventrad of L2 and L3. Thoracic segments Th2-3 have D1 and D2 setae

close to each other and SD1 close to SD2, all on same pinaculum, seta SV 1 on large pinaculum and

microseta MD1 also on pinaculum. Three L setae and two MSD microsetae without pinacula, one

additional sclerotized plate present behind D setae (Fig. 9). Abdomen has large pinacula separately

around D1 and D2 setae on segments Ab 1-7, position of seta D1 cephalad from D2 on segment

Ab9, setae SD1 and SD2 on same pinaculum on segments Ab 1-8 and larger pinacula around setae

Nota Lepi. 38(1): 47-58

53

Figure 5-8. Ethmia pyrausta: 5. Morphology of mouthparts, spinneret and labial palpi in dorsal view; 6. Hy-

popharynx from dorsal view; 7. Maxillae with maxillary palpi and sensilla of galeal and lacinial area; 8.

Labram and left mandible (scale bar = 0.1 mm).

54

Nupponen et al: Biology and distribution of the declining moth Ethmia pyrausta...

L3 and VI on segments Ab 1-7. Setae LI and L2 situated close to each other on segments Ab 1-9,

seta L3 present also on segment Ab9. Small pinaculum around seta L2 on segments Ab 1-5. Three

SV setae present on segments Ab 1-6, two SV setae on segments Ab 7-8 and one long SV and nu-

merous secondary setae on segment Ab9 (Fig. 10). Anal shield with D1 setae on level with SD2,

setal distance D2-D2 longer than D2-SD1 and small spines present between setae D2-D2 (Fig.

9). Seta D2 long on abdominal segments Abl-Ab9 but seta SD1 longer on anal shield. Crochets of

abdominal prolegs biordinal in mesal penellipse.

Larval habitus: Head smooth with pale green postclypeus, adfrons, dorsal part of frons and

narrow stripe from adfrons behind stemmata; head otherwise black. Sides of prothoracic shield

and pinacula of body black. Broad orange flecks in place of middorsal and spiracular lines, dorsal

zone between middorsal line and D2 setae dark greenish especially on thorax, but larva otherwise

dull white.

Notes on the biology

The habitats of E. pyrausta are open and sunny moist meadows, often located at the shore or

riverside (Fig. 3). Ovipositing females apparently prefer microhabitat with rather sparse, lower

vegetation and warmer microclimate than in the adjacent grassy areas. Usually such a habitat exists

as a narrow belt between forest and dense stands of Salix or Phragmites. The species has never

been found in forests, even in localities where the host plant is abundant in semi-shadowed open

patches within the forest. A common feature for localities of E. pyrausta is that they are open to

the southeast or east, and sunshine reaches the spots in the early morning.

The larva is oligophagous on Thalictrum species (Ranunculaceae). In Finland, the only recorded

host plant is Thalictrum flavum L. (Fig. 4), probably due to the fact that other species of Thalictrum

do not occur or are very rare in the region where E. pyrausta occurs. In Estonia, larvae have also been

found on Thalictrum aquilegiifolium L. (Nolcken 1871, Petersen 1924) and T. lucidum L. (E. & U.

Jürivete, pers. comm.). In an average season, larvae are of detectable size from late June and they

pupate in the first half of August. They feed on flower-buds, flowers and seeds. Full-grown larvae

feed also on leaves, but only when all seeds are eaten up. Larvae live singly and freely on the host

plant, although sometimes two to three larvae have been observed on one plant. Contrary to what is

stated in the literature (e.g. Emmet 1979) we did not detect any webs made by larvae on the plants.

The presence of larvae can be presumed from the evidence of partly eaten seeds. However, larvae of

some other Lepidoptera feed on seeds of Thalictrum too, and sometimes they occur sympatrically with

E. pyrausta , e.g., the geometrid Gagitodes sagittatus (Fabricius, 1787). Therefore, the occurrence of

E. pyrausta should always be confirmed by direct observation of a larva, not just by feeding damage.

Larval behaviour was studied three times: twice on the Aland Islands (19. vii. 2005 and 25.vii.2006)

and once in the south-eastern Estonia (24.vi.2007). The larva is predominantly nocturnal. On Aland,

three larvae were observed on 19.vii.2005 at 6 p.m. and 25 larvae from 1 1 :30 p.m. to 00: 15 a.m. (lo-

cal summer time, i.e. +3 h GMT). On 25.vii.2006 in the same locality, there were no signs of larvae

earlier in the day (3^1 p.m.), while about 1 50 almost full-grown larvae were observed at night from

1 1 :30 p.m. to 1 a.m. In SE Estonia, altogether more than 20 larvae of various ages were observed on

24.vi.2007 at dusk. Larvae become active at dusk, and climb onto the host plant to eat seeds. They

eat during a rather brief period (maximally half an hour), and then return to hide in the litter. The

larvae move rapidly and drop onto the ground very easily when disturbed. Later at night they are less

Nota Lepi. 38( 1 ): 47-58

55

Figure 9. Ethmia pyrausta : Chaetotaxy of head, thorax and anal shield, and left stemmatal ring.

Figure 10. Ethmia pyrausta : Chaetotaxy of abdomen.

56

Nupponen et al. : Biology and distribution of the declining moth Ethmia pyrausta...

active, possibly due to decreasing temperature and especially fog that often forms in such habitats

at night. During the second half of the night and daylight, larvae are mainly hiding in the soil, and

only occasionally visit their host plants to feed. Nolcken (1871) detected larvae on their host during

daylight, but he did not present further notes on the time of the records. Pupation takes place in a

dirty white or pale yellow cocoon in detritus on the ground. We did not rear any parasitoids from the

larvae. It is also possible that the behaviour of parasitized larvae changes and, thus, they cannot be

observed with the same methods.

The main flight period is in May with a peak about one to two weeks after budburst of birch. In

years with a late season and close to the seashore, the flight period starts later and extends even to

mid-June. Nolcken (1871) recorded adults in the period 28 April to 6 June (in 1865-1867), but the

annual flight period did not last more than two weeks. The adult of E. pyrausta is predominantly

diurnal. Occasionally, males fly at night too and come to light, usually on extremely warm nights.

In south-eastern Estonia, the behaviour of adults was studied on 12.V.2008. The previous night was

cold and the temperature decreased to +1°C in the second half of the night. In the morning, the sky

was clear and sunshine heated the wet vegetation from early morning. The first E. pyrausta male

was observed at 7:30 a.m., while the temperature was still low. Most males became active just after

8 a.m., and over 20 individuals were observed between 8 a.m. and 9 a.m. After that, flight virtually

stopped. During the active flight period, males were flying rather slowly and close to the ground

(height of flight about 1 m), probably searching for females. Sulcs and Sulcs (1978) recorded sim-

ilar ‘swarming’ of E. pyrausta in early morning. Males appear to re-activate at the middle part of

the day between 10 a.m. and 2 p.m., when the moths fly rapidly, straighter and higher (height of

2.5-3 m) than in the morning (Nolcken 1871; U. Jürivete unpublished; E. Öunap pers. comm.). It

is difficult to observe rapidly flying dark moths, and because of that, the species is seldom recorded

by chance. As far as we know, there are no records of an evening flight of E. pyrausta. Females do

not seem to fly much, but prefer to sit among the vegetation and presumably attract males.

Conservation

Ethmia pyrausta shows a highly sporadic distribution throughout its known range. It has apparent-

ly declined at least in the western parts of the range. For example, the occupancy of E. pyrausta

was systematically studied in 34 patches of T. flavum - including traceable previous findings -

throughout the Aland Islands in 2005 and 2006, but it was present in only one open and sunny

patch (Fig. 3) (Nupponen et al. 2007). On the other hand, the population in that patch was relatively

large. Based especially on the negative trends in various habitat characteristics, e.g. decreasing

amount and quality and high degree of fragmentation, the species has been rated as threatened both

in Finland (Kaitila et al. 2010; CR: criteria Bl ab(iii)c(iv)+2ab(iii)c(iv)) and Sweden (Bengtsson et

al. 2010; EN: criteria B2ab(i,ii,iii,iv,v)).

Ethmia pyrausta requires host plants that are growing in full sunshine. Therefore, the main reason

for the decline, at least in Finland and Sweden, is overgrowing of moist meadows after cessation

of grazing. All management activities should be performed late in the season, i.e. in August at the

earliest. This should ensure that E. pyrausta larvae have time to pupate before management starts.

Further, Thalictrum is highly vulnerable to grazing (Anonymous 2014; own observations) and only

late-season grazing or mowing can be recommended. Unfortunately, such late-season management

has almost ended in Sweden and Finland, even though plenty of herbivorous insect species are highly

Nota Lepi. 38(1): 47-58

57

dependent on it (Dahlström et al. 2008). In management, the first thing to take care of is to create

and maintain open sunny patches of moist meadow with plenty of host plants. Habitats occupied by

E. pyrausta are regularly dominated by Filipendula ulmaria (Rosaceae), and usually the main aim

is to reduce its abundance by mowing. If meadows are mown, only areas without Thalictrum should

be cut. However, in meadows that are grazed earlier than recommended, F. ulmaria protects plants

growing among it against grazing, so reasonable amounts of F. ulmaria in microhabitats with Thalic-

trum are beneficial in those cases (Anonymous 2014).

Acknowledgements

We thank the following colleagues for various help in preparing the present article: Bengt Â. Bengtsson (Färjestaden,

Sweden), Pavel Gorbunov (Ekaterinburg, Russia), Po vilas Ivinskis (Vilnius, Lithuania), Ene Jiirivete (Tallinn, Estonia),

Aleksander Lagunov (Miass, Russia), Elena Nupponen (Espoo, Finland), Timo Nupponen (Espoo, Finland), Vladimir

Olschwang (Ekaterinburg, Russia), Nils Ryrholm (Gävle, Sweden), Nikolay Savenkov (Riga, Latvia), Kimmo Silvonen

(Espoo, Finland), Ivars Sulcs (Riga, Latvia). Our thanks are also due to Lauri Kaila (Helsinki, Finland), Wolfram Mey

(Berlin, Germany) and Erik van Nieukerken (Leiden, Netherlands) for constructive comments on the manuscript. Âlands

landskapsregering (provincial government) supported our research on Aland Islands by permits and funding.

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highlands.html [accessed 8.L2015]

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G, Ryrholm N, Söderström B (2010) Fjärilar-Butterflies and moths-Lepidoptera. In: Gärdenfors U

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Nota Lepi. 38(1) 2015: 59-74 | DOI 10.3897/nl.38.8957

Multivariate indices as estimates of dry body weight for comparative

study of body size in Lepidoptera

Enrique Garci'a-Barros1

1 Departamento de Biologia, Facultad de Ciencias, Universidad Autônoma de Madrid, Campus de Cantoblanco,

C/ Darwin, 2, ES-28040 Madrid, Spain; garcia.barros@uam.es

http: //zoobank. org/4E9B42FA -BA 76-4 745-AFC7-E3DD9EC9 70E8

Received 16 November 2014; accepted 8 April 2015; published: 30 April 2015

Subject Editor: Jadranka Rota.

Abstract. Comparative studies on the size of adult Lepidoptera (moths and butterflies) frequently rely on

single linear estimates of body size, namely of forewing length or wingspan. As the shape of the wings of

these insects - in fact, of all body parts - differs from one taxon to another, such estimates of body mass may

not be adequate for comparisons across a wide taxonomic range. Using the length and width of the forewing,

thorax and abdomen, as well as the wing area of 375 species and their correlations with dry body weight,

several composite indices were determined that might be used in different circumstances. As the coefficients

of determination from the multivariate regression models were rather high (R2>0.96), the results are believed

to be reliable. A critical re-evaluation of the results indicates that important variations in the regression slopes

described here would be expected, if at all, only from species with unusual body shapes. Incidentally, the

bivariate relationships are in agreement with former comparative work on Lepidoptera and other terrestrial

insects in that the relationship between body weight and single linear measurements follows a slightly nega-

tively allometric trend, implying comparatively lighter bodies at the largest body sizes and relatively heavier

ones at the shortest body sizes.

Introduction

As one of the hyper-diverse insect taxa, the order Lepidoptera is well suited for comparative work

on subjects of broad biological relevance such as the evolution of body size and its correlation with

other traits (e.g., Nilsson and Forsman 2003; Simonsen and Kristensen 2003; Allen et al. 201 1;

Ribeiro and Freitas 201 1; Symonds et al. 2012). This requires an estimate of body size that is valid

across distantly related subtaxa, as a broad taxonomic coverage would be of interest for recovering

long-term evolutionary trends or patterns.

Although body mass, or weight, is generally accepted as an accurate measure of size for Lepi-

doptera (e.g., Miller 1977), adult body weight has been rarely used in comparisons across species,

and if so, only within a relatively narrow taxonomic framework (e.g., Agosta and Janzen 2005; Da-

vis et al. 2012). In fact, the published data on body weight cover a small number of the known moth

and butterfly species. This is largely due to the practical difficulties of obtaining live (fresh) adults

from a wide array of taxa and geographic regions for weighing in standard conditions. Most often,

the adult size of these insects has been estimated in one of two ways, depending on the purposes

of the study. The first consists of using body length or an alternative linear measure (such as head

60

Garcia-Barros: Multivariate indices as estimates of dry body weight...

width) to estimate body mass, based on the generally good correlations between those measure-

ments and fresh or dry body weight across large numbers of species of invertebrates (Sample et al.

1993; Hödar 1996 and references therein). This approach is frequently utilized in ecological stud-

ies on e.g. biomass production or on the diet of insectivore vertebrates (Hödar 1997; Heyman and

Gunnarson 2011; Legagneux et al. 2012) as well as in fresh water ecology (Benke et al. 1999). The

second context is that of ecological or evolutionary work on the Lepidoptera based on interspecific

comparisons of one linear measurement of the adult wings (generally well correlated to adult body

weight: Nylin et al. 1993; Miller 1977, 1997). Here, the most popular metrics are wingspan (the

distance between the tips of the forewings of a set specimen, or twice the distance between the tip

of one of the forewings to the center of the thorax) and forewing length (e.g., Hawkins and Lawton

1995; Beck and Kitching 2007; Hamback et al. 2007).

Wings are the most relevant structure of these insects to the human eye, and there are good rea-

sons for wing size to be correlated with body mass for functional reasons, as Lepidoptera are flying

insects. However, some degree of structural variation affecting the relationship between wing size

and body weight has been documented at several taxonomic levels including the intra-specific one

(Van Dyck et al. 1997; Tiple et al. 2009; Shreeve et al. 2009; Symonds et al. 2012). As already

stated by Miller (1977), the broad body architecture is likely to differ markedly between the mem-

bers of distantly related taxa of similar body weights, so that more precise estimates of body mass

of species in varied taxonomic positions require a more elaborate combination of linear measure-

ments. It is conceivable that a multivariate approach based on several variables correlated with

body weight might achieve this purpose.

The main objective of this study was to determine a composite index based on several linear

estimates that could predict accurately the dry body weight of set specimens (e.g., from mu-

seum collections or even scale illustrations) irrespective of the species phylogenetic position.

The reason for selecting dry body mass instead of fresh body weight is of a practical nature:

because these insects are usually preserved as dried samples in scientific collections, the possi-

bility to test and re-elaborate any results is far more feasible than obtaining reliable fresh (live)

weights from the same set of species. The second objective was to determine the sensitivity of

such an index to sample size (the number of species), taxonomic diversity and morphological

heterogeneity as a means to measure its robustness (if it is to be applied to species different

from those used to fit it).

Methods

To avoid heterogeneity caused by the patterns of sexual dimorphism in adult size, the comparison

was restricted to adult males from any available source, totaling 665 individuals from 375 species

distributed among 61 families. The selection emphasized the diversity of size within and across

families and included samples from any region in the world that could be processed.

Measurements

The measurements were performed on dry set (pinned or spread), complete male specimens. When

fresh adults were available, these were first dried in the position traditionally used for these insects

in entomological collections. The measures described below were taken in one of four ways: (a)

Nota Lepi. 38(1): 59-74

61

Figure 1. Slightly idealized representations of three typical adult Lepidoptera (left to right: Lasiocampidae,

Hepialidae, Gelechiidae) to illustrate the variables measured. The right side of the thoraces is represented as

devoid of the scale cover to make more evident the limits of this tagma. The three drawings are scaled to the

same forewing length. Linear measurements are indicated by bars and areas by a striped pattern. FWL = fore

wing length, FWW = forewing width, F WA = forewing area, F1WA = hind wing area, TL = thorax length, TW

= thorax width, AL = abdomen length, AW = abdomen width.

under a stereomicroscope with an ocular micrometer, (b) on a digitized scale drawing made with

an optical camera lucida adapted to a stereomicroscope (x 10 to x 40), (c) on a digital photograph

of the specimen taken together with a standard scale bar, taken either with a macro lens (up to 1:1)

or on a photo microscope at low magnification, or (d) with a Vernier caliper (exceptionally in the

case of some of the largest moths). The program Image J (Rasband 2012) was used to measure the

digitized images.

Six linear measurements (in mm) were taken (Figure 1): thorax length (TL), thorax width (TW,

taking the point of insertion of the fore wings as a reference), abdomen length (AL) excluding

terminal hair pencils or protruding genital appendages, abdomen width (AW, taken at the midpoint

of the line represented by AL), forewing length (FWL, from the insertion of the wing on its costal

margin to its apex including the fimbriae) and forewing width (FWW, the distance between edges

following a line perpendicular to FWL at its midpoint). In addition, the area of the fore- and hind-

wings (including the fringes) were recorded (FWA, HWA, as mm1 2). The mean species values are

available as Supplementary material (Suppl, material 1 : nexus format text).

Repeated measures and replicates

To estimate the magnitude of error measurement, the mean within sample and mean within species

coefficients of variation were calculated after replicated measurements taken on each individual

and between individuals within species.

(1) Every measurement was taken twice for each specimen using two different methods among

those detailed above (most frequently a, b and c), on two different dates.

(2) Whenever possible two male specimens of approximately the same size (judged from wing-

span by naked eye) of the species were processed. However, replications were not always

possible as data from single representatives of a number of species were included if this con-

tributed to an increase in the taxonomic or geographic coverage of the species selection.

62

Garcia-Barros: Multivariate indices as estimates of dry body weight...

Dry body weight

The insects were dried to a constant weight at 60° for 48 hours (72 h for the largest specimens).

The pins, if present, were removed carefully (but see below). The weight of the whole specimen

was determined to the nearest 0.01 mg in a Mettler AT261 balance (species of wingspan of ca. 15

mm or above) or in a Mettler Toledo XP6 microbalance with precision of 0.001 mg (individuals

smaller than that size).

Pinned specimens

Although medium or larger sized collection specimens can generally be de-pinned and remount-

ed without much difficulty, there is always some risk of damage. For a small number of loaned

specimens (ca. 20 individuals) the weight of the pins was estimated, then subtracted from that of

the dry mounted specimen. Samples of 10 individual pins from four different brands and num-

bers (gauges): 000, 00, 0 and 1 to 6 (all with nylon heads and 37 mm long) were measured and

weighed. The weights were taken to the nearest 0.01 mg, and the widths measured with a precision

of 0.0179 mm under a binocular microscope with an ocular scale line. The relationship between

the log-transformed weights and widths was highly consistent: log10 (pin weight in mg) = 2.339 +

1.908 log10 (pin diameter in mm), R = 0.997, P < 0.0001, n = 350.

Small moths

The smallest moths (broadly corresponding to the heterogeneous assemblage of the “microlepi-

doptera”) posed some special difficulties, which handicapped the use of reference collections as

sources of size data. These moths are fragile and very likely to be damaged if treated in the way de-

scribed above, and even though they are frequently mounted on smaller pins (‘minutiae’, weight-

ing 0.69-3.15 mg for widths of 0.10 and 0.20 mm respectively) the small variation in the length of

these tiny metal pieces represents an excessive error in terms of the specimen dry weight. More-

over, as the genital pieces are of interest for identification, collection specimens frequently lack the

abdomen or a large part of it as it was removed for identification. Finally, most of them cannot be

easily identified to species level without expertise. For these reasons the data from several families

in this category were obtained from a small reference collection at the author’s department. This

hosts expert-identified specimens collected two decades ago at a single site, so new samples were

taken at the same location during 201 1-2012 to reasonably cover the lower part of the size range,

although at the cost of low geographic variation.

Multivariate models

All the variables were transformed to their decimal logarithms. This facilitated comparisons with

results from earlier research (as most size-weight relations have been modelled using the equa-

tion weight = a x sizeb: Reiss 1989; Ganihar 1997), linear-regression approaches as well as some

demands of the comparative method adopted (described below). After log-transformation, all the

variables fitted reasonably to the normal distribution with Kolmogorov-Smimov test values of d<

0.049, P > 0.05 in all instances (Suppl, material 2: frequency distribution graph).

The multivariate models were fitted using the General Regression Models module of Statistica

(StatSoft 2004). For model selection, a manual iterative forward-backwards procedure was adopted

to exclude redundant variables.

Nota Lepi. 38(1): 59-74

63

Independent contrasts and phylogenetic hypothesis

The method of phylogenetically independent contrasts (Felsenstein 1985; Harvey and Pagel 1991)

was used to control for phylogenetic effects. The contrasts were calculated using the software

PDAP:PDTREE (Midford et al. 2009) integrated in the package Mesquite (Maddison and Maddi-

son 2011). Branch lengths were set to equal length (1.00), and the polytomies were estimated as

single contrasts, which were calculated after the original output.

The working hypothesis on phylogenetic relationships was built according to the classification

proposed by van Nieukerken et al. (201 1), with the relationships above the family level adapted after

the tree topologies from Kawahara and Breinholt (2014) complemented by Regier et al. (2009, 2013),

Mutanen et al. (2010), Bazinet et al. (2013) and Martijn et al. (2014). Further information was gath-

ered from other recent literature (details available in Suppl, material 3: documentation on phylogeny).

In the absence of any other references, the formal classifications of Fauna Europaea (Karsholt

et al. 2013) for the European species and of the Lepindex database (Beccaloni et al. 2013) for

other geographic regions was adopted. The tree was assembled manually; preference was given

to the most recent results, or to those with the highest statistical support, but keeping any former

hypotheses if these have not been contradicted. Thus, except in face of conflicting evidence the

formal taxa at the levels of superfamily, family, subfamily and genus were adopted even when

their monophyletic status had not been corroborated in all instances. The tree topology and data

are available from the Suppl, material 4 and 1 (4: tree topology, 1: tree nexus format). The result-

ing dendrogram showed high resolution (ca. 77%), which of course is overoptimistic in terms of

strictly phylogenetic criteria.

Regressions were done through the origin to estimate the correlations and slopes. After a mul-

tivariate regression model was obtained, Least Squares Regression was used to estimate the inter-

cept for the working data set keeping the evolutionary slopes already obtained.

Robustness of the models

The number of species and of supraspecific taxa available for this study was obviously small if

compared to the estimated number of existing species in the order Lepidoptera (more than 150,000

species: van Nieukerken et al. 2011). Thus, one further question can be posed - to what extent are

the results presented sensitive to the addition of new taxa? The relationship between the errors in

the predicted weight data and the diversity in body size, morphology (excluding body weight) and

taxonomy were determined. The underlying idea is that any sources of diversity that are positively

correlated to large errors in the predictions should denote species’ features liable to modify signifi-

cantly the models obtained.

The error in the predicted dry body weight (DBW) values were measured as the mean of the

absolute values of the residuals from the two best fit models (described below) calculated for

randomly selected subsets of n species, where n = 5, 10, 25, 50, 100, 150, 200, 250, 300 and 350.

Forty replicates were taken at each n plus one more sample consisting of the whole data set. The

taxonomic and structural diversities of each of such 40 1 species samples were estimated using the

following attributes:

(a) Species diversity: the number of species in each sample.

(b) Variation in dry body weight: the standard deviation of the log-transformed dry body weights.

64

Garci'a-Barros: Multivariate indices as estimates of dry body weight...

(c) Structural variation. This variable was intended to account for structural/anatomical variation

as reflected by the measurements taken, irrespective of body weight. To do this, each of the

eight variables were regressed on body weight, one at a time. The residuals of such bivariate

regressions were used as the new variables, now linearly independent of body weight. Ap-

plying Principal Component Analysis to this set of residuals (Bartlett’s Sphericity test X2 =

344.24, P < 0.001; KMO index = 0.72) resulted in three components accounting for 66.96%

of the variance (respectively 41.51%, 14.59% and 10.86%). The standard deviation in these

three components (weighted by the respective contribution of each component) was used as an

index of structural (body shape) diversity, linearly independent from dry weight.

(d) Taxonomic/phylogenetic diversity. This was tentatively estimated in four alternative ways: (1)

Number of clades (absolute number of supra-specific nodes). (2) Phylogenetic diversity (PH):

the number of clades or nodes represented in the sample minus one, plus the number of species

as defined by Faith (1992), with all branches set to 1.00. (3) Relative Phylogenetic Diversity

(RPD, the number of clades above the species level divided by the number of species). And

(4) Taxonomic Distinctness (Clarke and Warwick 1998; Allen et al. 2009); this was calculated

using the software PAST (Hammer et al. 2001) after simplifying the number of taxonomic

categories to 10 which included the suborders, superfamilies, families, subfamilies and genera

plus five intermediate levels.

As the relationships between the mean residuals and these variables tended to be asymptotic

rather than linear, the bivariate and multivariate regressions were performed using Generalized

Regression Models and the logarithmic link function.

Results

Size range

The dry body mass of the selected species covered a range of variation of nearly five orders of

magnitude, from 0.03 mg to more than 2 g, corresponding to forewing lengths of between 1.8 mm

and 1 10 mm (see Suppl, material 2 and 5; 2: frequency distribution; 5: mean by superfamily). The

lightest and smallest species belonged to the genus Stigmella (Nepticulidae, with one male weight-

ing 0.034 mg), while two males of the reputedly longest-winged moth, the Erebiidae Thysannia

agrippina (Cramer, 1776) (see e.g. Kons 1998) had dry weights of 916-1,300 mg and one male

of the Satumiidae Attacus atlas (L., 1758) weighed 1,126 mg. However the heaviest specimen

weighed belonged to the hawk-moth family ( Cocytius sp., Sphingidae, which exceeded 2. 1 grams).

The replicated measurements (Table 1) suggested that the forewing and thoracic linear dimen-

sions may reflect lower proportions of error than the abdomen length or width measurements when

taken of the same specimen. Although the estimates between pairs of individuals from the same

species differed to some extent, it was clear that the highest amount of variation was accounted for

by the abdomen data. Forewing length appeared to be even more constant than the thorax meas-

urements within individuals. This might reflect a bias in the observer’s abilities, although it is also

likely that the reference landmarks to measure wing length (the tegulae and the tip of the wing) are

more obvious than the other reference structures, especially when the body is coated by a dense

cover of hair-like scales.

Nota Lepi. 38(1): 59-74

65

Table 1. Estimate of measurement error for dry body weight and six linear measurements, measured as a

percentage of the mean. The values given are the mean coefficients of variation (100-CV) (± 1 SD) averaged

across individuals (from duplicated measurements on each specimen, n = 662) and from different replicates

of the same species (within species, n = 328).

Table 2. Relationships between dry body weight and the test variables based on the species mean values,

estimated both by bivariate regression (left four columns) and in a multivariate regression model (right three

columns; intercept = -0.489, multiple R = 0.983, adjusted R2= 0.965). The ß values represent the relative

contribution of each variable in the multivariate model.

Bivariate regressions and preliminary multivariate regressions

The results from bivariate regressions of DBW on the other variables as well as the full multivar-

iate results (with all the variables in the model) are presented in Table 2 (species means, all R >

0.92) and Table 3 (independent contrasts, all R > 0.82). The effects of the linear estimates of wing

size (FWL and FWW), although significant in the bivariate comparisons performed on the species

data, were outweighed by those of the forewing area (FWA) in the multivariate approach. Across

the contrasts, FWL had a significant but negative effect in the regression models suggesting a com-

plex relationship between body weight and wing size and shape.

Multivariate regression model selection

Several alternative models fit by stepwise regression were calculated with multiple R values above

0.979 in all instances. Models 1 and 2 (Table 4; Figure 2) are those with the highest multivariate

R based in the species raw data and in the independent contrasts respectively. These two models

included the effects of wing area, which may be more difficult to measure in spread specimens.

However, because of their highest fits they were used as the basis for the last/next step. Several

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Garci'a-Barros: Multivariate indices as estimates of dry body weight...

Table 3. Relationships between dry body weight and the test variables based on the independent contrasts,

estimated by bivariate regression (left three columns) and by multivariate regression (right three columns;

multiple R = 0.914, adjusted multiple R 2 = 0.833). All regressions were forced through the origin (no inter-

cept). The ß values represent the relative contribution of each variable in the multivariate model.

Table 4. The two multivariate models with highest R scores among those fitted using the species mean val-

ues (1) and the phylogenetically independent contrasts (2). The statistics given are the coefficients of the

intercepts and slopes (Coeff.), ß values (relative contribution of each variable after standardization) and P

(significance). The multivariate statistics are represented at the base of the table. The regression based on the

independent contrasts was done through the origin (without intercept, statistics in the two bottom rows); the

intercept given (-0.553) was fitted a posteriori for the species values in the data set using the slopes (coeffi-

cients) stated.

Model statistics

alternatives (Suppl, material 6: alternative models) should allow estimations of DBW in circum-

stances that are frequent in entomological collections such as specimens without abdomen or with

its distal end missing due to identifications based in the external genitalia.

Robustness of the models

The regressions of the estimated error of the predictions (measured as the mean of the absolute

value of the residuals) on the indicators of taxonomic, size and structural diversity led to the same

Nota Lepi. 38(1): 59-74

67

Figure 2. Dispersion plots illustrating the fit (predicted on observed weights) of the two multivariate models

of highest R2 scores based on the raw species data (above) and the independent contrasts (below) (respective-

ly, models 1 and 2 in Table 4).

Table 5. Sensitivity of the best models to several sources of diversity in the species selected. Relationships

between the deviations of the predicted data (mean absolute residuals from 401 subsets of 5-375 species)

based on the multivariate models 1 and 2 (from Table 4) and several alternative estimates of structural di-

versity (number of species, taxonomic and phylogenetic diversity, morphology and body weight), estimated

through multiple regression. The contributions of the variables are represented in the upper (Coeff. = coeffi-

cient, Wald = Wald’s statistic) and the multivariate statistics in the lower rows. The Ordinary Least Squares

(OLS) R2 values calculated a posteriori for the two multiple regression models are given for comparison. PH

= Phylogenetic diversity, RPD = Relative Phylogenetic Diversity.

results in the bivariate and multiple tests, irrespective of the data analyzed (species values or inde-

pendent contrasts); thus, for simplicity, only the multivariate results are presented in Table 5. Only

two of the variables had significant effects with opposite signs: morphological diversity (with a

positive coefficient) and the relative phylogenetic diversity (with a negative effect).

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Garcia-Barros: Multivariate indices as estimates of dry body weight...

Discussion

The results generally show high correlations between all linear dimensions of the Lepidopteran

body, or the wing areas, and total dry body weight. This is not surprising given the relatively

important range of sizes covered and, especially, because a functional link between the variables

measured and total body size should exist in insects that must be able to fly effectively such as the

male specimens of moth and butterfly species studied.

The results are consistent with the fact that the wings of Lepidoptera are thin structures (thus

relatively light even if comparatively broad and evident) while the largest proportion of the body

weight is determined by the weight of the main thoracic and abdominal structures. Forewing length

is a popular estimate of body size in butterflies and moths as it is easier to measure than other body

dimensions. However, this measure has by itself a lower predictive power of dry body weight than

the thoracic dimensions (length and width) or, depending on the method used, abdomen length.

Thus, wingspan, taken as the distance from the midpoint of the thorax to the tip of the forewing,

would in theory be more accurate than the length of the wing alone as it would partly account

for thorax width. However, as stated by Miller (1977) the estimate of ‘wingspan’ most widely

used in the specialized literature is the distance between the tips of the two forewings, where the

spreading technique is a potential source of error. Alternatively, some of the body dimensions,

especially the abdomen width, tend to be measured with lower accuracy than wing size. In spread

collection specimens, the abdomen is frequently deformed and contracted to different degrees, and

measurements made on the thorax may be hindered by the dense scale/hair clothing of some of

these insects. Under these circumstances a composed ‘body size index’ appears to be a practical

alternative measurement to body weight, particularly when different species are to be compared.

For the linear measurements that are more directly related to body length, such as the thoracic

and abdominal lengths, the slopes determined across the species means (2. 7-2. 8, see Table 2) are

exactly in the same range as those found for the relationship between body length and dry mass in

terrestrial and aquatic insects on a wider taxonomic scope (2.6 to 2.9: Rogers et al. 1976; Schoen-

ert 1980; Bugherr and Meyer 1997; Benke et al. 1999), or within the order Lepidoptera (Ganihar

1997). Hödar (1996) obtained slopes in the range 2. 8-2. 9 for the regressions of body weight on

head width for butterflies and moths. This supports the idea that dry body mass correlates to single

linear measurements such as body length following a slightly negative allometric trend (that is,

with a slope slightly below 3.0 which would be expected for the volume to length ratio), at least

if estimated by Least Squares Regression. Values of the slope based on the independent contrasts

tend to be more conservative (Table 3). However generalizing on these grounds remains difficult

since single linear surrogates of body weight may well vary among taxa (e.g. from 2.1 to 2.9 be-

tween two families of Lepidoptera; Miller 1977, 1997).

Among the several drawbacks of the present results is the fact that intraspecific variation has not

been controlled for, and cannot be distinguished from other sources of error. This may be accept-

able under the assumption that intraspecific variation in body weight is generally higher than inter-

specific variation for the same trait. Given this and the widespread phenomenon that intraspecific

allometric trends follow different (generally less steep) slopes than the interspecific trends in ani-

mal taxa (e.g. Harvey and Pagel 1991), one corollary is that the body mass indexes presented here

are probably not suitable for determining dry body weights accurately within a species. One further

Nota Lepi. 38(1): 59-74

69

limitation of the results presented concerns the estimation of dry body weight in living or fresh (not

dried) adults of Lepidoptera, because all the body parts experience some degree of contraction after

drying (including the wings; Van Hook et al. 2012); these effects are especially noticeable in the

abdomen. In such cases, a suboptimal model (Suppl, material 6: alternative models) could be used

as an approximation, or alternatively the bivariate relationships of body weight to forewing length

or area as given in Table 2.

Of course, it is likely that the predictive accuracy of the regression models selected can be

improved by spreading the selection of species. The results in Table 5 suggest that this would

neither be achieved simply by increasing the number of species compared nor by broadening their

variance in body weight; instead, it seems that the amount of error in the predictions is primarily

correlated with the proportion of morphological diversity of the species compared (irrespective of

their body weight) relative to their phylogenetic diversity. In other words, the results may be rela-

tively stable unless for species selections featured by extreme variations in wing and body shape,

from subtaxa of Lepidoptera not represented in the sample analyzed.

Although the comparative method of independent contrasts is statistically robust in the absence

of accurate estimates of branch lengths, the contrasts are calculated by dividing the differences be-

tween each pair of values at a node by the estimated evolutionary distances (derived directly from

the branch lengths; Felsenstein 1985). This is a source of uncertainty when the precise value of the

regression slopes is of interest. Further, the overall value for the slope of a relationship within a

large taxon may represent, in some instances, the average of several slopes featuring the different

subtaxa (e.g., for butterflies: Garcia-Barros 2002). Thus, although the formulae derived from the

independent contrasts might be suitable for the estimation of dry body weight in species from taxa

not prospected in this work, it may be subject to criticism and re-evaluation. The fact that their

fit to the data was slightly lower than that based on the raw species data may simply reflect some

degree of over-sampling on closely related species, but on the basis of the results and for species

similar to those selected preference is given to model 1 (Table 4), or alternatively to models 5 and

6 (presented in Suppl, material 6: alternative models).

Conclusion

The fact that the multivariate approaches presented here showed high R2 scores (> 0.94) for a much

wider range of size, morphology and taxonomic variety than that in any former comparable study

on Lepidoptera suggest that, although liable to be refined, they may represent a useful tool for

comparative work when a wide taxonomic scope is necessary.

Acknowledgements

I wish to thank Pascual Torres (SIDI, Universidad Autonoma de Madrid) for weighing most of the smallest specimens and

Mercedes Paris (Museo Nacional de Ciencias Naturales, Madrid) for the loan of selected specimens. Juan Pablo Berrocal

assisted during the initial stages of the study. Most problems related to the identification of the samples would not have been

resolved without the help of several colleagues, namely Antonio Vives Moreno, Gareth E. King, Joaquin Baixeras, José-Lu-

is Yela and Elisenda Olivella. Thanks are also due to D. Molina for his samples of Lepidoptera from Peru and Ecuador.

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Garcia-Barros: Multivariate indices as estimates of dry body weight...

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taxonomic affiliation, butterfly, and nectar flower morphology. Journal of Natural History 43: 855-884.

doi: 10.1080/00222930802610568

Van Dyck H, Matthysen E, Dhont A (1997) Mate-locating strategies are related to relative body length and

wing colour in the speckled wood butterfly Par arge aegeria. Ecological Entomology 22: 116-120. doi:

1 0. 1 046/j . 1 365-23 1 1 . 1 997.0004 1 .x

Van Hook T, Williams EH, Brower LP, Borkin S, Hein J (2012) A standardized protocol for ruler-based mea-

surement of wing length in monarch butterflies, Danaus plexippus L. (Nymphalidae, Danainae). Tropical

Lepidoptera Research 22: 42-52.

Nota Lepi. 38( 1): 59-74

73

Supplementary material 1

Nexus format text.

Authors: Enrique Gracia-Barros

Data type: Adobe PDF file

Explanation note: Tree topology for the phylogenetic hypothesis adopted, to be used as input in applications

reading nexus (requires some slight previous edition).

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

Supplementary material 2

Frequency distribution graph.

Authors: Enrique Garcia-Barros

Data type: Adobe TIF file

Explanation note: Frequency distribution of the dry body weight data (mg) across the species studied.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

Supplementary material 3

Documentation on phylogeny.

Authors: Enrique Garcia-Barros

Data type: Adobe PDF file

Explanation note: This is a list of references including the most relevant sources of information used to build

the hypothesis on phylogenetic relationships which were not quoted in the main text.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

Supplementary material 4

Tree topology.

Authors: Enrique Garcia-Barros

Data type: Adobe PDF file

Explanation note: Graphic display (dendrogram) to show the hypothesis on phylogenetic relations adopted in

this work, after the sources quoted in the main texta and in the file: Supplementary material 3.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

74

Garcia-Barros: Multivariate indices as estimates of dry body weight...

Supplementary material 5

Mean by superfamily.

Authors: Enrique Garcia-Barros

Data type: Adobe PDF file

Explanation note: Mean dry body weight and wing length by superfamily, and sample sizes.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

Supplementary material 6

Alternative models.

Authors: Enrique Garcia-Barros

Data type: Adobe PDF file

Explanation note: Alternative or suboptimal regression models derived from the species means or from the

independent contrasts.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.

org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to

freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the

original source and author(s) are credited.

Nota Lepi. 38(1) 2015: 75-88 | DOI 10.3897/nl.38.4621

Description of immature stages of Xestia brunneopicta

(Matsumura, 1925), with a key to the mature larvae of the European

species of Xestia ( Pachnobia ) (Lepidoptera, Noctuidae)

Matti Ahola1, Martti Kuisma2, Reima Leinonen3, Hannu Saarenmaa4,

Kimmo Silvonen5

1 Metsänreunantie 27 G, FIN-85900 Reisjärvi, Finland; mj .ahola@kotinet.com

2 Annalantie 107, FIN-14610 Lepaa, Finland; martti.kuisma@pp3.inet.fi

3 Rauhalantie 14 D 12, FIN-87830 Nakertaja, Finland; reima.leinonen@kajaani.net

4 University of Eastern Finland, SIB Labs Digitarium, P.O. Box 111, FIN-80101 Joensuu; hannu.saarenmaa@uef.fi

5 Pronssitie 28, FIN-02750 Espoo; silvonen@kolumbus.fi

http: //zoobank. org/53BD 7012-FBB9-4 708-BC9 7-88C4B35BFD44

Received 29 January 2015; accepted 16 March 2015; published: 12 May 2015

Subject Editor: Alberto Zilli.

Abstract. Immature stages of Xestia brunneopicta (Matsumura, 1925) are described and illustrated from an

ex ovo rearing. The female was collected during a Finnish-Russian expedition to the province of Chita in East

Siberia in 2013. Eggs were laid in a plastic jar at Chara Sands on the 7th of July. Larvae hatched between the

20th and 21st of July. Rearing of larvae was undertaken in Finland by four lepidopterologists. A key is given

that includes the known European larvae of the subgenus Pachnobia Guenée, 1852 sensu Lafontaine et al.

(1998), mostly based on the morphology and larval chaetotaxy. The closest relatives on the basis of larval

morphology are discussed.

Introduction

The early stages of Xestia brunneopicta (Matsumura, 1925) have remained undescribed in spite

of its wide distribution from Magadan to the East-Siberian Tuva in Russia (Kononenko 2005) and

rarely also in Kuusamo, Finland (Mikkola et al. 1989). Ahola and Silvonen (201 1) described larvae

found in nature in 1 982 (Kuusamo) and 1 992 (Kuusamo, Kuhmo) as X. brunneopicta on the basis

of their differences in chaetotaxy and habitus compared to those of X. gelida (Sparre-Schneider,

1883). However, through the recent findings, we can now state that the larvae ofX. gelida and the

real X brunneopicta are very distinct from each other, particularly in their outer appearance, as

shown in this paper. The Finnish larvae formerly reported as Xestia brunneopicta are now re-iden-

tified as those of X gelida even though some of their characters differ slightly from other individ-

uals of A. gelida examined. Egg, larva and pupa ofX. brunneopicta are described and illustrated.

Pachnobia was described by Guenée, 1 852 as a genus and based on the type-species Pachnobia

carnea, a misidentification of Noctua tecta (Hübner, 1808) (Poole 1989). In Poole’s catalogue

(1989) Pachnobia was included as a synonym in the large genus Xestia Hübner, 1818. Lafontaine

(1998) arranged Xestia into four subgenera: Xestia , Megasema Hübner, 1821 .Pachnobia and Rad-

76

Ahola et al. : Description of immature stages of Xestia brunneopicta...

dea Alpheraky, 1892. Two subgenera, Anomogyna Staudinger, 1871 and Schoyenia Aurivillius,

1883, were included in Pachnobia , because of larval characters. We follow here this opinion, al-

though some other concepts have been published later (for instance Beck 1 999, Fibiger and Hacker

2005, Ahola and Silvonen 201 1). A key to the larvae of 18 European species of Xestia {Pachnobia)

is provided and it is mainly based on their morphology and chaetotaxy.

Materials and methods

The larval material originates from a female collected during an expedition to the province of Chita

in East Siberia, N56. 87133, E118. 18302, at an elevation of 750 m.a.s.l. on 7.VÜ.2013 by Hannu

Saarenmaa. Seventy-two eggs were laid on the needles of Larix gmelini Rupr. (Pinaceae) in a

plastic jar at Chara Sands 7-10 July. The rearing of eggs was carried out in Finland by HS and 60

larvae hatched after two weeks on 20-2 1 July. All larvae were reared under different lamps from

1 8 to 24 hours of light every day, because the northern larvae grow faster in continuous daylight.

Twenty larvae grew to maturity during autumn and went into diapause instead of pupating. We fol-

low Beck (2000) in descriptions of cuticular ornaments. Two larvae were prepared by dry inflating.

Larval chaetotaxy nomenclature follows Hinton (1946) while pupal follows Patocka and Turcâni

(2005). The hypopharyngeal complex, mandibles and labrum were dissected and preserved on a

slide to study the morphology.

Descriptions of immature stages

Egg (Fig. 6): The eggs were laid on the needles of Larix gmelini. Shortly after laying they were

whitish grey, but fertile eggs darkened in two days. The micropyle area became dark reddish

brown, and the narrow zone around the micropyle was reddish brown

Morphology of full-grown larva (Figs 1-5): Spinneret flat, 2.5 x as long as wide, ventral lip

straight, dorsal lip short-fringed, longitudinal grooves present on dorsal surface. Base segment of

labial palp (Lpsl) about 2x as long as wide, second segment (Lps2) 1/5 as long as Lpsl, labial

palp seta Lpl slightly longer than Lps2, seta Lp2 shorter than Lpsl (3/5) and 2x as long as Lpl.

Hypopharynx with long spines on anterior surface above spinneret, distolateral spines slightly

longer and stouter than distomedian spines, spines on lateroposterior part forming row of 8-10

differentiated, triangular teeth, lateral surface above this row densely covered with tiny spinules,

posterior area of medial part bare. Stipular setae below spinneret shorter than seta Lp2 of labial

palp, situated in front of prementum. Mandible with two setae on outer surface, six teeth on cutting

margin, three ridges on inner surface terminating in low protuberances before cutting margin and

triangular tooth on first ridge. Maxillary palp three-segmented, second segment longer than galeal

lobe, sensillum styloconicum of galeal lobe as long as end segment of maxillary palp, three sensilla

trichodea present. Labrum with low, rounded notch. Epicranial suture slightly shorter than height

of frons. Six stemmata present, distance between second (Oc2) and third (Oc3) stemma greater

than those between Ocl-Oc2 or Oc3-Oc4, distance Ocl-Oc2 greater than Oc3-Oc4. Abdominal

prolegs on abdominal segments 3- 6 (Ab3-6) equal in size, crochets uniordinal, 17-20 on Ab3,

22-24 on Ab6 and 26-28 in Ab 10. Body without warts or other protuberances.

Chaetotaxy resembling that of other members of Pachnobia'. Setae of head and body rather long

when compared to the height of the spiracle on Ab8: PI on head 3. 1-3.3 x, D2 on Ab2 1 .6-1.9 x,

D2 on Ab7 1.3-1.5x and D2 on Ab8 2.1-2.2x height of spiracle 8. Seta SD1 hair-like on thorax and

Nota Lepi. 38(1): 75-88

77

Figures 1-5. X. brunneopicta, morphology of larval mouth parts. 1. A. Scheme of spines of hypopharynx,

from left: distoanterior, distomedial, distoposterior, distolateral, lateral tooth, posterior dorsolateral, posterior

medial and lateroposterior spines. B. Spinneret and labial palpi in dorsal view. C. Stipular setae in frontal

view. 2. Hypopharynx in dorsal view. 3. Left maxilla in dorsal view. 4. Left mandible in oral face. 5. Labrum

in dorsal view. Scale 0.1 mm.

78

Ahola et al. : Description of immature stages of Xestia brunneopicta...

Table 1. Relevant distances between setae of larva of Xestia brunneopicta. Ab = abdominal segment.

Ab9, tonofibrillary platelet present below seta SD 1 on meso- and metathorax and two SV setae on

Abl . Relevant larval setal distances are presented in Table 1.

First instar larva: Length about 2 mm. Head pale brown, body pale greenish grey with small dark-

brown setal bases. Prolegs on Ab3-6 well developed, those on Ab3^1 much smaller (Fig. 7).

Second instar larva: Length about 4 mm. Head brown. Dorsal region greenish, ventral region pale

yellowish green, setal bases dark brown. Narrow, whitish dorsal and subdorsal lines present, shields

on thoracic segments 1 (Thl) and Ab 10 pale brown (Fig. 8).

Third instar larva: Length about 7-9 mm. Head brown, stripes visible but weak. Shields pale

brown, dorsal lines yellowish, not sharp-edged on shields. Dorsal zone green with white, narrow,

middorsal and subdorsal lines without darker margins, setal bases small, pale green. Subdorsal zone

dark olive green, spiracular line yellowish white, broad. Pleural and ventral zones pale green (Fig. 9).

Fourth instar larva (Fig. 10): Length 14-18 mm: Head brown, netfields visible but translucent,

frons and anterior zone green, brown setal points on brown bases. Thoracic and anal shields pale

brown, dorsal and subdorsal lines present as yellowish white flecks on prothoracic shield but absent

on anal shield. Body green on dorsal region with pale green or with whitish elements, yellowish green

between abdominal segments, dark green on ventral subdorsal zone. Middorsal line nearly white,

broad, continuous, subdorsal line slightly narrower, broken into spots. Spiracular line broad, yellow-

ish white, sharply bordered above dark green ventral subdorsal zone. Ventral region pale green. Setal

points black on dorsal region with dark green dorsal/whitish ventral bases.

Penultimate and last instar (Figs 1 1-12): Length of last instar larva 35^10 mm. Head brown, stripes

brown, reticulate structure with brown bands and pale brown, weakly visible netfields. Frons and

anterior zone greenish, adfrons brown, ocellar zone pale yellowish brown, setal points dark brown.

Prothoracic shield darker green than body, with narrow pale grey middorsal line, subdorsal line not

visible, shield caudally bordered with narrow, blackish grey colour. Anal shield greenish brown, lines

not visible, sutures brown, setal points blackish brown. Dorsal and ventral regions of body green,

middorsal and subdorsal lines white, narrow, short and broken, not visible on Ab9-10, both lines

with dark-green margins. Dorsal part of spiracular line visible, white, narrow, dorsally dark violet-green

border, ventrally no border. Dorsal and subdorsal zones mottled by small, white elements and longitudinal

violet-green colour elements; dorsal zone with diffuse wedge-shaped diamond figures; setal points of D1 and

Nota Lepi. 38(1): 75-88

79

Figures 6-8. Egg and small larvae of X. brunneopicta. 6. Egg on needle of Larix gmelini (Photo: M. Ahola).

7. The 1st instar larva on needle of Larix sibirica (Photo: M. Ahola). 8. The 2nd instar larva on needle of Larix

sibirica (Photo: M. Ahola).

D2 black with small white bases, microsetae MD 1 and MD2 on common large and white base on metathorax,

other MD1 bases also white but small. Spiracles yellowish with black edges. Thoracic legs with green coxae

and pale brown tibiae, prolegs green.

Pupa (Figs 13-14): Dark brown. Frons without tubercles or projections, labium and labial palpi visible,

proboscis exceeding caudal margin of Ab4, prothoracic femora visible, thoracic legs adjacent to antennae.

Abdominal spiracles narrow, Ab5-7 without elevated transverse ridge in front of spiracles, without transverse

row of spines or lateral spines; punctuation present close to bases of Ab4-7. Cremaster short, flat, quadrangu-

lar, with transverse dorsal and ventral furrows and three pairs of setae, D2 close together, short and hook-like,

D1 longer than D2, stout, LI short and stout, situated beside D2 on caudal margin of cremaster. Pupation after

hibernation without feeding in flimsy weak cocoon.

Observations on rearing and host plants

Larvae hatched on July 20 and 21 . They were in a plastic jar where they could choose between

plants Larix sibirica Maxim. (Pinaceae), Vaccinium myrtillus L. (Ericaceae) and Polygonum avic-

ulare L. (Polygonaceae). About 40 of the 60 larvae chose L. sibirica or V. myrtillus and the rest

chose P. aviculare. Five larvae died during the first week for unknown reasons. After about a week

the group of larvae was divided amongst four Finnish lepidopterists. It appeared later that young

80

Ahola et al. : Description of immature stages of Xestia brunneopicta...

Figures 9-12. X. brunneopicta larvae. 9. The 3rd instar larva on Andromeda polifolia L. (Photo: K. Silvonen).

10. The 4th instar larva on Larix decidua Miller (Pinaceae) (Photo: K. Silvonen). 11. Last instar larva with darker

brown head (Photo K. Silvonen). 12. Mature larva on Salix phylicifolia. (Photo: P. Puntila).

larvae could feed also on Poa annua L. (Poaceae) and Salix phylicifolia L. (Salicaceae). Full-

grown larvae were rather polyphagous in laboratory conditions, feeding also on Rubus idaeus L.

(Rosaceae),v4/ftws incana (L.) Moench (Betulaceae), Salix sp. (Salicaceae) and Lonicera xylosteum

L. (Caprifoliaceae). Larvae were mostly reared under a lamp.

Notes on natural environments

The collecting site of the female in Chara Sands is a peculiar dune habitat with occasional springs,

bogs and coniferous tree patches. It does not represent the typical habitat of the species. During

the trip (3-12 July) other localities near Chara were investigated, for which another article on the

results of the expedition is under preparation (Saarenmaa et al., in prep.). X brunneopicta was

common across all sites, but it seemed to be more frequent in the lowlands than in the mountain

valleys. Typical habitat for the species is forested bog with Larix gmelini (Fig. 15). However, it

was most numerous in low Salix and Alnus vegetation on the banks of the Chara River. Other spe-

Nota Lepi. 38(1): 75-88

81

Figures 13-14. X brunneopicta pupae. 13. Pupa in ventral view. 14. Cremaster of pupa in dorsal view (Pho-

tos: M. Ahola).

cies typical for these localities include several other Xestia ( Pachnobia ) species such as X. atrata

(Morrison, 1874), and also Folia altaica (Lederer, 1853), P. conspicua (A. Bang-Haas, 1912), P.

vespertilio (Draudt, 1934), P. vesperugo Eversmann, 1856, and the arctiine Borearctia menetriesii

Eversmann, 1846.

Key to the larvae of subgenus Pachnobia

Larvae of the subgenus Pachnobia differ from subgenera Megasema and Xestia in chaetotaxy and

morphology. Dorsal setae are long in Pachnobia , seta D2 on Ab8 is more than twice as long as

height of spiracle of same segment, whereas it is about as long as height of spiracle in subgenera

Megasema and Xestia. Also, setal distances P1-P2 (head) and VI -VI (Ab7) differ on average

(Table 2). Spinneret of larvae in subgenus Pachnobia is long, about 1.5-5. Ox as long as wide, flat

(except X liquidaria), dorsal margin with short fringes (without fringes in X liquidaria) and ven-

tral margin straight. Megasema and Xestia larvae have a short spinneret, 1 .0—1 .5 x as long as wide

(except 2-3 x in X collina, castanea and agathina), dorsal margin with longer fringes and ventral

margin more or less bilobed. Body colour and pattern vary a lot. Larva of X liquidaria is peculiar

with a tubular spinneret but long dorsal setae as in subgenus Pachnobia.

Larvae of European species of Xestia ( Pachnobia ) albonigra (Kononenko, 1981) andX. ( Pach-

nobia) thula Lafontaine & Kononenko, 1983 are still unknown, and are not included in the fol-

lowing key.

1 Netstructure of head negative, netfields darker than bands, spinneret tubular, distal

part of hypopharynx bare, without spines or granules, subdorsal line wide, white, spir-

acular line absent. Ratio of setal distance L1-L3/L1-L2 on Th3 varies 0. 9-1.0, mean

1.1, N = 2 and on Ab2 ratio of setal distances SD1-SD2/SD2-ST2 varies 4. 5-5. 3,

mean 4.9, N = 2 Xestia {Pachnobia) liquidaria (Eversmann, 1848)

Netstructure of head positive, netfields paler than bands, spinneret flat, distal part of hy-

popharynx covered with spines or granulated, lines of body vary. Ratio of setal distance

L1-L3/L1-L2 on Th3 varies 1.2-3. 3, mean 1.8, N = 146 and ratio of setal distances

SD 1-SD2/SD2-ST2 on Ab2 varies 1.1-4.3, mean 2.2, N = 137 2

82

Ahola et al.: Description of immature stages of Xestia brunneopicta...

Figure 15. Typical habitat ofX. brunneopicta and the other species mentioned in the article. Chara River with

richer vegetation on its banks is 100 meters to the left (Photo: H. Saarenmaa).

Table 2. Differences in chaetotaxy between larvae of Pachnobia and Megasema + Xestia. Ab = abdominal

segment, ST = spiracle.

2 Mandible without inner teeth or sometimes low swelling present on first ridge, transverse

cleft of hypopharynx not visible, setae very long, length of seta D2 on Ab8 more than 3x

height of spiracle on same segment (mean 3.4, range 2.2-5. 0, N = 27), X borealis differs

(2.2-2. 5) 3

Mandible with 1-2 triangular teeth, hypopharyngeal transverse cleft usually present, se-

tae shorter, length of seta D2 on Ab8 about 2x height of spiracle on same segment (mean

2.3, range 1. 4-4.2, N = 109),X. sincera differs (3. 5^1. 2) 9

3 Mandible with low swelling on first ridge, location of pore XDc close to seta XD2 on

prothoracic shield (ratio XD 1 -XDc/XD2-XDc ranges 2. 1-6.0, mean 3.3, N = 7), on anal

shield setal distance D1-SD2 longer than D1-D2 (ratio D1-SD2/D1-D2 ranges 1.1-1. 5,

Nota Lepi. 38(1): 75-88

83

mean 1 .3, N = 7). Head small, thorax tapered towards head, setal bases D1 and D2 as

white, sharp-edged full-bases 4

Mandible without swelling on first ridge, location of pore XDc more distant from seta

XD2 on prothoracic shield (ratio XDl-XDc/XD2-XDc ranges 1. 4-3.0, mean 2.1, N =

20), on anal shield setal distance D1-SD2 mostly shorter than D1-D2 (ratio D1-SD2/

D1-D2 ranges 0.8-1. 3, mean 0.9, N = 18). Thorax not tapering towards head, setal bases

D 1 and D2 yellowish, greyish or absent and not sharp edged 5

4 Dorsal zone pale violet whitish with darker arrow-chevron figure, subdorsal zone black-

ish, subdorsal line yellowish white ventrally bordered by black diagonal bands, spiracu-

lar line pale brownish beige, paler than greyish-brown pleural zone

Xestia ( Pachnobia ) fennica (Brandt, 1936)

Dorsal zone uniformly red-brown without arrow-chevron figure, subdorsal zone dark

red-brown dorsally, subdorsal line whitish broken to few flecks, spiracular line reddish

beige like pleural zone Xestia (Pachnobia) rhaetica (Staudinger, 1871)

5 Hypopharynx with row of long posterior lateral spines, about as long as distal lateral

spines, distance between ocelli Ocl-Oc2 shorter than that of Oc2-Oc3 (ratio Ocl-Oc2/

Oc2-Oc3 ranges 0.6-0. 8, mean 0.7, N = 7), seta PI on head shorter than 1 .5x seta D2 on

Ab2 (ratio P1/D2 range 1.2-1. 5, mean 1.4, N = 7), on anal shield distance Dl-Dl more

than 3x that of D2-D2 (ratio D1-D1/D2-D2 range 2.9-4. 1, mean 3.4, N = 7). Frons and

stripes black or blackish brown on head, coxae of thoracic legs black, subdorsal lines

yellowish on prothoracic and anal shields 6

Hypopharynx with row of shorter posterior lateral spines, shorter than distal lateral

spines, or without differentiated spines in this area, distance between ocelli Ocl-Oc2

about as long as that of Oc2-Oc3 (ratio Ocl-Oc2/Oc2-Oc3 ranges 0.8-1. 7, mean 1.1, N

= 13), seta PI on head longer than 1.5x seta D2 on Ab2 (ratio PI /D2 range 1. 5-2.0, mean

1 .7, N = 7), on anal shield distance Dl-Dl about 2x that of D2-D2 (ratio Dl-Dl /D2-D2

range 1 .2-2.9, mean 2. 1 , N = 12). Head with brownish or grayish frons and stripes, coxae

of thoracic legs paler, subdorsal lines whitish or grayish on shields 7

6 Spinneret about 1.5x as long as wide. Dorsal zone of larva darker reddish grey, black,

wedge-shaped flecks above subdorsal line wide, coming into contact with both D 1 and

D2 bases on Ab8, this line sharp also on shields, pinacula at bases of D setae visible only

on Ab9 Xestia (Pachnobia) laetabilis (Zetterstedt, 1839)

Spinneret about 2x as long as wide. Dorsal zone of larva pale grey with reddish tinge,

black wedge-shaped flecks above subdorsal line narrow, coming into contact only with

D2 bases, this line obscure, broken into spots on shields, pinacula at bases of D setae

present on Ab 1-9 Xestia (Pachnobia) distensa (Eversmann, 1851)

7 Differentiated posterior lateral spines absent on hypopharynx, seta Lp 1 of labial palpus

unusually long, about 4x length of second segment of labial palpus, seta PI on head

shorter than epicranial suture (ratio Pl/Es range 0. 8-0.9, mean 0.8, N = 3), setal distance

SD1-SD2 about 2x as long as distance between seta SD2 and spiracle on Ab2 (ratio

SDl-SD2/SD2-spiracle ranges 1. 8-2.1, mean 2.0, N = 3). Subdorsal zone lichen pat-

terned with blackish grey and whitish elements, spiracular line yellowish white, broad,

84

Ahola et al. : Description of immature stages of Xestia brunneopicta...

widely broken below spiracles by ground color and blackish

Xestia ( Pachnobia ) borealis (Nordstrom, 1933)

Differentiated posterior lateral spines may be weak but present on hypopharynx, seta

Lpl of labial palpus shorter, at most 2x length of second segment of labial palpus, seta

PI on head longer than epicranial suture (ratio Pl/Es range 1. 1-2.0, mean 1.5, N = 10),

setal distance SD1-SD2 about 3x distance between seta SD2 and spiracle on Ab2 (ratio

SDl-SD2/SD2-spiracle ranges 2. 3-3. 7, mean 2.8, N = 10). Subdorsal zone not lichen

patterned, spiracular line obscure or absent 8

8 Labial palpus with seta Lp2 as long as first segment, setal bases without pinacula on ab-

domen, number of crochets on Ab6 29-33, distance between setae D2-D2 short on anal

shield (ratio D1-D1/D2-D2 ranges 1.9-2. 9, mean 2.5, N = 5). Larva with prominent,

white middorsal and subdorsal lines on dorsal region, subdorsal line bordered by black

wedge-shaped dorsal flecks Xestia ( Pachnobia ) lyngei (Rebel, 1923)

Labial palpus with seta Lp2 shorter than first segment, setal bases with pinacula on ab-

domen, number of crochets on Ab6 1 8-20, distance between setae D2-D2 long on anal

shield (ratio D1-D1/D2-D2 ranges 1.2-1. 7, mean 1.5, N = 5). White dorsal lines narrow,

not prominent, and black wedge-shaped flecks absent on dark brown dorsal region

Xestia (. Pachnobia ) quieta (Hübner, 1813)

9 Skin of distal region of hypopharynx granulated with a few spines or bare, mandible with

two inner teeth, setal distance D2-SD2 2x as long as XD2-SD2 on prothorax (range

1 .7-2.8, mean 2. 1 , N = 15), length of seta D2 on Ab8 shorter than 2x as long as height of

spiracle (length D2 Ab8/height of spiracle ST8 varies 1 .4-2.0, mean 1 .7, N = 1 5). Frontal

stripe of head pale greyish brown, much paler than cervical stripe, and spiracular line of

abdomen absent or obscure 10

Skin of distal region of hypopharynx smooth and densely covered with spines, mandi-

ble with 1-2 teeth, setal distance D2-SD2 slightly longer than XD2-SD2 on prothorax

(range 1.0-2. 7, mean 1.4, N = 119), length of seta D2 on Ab8 longer than 2x as long as

height of spiracle (length D2 Ab8/height of spiracle ST8 varies 1. 7-5.0, mean 2.6, N =

121). Frontal stripe of head mostly of same colour as cervical stripe, and spiracular line

usually visible 11

1 0 Distance between setae D2-D2 on anal shield longer than height of spiracle of Ab8 (ratio

D2-D2/height of spiracle ranges 1.0-1. 3, mean 1.1, N = 6), setal distance Dl-Dl on anal

shield about 2x as long as D2-D2 (ratio D1-D1/D2-D2 ranges 1.7-2. 3, mean 2.0, N =

6). Larva dark reddish brown, middorsal line weak, whitish, mostly covered by blackish

brown margins Xestia (Pachnobia) alpicola (Zetterstedt, 1839)

Distance between setae D2-D2 on anal shield shorter than height of spiracle of Ab8 (ra-

tio D2-D2/height of spiracle ranges 0.7-0. 8, mean 0.8, N = 3), setal distance Dl-Dl on

anal shield about 3x as long as D2-D2 (ratio D1-D1/D2-D2 ranges 2. 7-3.0, mean 2.9,

N = 3). Larva yellowish brown, middorsal line whitish, more visible because of narrower

margins Xestia (Pachnobia) albuncula (Eversmann, 1851)

11 Mandible with two inner teeth, setal distance SD1-SD2 mainly less than 2x that of

SDl-spiracle on Ab2 (ratio SD1-SD2/SD1 -spiracle varies 1. 1-3.3, mean 1.8, N= 48),

X atrata with longer SD1-SD2 (range 2. 1-3.3, N = 5). Larva brown, subdorsal line not

touching bases of setae D2 12

Nota Lepi. 38(1): 75-88

85

Mandible with one triangular inner tooth, setal distance SD1-SD2 mainly more than 2*

that of SDl-spiracle on Ab2 (ratio SD1-SD2/SD 1-spiracle varies 1.4^1. 3, mean 2.3,

N= 44), X gelida andX. brunneopicta with short SD1-SD2 (1. 4-2.4, N = 16). Ground

colour varies, subdorsal line usually touching bases of D2 setae 14

12 nner tooth on first ridge of mandible quadrangular, distance between ocelli Ocl-Oc2 on

head longer than that of Oc2-Oc3 (ratio Ocl-Oc2/Oc2-Oc3 ranges 1.0-1. 4, mean 1.2,

N = 5), seta D2 closer to spiracle on Ab2 (ratio SDl-SD2/SD2-spiracle ranges 2. 1-3.3,

mean 2.7, N = 5). Dorsal and ventral regions of body of same color, spiracular line indis-

tinct Xestia ( Pachnobia ) atrata (Morrison, 1 874)

Inner tooth of mandible triangular, distance between ocelli Ocl-Oc2 on head shorter

than that of Oc2-Oc3 (ratio Ocl-Oc2/Oc2-Oc3 ranges 0. 5-1.0, mean 0.8, N = 44), seta

D2 more distant from spiracle on Ab2 (ratio SDl-SD2/SD2-spiracle ranges 1.1-2. 5,

mean 1 .7, N = 43). Ventral region of body paler than dorsal region, spiracular line visible

and bordered sharply against subdorsal zone... 13

13 Frontal stripe of head pale brown or pale greyish brown, paler than cervical stripe, bases

of setae D1 and D2 of abdomen ventrally yellowish white, spiracular line with yellowish

white dorsal part and mottled by reddish brown elements

Xestia ( Pachnobia ) speciosa (Hübner, 1813)

Frontal stripe dark greyish brown like cervical stripe, bases of D1 and D2 of abdomen

ventrally whitish, small spiracular line without differentiated white dorsal part and not

mottled by brown elements Xestia (Pachnobia) viridescens (Turati, 1919)

14 Larva green, mottled with small, white elements, without prominent pattern, dorsal, sub-

dorsal and spiracular lines white, narrow. Setal distance D1-D2 of Ab9 about Vi distance

D1-SD1 (ratio D1-D2/D1-SD1 range 0.5-0.7, mean 0.6, N = 2)

Xestia ( Pachnobia ) brunneopicta (Matsumura, 1925)

Larva not uniformly green, dark dorsal pattern present on body, lines variable. Setal

distance D1-D2 of Ab9 about as long as distance D1-SD1 (ratio D1-D2/D1-SD1 range

0.7-1. 3, mean 1.0,N = 42) 15

15 Spinneret long, more than 3x as long as wide and tapered apically; setal distance SD1-

SD2 less than twice as long as that of SD2-spiracle on Ab8 (except aequaeva 2.5-3. 1 x),

range 1 .4-3.1 , mean 1 .8, N = 19. Ground colour of larva distinctive dark grey with dorsal

pinacula or dorsal zone pinkish cream and subdorsal zone blackish 16

Spinneret shorter, less than 3x as long as wide with parallel sides; setal distance SD1

SD2 more than 2x as long as that of SD2-spiracle on Ab8 (except sincera 1.5-1. 9 x)5

range 1.5^1. 5, mean 2.5, N = 102. Ground colour of larva different 17

1 6 Larva dark grey with large black pinacula on dorsal region, spiracles black, number of

crochets on Ab 10 varies 17-24 (N = 2), setal distance L1-L3 on metathorax about 3x

that of L1-L2 (range 2. 6-3. 3, mean 2.9, N = 2)

Xestia (Pachnobia) aequaeva (Benjamin, 1934)

Pinacula absent on dorsal region, ground colour different, spiracles yellowish, number

of crochets on Ab 10 varies 26-43 (N = 19) setal distance L1-L3 about 1.5x that of Ll-

L2 (range 1. 2-2.0, mean 1.5, N = 19) on metathorax. Dorsal zone of larva pale pinkish

cream, white, usually only ventrad from seta D2 visible subdorsal lines with blackish

86

Ahola et al: Description of immature stages of Xestia brunneopicta...

dorsal border, subdorsal zone blackish brown

Xestia (. Pachnobia ) gelida (Sparre-Schneider, 1883)

17 Tiny spinules present distally on middle of posterior part of hypopharynx and partly

forming transverse rows, inner tooth of mandible small, seta SD1 on abdominal seg-

ments more distant from spiracle (on Ab8 ratio SD1 -spiracle/height of spiracle ranges

1.9-3. 5, mean 2.3, N = 5). Body blackish grey and whitish, lichen patterned, middorsal

line white, enlarged on posterior parts of abdominal segments, subdorsal line whitish,

enlarged towards seta D2, spiracular line broad, white with dark breaks below spiracles

and dorsally bordered by black wavy margin

Xestia ( Pachnobia ) sincera (Herrich-Schäffer, 1851)

Spines absent on middle posterior part of hypopharynx, inner teeth of mandible robust,

seta SD1 on abdominal segments closer to spiracle (on Ab8 ratio SD1 -spiracle/height of

spiracle ranges 0.9-1. 7, mean 1.3, N = 26). Body brown or yellowish-brown, not lichen

patterned, lines vary but spiracular line not prominent and its dorsal margin straight 18

18 Seta Lp2 of labial palpus short, 2/5 of length of base segment, spinneret slightly shorter

than 2x as long as wide, setal distance XD1-XD2 about 2x as long as XD2-SD2 on

prothorax (ratio XD1-XD2/XD2-SD2 range 2. 0-2. 5, mean 2.1, N = 10), on Ab2 dis-

tance SV1-SV3 longer than SV1-SV2 (ratio SV 1-SV3/SV 1-SV2 range 1.0-1. 9, mean

1.3, N = 10). Larva reddish brown, dorsal lines whitish, narrow, both bordered with

blackish fleck at anterior parts of segments, spiracular line yellowish white, sharp edged,

figures of dorsal zone obscure Xestia (. Pachnobia ) tecta (Hübner, 1808)

Seta Lp2 of labial palpus long, about 2/3-1 x length of base segment, spinneret 2-2.5 x

as long as wide, setal distance XD1-XD2 about 1 .5x that of XD2-SD2 on prothorax (ra-

tio XD1-XD2/XD2-SD2 range 1.2-1. 9, mean 1.5, N = 17), on Ab2 distance SV1-SV3

shorter than SV1-SV2 (ratio SV 1-SV3/SV 1-SV2 range 0.5-1. 1, mean 0.8, N = 17).

Larva yellowish or grayish brown with broad dorsal lines of same colour, bordered with

sharp, black, narrow margins, subdorsal line without ventral margin, markings of dorsal

zone like thin arrow-head chevron figure on Ab 1-8, spiracular line of same colour, dor-

sally bordered with sharp, blackish margin

Xestia ( Pachnobia ) lorezi (Staudinger, 1891)

Discussion

The appearance of the larval stages of X brunneopicta differs greatly from those of X gelida and

X fabulosa (Ferguson, 1965). This is a surprise considering the adult of A brunneopicta has been

considered to be closely related to them (Lafontaine et al. 1998). The larva of A. fabulosa is similar

to that of X gelida and therefore we supposed that the larva of X brunneopicta could resemble this

species as well. Ahola and Silvonen (201 1) described larvae close to A. gelida as possible A brun-

neopicta from Kuusamo and Kuhmo in Finland. These three larvae differ from A. gelida in having a

paler dorsal zone, shorter visible part of the middorsal line, the subdorsal line is broken into flecks,

has a narrower black dorsal margin of subdorsal line, is not enlarged on Ab7-8 and has a wider

white dorsal part of the spiracular line. Also, setal positions are slightly different, and SD1 is more

distant from the spiracles. However, we now rather see these differences as variation in A. gelida.

Nota Lepi. 38(1): 75-88

87

Many Noctuidae have green larvae. For example, in Europe there are more than 140 species

with such larvae. Larvae with green bodies and narrow white dorsal and subdorsal lines are not so

common, but still about 40 species have such larvae. However, a quarter of them can occur in the

same northern areas withX. brunneopicta. In Finland larvae of Orthosia gothica (Linnaeus, 1758)

and O. incerta (Hufnagel, 1766) resemble those of X brunneopicta. The brown head and position

of the spiracles above the spiracular line on Ab7 separates X brunneopicta readily from Orthosia

species. Xestia includes also two European species with green larvae, namely X ochreago (Hüb-

ner, 1790) and some variations of X castanea (Esper, 1798). They have, however, short dorsal

setae, and the head is green.

DNA barcodes ofX. brunneopicta differ from those of X gelida andX. fabulosa by a minimum

of 6.47% and 6.73% genetic distance, respectively (Marko Mutanen, pers. comm.), also suggest-

ing that X. brunneopicta is perhaps not a very close relative of these species. Based on the COI

sequences from one Finnish and one Russian specimen of X brunneopicta , the closest relatives

ofX. brunneopicta are A lorezi (4.44%), X sincera (4.60%) andX. ursae (McDunnough, 1940)

(4.61%), but many other Xestia species show less than 6% divergence as well. Based on DNA bar-

codes, no other Xestia species is a very close relative of X brunneopicta , and based on both larval

morphology and DNA barcodes, its sister species remains unclear.

Acknowledgements

We thank the Ella and Georg Ehmrooth Foundation for their support of the expedition, and the members

of the expedition Pekka Alestalo (Finland), Andrei Biksaleyev (Russia), Oleg Korsun (Russia) and Jukka

Tiittanen (Finland) for fieldwork in the Chara area. The authors are also grateful to Vladimir V. Dubatolov

(Russia) for identification of the other moth material, to Marko Mutanen (Finland) for comments on DNA

barcoding and to Pekka Puntila (Finland) for the photographs of the mature larva. Our thanks are also due to

Kari Nupponen (Finland) for helping with the manuscript, Lauri Kaila (Finland) for valuable comments and

suggestions and Don Lafontaine (Canada) for reviewing this paper and for his important comments which

helped to improve and clarify the text.

References

Ahola M, Silvonen K (201 1) Pohjoisen Euroopan yökkösten toukat. Larvae of Northern European Noctuidae.

Osa 3, volume 3. Viestipaino Oy, Tampere, 600 pp.

Beck H (1960) Die Larvalsystematik der Eulen. Abhandlungen zur Larvalsystematik der Insekten 4. Berlin, 406 pp.

Beck H (1974) Zur Beschreibung der Zeichnung (Ornamentik) von Insektenlarven. Eine Anleitung am

Beispiel von Noctuidenlarven. Atalanta 5: 121-143.

Beck H (1999-2000) Die Larven der Europäischen Noctuidae. Revision der Systematik der Noctuidae (Lep-

idoptera, Noctuidae) Vol I-IV. Herbipoliana 5. Marktleuchten, 859 + 447 pp.

Fibiger M, Hacker HH (2005) Systematic list of the Noctuoidea of Europe (Notodontidae, Nolidae, Arctiidae,

Lymantriidae, Erebiidae, Micronoctuidae and Noctuidae). Esperiana 1 1: 93-205.

Hinton HE (1946) On the homology and nomenclature of the setae of lepidopterous larvae, with some notes

on the phylogeny of the Lepidoptera. Trans. R. ent. Soc. Lond. 97: 1-37.

Kononenko VS (2005) An annotated check list of the Noctuidae (s.l.) (Insecta, Lepidoptera) of the Asian part

of Russia and the Ural region. Noctuidae Sibiricae, Vol. I. Entomological Press, Soro, 243 pp.

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Lafontaine JD (1998) Noctuoidea, Noctuidae (part - Noctuini). In: Dominick RB et al. The Moths of

America North of Mexico, fasc.27.3: 1-348.

Lafontaine JD, Mikkola K, Kononenko VS, Ahola M (1998) Subgenus Pachnobia Guenée. In: Lafontaine

JD. Noctuoidea, Noctuidae (part - Noctuini). In: Dominick RB et al. The Moths of America North of

Mexico, fasc.27.3, 139-165.

Mikkola K, Sinervirta M, Vaalamo K (1989) Xestia brunneopicta (Matsumura) new to Europe (Lepidop-

tera, Noctuidae). Baptria 69: 33 -37.

Patocka J, Turcâni M (2005) Lepidoptera Pupae. Central European Species. Text volumes. Apollo Books,

Steenstrup, 542 pp.

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Nota Lepi. 38(1) 2015: 89-102 | DOI 10.3897/nl.38.4816

In Memoriam: Niels Peder Kristensen (1943-2014)

Thomas J. Simonsen13, Ole Karsholt2, Malcolm J. Scoble1

1 Natural History Museum, United Kingdom, London, UK

2 The Natural History Museum of Denmark, Copenhagen, Denmark

3 Current address: Natural History Museum Aarhus, Aarhus, Denmark

http://zoobank.org/F762FCAl-0D5A-4CDF-9EB3-93A 7800752FD

Received 3 March 2015; accepted 18 March 2015; published: 12 May 2015

Subject Editor: Jadranka Rota.

Niels Peder Kristensen, Honorary Member and

former president of SEL, passed away on Satur-

day December 6th 2014 in Copenhagen. While his

death was not unexpected, its timing came earli-

er than we had thought or hoped. His loss is felt

widely and intensely.

Bom on March 2nd 1943, Niels was the second

child of Thorkil and Ellen Christine Kristensen

(nee Nielsen). His father was an academic, politi-

cian and thinker who served as Minster of Finance

in two different government cabinets, and later as

General Secretary of the OECD. Growing up in

such an environment undoubtedly had a profound

influence on Niels’ own world view, one which

was powerfully international in its expression, yet

retaining a strong interest and deep concern for

Danish issues - local and national.

Niels developed an interest in entomolo-

gy and lepidopterology in particular at an ear-

ly age, and once told TJS about the first time,

when eight years old, he visited the Entomology

Department at the Zoological Museum in Copen-

hagen (ZMUC) ‘clutching his father’s hand’. After completing high school at Birkerod Statsskole

in 1961, Niels enrolled as a biology student at the University of Copenhagen, and quickly became

a regular visitor to the Entomology Department of the Museum, where he had already started as a

volunteer during his last years at high school. In 1965, while still a student, he published his first

paper, which was on the faunistics of Danish cicadas. From the very start of his scientific career, one

of Niels’ abiding interests was the evolution (particularly evolutionary morphology) of primitive

Lepidoptera. Indeed, the work for his Mag. Scient, degree was on the comparative morphology

of the primitive glossatan family, Eriocraniidae. During this study, Niels spent the academic year

1966-67 at the University of Bristol, working with the eminent and extremely knowledgeable Brit-

Figure 1. Niels Peder Kristensen, March 2nd 1943

- December 6th 2014 (photo: Birgit Nielsen).

90

SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014)

ish entomologist Howard E. Hinton, who was at that time pioneering the use of scanning electron

microscopy in entomology. It was while working with Hinton that Niels came to appreciate the

value of SEM in comparative morphology. Through it, he made the surprising discovery (published

in 1970) that the most primitive Lepidoptera have flat, solid wing scales (i.e. lacking an internal lu-

men), a condition contrasting strongly with the hollow wing scales generally found in Lepidoptera.

After obtaining his Mag. Scient, degree in 1968, Niels was offered a tenure-track position at

the ZMUC as Amanuensis (Assistant Professor). He was promoted to Associate Professor in 1972

and appointed as Full Professor of Entomology in 1995. In 1970, Niels visited one of Europe’s

foremost comparative invertebrate morphologists, Jean Chaudonneret, at the Université de Dijon

where he enhanced considerably his skills in insect histology and semi-thin sectioning. Working

with Hinton, Chaudonneret and Karl G. Wingstrand, the professor of comparative morphology at

the University of Copenhagen, unquestionably had a profound influence on Niels’ development

as a scientist. He often referred to the effect these three mentors had on his career. Niels was also

deeply interested in the analytical methods used in evolutionary research. Together with fellow

entomologist Nils Möller Andersen and the palaeontologist Niels Bonde, he was a pioneer in Den-

mark, and more widely in Scandinavia, of Hennig’s phylogenetic systematics, and his cladistic

analyses of the higher-level relationships of butterflies in 1976 remained the standard work on the

subject until the study by de Jong et al. (1996) 20 years later.

From the very start of his career, Niels was deeply interested in the morphology and phylogeny

of the higher insects. In 1975 he published (Z. zool. Syst. Evolut. -forsch. 13, pp. 1^14) one of his

most influential papers: “The phylogeny of hexapod ‘orders’. A critical review of recent accounts”.

Thirty years later, Grimaldi and Engel (2005, p. 144) referred to this work as “perhaps the single most

important paper in systematic entomology”. This publication formed the basis of his five updated

reviews of the subject. The last of these was published in Eur. J. Ent. in 1999, while perhaps the most

notable of them is the 1991 text-book chapter “Phylogeny of extant hexapods” in “The Insects of

Australia”, which should be mandatory reading for all students of systematic entomology. Over the

years Niels authored or co-authored a number of papers on higher Hexapod relationships especially

on the lower Hexapod orders, Trichoptera, the enigmatic New Zealand mecopteran family Nanno-

choristidae, Neuroptera, and of course Mantophasmatodea — the first new insect order to be described

for 90 years, the description of which he co-authored in 2004.

It was, however, the Lepidoptera that remained Niels’ main interest, and the majority of his publi-

cations are on that order. They range in scope from nomenclatural and faunistic notes to higher-lev-

el phylogenetics and to the exceptionally detailed, comparative morphological studies of primitive

Lepidoptera. More than anything else, these exquisite studies became his professional hallmark. His

early enthusiasm for scanning electron microscopy and histology were combined with transmission

electron microscopy and became methodological cornerstones in his work throughout his working life.

Much of his productivity, particularly in the first half of his career, led to highly detailed studies of lit-

tle-understood structures and organ systems of primitive Lepidoptera, including overall head and neck

anatomy, mouthpart morphology, anatomy of the alimentary canal, structure of the trachaea system,

comparative morphology and anatomy of male and female genitalia, and wing scales and vestiture.

Niels’ work on primitive Lepidoptera morphology and anatomy was always embedded in the context

of higher Lepidoptera evolution, and his ultimate goal was to establish the early evolutionary patterns

within the order, thereby creating a sound basis for further studies higher up the lepidopteran tree. In

1978 and 1979 he also described two new families of primitive Lepidoptera, the basal hepialoid family

Nota Lepi. 38(1): 89-102

91

Figure 2. Niels studying his beloved homoneuran Lepidoptera (photo: TJS).

Neotheoridae and the non-ditrysian family Heterobathmiidae (the latter in collaboration with the late

Ebbe S. Nielsen). His work on primitive Lepidoptera phylogeny and comparative morphology culmi-

nated in his Dr Scient, dissertation “Studies on the morphology and systematics of primitive Lepidop-

tera” published in Steenstrupia in 1984. Until the modifications introduced by the very recent advent

of phylogenomic studies and especially the surprising discovery of a new primitive moth family from

Australia, this remained the standard work on the evolution of the homoneurous Lepidoptera.

In the early 1990s Niels was appointed the editor-in-chief of the two Lepidoptera volumes of

the Handbook of Zoology. This immense undertaking was to dominate his professional life for the

following decade. The two volumes, which were published in 1998 and 2003, defined the latter part

of his career as much as his work on higher Hexapod phylogeny and comparative Lepidoptera mor-

phology had shaped his early and mid career, although he continued his work on these topics until

illness forced him to stop just weeks before his death. Niels had anticipated writing or co-authoring

a substantive part of the first volume. He did not, however, expect to have made a similar input to

the second volume, which was on morphology and physiology. Having to do so resulted in a much

greater effort on his part than he had intended: moreover, it required him to write about subjects on

which he did not consider himself an expert. The result, nevertheless, stands as a landmark publi-

cation and a tribute to Niels’ capacity and breadth of knowledge. The Handbook would have been

more than enough of a mega-project for most of us, so it is remarkable that Niels also spent much

time and effort during his last years editing a book on the insects of Greenland instead of completing

some of his own research projects. While he certainly believed in the value of the Greenland work,

his resolve was propelled by that innate sense of responsibility and conscientiousness that were so

evident in his personal makeup.

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SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014)

After the mammoth task of completing the Handbook, Niels returned to his work on primitive

Lepidoptera — at least as much as his administrative duties permitted. At the time of his death,

he was involved in long-term studies of several groups: Micropterigidae with G. W. Gibbs and D.

L. Lees; Mnesarchaeidae with G. W. Gibbs; Hepialoidea with TJS. A phylogenomic study of the

non-ditrysian lineages (part of the LepTree project), of which he was a senior co-author, was sub-

mitted just weeks after his death (and was dedicated to him). But his most significant contribution

in the last stage of his career was the discovery and description of an extraordinary homoneuran

family from Kangaroo Island, Australia (the so-called Kangaroo Island Moth, or KIM) and its

significance for modifying our understanding of early Lepidoptera evolution. The paper was, to his

great pleasure, accepted for publication before his death.

Besides being a leading research scientist, Niels was a highly engaging and inspiring teacher

and supervisor. Early in his career, he wrote, in Danish, a detailed yet concise compendium of

systematic entomology (“Systematisk Entomologi” 1974), which for years was the standard text-

book on the subject at the University of Copenhagen. For over two decades he was a driving force

behind an advanced course in systematic entomology and insect morphology, which was taught

biennially at the University. Niels’ lectures displayed not only the depth of his learning, but were

rich in subtle humour, a quality of which he was a master. TJS recalls a particular entomology

lecture (in 1996) in which Niels was explaining the morphology of the thorax, including flight

mechanisms. To ensure that the students understood the complex ways in which insects move their

wings to minimize drag, he demonstrated by lying face down on a table, still clad in jacket and tie

and with his feet sticking out, waiving his arms in the air! While this performance was not charac-

teristic of most university professors, it worked — TJS resolved there and then to do his graduate

studies under Niels’ supervision.

Throughout his career, Niels supervised several Masters and PhD students and postdoctoral fel-

lows. He took a deep interest in their well-being, both professional and personal, and derived im-

mense pleasure from their subsequent successes while keeping in close contact with them after they

graduated. (After graduating and moving away from Copenhagen, TJS spent numerous hours on

the phone with Niels discussing his own work, Niels’ work, the world in general and entomology in

particular.) It was therefore also with great sadness and regret that he found himself writing obituaries

for two of his most talented PhD students, the coleopterist Michael Hansen and the lepidopterist Ebbe

S. Nielsen, both of whom died prematurely in the year 2000. Niels considered the more sociological

aspects of entomology and lepidopterology to be integral parts of the (informal) training of a student.

TJS recalls numerous meetings with Niels, intended to be brief, but often extending to a couple of

hours, and invariably covering a wide range of aspects such as the history of science, the works (pres-

ent and past) of other entomologists, anecdotes, amateur entomology, entomology and society. One

of Niels’ great qualities as a supervisor was that hallmark of all top supervisors — he had an intuitive

understanding of the level and extent of supervision needed to fit the individual student, endeavouring

always to bring out the best in him or her.

Niels started collecting butterflies and moths as a schoolboy, and although he never built up a

large collection, this activity influenced his choice to become a biologist and a specialist in Lep-

idoptera. At that time there was no tradition for lepidopterology at the ZMUC, and Niels was the

first academically trained lepidopterist at the museum — despite being advised by the head of the

entomology department at that time, S. L. Tuxen, to find a more scientific group! Later he often

Nota Lepi. 38( 1): 89-102

93

defended collecting Lepidoptera, arguing that is an important way to get young people interested

in entomology.

Niels appreciated deeply (serious) amateur lepidopterists, being well aware that major parts of

the Lepidoptera collections in larger museums had been collected by them. He valued their efforts,

spoke positively and warmly about them and did much to help them, for example through his work

for societies with large amateur memberships, by advising on scientific matters and in providing

help to get collecting permits.

While Niels enjoyed experiencing Lepidoptera and other insects in nature, and he would often

run a mercury vapour light at his summer cottage, he was not primarily a field worker. He felt, and

indeed demonstrated, that he could serve the study of Lepidoptera best by focusing his exceptional

skills on the study of key taxa at the museum bench.

Besides his research and teaching, Niels shouldered a substantial administrative burden at the

ZMUC, at which he spent his entire career. This included several stints as Chair or Deputy Chair

of the Entomology Department, two periods as Deputy Director of the museum, and three years as

Director. He was also Head of Zoology at the newly designated Natural History Museum of Den-

mark from 2004-2006. Although his heart remained in his research, he carried out these time-con-

suming administrative responsibilities conscientiously and with a great sense of love and concern

for the museum.

Throughout his career, Niels was deeply involved in entomological and lepidopterological soci-

eties. He was President of the Danish Entomological Society from 1989 to 1999, Council member

of the International Congress of Entomology from 1988 to 2004 (Deputy Chair 2000-2004), and of

course President of the SEL from 1998 to 2007, and Chair of the SEL congress in Korsor, Denmark

in 2002. One of Niels’ long-standing ambitions was to hold a joint European-North American Lepi-

doptera Congress. Although this did not take place during his own tenure as President, he was very

pleased to see the first joint meeting of the Lepidopterists’ Society and SEL in Denver, Colorado in

2012 (even if he could not attend the meeting himself).

During his career, Niels received many honours and awards, a testimony to his achievements and

pre-eminence in his field. He was a member of the Danish Academy of Natural Sciences, a member

of the Royal Danish Academy of Sciences and Letters, a corresponding member of the Finnish En-

tomological Society, an honorary ‘Foreign member’ of the Linnean Society of London (1998), an

honorary member of Sociedad Hispano-Luso- Americana de Lepidopterologia (SHILAP), an honor-

ary member of the Danish Entomological Society, an honorary research fellow at the Natural History

Museum, London, an honorary member of Gesellschaft für Biologische Systematik, an honorary

member of Societas Europaea Lepidopterologica, an honorary fellow of the Royal Entomological

Society, and an honorary member of the Russian Entomological Society. In 1988 he was awarded the

‘Karl Jordan Medal’ (Lepidopterists’ Society) for “outstanding original research in lepidopterology”,

in 1999 he received the Joachim Jungius-Medaille (J.J.Gesellschaft der Wissenschaften, Hamburg)

for “herausragender Leistungen in Wissenschaft und Forschung”, and in 2014 shortly before his

death he gained the 2014 Linnean Medal (Zoology).

Niels cared deeply about the future of European entomology and lepidopterology and was dis-

mayed by the progressive decline of staff numbers and funds at several major research institutions

(including his own in Denmark). He believed firmly in the need for basic research, and that in

publicly funded institutions such as museums it should be possible for researchers to focus on

94

SiMONSEN et al.\ In Memoriam: Niels Peder Kristensen (1943-2014)

academically interesting questions that do not necessarily have immediately obvious economic,

social or medical benefits.

Unsurprisingly, Niels’ standing, awareness and understanding of wider socio-political issues in

science led to him being asked to act as Director of the ZMUC. While he had an interest in uni-

versity politics and administration, his real love was for his research, but he accepted the position

(doing two terms of service) partly from a sense of duty, and partly because he felt that the position

should be held by an acknowledged researcher rather than a mandarin. Having high ethical stand-

ards, Niels rarely sought the easiest solution to any problem, but rather the one that he thought to

be right. He was a conscientious leader, made great efforts to keep abreast of relevant matters and

always made time for his colleagues, whatever their level in the organization. He undoubtedly

suffered during his extended directorship, both as a result of these personal qualities and through

the loss of most of his time for research. Alas, he had to endure more frustration due to seemingly

endless cuts to the museum’s funds. But when the Ministry of Education and Research ordered cuts

of several positions at the museum, he felt he could no longer accept the responsibility for running

the institution and stepped down in protest.

Niels was critical of the plans for a new natural history museum in Copenhagen, which would

have resulted in newer but reduced facilities. He was particularly disturbed about the idea of de-

molishing the ZMUC building, which had been purpose built and which he considered to be still fit

for purpose. During his later years he spoke and wrote against the idea, and was disappointed that

the management of the Natural History Museum of Denmark, and many of his colleagues, did not

agree with him. He also expressed concern about the appointment at the museum of scientists with

little experience of collections-based research.

At the time of his death, Niels had settled into a productive retirement: relieved of administrative

responsibilities, it was a phase of his life that he was enjoying thoroughly. So it is heart-breaking that

he missed the prolonged and active retirement he would have found so fulfilling. Moreover, it leaves

the scientific community bereft of the many works that would surely have been produced by him. The

entomological world has become a much poorer place without Niels’ profound knowledge and in-

sight, his generosity of spirit, his conscientiousness and his quiet humour. For all these qualities he is

and will continue to be missed deeply. He was also a loving family man and our deepest condolences

are extended to his wife Else and their daughters.

HAVAMAL (Our translation)

Livestock die

Kinsman dies

We all die just the same

Only one thing I know which never dies

The judgment of a dead man’s life.

HAVAMAL (Danish)

Fæ dor

Frænde dor

Dor selv pâ samme vis

Kun et ved jeg som aldrig dor

Dommen over dod mands liv

Nota Lepi. 38(1): 89-102

95

Acknowledgements

We wish to thank Lars Vilhelmsen, Natural History Museum of Denmark for sharing biographical data he had collected. We

thank Birgit Nielsen, Frederiksværk, Denmark for permission to use the portrait photo of Niels P. Kristensen.

References

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and prospects. Entomologica scandinavica 27: 65-101. doi: 10.1163/187631296X00205

Grimaldi D, Engel MS (2005) Evolution of the Insects. Cambridge University Press, Cambridge, UK, 755 pp.

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156. Karsholt O, Kristensen NP, Simonsen TJ, Ahola M (in press) Lepidoptera (moths and butterflies). 51 pp.

In Böcher J, Kristensen NP, Pape T, Vilhelmsen L (Eds). The Greenland entomofauna. An identification

manual of insects, spiders and their allies. E. J. Brill, Leiden.

155. Kristensen NP, Hilton DJ, Kallies A, Milia L, Rota J, Wahlberg N, Wilcox SA, Glatz RV, Young DA,

Cocking G, Edwards ED, Gibbs GW, Halsey M (2015) A new extant family of primitive moths from

Kangaroo Island, Australia, and its significance for understanding early Lepidoptera evolution. Systematic

Entomology 40: 5-16. doi: 10.111 1/syen. 12115

154. Kristensen NP, Rota J, Fischer S (2014) Notable plesiomorphies and notable specializations: head struc-

ture of the primitive “tongue moth” Acanthopteroctetes unifascia (Lepidoptera: Acanthopteroctetidae).

Journal of Morphology 275: 153-172. doi: 10.1002/jmor.20205

153. Kristensen NP (2013) Intraspecific variability in gross design of moth brains: A caveat concerning ex-

pectedly ‘good’ characters’ in systematic entomology (Insecta: Lepidoptera). Zoologischer Anzeiger 253:

112-113. doi: 1 0. 1 0 1 6/j .j cz.20 1 3 .09.002

152. Engel MS, Kristensen NP (2013) A history of entomological classification. Annual Review of Entomol-

ogy 58: 585-607. doi: 10.1 146/annurev-ento- 12081 1-1 53536

151. Hünefeld F, Kristensen NP (2012) The female postabdomen and genitalia of the basal moth family

Heterobathmiidae (Insecta: Lepidoptera): Structure and phylogenetic significance. Arthropod Structure &

Development 41: 395^107. doi: 10. 1016/j.asd.2012.05.001

150. Beutel RG, Kristensen NP (2012) Morphology and insect systematics in the era of phylogenomics. Ar-

thropod Structure & Development 41: 303-305. doi: 10.1016/j.asd.2012.05.003

149. Hünefeld F. Kristensen NP (2012) Two new heterobathmiid moth species with distinctive female genital

configurations (Lepidoptera: Heterobathmiidae). Zootaxa 3281: 61-68.

148. Kristensen NP (2012) Molecular phylogenies, morphological homologies and the evolution of moth

‘ears’. Systematic Entomology 37: 237-239. doi: 1 0. 1 1 1 l/j. 1 365-3 1 13.201 2.006 1 9.x

147. Gibbs GW, Kristensen NP (2011) Agrionympha, the long-known South African jaw moths: a revision

with descriptions of new species (Lepidoptera, Micropterigidae). Zootaxa 2764: 1-21 .

146. Djemæs M, Kristensen NP (2011) Derived morphology in a basal moth: The uniquely specialized

sternum V glands of Agathiphaga. Arthropod Structure & Development 40: 559-569. doi: 10.1016/j.

asd.201 1.06.001

145. Rota J, Kristensen NP (2011) Notes on taxonomic history, thoraco-abdominal articulation, and current

placement of Millieriidae (Insecta: Lepidoptera). Zootaxa 3032: 65-68.

144. van Nieukerken E, Kaila L, Kitching IJ, Kristensen NP, Lees, DC, Minet J, Mitter C, Mutanen M, Regier

JC, Simonsen TJ, Wahlberg N, Yen S-H, Zahiri R, et al. (38 additional authors) (2011) Order Lepidoptera.

96

SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014)

In: Zhang Z-Q (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxono-

mic richness. Zootaxa 3148: 212-221.

143. Kristensen NP (2011) Michael Fibiger 29. juni 1945 - 16. februar 2011. Entomologiske Meddelelser 79:

153-165.

142. Hünefeld F, Kristensen NP (2010) The female postabdomen and internal genitalia of the basal moth

genus Agathiphaga (Insecta: Lepidoptera: Agathiphagidae): Morphology and phylogenetic implications.

Zoological Journal of the Linnean Society 159: 905-920. doi: 10.1 111/j. 1096-3642.2009. 00590.x

141. Kristensen NP, Gaedike R (2010) Extraordinary moths and an extraordinary moth researcher: An essay

review of G. S. Robinson’s Biology, distribution and diversity of tineid moths. Nota Lepidopterologica

33: 3-8.

140. Lees DC, Rougerie R, Zeller- Lukaschort C, Kristensen NP (2010) DNA mini-barcodes in taxonomic

assignment: a morphologically unique new homoneurous moth clade from the Indian Himalayas described

in Micropterix (Lepidoptera, Micropterigidae). Zoologica Scripta 39: 642-661. doi: 10.1 111/j. 1463-

6409.2010.00447.x

139. Kristensen N P, Nielsen PS (2010) Et Hoffmeyer-manuscript om vore ’stribede’ kollesværmere. Lepi-

doptera 9: 289-307.

138. Beutel RG, Kristensen NP, Pohl H (2009) Resolving insect phylogeny: The significance of cephalic

structures of the Nannomecoptera in understanding endopterygote relationships. Arthropod Structure and

Devopment 38: 427^160. doi: 10.1016/j.asd.2009.05.002

137. Kaaber S, Kristensen NP, Simonsen TJ (2009) Sexual dimorphism and geographical male polymorphism

in the ghost moth Hepialus humuli (Lepidoptera: Hepialidae): Scale ultrastructure and evolutionary as-

pects. European Journal of Entomology 106: 303-313. doi: 10.14411/eje.2009.036

136. Béthoux O, Kristensen NP, Engel M (2008) Hennigian phylogenetic systematics and the ‘groundplan’

vs. ‘post-groundplan’ approaches: A Reply to Kukalovâ-Peck. Evolutionary Biology 35: 317-323. doi:

10. 1007/sl 1692-008-9035-6

135. Kristensen NP (2008) Natsværmeren ‘humleæder’, dens flyveaktivitet, vingeskæl-struktur og variation I

Nordeuropa. Dyr i Natur og Museum 2008/2: 22-25.

134. Kristensen NP (2008) Early Lepidoptera evolution. Gesellschaft für Biologische Systematik Newsletter

20: 50-55.

133. Kristensen NP, Scoble MJ, Karsholt O (2007) Lepidoptera phylogeny and systematics: the state of inven-

torying moth and butterfly diversity. Zootaxa 1668: 699-747.

132. Kristensen NP (2007) Nils Möller Andersen. Dansk Naturhistorisk Forening Ârsskrift 16/17: 50-51.

131. Kristensen NP (2007) Leif Lyneborg - 3. januar 1932 - 10. September 2006. Dansk Naturhistorisk Foren-

ings Ârsskrift 16/17: 68-75.

130. Faucheux MJ, Kristensen NP, Yen S-H (2006) The antennae of neopseustid moths: Morphology and

phylogenetic implications, with special reference to the sensilla (Insecta, Lepidoptera, Neopseustidae). Zo-

ologischer Anzeiger 245: 131-142. doi: 10.1016/j.jcz.2006.05.004

129. Kristensen NP (2006) Nils Möller Andersen 21. november 1940 - 12. maj 2004. Det Kongelige danske

videnskabemes Selskab. Oversigt over Selskabets Virksomhed 2004-05: 217-224.

128. Kristensen NP (2005) Jens Bodtker Rasmussen - Obituary. Tropical Zoology 18: 149-149. doi:

10.1 080/03946975 .2005 . 10531217

127. Krenn HW, Kristensen NP (2004) Evolution of proboscis musculature in Lepidoptera. European Journal

of Entomology 101: 565-575. doi: 10.1441 l/eje.2004.080

126. Kristensen NP (2004) Om indsamling og udskillelse. Nogle naturhistoriske Perspektiven Danske Museer

17/5:20-21.

125. Kristensen NP (2004) Paul Johannes Holst-Christensen. Dansk Naturhistorisk Forening Ârsskrift 14:

86-89.

Nota Lepi. 38(1): 89-102

97

124. Kristensen NP (2003) Resolving the basal phylogeny of Lepidoptera: morphological evidence. Entomo-

logische Abhandlungen 61:1 67-169.

123. Kristensen NP (2003) Reproductive organs. Pp. 427-447 in N.P.Kristensen (Ed.) Lepidoptera: Moths and

butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de Gruyter, Berlin & New York.

122. Akai H, Hakim RS, Kristensen NP (2003) Labial glands, silk and saliva. Pp. 377-388 in N.P.Kristensen

(ed.) Lepidoptera: Moths and butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de

Gruyter, Berlin & New York.

121. Warrant E, Kelber A, Kristensen NP (2003) Eyes and vision. Pp. 325-359 in N.P.Kristensen (ed.) Lep-

idoptera: Moths and butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de Gruyter,

Berlin & New York.

120. Barbehenn RV, Kristensen NP (2003) Digestive and excretory system. Pp. 165-187 in N.P.Kristensen

(ed.) Lepidoptera: Moths and butterflies 2. Handbuch der Zoologie/Handbook of Zoology' IV/36: Walter de

Gruyter, Berlin & New York.

119. Hasenfuss I, Kristensen NP (2003) Skeleton and muscles: immatures. Pp. 133-164 in N.P.Kristensen

(ed.) Lepidoptera: Moths and butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de

Gruyter, Berlin & New York.

118. Kristensen NP (2003) Skeleton and muscles: adults. Pp. 39-131 in N.P.Kristensen (ed.) Lepidoptera:

Moths and butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de Gruyter, Berlin &

New York.

117. Kristensen NP, Simonsen TJ (2003) ‘Hairs’ and scales. Pp. 9-22 in N.P.Kristensen (ed.) Lepidoptera:

Moths and butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de Gruyter, Berlin &

New York.

116. Chauvin G, Kristensen NP (2003) Integument. Pp. 1-8 in N.P.Kristensen (ed.) Lepidoptera: Moths and

butterflies 2. Handbuch der Zoologie/Handbook of Zoology IV/36: Walter de Gruyter, Berlin & New York.

115. Kristensen NP, Simonsen TJ (2003) Scale length/wing length correlation in Lepidoptera. Journal of Nat-

ural History 37: 673-679. doi: 10.1080/00222930110096735

114. Karsholt O, Kristensen NP (2003) Plesiozela , gen. nov. from temperate South America: apparent sis-

ter-group of the previously known Heliozelidae (Lepidoptera: Incurvarioidea: Heliozelidae). Invertebrate

Systematics 17: 39-46. doi: 1 0. 1 07 1/IS02047

113. Karsholt O, Kristensen NP (2003) Kastaniemollet: et kont nyt skadedyr i Danmark. Dyr i Natur og Mu-

seum 2003/1: 9-11.

112. Kristensen NP (2002) Mantophasmatodea: en nyopdaget orden af nulevende insekter. Dyr i Natur og

Museum 2002/2: 24-27.

111. Klass K-D, Zompro O, Kristensen NP, Adis J (2002) Mantophasmatodea: a new insect order with extant

members in the Afrotropics. Science 206: 1456-1459. doi: 10. 11 26/science. 1069397

110. Kristensen NP (2002) Ebbe Schmidt Nielsen. Dansk Naturhistorisk Forening Ârsskrift 12: 58-60.

109. Kristensen NP (1994—2002) Contributions to ‘ Danmarks Nationalleks ikon Y Den Store Danske Encyklopce-

di’ (vols 1-20, Gyldendal, Copenhagen), including ‘Dagsommerfugle’ [butterflies], ‘Insekter’ [insects], ‘Led-

dyr’ [arthropods], ‘Sommerfugle’ [Lepidoptera], ‘Urinsekter’ [apterygotes] and numerous shorter (< 50 lines)

articles on zoological (mostly entomological) subjects, including biographies of zoologists.

108. Simonsen TJ, Kristensen NP (2001) Agathiphaga wing vestiture revisited: evidence for complex ear-

ly evolution of lepidopteran scales (Lepidoptera: Agathiphagidae). Insect Systematics and Evolution 32:

169-175. doi: 10.1163/187631201X00128

107. Klass K-D, Kristensen NP (2001) The ground plan and affinities of hexapods: recent progress and open

problems. Annales de la Société Entomologique de France (N. S.) 37: 265-298.

106. Kristensen NP (2001) Ebbe Schmidt Nielsen 7 June 1950 - 6 March 2001. Nota Lepidopterologica 24/3:

3-9.

98

SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014)

105. Kristensen NP (2001) Henning Anthon. Entomologiske Meddelelser 69: 65-68.

104. Kristensen NP (2001) Michael Hansen. Dansk Naturhistorisk Forening Ârsskrift 1 1 : 67-69.

103. Krenn HW, Kristensen NP (2000) Early evolution of the proboscis of Lepidoptera (Insecta): external mor-

phology of the galea in basal glossatan moth lineages, with remarks on the origin of the pilifers. Zoologisher

Anzeiger 239: 179-196.

102. Kristensen NP (1999) Phylogeny of endopterygote insects, the most successful lineage of living organ-

isms. European Journal of Entomology 96: 237-253.

101. Dugdale JS, Kristensen NP, Robinson, GS, Scoble MJ (1998) The smaller microlepidoptera-grade super-

families. Pp. 217-232 in N.P.Kristensen (ed.) Lepidoptera: Moths and butterflies 1. Handbuch der Zoologie/

Handbook of Zoology IV/35(1999): 51-63. Walter de Gruyter, Berlin & New York.

100. Edwards ED, Gentili P, Horak M, Kristensen NP, Nielsen ES (1998) The Cossoidea and Sesioidea. Pp.

181-197 in N.P.Kristensen (ed.) Lepidoptera: Moths and butterflies 1. Handbuch der Zoologie/Handbook of

Zoology IV/35: (1999) 51-63. Walter de Gruyter, Berlin & New York.

99. Dugdale JS, Kristensen NP, Robinson GS, Scoble MJ (1998) The Yponomeutoidea. Pp. 119-130 in

N.P.Kristensen (ed.) Lepidoptera: Moths and butterflies 1 . Handbuch der Zoologie/Handbook of Zoology

IV/35(1999): 51-63. Walter de Gruyter, Berlin & New York.

98. Kristensen NP (1998) The homoneurous Glossata. Pp. 51-63 in N.P.Kristensen (Ed.) Lepidoptera: Moths

and butterflies 1 . Handbuch der Zoologie/Handbook of Zoology IV/35(1 999): 5 1-63. Walter de Gruyter, Ber-

lin & New York.

97. Kristensen NP (1998) The non-glossatan moths. Pp. 41^49 in N.P.Kristensen (Ed.) Lepidoptera: Moths and

butterflies 1. Handbuch der Zoologie/Handbook of Zoology IV/35: 51-63(1999). Walter de Gruyter, Berlin

& New York.

96. Carter D, Kristensen NP (1998) Classification and keys to higher taxa. Pp. 27—40 in N.P.Kristensen (ed.)

Lepidoptera: Moths and butterflies 1. Handbuch der Zoologie/Handbook of Zoology IV/35(1999): 51-63.

Walter de Gruyter, Berlin & New York.

95. Kristensen NP, Skalski A (1998) Palaeontology and phylogeny. Pp. 7-25 in N.P.Kristensen (ed.) Lepidop-

tera: Moths and butterflies 1. Handbuch der Zoologie/Handbook of Zoology IV/35(1999): 51-63. Walter de

Gruyter, Berlin & New York.

94. Kristensen NP (1998) Historical Introduction. Pp. 1-5 in N.P.Kristensen (ed.) Lepidoptera: Moths and

butterflies 1. Handbuch der Zoologie/Handbook of Zoology IV/35(1999): 51-63. Walter de Gruyter, Berlin

& New York.

93. Kristensen NP, Nielsen E S (1999). Heterobathmia valvifer n.sp.: A moth with large apparent “ovipositor

valves” (Lepidoptera, Heterobathmiidae). Steenstrupia 24(1998): 141-156.

92. Kristensen NP (1997) Early evolution of the Lepidoptera + Trichoptera lineage: phylogeny and the ecologi-

cal scenario. Mémoires du Muséum National d'histoire Naturelle 173: 253-271 . [Japanese translation, with

minor differences: pp. 182-200 in T. Yasuda (Ed.) (1988): Biology of Microlepidoptera]

91. Kristensen NP (1997) The ground plan and basal diversification of the hexapods. Pp. 281-293 in Fortey,

R. A. & Thomas, R. H. (eds), Arthropod Relationships, Chapman & Hall, London.

90. Kristensen NP (1997) Japetus Steenstrup - 100 âr efter. Dyr i Natur og Museum 1997/2: 21-25.

89. Kristensen NP (1997) Myrelover i Danmark. Dyr i Natur og Museum 1997/1: 24—27.

88. Nielsen ES, Kristensen NP (1996) The Australian moth family Lophocoronidae and the basal phylogeny

of the Lepidoptera-Glossata. Invertebrate Taxonomy 10: 1199-1302. doi: 10.1 07 1/IT996 1 1 99

87. Davis DR, Karsholt O, Kristensen NP, Nielsen ES (1995) A revision of the genus Ogygioses (Lepidoptera:

Palaeosetidae). InvertebrateTaxonomy 9: 1231 1263. doi: 10.1 07 1/IT995 1231

86. Kristensen NP (1995) Forty years’ insect phylogenetic systematics. Hennig’s “Kritische Bemerkungen...”

and subsequent developments. Zoologische Beiträge (N. F.) 36: 83-124.

85. Kristensen NP (1995) Sydbogemollene. Dyr i Natur og Museum 1995/1: 30-31.

Nota Lepi. 38(1): 89-102

99

84. Kristensen NP (1994) Den naturhistoriske samling og forskningen. Nordisk Museologi 2: 47-56.

83. Karsholt O, Kristensen NP, Kozlov MV (1994) Eriocrania cicatricella (Zetterstedt, 1839) the correct

name of the moth currently known as Eriocrania haworthi Bradley, 1966 (Lepidoptera: Eriocraniidae).

Entomologiske Meddelelser 62: 91-93.

82. Melzer RR, Kristensen NP, Paulus HF ( 1 994)The larval eye of nannochoristid scorpionflies (Insecta, Me-

coptera). Acta Zoologica 75: 201-208. doi: 10.1 111/j. 1463-6395. 1994.tb0 1207.x

81. Kristensen NP, Nielsen ES (1994) Osrhoes coronta Druce, the New World palaeosetid moth: A reap-

praisal, with description of a new type of female genital apparatus (Lepidoptera, Exoporia).Entomologica

scandinavica 24: 391-406. doi: 10.1163/187631293X00181

80. Kristensen NP (1994) Mä man gerne samle pä insekter? Dyr i Natur og Museum 1994/1: 7-10.

79. Kristensen NP (1994) Bent W. Rasmussen. Entomologiske Meddelelser 62: 95.

78. Kristensen NP (1994) Karl Georg Wingstrand. Det Kongelige danske videnskabemes Selskab. Oversigt over

Selskabets Virksomhed 1992-93: 155-166.

77. Kristensen NP (1993) En enestâende tilgang til Zoologisk Museums sommerfuglesamling. Dyr i Natur og

Museum 1993/2: 20-22.

76. Kristensen NP (1993) Biodiversitetens dimensioner: kvantitet og ’kvalitet’. Naturens Verden 1993:

163-179. [with contributions from Andersen, P. F., Coull, B., de Kalin Arrogo, M., Friis, I., Greuter, W.,

Ihlenfeldt, H.D. & Strid, A.]

75. Kristensen NP, Rasmussen JF (1993) Biodiversitet som videnskabelig og samfundsmæssig udfordring.

Naturens Verden 1993: 197-208. [with contributions from Ehrlich, P. R., Fjeldsâ, F., Janzen, D. H., Strid,

A. & von Bothmer, R.]

74. Kristensen NP, Larsen K (1993) Biodiversitet i en verden under forandring. Naturens Verden 1993: 161-162.

73. Kristensen NP (1991) Phylogeny of extant hexapods. Pp. 125-140 in CSIRO (ed.): The Insects of Australia.

Carlton: Melbourne University Press. [Reprinted with minor alterations in I.D. Naumann (ed.) Systematic and

Applied Entomology. An Introduction (1994). Carlton: Melbourne University Press.

72. Kristensen NP (1990) Morphology and phylogeny of the lowest Lepidoptera-Glossata: Recent progress

and unforeseen problems. Bulletin of the Sugadaira Montane Research Center 1 1 : 105-106.

71. Kristensen NP (1990) The trunk integument of zeuglopteran larvae: One of the most aberrant arthropod

cuticles known (Insecta, Lepidoptera). Bulletin of the Sugadaira Montane Research Center 1 1 : 101-102.

70. Kristensen NP (1990) Den ‘eksploderede’ goliathbille. Dyr i Natur og Museum 1990/2: 29.

69. Kristensen NP (1990) Torben W. Langer *7.6.1924 ^ 1 3.4. 1 988. Entomologiske meddelelser 58: 95-96.

68. Nielsen ES, Kristensen NP (1989) Primitive Ghost Moths. Monographs on Australian Lepidoptera, 1,

1-206, CSIRO, Canberra.

67. Kristensen NP (1989) Insect phylogeny based on morphological evidence. Pp 295-306 in Femholm,

B. et al. (eds): The Hierarchy of Life. Molecules and Morphology in Phylogenetic Analysis. Amsterdam:

Elsevier.

66. Kristensen NP (1989) The New Zealand scorpionfly {Nannochorista philpotti comb.n.): wing morphology

and its phylogenetic significance. Zeitschrift fur zoologische Systematik und Evolutionsforschung 27:

106-114. doi: 10. 1 1 1 1/j. 1439-0469. 1 989.tb00335.x

65. Kristensen NP (1988) Biography and contributions of prof, dr Jean Chaudonneret, 1984 Recipient of The

Distinguished International Award in Insect Morphology and Embryology. International Journal of Insect

Morphology and Embryology 17: 171-176. doi: 10.1016/0020-7322(88)90034-7

64. Kristensen NP (1986) Nachruf Anker Nielsen. Trichoptera Newsletter 13: 4-6.

63. Kristensen NP (1985) De forste snabelsommerfugle. Dyr i Natur og Museum 1985/2: 10-12.

62. Kristensen NP (1985) Sommerfuglenes tidligste udvikling. Dyr i Natur og Museum 1985/1: 14-18.

61. Kristensen NP (1985) Sommerfuglenes storsystematik/The higher classification of Lepidoptera. In: Sch-

nack, K. (ed.): Katalog over de danske Sommerfugle. Entomologiske Meddelelser 52/2-3: 6-20.

100

SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014)

60. Kristensen NP (1985) Anker Nielsen: 21. februar 1907 - 9. december 1984. Videnskabelige Meddelelser

fra dansk naturhistorisk Forening 146: 115-124.

59. Kristensen NP (1984) Studies on the morphology and systematics of primitive Lepidoptera (Insecta).

Steens trupia 10: 141-191.

58. Kristensen NP (1984) The male genitalia of Agathiphaga (Lepidoptera, Agathiphagidae) and the lepi-

dopteran ground plan. Entomologica Scandinavia 15: 151-178. doi: 10.1163/187631284X00127

57. Kristensen NP ( 1 984) The larval head of Agathiphaga (Lepidoptera, Agathiphagidae) and the lepidopteran

ground plan. Systematic Entomology 9: 63-81. doi: 10.1 1 11/j. 1365-3113. 1984.tb00502.x

56. Kristensen NP (1984) The pregenital abdomen of the Zeugloptera (Lepidoptera). Steenstrupia 10: 113—

136.

55. Kristensen NP (1984) Respiratory system of the primitive moth Micropterix calthella (Linnaeus) (Lepi-

doptera: Micropterigidae). International Journal of Insect Morphology and Embryology 13: 137-156. doi:

10.101 6/0020-7322(84)90022-9

54. Kristensen NP (1984) Skeletomuscular anatomy of the male genitalia of Epimartyria (Lepidoptera: Mi-

cropterigidae). Entomologica Scandinavia 15: 97-112. doi: 10.1163/187631284X00091

53. Kristensen NP (1984) S. L. Tuxen. International Journal of Insect Morphology and Embryology 13: 311—

314. doi: 10.1016/0020-7322(84)90007-2

52. Kristensen NP (1984) S. L. Tuxen. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 144:

157-170.

51. Kristensen NP, Nielsen ES (1983) The Heterobathmia life history elucidated: Immature stages contradict

assignment to suborder Zeugloptera (Insecta, Lepidoptera). Zeitschrift für zoologische Systematik und

Evolutionsforschung 21: 101-124. doi: 10. 1 1 1 1/j. 1439-0469. 1983.tb00280.x

50. Kristensen NP, Nielsen ES (1982) South American micropterigid moths: two new genera of the Sa-

batinca-group (Lepidoptera: Micropterigidae). Entomologica Scandinavia 13: 513-529. doi:

10.1163/187631282X00318

49. Henriksen HJ, Kristensen NP (1982) Dagsommerfuglen Colias alf acariens is, en nyopdaget strejfgæst i

Danmark (Lepidoptera: Pieridae). Entomologiske Meddelelser 49: 123-131.

48. Kristensen NP. (1982) Splittng or widening: remarks on the taxonomic treatment of paraphyletic taxa.

Annales Zoologici Fennici 19: 201-202.

47. Kristensen NP, Nielsen ES (1981) Double-tube proboscis configuration in neopseustid moths (Lepidop-

tera: Neopseustidae). International Journal of Insect Morphology and Embryology 10: 483^186. doi:

10.101 6/0020-7322(8 1 )90027- 1

46. Kristensen NP, Nielsen ES (1981) Intrinsic proboscis musculature in non-ditrysian Lepidoptera-Glossata:

Structure and phylogenetic significance. Entomologica Scandinavia Supplement 15: 299-304.

45. Kristensen NP, Nielsen ES (1981) Abdominal nerve cord configuration in adult non-ditrysian Lepi-

doptera. International Journal of Insect Morphology and Embryology 10: 89-91. doi: 10.1016/0020-

7322(81)90015-5

44. Kristensen NP (1981) Phylogeny of insect orders. Annual Review of Entomology 26: 135-157. doi: 10.1146/

annurev.en.26.0 1 0 1 8 1 .00 1 03 1

43. Kristensen NP (1981) Amphiesmenoptera. Trichoptera. Lepidoptera. Pp. 325-330, 412-415 in Hennig, W.:

Insect Phylogeny (Pont & Schlee eds). Chichester, New York, Brisbane, Toronto: John Wiley & Sons.

42. Kristensen NP (1980). Sesia andrenaeformis Laspeyres, 1801 (Insecta, Lepidoptera): Proposed conserva-

tion. Z.N.(S) 2139. Bulletin of Zoological Nomenclature 37: 156-157.

41. Kristensen NP (1980) Sphinx tipuliformis Clerck, 1759 (Insecta, Lepidoptera): Proposed conservation.

Z.N.(S) 2138. Bulletin of Zoological Nomenclature 37: 154-156.

Nota Lepi. 38(1): 89-102

101

40. Kristensen NP, Nielsen ES (1980) The ventral diaphragm of primitive (non-ditrysian) Lepidoptera. A

morphological and phylogenetic study. Zeitschrift für zoologische Systematik und Evolutionsforschung

18: 123-146. doi: 10.1 1 1 1/j. 1439-0469. 1 980.tb00734.x

39. Kristensen NP, Nielsen ES (1979) A new subfamily of micropterigid moths from South America. A con-

tribution to the morphology and phylogeny of the Micropterigidae, with a generic catalogue of the family

(Lepidoptera: Zeugloptera). Steenstrupia 5: 69-147.

38. Kristensen NP (1979) Wilhelm van Deurs in memoriam. Lepidoptera NS 3: 191-192.

37. Heath J, Kristensen NP Nielsen ES (1979) On the identity of Tinea tunbergella Fabricius, 1987 and Tinea

thunbergella Fabricius, 1794 (Lepidoptera: Micropterigidae, Gracillariidae). Entomologica Scandinavia

10: 9-12. doi: 10.1163/187631279X00358

36. Kristensen NP (1979) H. Bruce Boudreaux: Arthropod Phylogeny with Special Reference to Insects.

Systematic Zoology 28: 638-643.

35. Kristensen NP (1979) The Mnesarchaea proboscis, a correction. Entomologica Generalis 5: 267-268.

34. Kristensen NP ( 1 978) A new familia of Hepialoidea from South America, with remarks on the phylogeny

of the subordo Exoporia (Lepidoptera). Entomologica Germanica 4: 272-294.

33. Kristensen NP (1978) Ridge dimorphism and second-order ridges on wing scales in Lepidoptera: Ex-

oporia. International Journal of Insect Morphology and Embryology 7: 297-299. doi: 10.101 6/0020-

7322(78)90010-7

32. Kristensen NP (1978) Observations on Anomoses hylecoetes (Anomosetidae), with a key to the hepialoid

families (Insecta, Lepidoptera). Steenstrupia 5: 1-19.

31. Kristensen NP (1978) Obituary: Henning Lemche. Bulletin of Zoological Nomenclature 35: 5-6.

30. Kristensen NP (1978) Phylogenetic methodology in hexapod high-level systematics: Results and perspec-

tives. Norwegian Journal of Entomology 25: 84-85.

29. Kristensen NP (1976) A redescription of the male genital morphology of Paramartyria immaculatella (In-

secta, Lepidoptera, Micropterigidae). Steenstrupia 4: 27-32.

28. Kristensen NP (1976) Remarks on the family-level phylogeny of butterflies (Insecta, Lepidoptera, Rhopa-

locera). Zeitschrift für zoologische Systematik und Evolutionsforschung 14: 25-33. doi: 1 0. 1 1 1 1/j. 1439-

0469. 1976.tb005 15.x

27. Rothenborg HW, Sjolin K-E, Kristensen NP (1976) Sandlopper. Souvenirs fra tropeferien. Ugeskrift for

Læger 1976: 2437-2440.

26. Kristensen NP (1975) The phylogeny of hexapod “orders”. A critical review of recent accounts. Zeitschrift

für zoologische Systematik und Evolutionsforschung 13: 1^14. doi: 1 0. 1 1 1 1/j. 1439-0469. 1 975.tb00226.x

25. Kristensen NP (1975) On the evolution of wing transparency in Sesiidae (Lepidoptera). Videnskabelige

Meddelelser fra Dansk naturhistorisk Forening 137: 125-134.

24. Fibiger M, Kristensen NP (1974) The Sesiidae (Lepidoptera) of Fennoscandia and Denmark. Fauna En-

tomological Scandinavia 2: 1-91.

23. Birket-Smith S, Kristensen NP (1974) The skeleto-muscular anatomy of the genital segments of

male Eriocrania (Insecta, Lepidoptera). Zeitschrift für Morphologie und Ökologie der Tiere 77: 1 57-174.

doi: 1 0. 1 007/BF003742 14

22. Achtelig M, Kristensen NP (1974) A re-examination of the relationships of the Raphidioptera (Insec-

ta). Zeitschrift für zoologische Systematik und Evolutionsforschung 11: 268-274. doi: 10.1 11 1/j. 1439-

0469.1973.tb00147.x

21. Karsholt O, Kristensen NP (1974) Undersogelser over sommerfuglefaunaen pä Hesselo. Entomologiske

Meddelelser 42: 33-47.

20. Kristensen NP (1972) Et fund af Dysgonia algira i Danmark (Noctuidae). Lepidoptera (N. S.) 2: 106-107.

102

SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014)

19. Kristensen NP (1972) Sommerfuglenes stilling i insektsystemet. Lepidoptera (N. S.) 2: 61-67.

18. Kristensen NP, Jelnes JE (1972) Om navngivning af ’’aberrationer”. Flora & Fauna 78: 25.

17. Kristensen NP (1971) Dagsommerfuglenes storsystematik. En oversigt over nyere undersogelser. Ento-

mologiske Meddelelser 39: 201-233.

16. Kristensen NP (1971) Sikre bestemmelseskarakterer hos hunneme af Adopaea lineola og A. flava (Lep.,

Hesperiidae). Entomologiske Meddelelser 39: 133-136.

15. Kristensen NP (1971) Et sjællandsk eksemplar af Nymphalis xanthomelas (Lep., Nymphalidae). Entomol-

ogiske Meddelelser 39: 129-132.

14. Kristensen NP (1971) The systematic position of the Zeugloptera in the light of recent anatomical inves-

tigations. Proceedings of the XHIth international Congress of Entomology in Moscow, 2-9 August 1968

1:261.

13. Kristensen NP, Kaaber S, Wolff NL (1971). Europas dagsommerfugle. 266 pp. Gads Forlag, Kobenhavn.

[Translated and revised edition of: L. G. Higgins & N. D. Riley (1970). A field guide to the butterflies of

Britain and Europe].

12. Kristensen NP (1970) Systematisk Entomologi. Munksgaards Forlag, Copenhagen. 173 pp.

11. Kristensen NP (1970) Morphological observations on the wing scales in some primitive Lepidoptera (In-

secta). Journal of Ultrastructure Research 30: 402^110. doi: 10.1016/S0022-5320(70)80071-5

10. Kristensen NP (1968) The anatomy of the head and the alimentary canal of adult Eriocraniidae (Lep.,

Dacnonypha). Entomologiske Meddelelser 36: 239-315.

9. Kristensen NP (1968) The skeletal anatomy of the heads of adult Mnesarchaeidae and Neopseustidae (Lep.,

Dacnonypha). Entomologiske Meddelelser 36: 137-151.

8. Kristensen NP (1968) The morphological and functional evolution of the mouthparts in adult Lepidoptera.

Opuscula Entomologica 33: 69-72.

7. Kristensen NP (1967) A note on Chapmania kaltenbachi sensu Hering 1932 (Lep., Eriocraniidae). Ento-

mologiske Meddelelser 35: 346-348.

6. Kristensen NP (1967) Erection of a new family in the Lepidopterous suborder Dacnonypha. Entomologiske

Meddelelser 35: 341-345.

5. Kristensen NP (1966) Om sæsondimorfien hos Plusia chrysitis (L.) (Lepidoptera, Noctuidae). Flora &

Fauna 72: 155-158.

4. Kristensen NP (1966) Notes on Sterrha ochrata, a moth new to the Danish fauna (Lep., Geometridae).

Entomologiske Meddelelser 34: 214-220.

3. Kristensen NP (1966) On the subgeneric position of Orthosia porosa (Lep., Noctuidae). Entomologiske

Meddelelser 34:211-213.

2. Kristensen NP (1965) Cikaden Eupteroidea stellulata (Burmeister 1841) i Danmark. (Hemiptera, Cicadellidae).

Flora & Fauna 71 : 81-82.

1. Kristensen NP (1965) Cikader (Homoptera auchenorrhyncha) fra Hansted-reservatet. Entomologiske Med-

delelser 30: 269-287.

Nota Lepi. 38(1) 2015: 103-105 | DPI 10.3897/nl.38.5083

Book Review: Die Widderchen des Iran [The burnet moths of Iran]

Raymond Guenin1

1 Grauholzweg 14, CH-3084 Wabern, Switzerland; raymondguenin@bluewin.ch

Received 1 1 April 2015; accepted 17 April 2014; published: 12 May 2015

Subject Editor: Jadranka Rota.

T. Keil 2014: Die Widderchen des Iran. 17. Beiheft der Entomologischen Nachrichten und Berichte. 461 pp.

ISSN: 0232-55535. Price: €238.

The first research on the zygaenid fauna of Iran goes back to the first half of the 1 9th century with

the description of the two taxa, Zygaena cuvieri Boisduval (1828) and Z. haematina Kollar, 1849.

Only a short time later new material came to Europe which had been collected by Joseph Haber-

hauer and Hugo Christoph under partly very adventurous circumstances during their expeditions

to Iran. This led to the description of another five taxa. Just before the Second World War, Fred

Brandt brought spectacular material to Europe. He had travelled alone in 1936 through the northern

Iran (Elburz, Kuh-e Binaloud) and in 1937 and 1938 to then hitherto unknown regions of southern

and eastern Iran as far as the Afghan and Pakistan borders where he collected exceptional material

that included many new species. Although the interest in the zygaenid fauna of Iran increased after

the Second World War and also the habitats and the biology and ecology of some species were

described, it was not until about 20 years ago that a new area of research on the Iranian Zygaenidae

began. Axel Hofmann, Bernard Mollet, Clas Naumann, Gerhard Tarmann, W. Gerald Tremewan

and Thomas Keil are the most important contributors to this new development of research. Thom-

as Keil visited Iran alone more than 30 times during the last 17 years. Consequently, the number

of relevant publications increased significantly. Most papers were published in various journals.

Except for contributions by Axel Hofmann and W. Gerald Tremewan as well as the papers by Mo-

hmoud Karami, Clas Naumann and W. Gerald Tremewan on the genus Zygaena Fabricius, 1775,

there was no comprehensive work available, especially not for the subfamily Procridinae. This gap

has now been closed by Thomas Keil’s book “Die Widderchen des Iran”.

This book is impressive by its size alone and by its generously designed binding in a linen

hard cover. That the text is published in two languages is not a great novelty. However, that the

German text is completely translated into Farsi by Maryam and Hossein Rajaei and that also the

book is published completely in Roman letters (German text) and in Farsi calligraphy (Iranian

text) is exceptional. It is an acknowledgement to the hospitality of Iran’s people and shows respect

to Iran’s environment. Moreover, Iranian scientists can now rely on a profound standard work on

Zygaenidae which documents in an impressive way the diversity and unique status of the Iranian

zygaenid fauna. It is to be hoped that this work will influence future decisions on preserving the

environment of Iran.

The book begins with a general part that contains information about the geography of Iran, its

climate and its biotopes. This is followed by an attempt to associate the different Zygaenidae spe-

cies to faunistic subunits, followed by comments on endangered taxa and a list of the terminology

of morphological characters mentioned in the book. In an annotated list of species, 25 Procridinae

104

Guenin: Book Review: Die Widderchen des Iran [The bumet moths of Iran]

(22 of them confirmed for Iran) and 42 Zygaenidae (41 of them confirmed for Iran) are mentioned.

The remaining four species (not yet confirmed for Iran) can be found most probably also in Iran

based on the present knowledge of their distribution. Not only the high number of taxa but also

the high number of 30 endemic species shows the exceptional character of Iran’s zygaenid fauna.

In the systematic part 67 species are mentioned. The text is presented in a concise form and

gives information about the distribution of each species (global and in Iran), the morphology (de-

scriptions of imagines and early stages), and the ecology (habitats, larval host-plants, habits). The

text is accompanied by colour plates of aesthetically superb quality which also include the imagi-

nes as for the early stages (in most cases). Some of the larvae are figured several times in different

instars and colour variations. The selected format of the images with 8.5 x 12.5 cm allows the rec-

ognition of even the smallest details. Some of the magnifications in large format are exceptional,

e.g. Zygaenoprocris (Keilia) minna (Efetov, 1991) on p. 105, Zygaena (Mesembrynus) nocturna

nocturna Ebert, 1974, on p. 161 . The author of this review had the opportunity to collaborate in the

ongoing Swiss book series ‘Schmetterlinge und ihre Lebensräume’. He therefore knows very well

what an enormous amount of work with lots of associated frustration is required to produce such

a documentation of pre-imaginal stages, especially of the Zygaenidae. This group has especially

complicated diapause rhythms that are a real challenge for anyone who tries to rear species from

egg to imago. It is amazing to note how great Thomas Keifs tolerance against such frustrations

must have been to be able to achieve such an impressive result.

The following chapter is devoted to the genitalia structures. These are essential especially for

the identification of the Procridinae. The genitalia of males and females are figured in black and

white photography. The size of the figures is well chosen and allows recognition of the relevant

characters clearly. Only in a few cases in the male genitalia (e.g. Zygaenoprocris rjabovi (Al-

berti, 1928) on p. 340 and Z. khorassana (Alberti, 1939) on p. 341) will the reader have some

difficulties to recognise the form of the comuti on the vesica in the phallus. The main reason

for this seems to be the fact that the sclerotisation of the phallus is of very different intensity.

Perhaps in such cases line drawings could have given a better result. The female genitalia are

well recognisable. However, in some cases it could have helped to figure them in different views

(dorsal, lateral, ventral).

Another highlight of this book are the aquarelle paintings produced by Anja Spindler. These are

of superb quality and can be compared with those of Frantisek Gregor (e.g. in Tarmann, G. M., Zy-

gaenid Moths of Australia) or Peter Wymann (e.g. in Lepidopteren- Arbeitsgruppe, Schmetterlinge

und ihre Lebensräume, vols 1-3). The work contains 31 colour plates with these paintings illus-

trating 1 86 specimens that are reproduced three times their natural size. This allows an impressive

overview and is an aesthetic highlight.

The next chapter deals with the distribution. For 66 species distribution maps are provided. The

localities are listed under ‘Verbreitung im Iran’ with the treatment of the respective species in the

systematic part.

Under the headline ‘Nahrungspflanzen der Raupen’ there follows on 16 colour plates that are a

compendium of larval host-plants. The whole of the information is based on personal research by

Thomas Keil. We can see that although many larval host-plants can be already identified to spe-

cies level there is still a significant number left where only the genus is known. There is still a lot

research to do. This chapter is followed by a comprehensive picture gallery of Iranian habitats for

zygaenids. It is almost a dream for a European entomologist to see these exceptional habitats in Iran.

Nota Lepi. 38(1)2015: 103-105

105

Last but not least an extensive list of literature and the contents list are provided. Moreover, a

summary of Thomas KeiTs scientific activities that were often accompanied by his wife Christine

Keil provides the book with a nice ending.

Some small recommendations for additions should a second edition ever be published are: men-

tion the sexual dimorphism in some species; introduce a paragraph dealing with ‘similar species’

for easier identification; give chorological data; explain genitalia characters for a differential diag-

nosis (especially for Procridinae).

This work is the result of extraordinary dedication and is a milestone in Zygaenidology. It will

be a solid base for any future research.

(Translated from German by Gerhard Tarmann.)

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SMITHSONIAN LIBRARIES

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androconial scales and new distributional data

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In Memoriam: Niels Peder Kristensen (1943-2014)

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Book Review: Die Widderchen des Iran [The burnet moths of Iran]

Raymond Guenin

http://nI.pensoft.net